VASCULAR PLANT SYSTEMATICS

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VASCULAR PLANT SYSTEMATICS

by

ALBERT E . RADFORD

HILLIAH C. DICKISON, JIMHY R . 1'1ASSEY, C. RITCHIE BELL

with

CONTRIBUTIONS

by

Ben H . Smith, Kenneth M. Becker , Theodore J . Cravel lo , Stanwyn C . Shetler,

Laur i e S . Radford , Cliffo r d R . Parks, Norton G. Hil l er , J . Kenneth Moore,
Hajor H. Goodman , Theodore M. Ba rkl ey , James H. Ha r din , Robert F . Thorne,
Leo J . Hickey, John T . Mickel, Lytton J. Husselman, Robert L . lHlbur and
John N. Herr , Jr .

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"ASPECTUS"

ILLUSTRATIONS
(Except Appendix)

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MARION S. SEILER

HARPER & ROt-!, PUBLISHERS

NEI'] YORK , EVANSTON , SAN FRANCISCO, LONDON
 22 451

Chapter 22 . ANALYSIS OF CHARACTER VARIATION IN SYSTEr.IATICS*

Knowingly or not, all systematists are engaged in the analysis of charac-

ter variation. The purpose of this paper is to outline approaches to the study
of character variation in systematics . Perhaps this chapter will be of greatest
value if it motivates us to get out of our usual "thought-ruts." Some peo ple
have not re-eva!uated their ideas, purposes, or methods of analysis since their
graduate days . For example , many people still use Anderson ' s pictorialized
scatter diagram as the only approach to the study of character variation due
to introgression . As creatures of habit, we tend to think along the path of
least resistance. So, if one explanation is given for a set of phenomena, it
is easiest from that point on not to keep searching for novel explanations .
The same concept is t r ue , to some degree , in methodology: while I have tried
to include all major areas of systematics in this review, I am aware that my
own background and i nterests influence the emphasis given to each area and
method . Hopef ully , a glimpse at the many methods of studying character varia-
tion used in systematics today will have a stimulating effect and permit the
development of ne," paths of thought and research . One cannot help being im-
pressed with t he potential benefits to both the descriptive and explana tory
phases of systematics when its workers consider the use of the contents and
methods of each other ' s fields as well as those developed in ecology, other
areas of biology and areas outside of biology, e . g . , psychology , business ,
and engineering .

I find it interesting that with the proposed Flora No r th America Program

and a growing number of projects to "computerize herbaria" (see Crovello ,
1972 for references), botanists have forged ahead of zoologists in the use of
computers for information retrieval. Unfortunately, plant systematists a re
far behind in the use of computers to analyze character variation. Perhaps
this book will help to alleviate the situation .

NECESSARY PRIHITIVE NOTIONS

To minimize misunderstanding, certain essential concepts, or primitive

notions, must be agreed upon before any comparison of approaches can be made.
An object is an individual organism or a col l ection of organisms grouped by
some conwon property . Sokal & Sneath (1963) designated the object an opera-
tional taxonomic unit, i . e., the unit to be classified . A system is a collec-
tion of objects, their properties, the interactions among them , and all
"environmental" forces acting upon them . A character is a property of an
object or of a system that cannot logically be subdivided further . This com-
pares with Sakal & Sneath ' s (1963) operational definition of charac t er . A
character state is the particular expression of a character of a given objec t
or of a system . For example, in a taxonomic study of willows, each taxospecies
may arbitrarily be designated as an object . Each has the character s t amen num-
ber . The charac t er state for object (species) A may be one stamen ; for object
(species) B it may be fLve stamens , e t c., depending on the indivi duals observed
in each taxospecies .

The choice of objects , of systems, of char acters, and of character sta tes
is arbitrary , and the choices vary with the investigator and with the purpose

*Contributed by Theodore J. Crovello, University of Notre Dame .

 22 451

Ghapter 22. ANALYSIS OF GHARAGTER VARIATION IN SYSTENATICS'~

Knowingl y or not, all systematists are engaged in the analysis of charac-

ter variation . The purpose of this paper is to outline approaches to the study
of character variation in systematics. Perhaps this chapter \..rill be of greatest
value if it motivates us to get out of our usual "thought-ruts. " Some people
have not re-evaluated their ideas, purposes. or methods of analysis since their
graduate days. For example, many people still use Anderson ' s pictorialized
scatter diagram as the only approach to the study of character variation due
to int r og r ession . As creatu r es of habit , we tend to think along the path of
least resistance. So, if one explanation is given f or a set o f phenomena, it
is easiest from that point on not to keep searching for novel explanations .
The same concept is true, to some degree, in methodology : while I have tried
to include all major areas of systematics in this review, I am aware that my
own background and interests influence the emphasis given to each area and
method . Hopefully, a glimpse at the many methods of studying character varia-
tion used in systematics today will have a stimulating effect and permi t the
devel opment of new paths of thought and research. One cannot help being im-
pressed with the potential benefits to both the descriptive and explanatory
phases of systematics ~..rhen its workers consider the use of the contents and
methods of each other ' s fields as well as those developed in ecol ogy, other
areas of biology and areas outside of biology, e . g ., psychology , business,
and engineering .
,
I find it interesting that with the propos e d Flora North America Program
and a growing number of projects to " computerize herbaria" (see Grovello ,
1972 for references), botanists have forged ahead of zoologis ts in the use of
computers for information retrieval . Unfortunately, plant systematists are
far behind in the use of computers to analyze character variation . Perhaps
this book wil l help to alleviate the s it uation .

NECESSARY PRINITIVE NOTIONS

To minimize misunderstanding , certain essentia l concepts , or primitive

notions, must be agreed upon before any comparison of approaches can be made .
An object is an indivi dual organism or a collection of organisms grouped by
some common p r operty. Sokal & Sneath (1963) deSignated the object an opera-
tional taxonomic unit, i . e. , the unit to be classified . A system is a collec-
tion of objects, their properties , the interactions among them, and all
"environmental" forces acting upon them . A character is a property of an
obje c t or of a system that cannot logically be subdivided further. This com-
pares with Sokal & Sneath's (1963) operational definition of character. A
character state is the particular expression of a character of a given object
or of a system . For example, in a taxonomic study of \..rillows , each taxospecies
may arbitrarily be designated as an object . Each has the character stamen num-
ber . The character state. for object (species) A may be one stamen; for object
(species) B it may be five stamens, etc ., depending on the individuals obse r ved
in each taxospecies .

The choice of objects , of systems, of characters , and of character states

is arbitrary, and the choices vary with the investigator and with the purpose

*Con tributed by Theodore J. Grovello , University of Notre Dame .

452 22

of the study. Perhaps the areas of sys tematics that would most st rongly
deny a r bi trariness are those using charact ers of organisms whose genetic
inheritance has been worked out. But even here , one does not use all
genetical l y controlled characters , nor does the c hoice of cha r acter states
that co incide with genetic f act mean that this is the most useful choice .
For example , i n a systematic study of North American Cardami ne , it may be
of more value t o recognize only two c haracter states of the character leaf
pubescence : pubescent an d not pubescent, without regard for t he several
genetical l y differ en t t yp e s of pubescence that are possib l e .

We must also consider the two primitive no tions of character space

and ob j ect space . Character s pace is an abstraction t hat describes a
system. each d imension of which is a character of the system or a f unc -
tion of one or more charac t ers of the system. Object spac e describes a
system. each dimension of which is an object of the sys tem o r a function
of one or more objec t s of the system . In both spaces if more than three
dimens ions are i nvol ved . the space is considered a hyperspace .

I sha l l adopt the view t hat any systemat i st wo rks with a system.
Taxonomists work with sys tems . To some , t he system consists only of ob-
j ects . the specimen s or populations being classif ied. For o th ers . the
specimens are linked due to e volutionary processes . The biosystematist
inte rested in pollination systems o r in a comparison of l ife tables of
populations of Trillium cer tainly works with a sys t em . Thus a ll are
i n terested either in t he desc r iption of the system o r in the explanat ion
of how the syst em , wi th it s particu l ar pattern, arose and maintains itself.
Sometimes a system is so well known or cons id ered in such a simple way
that no formal systems descr iption is under taken. I shall cons i der the
analysis of character variation as first genera t ing a description of a
system . e.g ., a s pecies complex , and then contributing to an explanation
of it s origin and main tenance .

1. THE CHARACTER VARIATION ANALYS I S PROCESS

Being a funct ion of the inve st igator ' s purpose , of his resources . and
of the specific biological system under study, the procedure used to ana ly ze
character va r iation will vary with the investigation . Nevertheless . eac h
character variation anal ys i s can be considered as one i nstance de rived from
a general . mult istage process. This sec tion presen t s t he general procedure
in detail . It emphasizes tha t the procedure is a multistage decision pro-
cess and that each stage (and its set of substages) requires quite oft en
arbitrary dec isions as to how to carry out the give n stage. Table 22 - 1
li sts the steps of the character varia tion analysis process.

A. Purpose of the Ana l ysis

1. Clea r s ta tement of the purpose . A concise, c lear sta t ement of the

purpo se of the planned character varia tion analysis is essen-
tial . Frequen tly, investigators assume that their reasons for
studyin g charact er variation are both obvious and clearcut .
Deci sions as to the "best " method for the study of characte r
var iat ion and the "best" characters to use are partly a func -
tion of one ' s p urpose a nd may c hange wi t h d ifferent purpose s .
Having stated the purpose of the study , the analyst of cha r ac-
ter variation then must cons i der the following points.
22 453

TABLE 22 - 1. The Character Variation Analysis Process

Stage Substage s or Hethod s

A. Purpose of t he analysis 1. Clear statement of the purpose

2. Value of the study to science
3. Relevance of t he study to society
4. Feasibility of the charactex
variation analysis
S. Potent i al of t he s tud y to fulfill its
stated purpose

B. Data accumulat ion l. Relevant information search

2. Choice of the sys tem
3. Choice of objects
4. Choice of c h aracter s
5. Choice of paramete r s of the
character s
6. Choice of character states
7. Cho ice of character state codes
8. Filling in the basic data matrix :
Sampling designs
Design of experi ments ,
9. Veri f i cation of the basic data matri x

C. Mathematical analysis and 1. Statis tics

s ummarization 2. Similarity estimation
3. Description of st ructure in a reduced
(See Table 22-2 for detai ls) number of dimens ion s
4. Decisions and op timi zation
5. Geome try

D. Summarization graphics 1. Frequen cy distributions

2. Scatter diagrams
(See Tab l e 22-3 for details) 3. Directed graphs
4. Und irected graphs
5. Geogr aphic maps
6. Niscellaneous

E. Evaluation of resul ts and Extensiona l expansion of data

r eanalys is Intensional expan s ion of data

F. The dynamic data bank An individual's data bank

A network of data banks
 454 22

2. Value of the study to science. Many argue that any knowledge is

of value to the advancement of science. But two facts must
be considered: some kinds of knowledge are more valuable
than others at any given pOint in the development of science,
and only a finite amount of human and other resources are
available for systematics. As a result, decisions must be
made as to \..rhat studies should be carried out. But, as is
true for most of the stages and substages of the character
variation analysis process, an objective, nonarbitrary deci-
sion function does not exist to determine the relative values
to science of different character variation studies in a given
field or subfield.
3. Relevance of the study to society . As evolutionary problems in
systematics and ecology become more important to the future
of society , there is pressure to make every study socially
relevant. This demand has two distinct sources . The first
is economic. If granting agencies give money only for proposed
character variation studies that are relevant to society's
problems, then most researchers who want to remain active
will move to such research. Secondly. some potential students
of systematics will refuse to enter this career unless its
relevance to society is made clear.
4. Feasibility of the character variation anal ysis. Most problems
in systematics are complex and cannot be separated into iso-
lated components for analytical laboratory study as easily as
some problems concerning systems in molecular and cell biology .
, It is often difficult to know in advance the amount of time,
equipment , and manpower that will be necessary. Nost proj ec ts
require years to complete--e . g., a study of strategies of evo-
lution within and among genera of the North American Brassica-
ceae or a monograph of the genus Ranunculus. Granting agen-
cies and administrators usually demand frequent reports and do
not fund projects for long periods of time all at once . This
is unfortunate if it splinters the r esearch effort, but on the
other hand it compels the principal investigator to stop peri-
odically to rethink his project and to justify its value ,
feasibility , and relevance .
5. Potential of the study to fulfill its stated purpose . At least two
aspects of the substage must be considered, the quality and the
quantity of the proposed character variation analysis. By the
quality of the research I mean the ability of the results to
provide a definite choice between the alternative hypotheses
formulated for the purpose of the study. That is, will the
research be germane to the purpose, or wil l it just give us
more information of uncertain value about the system under
study? By the quantity of research I mean whether , although
it is germane research , the character variation analysis pro-
gram will have sufficient pOt"er to discriminate between alterna-
tive hypotheses or whether it will result only in inconclusive
evidence about the hypotheses under test.

B. Data Accumulation
1. Relevant information search. Significant research in systematics
requires accurate knowledge about the biological system or
phenomena chosen for study (e.g .• the family Brassicaceae, or
22 455

strategies of weed populations) , and at least a t.,rorking knowledge of

the current methods used to study such systems.

It is probably not possible to list all i n f ormation sources of

value to systematists . Also, our information sources are undergoing
rapid change--the amount and kinds of in f ormation are increasing, and
the available methods of retrieval and analysis of such sources are
improving constantly, thanks to advances in computer science. The
following comments are made in the light of what could be available to
workers on a widespread scale, because the technology exists today.

Let me refer to the information sources collectively as the sys-

tematic data bank. I am concerned with the structure and content of
the data bank. The actual computer hardware (remote terminals, cen-
tral processors) and soft~"are (special information retrieval programs,
simulation languages) exist today to handle the data bank.

The data bank will obtain input mostly from four . sources: 1. the
literature, 2. museum and herbarium collections, 3 . botanical gardens
and arboreta, and 4. raw data . Every scientist has a set of file
drawers overflowing with index cards that contain references that he
thinks may be of value to him . They may have additional descriptors
or key words beyond the usual information (author, date, title, cita-
tion). A new card file quickly enters into an exponential rate of
growth, but this is short-lived since the worker realizes that to keep
up with the relevant literature in detail, he would have no time left ,
for anything else. Often at this stage, the original set of important
references is adulterated by the addition of references that are not
too valuable to his research, but that were sent to him as unrequested
reprints that he feels compelled to catalog. The result is an unwieldy
file. What could be the best summary of the past literature in a field
to incoming graduate students becomes quite useless.

The above problem can be attacked from two extremes by using com-
puterized data processing. Attempts can be made to capture the entire
biological literature in a form sui t able for electronic data process-
ing. This data bank then can be searched, usually with the aid of a
researcher's interest-profile. to produce a current or a cumulative
list of germane articles. Such a service must be performed by a large
commercial , government. or non-profit scientific organization. (Note
scope and use of Biological Abstracts in Chapter 30.)

At the other extreme , a worker can simultaneously type the infor-

mation about a reference on his file card and produce a punched paper
tape or, more recently, a magnetic tape that can be read into a com-
puterized data bank. The results can include a periodically updated
alphabetical listing, a key-word-in-context permutation of all or
selected title ~ords of all or selected articles, etc. He will have
immediate and useful recall of his extensive bibliography, with the
possibility of it being rearranged in any of several different, pre-
viously expensive ways. For example, my current personal bibliography
file is on magnetic tape and contains 7,000 re f erences of interest to
me . Each reference has attached to it one or several deep-index sub-
ject numbers reflecting my interest (e.g., 60 is numerical taxonomy,
70 is evolution, 93 is flowering plants, 99 indicates that I have the
456 22

reprint) . Simply by key punching on a search request card the numbers

60 and 93 i n the proper way I get a printout of just those references
dea ling with numerical taxonomy of flowering plants . It takes less
than one minute of computer time to search these 7,000 references and
to print out the " hits" in whatever order I request . I f someone else
were doing this we ought to be able to trade bibliography data banks
very easily, thus ex panding our retrospective literature searches.

The second major source of input to a data bank of value to sys -

tematists is specimen data from herbaria and museums. All coll ections
are arranged according to one primary organizi.ng character, usually the
taxon name. This is best for most systematic \wrk. But biosystema-
tists . evolutionists and ecologists want information arranged in a dif-
ferent \"ay . For example. an evolutionist might be interested in con-
vergent patterns of evolution of late fall f lowering time, which would
require him to look through all spec imens in the collections. Or, a
plant geographer might ask \"hat specimens in a 1,000,000 specimen her-
barium are from the Channel Islands off California . An ecologis t wo rk-
ing with a regional planner might wish to know what plants used to grow
in the Wabash Valley before the beginning o f heavy industry and e xact ly
where they might be collected no\,' . A bio sy :;;tematist may desire a list
of all previous collections of a taxon. A computer printout from d i f ··
ferent herbaria would save considerable time . He can provide such a
service now for the 65,000 specimens in the Greene Herbarium at Notre
Dame (Crovello , 1972) . Similar projects are underw<ly at Boulder, Cape-
town (Hal l, 1972) , Nexico City (Gomez-Pompa and lIlevlin.!S. 197 3), Ott<1\,<1
and Saskatoon (Argus and Sheard, 1972) and Washington.

Deciding what collections and hOt" much information to capture ,

and how to do it, is itsel f a complex multistage decision process that
can only be mentioned here . Suffice it to say that systematists are
active in plann i ng and implementing the computerization of specimen
collect ions and systematic literature. In addition to those mentioned
above , Crovello & MacDonald (1970) list over 40 such projects.

The third data source are botanical gardens and arboreta . Natural
history preserves i n urbanized areas perhaps should also be included
here rather than in the next category. A good example is the work of
Lindsey et al . (1969) . Botanical gardens and arboreta have been a
source of seed and other living material for centuries. The Amer i can
Horticultural Society presently supports a computerized Plant Records
Center. Soon this should a llow a systematist to make just one inqui ry
and receive info r mation on the availability of plant material of a
given taxon in all of the major botanical gardens and arboreta i n the
United States (see discussion in Chapter 32) .

The fourth major input source to the data bank involves raw experi-
mental data and data from any natural system . They may be data from a
relatively undisturbed woods or t ram sicle\~a l k samples f rom a metropoli-
tan area . The raH data tha t fOT Ui t t'e basis f or the summary tables a nd
conc l usion s ui j ournal articles nr e j o s t, p jthe r imm" dia te ly bec a us e t he
author does not keep dat a records ve r y 101l.1! OT. Hlrc f req uently . because
on his retiremen t the data recot"ds a r e s o old :la d the d e tails so vaguely
remembered that they are d iscarded as use l ess.
22 457

The preservation of raw data in easily retrievable form has impor-

tant uses. These include the reanalysis of the same data for the same
or a different purpose with better or more recent methods , and as a
base to allow building more firmly on the earlier work of others. For
example, Cravello (1968a) measured over 200 characters on over 500
specimens of willow from California and analyzed this bank of data to
make formal taxonomic decisions. These data are deposited on punched
cards in the Herbarium of the University of California at Berkeley.
A future worker may wish to reconsider the taxonomy of the group or
to analyze the genus to detect evolutionary phenomena. The original
200 by 500 data bank may serve his purpose in itself or, more probably,
it can supplement the data that he will collect. In other words, his
conclusions will be based on the detailed data accumulated by an
earlier worker. Hopefully the new conclusions will be sounder, since
they are derived from an expanded data base.
2. Choice of the system. Choice of a system is sometimes dictated by a
supervisor, by interest, by its relative availability for manipula-
tion and testing, by social and political relevance, or by a combina-
tion of these. In this substage, as in the next four, the analyst of
character variation must ask continuously what effect the choice of
different systems will have on the final outcome. At one extreme, a
chosen system may be unable to provide any answers to satisfy the pur-
pose. And only one system can serve the purpose. On the other hand,
the purpose may be such that any system will provide answers. Then
the problem is to decide how general the results are for other systems.
3. Choice of objects. The problem that confronts the investigator here is to
,
decide at what l evel of organization to analyze objects . In taxonomy
should one consider each plant, each population, or each species as an
object?

Such decisons are examples of perhaps the most common compromise

function in systematics. The compromise is between accuracy (or reso-
lution) and practicality (or summarization). For example, presenta-
tion of almost any hierarchy, not just the taxonomic one, demands a
compromise between the accuracy with which relations among the objects
are presented and the degree of summarization that is thought necessary
to permit a biolog ist to understand the salient relationships among
groups of objects. Any time different objects are grouped together
and treated as similar, inaccuracy is present.
4. Choice of characters. In some disciplines this has produced heated debate.
Smith (1965) called numerical taxonomists existentialists because they
will use any character for which data are available. Other taxonomists
would rather choose some subset of the total available characters
according to some criterion such as alleged phylogenetic importance,
relative closeness to the DNA level (protein characters versus mor-
phological character), or discriminating value. Actually, numerical
taxonomists also choose characters but they do not add the additional
criteria above, so they have a greater number available for use.

What criteria are used to select characters besides those in the

definition given earlier? The characters must be measurable with the
available resources, be present in at least a majority of the objects,
and be germane to the purpose of the study.
458 22

5. Choice of parameters of the characters . Two parameters , the mean and the
variance (or the standard deviation) , appe ar most fre quen tly in charac-
t er variation analyses. Neans are estimates of centra l tendency , or of
location of the sample, wh ile the variance is an estimate of the degree
of variation abou t the po int of central tendency . Use and comparison
of means should be only a first step in the analysis . Comparison of the
degree of variat i on among samples , using the var iance or any other such
esti mate, should allo\" the analyst to formulate and to try to answer a
completely different set of meaningful character variation questions.
For exampl e , geographic variation studies using the mean demon-
strate clinal patterns . This is explained by the different forces
presen t from one end of the transect to the other . But what if the
variance also shows a cline? Hhat if the peak of the variance cline
is geographically different from that of t he mean? \fuat would th i s
mean to such biologically important charactcl.·s a s selective pressure?
Population and ecol ogical geneticlsts have used the vari.an ce as an
essential part a nd tool of their research analysis. Worker!. in syste-
matics should at least consider using the variance as they have the
mean . The next s tep i s to genera lize and ask what other parameters,
statistical or not, could be used with value in a character variation
analysis.
6. Choice of character s tat es . If a charact e r is to be measured , what set of
values should be used to r ecord the s tat e of the character i n a given
individual? For continuous characters such as leaf length, it might
b e the nearest millimet er or the nearest centimeter . Nore difficult
dec i sions i nvol ve qualitative attributes , such as l ea f shape . Here,
an investigator ' s operational procedure would be simply to recognize
those patterns as dis t i nct that best serve the purpose of his anal ysis .
This rule sound s terribly subjective and unscientific, but, to deter-
mine characte r states fo r qualitative characters , no f ormal decision
function exists that considers one ' s material and purpose any better.
7. Choice of character state code s . This is a p r actical procedure that is
becoming increaSingly important. As an example , suppose samples are
collected on different days ove r severa l years , as are the millions of
plant spec i mens in our herbaria. Date of collection is a valuable
character for analyses of phenology and should be considered as a pos-
sib l e source of c onfounding variation . Assumin g that everyone records
the day, month, and year, should it be r ecorded as 20 November , 1972 ,
or November 20 , 1972, or 11/20/72, or 20/l1/72? It is relatively unim-
portant as l ong as o ne is \wrking in isolation . But if you \.;ri sh to
swap data banks \"ith a colleague , other programming may be n e cessary .
Still the problem i s not serious . Howeve r , consider the automated
aspects of the proposed Flora North America Program. Its data ban k
woul d easil y number i n t he millions of pieces of in formation. Char-
a c ter state code compatibility then takes on an e xpensive and time-
consuming aspect . By recordin g t he da te as year, month , day, with two
digits for each , the compute r could arrange them in c hrono logica l order
by simply sorting all six col umns at once , rather than having t o make
several passes .
8. Filling in the basic data matrix . On ce all of the nece ss ary decisions
have been made in the above substages, the c harac ter by object basic
data matrix can be filled in . As underst ood by numerical taxonomists,
this is a two dimen s ional table , \·lith on e r ead in g per cha r ac ter per
object. I think it valuable. however, t o consider expansion into
higher dimen s ions by consider ing more than one parameter. Although
 458 22

5. Choice of parameters of the characters . Two parameters, the mean and the
variance (or the standard deviation), appear most frequently in charac-
ter variation analyses . :Heans are estimates of central tendency , or of
location o f the sample, while the variance is an estimate of the degree
of variation about the point of central tendency . Use and comparison
of means should be only a first step in the analysis. Comparison of the
degree of variation among samples, using the variance or any other such
estimate, should a11m" the analyst to formulate and to try to answer a
completely different set of meaningful character variation questions.
For example, geographic variation studies using the mean demon-
strate clinal patterns. This is explained by the different forces
present from one end of the transect to the other. But what if the
variance also shows a cline? lfuat if the peak of the variance cline
is geographically different f rom that of the mean? \fuat would this
mean to such biologically important characters as se lective pressure?
Population and ecological geneticists have used the variance as an
essential part and tool of their res02arch analysis. Workers in syste-
matics should at least consider using the variance as they have the
mean. The n ext step is to generalize and ask what other parameters,
statistical or not, could be used with value in a character variation
analysis .
6. Choice of character states. If a character is to be measured, what set of
values should be used to record the state of the character in a given
individual? For continuous characters such as leaf length, it might
be the nearest millimeter or the nearest cent imeter. Mor e difficult
decisions involve qualitative attributes , such as leaf shape. Here,
, an investigator's operational procedure would be simply to recognize
those patterns as distinct that best serve the purpose of his analysis.
This rule sounds terribly subjective and unscientific, but, to deter-
mine character states fo r qualitative characters, no formal decision
function exists that considers one ' s material and purpose any better.
7. Choice of character state codes . This is a practical procedure that is
becoming increasingly important . As an example, suppose samples are
collected on different days over several years, as are the millions of
plant specimens in our he rbaria. Date of collection is a valuable
character for analyses of phenology and should be considered as a pos-
sible source of confounding variation. Assuming that everyone records
the day , month, and year , should it be recorded as 20 November, 1972,
or November 20, 1972, or 11/20/72, or 20/ll/72? It is relatively unim-
portant as long as one is \·,rorking in isolation . But if you wish to
swap data banks with a colleague, other progranuning may be necessary .
Still the problem is not serious . Hm-lever , consider the automated
aspects of the proposed Flora North America Program. Its data bank
would easily number in the millions of pieces of informa tion. Char-
acter state code compatibility then takes on an expensive and time-
consuming aspect. By recording the date as year, month, day, with two
d ig its for each, the computer could arrange them in chronological order
by simply sorting all six columns at once, rather than having to make
several passes.
8. Filling in the basic data matrix. Once all of the necessary decisions
have been made in the above substages, the character by object basic
data matrix can be fillE'd in . As under stood by numerical taxonomists,
this is a t\<lO dimensional table , \.;rith one reading per character per
object . I think it valuable, hQ1;<lever, to consider expansion into
higher dimensions by considering more than one parameter. Although
22 459

men tioned above in a different context, the immediate val ue of us ing

a three-dimensional basic data matrix wOlild be in a better descrip-
tion of the objects and a more reliabl e est ima te of their phenetic
relationship. Similarly, the more parameters available, the bet ter
the basic data matrix should be for systems models such as pollina-
tion simulations .

Expa nsion of the basic data matrix into other dimensions also
could provide new data on the same objects but at different times
and could accommodate data for the same objects but from differ ent
places. The latter is a function of the scope of the objec ts. If
they were specimens or site samples . they wou l d be irrelevant . But
if f or some reason one c ho se species as objects, then they would be
germane. This is an important concept in the fo r mation of basic
da t a matrices . A basic data mat r ix based on a given n umber of
characters and objects can always be reduced in dimensions and in
the magnitude of any dimension (e . g ., the number of columns can be
reduced) . Either type of reduction can occur in at least two wa ys :
either by simpl e deletion , or by combining according to some fun c-
tion or qualitative rule. For example, cons ider a numerical taxo-
nomi c study where the objects a r e individ ual s i te co llect ions of
one speCies and information is pr esent on the mean , variance, and
range for many sites of the sp ecies over several years. For a
certain analys is, all of this information for each species may be
expressed as a simple numb er , resultin g in a simple two- dimen sional
char acter by spe cies basic data matrix . The po i nt to understand is
that such con densation of the matrix is poss ible and s hould be use -
fu l for given purposes . To pa raphrase , "Ie can change the s truc -
ture of the basic data mat rix, but this change is a l ways in one
dir ection. '.fe cannot begin with a character by species mat r ix
and obtain a character by site matri.x with real data . (He could
generate random sites if the mean and v~rianc e vectors \,rere avail -
able for each species , but this presuppo ses ear lier informa tion. )
So , the analyst of character variat ion must be sure that the
o rig inal ob jects and characte r s are s uffic iently homogeneous t o
answer all of the questions and ~urpos e s fo r which t hey a r e i n t end ed .

After the objects, such as loc al populations of a spec ies,

are chosen, the design of exper imen ts must be con sidered before
sampling and actual data Acc umulation begins . Books on designs
involving analys is of variance are available ; e . g ., Cochran and
Cox (1957). Unfortunately, designs for other statistical pro-
cedures are rarely di scussed . Fo r example , one can read how to
car ry out a general sys tems ana l ysis , b ut details often are l earne d
only by trial and error.

After a design is chosen , sampling designs must be carried

out. Most character variation analyses require sampling at several
levels of organiza t ion, and different sampling procedur es may be
required a t differen t levels of the sampl ing hierar chy. For exam-
ple . to measure gene flow within a flowering-p l ant species by
po llen-tagging experiments, one first chooses the local popula-
t i ons to be investigated . perhaps by a stratified sampling design .
Hithin each site , c hoice of ind ividual plants to monitor may be by
random samp ling .
460 22

Probably no extensive basic data matrix will ever be free

from the problems of missing data. Even when each cell is repre-
sented by at least one value, some will be more reliable than
others . Few studies in any area of systematics have evaluated
the effect of missing data. In numerical taxonomy. Cravello
(1968b) compared estimates of similarity based on a 43 character
by 30 species matrix of California willows ; In one analysis, a
comparison was made between a full data matrix and the same matrix
from which data had been removed in the same pattern as shown by
Hunz's flora. The OTU by OTU relevance of the latter was only 48
percent . Yet the resultant species by species similarity matrices
were highly correlated (0.887) and did not differ significantly in
their means and variances. This suggests that taxonomic data
matrices with missing data still are valuable.
9. Verification of the basic data matrix. ffilether the data are obtained
from experiments by the analyst himself. from some data bank. or
from the literature. some degree of effort is necessary to verify
the validity of the values of the resulting matrix. This may range
from simple verification of experimental results. which have been
punched on tab cards or recorded in code on magnetic tape. to ex-
tensive checki ng of the reliability of data obtained from other
sources.

C. Hathematical Analysis and Sununarization

Methods used in mathematical analysis are not the same in all phases
of systematics . Methods vary most between descriptive stages (taxonomy)
and explanatory stages (studying the dynamics of populations or the pro-
cesses of evolution) .

Table 22-2 contains an expanded listing of methods of mathematical

analysis and summarization that have been used or have potential in charac-
ter variation analyses. They are grouped into somewhat artificial catego-
ries. Other groupings are possible. but they are probably less satisfac-
tory than the one chosen. I am unable to produce an absolutely satisfac-
tory grouping because the subfields of mathematics are related in a com-
plex pattern. Even the simple two-level hierarchy of Table 22-2 causes
distortions in relationship among the entries.

Even a brief sketch of each of the basic entries of Table 22-2 would
require the space of a book, not an article. Thus. in the following para-
graphs I shall not consider items whose use is obvious and agreed upon.
References in Tables 22-2 and 22- 3 were selected if they were recent . com-
prehensive. unique, botanical, or readily available . They are not meant
to be complete. Rather. they should serve as examples of recent uses of
the technique and as basic guides to the literature . Extensive bibliog-
raphies of a broad field like character variation analysis in systematics
must await computerized information retrieval. References to major reviews
and introductions appear next to the titles of the major categories. Some-
times the reader may have to cross into other fields to find a suitable
treatment. Within each category the number of references will vary. Some
readers may find it interesting to compare these tables with their analogs
in Crovello (1970). Those in the 1970 paper have more items and more uses.
indicating perhaps that zoological systematists are further into quantita-
tive character analysis than are botanists.
22 461

TABLE 22 - 2. Me thods of :-lat hemat i cal Analysis and Summarization

- -- - -A.- -- - ---" ==.======:;::::==:==0=== ==
Sta ti ~L ics (B1aeklth and Reyment. 1971 ; Sokal and
Rohlf , 1969)
- - - - - --
Transfo rmat ions l1u1tiple r egression
3as i c statistic s General polynomi al r e gression and
Nonparamet ric methods trend surface analysis
Analysis of frequencies Analysis of var iance
Linear correlation Analysis of covariance
Curvilinear correlation Multivariate analysis of variance
Hultiple correlation Evolutionary and population allometry
Linear regression

B. Similarity estimation (Blackith and Reyment. 1971; Mayr, 1965;

Sneath and Sokal, 1973)

Association and correlation coefficients: Anderson, 1949; Rhodes , et a1. ,

1968)
Distance coefficients (Crove110, 1968a, 1969)
Information theory (Hawksworth , et a1., 1968)
Cladistics (Dayhoff, 1969; Fitch and }illrgo1iash, 1967; Kluge and Farris, 1969)
C. Description of structure in a reduced number of dimensions (Cole , 1969;
Morrison, 1967; Sneath and Sokal , 1973; t.Jilliams and Dale, 1965)

Visual r earrangement of a matrix:

Character by object basic data matrix ,
Similarity or distance matrices (Figure 22-1)
Principal component analysis (Prance et al. , 1969)
Fac tor analysis
Canonical analysis
Hultidimensional scaling (B1ackith and Reyment, 1971)
Cl uster analysis (Jackson and Crovello, 1971)
Hyperfootballs (Reyment , 1969)
Charact er analYSis (Crovello, ,Q68d)
- - - " ~~-,;--;-~-:-c;--;= ___-
D. Decis ions and op timization (Blackith and Reyment, 1971)

Decision t heo r y, ;l.'1tt-=-·, ,~ r ecog;\it-lon :

!)isc r iminant f u n:: t j (m s (TIC1 _':lahy , 1"(.1) ; Goodman , 1967)
Dec i.s ion tree s
Taxonomi c keys (Bc s <: e Tt _ 1969; G()cQ :-. l 1, 1968 ; Hall , 1970; Hor s e et a1 ,
1968)
Optimiz.:1tion th eory (Rosen, 1967; Ruben, 1967)

E. Geometry (Busacker and Saaty, 1965)

Euclidean geometry
Non-Euclidean geometry
Graph theo ry and networks (Gabriel and Soka1, 1969 ; Hoss, 1967; Prance et a1.,
1969; Wirth et al., 1966)
Topology (Sneath , 1967; Thompson , 1942)
 462 22

1. Statistics. Today sta t i stics enjoys wide acceptance among sys tematists .
The concurrent development of digital computers and of improved method s
of automated data accumulation are causing an even wider use of statis-
t ics and of mor e involved statistical methods . Interestingly , para-
metric statistics as we know it may s oon be succeeded by the methods
of data analy s i s (Sakal and Rohlf, 1969 : Chap. 17). To give one exam-
ple , severa l years ago no c haracte r variation analy s t tJi th extensive
data on 10 c ha racte rs would cons truc t the 45 scatter d i agrams of each
pair, l e t alon e r epe at the procedur e using several transformations .
Today, even a second - generation compu ter can comple te the task in
minutes .
To use sta tistics int elligently i n plant sys t ematics requ i res
serious study. Fortunately several c l ear t exts are available which
contain more than " cookbook" procedur es . Except fo r basic descrip-
tive and inferential statisti cs , plant systematists have not used
more advan ced techniques of Category A. This is unfortunate because
such methods can provide f r esh insights into complex systems , and
plant pop ulation s eries are complex .
The fina l item in Ca t egory A of Tabl e 22 - 2 is Allometry. Perhaps
this is out of place here , but it is di f fic ult to f it it anywher e ,
since it is not a method but an area of systematics. Yet i t comb ines
mathematical methods in such a way and has such grea t promise fo r the
f uture t ha t it deserves mention .
2. Similarity Estimation. This has been used mostly by taxonomic workers,
but it has great potential for evolutionary systems analysts t"hene ver
they wish to summarize relations hips among many characters, objec t s ,
or components of a system.
, The normal use of similarity estimation begins with a character by
object bas i c data matrix . Some coeff i cient , or a distan ce measure in
a cho sen me tric, is calculated between all pairs of objects and resul ts
i n an object similarity (or dis t ance ) matrix. In t axonomy each objec t
(or OTU) is a t axon, or even a l ocal popul ation or a n organism . While
such s i mi l ari ty es timates are of grea t va l ue in summarizing re l a tion-
ship s among object s , th ey are ave r ages or summations of some sort . This
means that the part icular characters in which two objects differ are not
r evealed , nor is there any indication as to which pairs of objects differ
only slightly in many chara cter s and which pairs differ consider ably,
but in only a few characters.
3. Desc r iption of Structure in a Reduced Number of Dimensions. This approach
attempts to summa rize relationships among object s in fewer than the
original numbe r of the smaller of the two di mensions of the basic data
matrix . Fo r example , in a study of 50 characters and 20 objects, t he
various methods listed i n Category C of Table 22-2 would seek to mini-
mize di stor tion of the relationsh ip s in a new coordinate sys tem with
less than 20 indep endent dimensions (Gower, 1966) .
Cho ice of a particular method from Category C is important since
di fferent methods will give somewhat different estimates of re l ation-
ships among the objects . Usually, an object simila r i ty mat rix obtained
from some method of Category B, a character by character co rrelation,
or a variance-covariance matrix is required f or analysis by a method
of Ca t egory C. Exceptions include those studies t ha t simply rearrange
the charac t e r by object basic da ta matrix.
He can gr oup the me thods o f Ca tegor y C of Table 22- 2 by severa l
criteria. The methods easi l y se para t e i n to those t hat r educe t he
22 463

Figure 22 - 1 . Similarity matr ix

shaded to ShOH the percent of 100 %
similarity hetHeen pairs of 16
701099%
OIU ' s . [\·lalraven , H. C. 1970.
A statistical analysis of six- 501069%
teen taxa of R.hynchosia (Legumi - 30 10 49%
nosae) in the United States . 1 51 02 9%
Rrittonia 22 : RS·-92. Used Hith 010 14 %
permission. J

• "
• • .. "
" , •
"
•
, "
"

"

Figure 22-2 . 3-dirnens i onal graph a nd PRIM netl'JQrk based on 29 characters .

[From Jackson , R. C., and T. J . Cravello . 1971 . A compar ison of numerical
and biosystematic studies i n Haplopappus . Brittonia 23 : 5/,-70 . Us ed I.,rHh
permission . ]
 464 22

o riginal high number of dimensions to one mean ingful dimension , and those
that retain more than one meaningful dimension . }mny phenograms , claclograms .
and other simple graphs fall into the forme r category. Cluster analysis is a
term used to define one group of such approaches (Sakal and Sneath . 1963 ). In
contrast , methods such as principal components analysis usually produce more
than one meaningful dimension. If we begin \.;ith an. original character space
with a number of dimensions equal to the number of characters, the result of
a principal compon ents a nalysis will be a new se t of axes in the same charac-
te r space , but with a reduced number of independent axes along which signifi-
cant variation occurs. We can refer to the space of this new set of axes as
reduced character space . Similarly , if we carry ou t a principal componen ts,
or other, analysis on an object similarity table (through the derived corre-
lation or variance-covariance matrix) the reduced space can be referred to as
reduced object space .

Result s of the above typ es of analysis may yield a phenogram , cladogram,

or set of reduced axes in which objects can be positioned. Usually, such
summaries are then inspected visual l y and , from the observed pattern of vari-
ation, taxonomic or other decisions are made .

Two final approaches in Ca t egory C are of interest because the ir purpose

is often different from that of numerical taxonomy. Reymen t (1969) has cham-
pioned methods that analyze variation and covariation around some multivariate
point or points of central tend ency . Consider a bivariate scatter diagram
containing a set of observed points. Using a multivariat e test for the loca-
tion of the multivariate parametric mean, we can con struct an equa l frequency
confidence region at a given probability level (Sakal and Rohlf , 1969). It
will be elliptical in shape . tHth three or more characters, this ellipse
becomes a sphere or hype rs phere. Staying with two- dimensional ellipses for
Simplic it y , imagine the 95% confidence regions (our ellipses) of two popula-
t i ons . The ch ances are grea t that the origin o f each ellipse will be located
at a different point in this two-dimensional charac ter space, that the size
of each ellipse will differ, and that the orientation of the major axis ~"ill
differ . The important point is to discover what this means at the biological
level. For example, if the e llip ses from t~w populations differ in size but
not in orientation, does this mean there are similar selective forces at dif-
ferent intensities? Or, if the size is the same but the ellipses are oriented
in different directions, does this mean there a re different selective forces
working at the same intensity? Other combinations are possible, even in two
dimensions, but the possibilities increase as the number of characte rs (or
perhaps principal axes, etc.) increase.

Inverse or character analysis is a term that refers to studies similar to

the above but inverted; i.e ., the main concern is not to study relationships
among objects but among characters. Hany uses of factor analysis in system-
atics obtain this as an intermediate by-product of normal analysis. But
whereas the taxonomist is interested in object relationships, when he changes
hats and becomes an evolutionist, this previous by-product can be of prime
value to him in gaining insights into the causes of the pattern of variation
previously described. For example, why do cha racters c luster? One reason is
that they have been se l ected over time to meet an environmental se lec tive
force. As a by-product , then. one should be able to see the sa lient adaptive
complexes of characters at a fairly efficient ra t e . A character corre l ation
comp l ex actually may consist of a minimum of one adaptive compl ex , bu t 'if two

I
22 465

e nvironmental selective forces are correlated in their i ntensities over severa l

sites , then the evolutionist ' s task is to assign the characters in a cluster to
their proper selective force (Blackith , 1960; Cravello, 1968d). The potential
of this method for inter-taxon comparisons is immense .

This is a good time to cons i der the misuse of some of the me thods of
Category C of Table 22-2. Seal (1964) states several times in his text tha t
numerica l taxonomists part icularly have misused such methods as principal
components analysis . His argument is that the assumptions of this technique
require that the characters' values, the basic data matrix , be sampled from
one multivariate normally distributed population . Clearl y , in a taxonomic
study involving many different species and some discontinuous and non- normally
di stributed cha racters , this doe s not hold . Similarly , Seal and others criti-
c i ze the use of a distance coefficient t hat do es not weigh t each characte r
according to its co rrelation with other charac ters. Host coeffic i en t s in use
in numerical taxonomy do not consider relative correlation . Hahalonobis dis-
tance, D2, is suggested as being more valuabl e , since it takes correlation
i n to account , as well as the covariation within groups .

Given such criticisms on mathematical statistical grounds , how can anyone

decide to use principal components analysis in a st udy of many species or to
use a distance coefficient that does not consider co rrelations among cha rac-
ters? I think this decision i n the multistage decision process in systema-
tics is largely a function of the purpose of th e character variation analysis .
Seal ' s critic ism of the use of principal components analysis in systema tics is
valid if the worker tJants to indulge 1n statistical inferences about the re-
sult s (e.g . , i f he desires to establish probabilistic frequency ellipses about
each object or each character ). But much will r emain outside the realm of
parametric sta t istics since it seems easier to use randomization tests and
other nonparametric methods with the comput er to determine probability l evels.
If we see principal components study as another way of looking at our systema-
tic system and if it gives biological insight at a high rate, then I maint ain
that its use is justified . Of course , one does not use principal components
analysis alone . It is complemented by phenograms , which are most accurate at
the tip s (versus principal components results. \~hich are better in expressing
r elationships among clusters of objects), and by networks superimposed on the
first three principal components axes .

The use of distance coefficients that do not conside r character correla-

tions should depend on one ' s biological purpose , not on Hhether there exists
a readily ava ilable r epertoire of significance tests. Perhaps because of
training , most systematists think about rel ationships , i ncluding distances,
in a Euclidean space, wher e the Phythagorean theorem holds true. They also
think about one character per dimension. Now , if one's purpose is to estimate
the distance between objects in the context of a given set of characte rs, then
a distanc e measure that does not take corre l ations into account may be more
useful. Of course then there is a large amount of redundancy of information
in terms of information theory , but for some purposes , such redundancies are
important .

I sugges t the following philosophy fo r many stages of our character vari-

ation decision process: fo r any stage \Jith two or more alternative methods ,
always use as many methods as possible . My reason is that every technique
provides some new, non- redundant information . If method X gives five units
466 22

of biological insight i nto the s tudy and Y gives one uni t, but a di f -
fere nt one, I liouid be an inefficient sci en tist if I ignored Y. So,
in the case disc us sed above , my decision would be to calculate 0 2 , as
well as a distance coe ffi cient that does not take correl ations into
account . In general, there has been too little comparison of differ-
ent me thods, espec ially stat istical comparisons (as opposed to visual
e xamination of phenograms, etc.) . \-1ork that has been done (e . g. ,
Cr ovello , 1968c , 1 969; Gower , 1967 ; Noss, 1967; Prance et 81.. 1969)
indicates that changes in the index or the c luste ring method used do
produce different , but not unrelated , results. These results dis-
play roughly the same relationships. For example , severa l available
studies analyzed data via a character factor analysis and by the use
of an unweighted distance coefficient (Crovello, 1968c; t-loss , 1967) .
Then if one also calculates the unweighted distance between objec t s
even over just the first three factors , the resulting distance matrix
is highly correlated with that using distances based on all of the
characters . Using 131 morphological cha rac ters , Crove llo ( 1968c)
obtained correlation values of .842 to . 870 in a study of 30 taxo -
species of wi llow .
4. Decisions and Optimization . These are a concern of systematists inter-
ested in description or in modeling an evolutionary system. The par-
ticular decision or optimiza tion methods chosen themselves consti tute
a dec ision pro cess . Decisions are necessary in taxonomy in the formal
delimitat ion of taxa (classification) and, later, as a separate pro-
cedure in identifi cation of unknown specimens . There exi st few rules
on the classification of organisms--I~hich may be a good thing ! Host
taxonomists will recognize two se t s of organisms as distinct if t~e
members of each are relatively close to each other with respect to
their position in a characte r space and are relative l y removed from
the members of the other set. I n other words, a set of organisms is
put into one class if the within-group variance is relatively low and
the among- groups variance is relatively high . Numerical taxonomy ,
especia lly c l uste~ analysis and methods to di scover relationships in
reduced character or object space , is a valuab le aid in gain i ng in-
sights into the differential density of the set of objects in the ori-
ginal character space and in deciding on class bounda rie s .
Every human being is a taxonomist . Host do not realize it and
o thers do not care to admit it. Every day people g roup objec t s and
even other people into clusters and make decisions based on it. During
the last few years we have come to realize that Iwrkers 1n diverse
fields who us e computers to build sounde r c lassifications have created
methods that either are very similar or are di ffe rent but very appli -
cab l e in other fields. Thus it is refreshing to attend the annual
meeting of the interdisciplinary Classification Society and hear about
the methods used to clust er United Airlines passengers , languages,
countries and even b reak fas t ce r e al s ! Such meetings ce rtainly st imu-
late us to get out of our normal thought-rues.
Once classes have been decided upon, s ub sequen t organisms must be
assigned to one of the c l asses. This is the interest of the interdis -
c i plinary Pa tt e rn Recognition Society .
Discriminan t functions (Blackith and Reyment, 1971 ; Sea l, 1964)
were developed by Fisher in the 1930's when Edgar Ande rson presented
him with the problem of hO\~ best to discriminate betl~een species of
22 467

Iris . The procC!dure assigns c harac ter s dif f e r en tial weight s accorcl-
ing to th ei r relative ability to discrimina t e among c la sses . That
is, given a large basic data matrix with objects partitioned i nto
classes . t he procedure locates the optimal charac te rs to us e in id e n-
tif ica tion . Anderson's (1 949 ) hybrid i ndex is a c rude method i n terms
of sta tis tics , but as Goodman (1967) found recently, for given situa-
tio n s it may b e just as u se ful as discriminant funct ions . Jus t why
t he hybrid index and its comp l emen tary technique of the pic t or iali zed
bivariate scatt e r diagram a r e suffic ie n t to separa te close l y relate d
classe s is an inte r e sting bio l ogical qu es tion i n terms of th e minimum
numbe r of characters necessary to separate se t s of organisms tha t
occupy different niche s . Of course, if the number of characters or
classes increases above 10 , or 3 , respectively , Anderson ' s methods
are difficult to use .
5. Geomet r y . Nany r elation ships can bet ter be described and a nalyzed by
geomet ri c method s . al t hough algeb raic representat ion of the same
system is lLsually possible . Graph theo r y in general an d nett.,tQrks in
particular a re receiving increased att ention as activities in numeri-
ca l taxonomy i nc r ease . A graph is a geomet ri c st ruct ure con sis tin g of
point s (vertices) in space connected by a system of c urves ( edge s). A
dir(!;cted graph is o ne in 11hich e very edge has been ori e nted, or gi ve n
a direc t ion . An undirected graph lacks such orien tation . Fi na lly, 3
gra ph i s open if one canno t return to a vertex wi t hou t using at least
on e edge twice . In contrast , a closed graph is one wherein one can
r eturn to at l east one vertex without traversing any edge more than
once . Readers may expand their knotlledge of graph t heo r y beyond these
simplified definitions by reference to Busacker and Saaty (1965) . I n
the a bove terms most evolutionar y cro ssing polygons are closed , un-
directe d g raphs ; most phenograms and cladograms (without reticu lat e
evolution) a re open, directed graphs , and some nearest - ne i ghbor
grap hs are closed , und irec ted graphs . A network is a directed graph
without any loop s . A shortest spannin g tree is any s hortest possible
netwo r k connectin g a set of roints . Tabl e 22-3 inc l udes r efe r ences to
each of the four types of g raph. Why try to exp res s r ela tion ship s in
dif fe rent forma l sys tems, s uch as graph or set theory ? One r eason is
that the simp l e r ep r esentat ion in the new format is efficie nt. It
a lso provides a valuable insight into othend se ahstruse r elation ships .
Anothe r area of geomet ry that sho uld re ceive more attention i s
topology , or " rubb er sheet geomet ry . " D' Arcy Thompson (1942) demon-
strate d it s value to systema tic s when, by r egular transfo rma t ions of
points on a g rid placed ove r a dral-ling of one species o f fi sh , for
example , he co uld produce exactly the shape of another species of
fi sh . Recently Snea th (1 96 7) has expanded th is work in two directions
by conside r ing three dimens ion s Simult an eously instead of two , and by
avoid ing the use of an arbi trary s tanda r d " species " like Thompson' s .
The availability of digital compute rs and inc r e mental plo tte rs elimi-
nates the need for selecting a standard. All combinat ions can be tried
quickly. Coupled with s tudies of allometry, this techn ique i s j u st
wait i ng t o be uS,ed by plant syst e matists to help to underst a nd what it
takes developmentally to get from one plant habit to another. o r a
flower or leaf shape o f one species to t hat of another .
22 467

Iris. The procedure assigns charac t ers dif f eren t ial weigh t s accord-
ing to their rela tive ab il ity to d iscriminate amon g c la sses . Tha t
is , given a large basic data matrix \... ith objects partitioned in to
classes . the procedure loca tes the optimal charact ers to u se in i den-
tification . Anderson's (1949) hybr i d index is a cr ude method in terms
of sta tis tics , but as Goodman (1967) found recen tly, for given si t ua-
tions it may be just as usefu l as discriminant f unct ions . Just why
the hybrid index and its complemen tary technique of the pictoria li zed
bivaria t e sca tter diagram are sufficient to separate closely re l ated
classe s i s an inte resting biological quest i on i n term s of the minimum
number of characte rs necessary to separate sets of organisms tha t
occupy different niches . Of course , if the numbe r of characters or
cla sses increases above 10 , or 3 , respe cti vely , Anderson 's methods
are difficult to use .
S. Geometry . 1·la ny relationships c an better be descr ibed and anal yzed by
geometric methods , al t hough a l gebraic r ep rese ntation of the same
sys tem is usually possible . Graph theory in gene ral and netHorks in
particular a re receivin g increased attention as activities in numeri-
cal taxon omy i ncrease . A g r aph is a geome tri c s tructure consistin g of
points (vertices) in space conne c ted by a system of curves ( e dges). A
direc ted graph is one in Ivhich e very edge ha s bee n oriented, or given
a direction . An undirected graph lacks such orientation . Finally, a
graph is open if one canno t return to a vertex without usin g a t l east
one ed ge twice . In contrast , a closed graph is one wherein one can
r eturn to at least one vertex without traversing any edge more than
once . Readers may expand their knmvledge of graph theo r y beyond these
simplified defini t ions by reference to Busacke r and Saa ty (1965) . I n
the ab ove terms most evolutionary cro ssing polygons are closed , un-
direc ted graphs ; most phe nogr ams and cladograms (witho ut retic ulat e
evolution ) are open , d irecte d g raphs, and some ne arest -nei ghbor
graphs are closed , undirec ted graphs . A network is a di rected graph
withou t any loops . A shorte s t spannin g t re e is a ny s hortest possible
network connectin g a set of poin ts . Tabl e 22 - 3 includes r eferences to
each of the f our types of graph . Why t r y to express relationships in
diffe rent formal sys tems , s uch as graph or s et theory? One r e ason is
that the simple r ep r esentation in the new format i s eff icient . It
al so p r ovides a valuable insight into othe nvi se ahs tru se r e l a t ionships .
Another area of geomet ry that should rec eive more attention is
topology, or "rubber sheet geometry." D' Arcy Thompson (194 2) demon-
stra t ed i ts val ue to systema tics when, by regular t r ansforma tions of
points on a g r i d placed ove r a dral-li ng of one species of fish , for
example, he could produce exactly the shape of another species of
fi sh . Recent l y Snea th (1967) has expanded thi s work in two directions
by con sidering three dimens ions Simultan eously ins t ead of two , and by
avoiding the use of an arbi trary standard " species " l ike Thompson ' s.
The ava ilabilit y of d i gital comput ers and incrementa l plott e r s elimi-
nates the need for selecting a standard. All combinations can be tried
qu ick l y . Co upled with s tudies of al l ometry , th is t echnique is just
Ivaiting t o b e us ed by plant sy stematists to help to understand \-Ihac it
takes developmentally to get from one plant habit to a no ther , or a
f l ower or leaf shape o f one species to t ha t of a no the r.
468 22

Computers are responsible for the recent rise of activity in multi -

varia t e character variat i on . Hany of the st udies referenced in this a r ticle
would be impossible without them. Ste rling and Pol l ack (1968) provide a good
introduction to the digital computer and describe some of its uses . I n syste-
matics the digital computer i s being use d in two maj or ways : to carry out
calculations and then. often graphically , to summarize the r es ults ; they are
also used to retrieve information from large data banks s uch as described
earl i er under data accumulation . Analog computers are the other main type of
computer . These are special ly designed to so lve systems of differential equa -
tions. Their major advantage over digital computer s is that the user can
observe the effect of change of parameters on the system simply by changing
the val ue of any of severa l potentiometers. Choice of the type of computer
to use is a function of the model itself, of the purpose , of the amount of
data , and other considerations . Hybrid computers , combinations of digi tal
and analog computers, are available. Often, qualitative scanning of res ults
of a systems analysis is made on an analog computer and then, \-Then more accuracy
is desired for a specific combination of parameter values , the digital com-
puter is employed.

D. Summarization Gr aphiCS
Table 22 - 2 was diffic ult eno ugh t o present, but Table 22-3, contain-
ing the types of s ummarization graphics used in character variation analy-
sis , is even more so. Ideally, eac h type listed in Table 22 - 3 should also
be represented by a f igure in the text, but the cost and space needed
would be prohibitive . Several examples are given , howeve r, both he r e and
in chapter 21. The references provided next to each item in Tabl e 22-3
Here chosen primarily from publications that should be on the shelves of
any systematist, or in his institution ' s library . Every entry is given a
referenc e because al l entries could be used by plant systemat ists . Also ,
unless one looks at an actual figure, it is difficult to understand its
classification as to type and also how it differs from othe r s.
1 . Frequency Distributions. Fre quency dist r ibutions (category A of Table
22 - 3) are among the basic tools of any systematist . They a r e
graphs whose respons e variable is either the relative or absolute
frequency of the different states of the charact ers on one or more
other axes. For convenience they can be separated primar ily into
univari ate , bivariate, and multivariate frequency distributions.
Univariate distributions are f urther recognized in Table 22- 3 as
discontinuous or continuous, depending on the nature of the char-
acter being anal yzed . The most common t ype of discontinuous,
univa r iate frequency distribution is the bar diagram.
When we turn to continuous univariate frequency distributions,
histograms and freque ncy polygons a r e most familiar . A frequency
polygon is a frequency distribution \.;rith the frequency of each
recognized state plotted as a point on the graph and the set of
such points connected to its nearest neighbor on each side by a
straight line or by a higher degree c urve . Types of freque ncy
polygons impo rtant in systematics include relative frequency dis-
tributions (probability density functions), frequency distribu-
tions of the s tates of an index or other compound character, such
as an environmenta l gradient or discriminant fun c tion .
A Dice-gram is a type of f requency distribution that has been
compressed into one dimension or otherwise topologicall y t r ans-
formed. Sokal (1965) has commente d on the visual bias involved in
such graphs.
22 469

TABLE 22-3 . Summarization Graphics*

A. Frequency Distributions

Univariate frequency distributions:

Discontinuous :
Bar diagrams (Davis and Heywood , 1963 : 326)
Sets of horizontal lines of uniform thickness . one per species
(NacArthur and Connell , 1966 : 167)

Continuo us :
Histograms (Hayr , 1963: 1 88; Saka l and Snea th , 1 963 : 214)
Frequency pol ygons (Sokal and Rohlf. 1969 : 30)
Sets of shaded freq uency distributions (Odurn, 1956: 152)

Bivariate frequency distributions :

Hith isolines , hut flat (Seal, 1964: 102)

tolith different-sized circl es . but flat ( Sokal and Sneath , 1963: 214)
\.Jith isolines . but in perspective (Sakal and Rohlf . 1969: 501)

Hultivariate frequency diagrams

,
B. Scatter Diagrams

Unreduced axes :

Biva riate scatter diagrams:

Plain (Mayr, 1969: 172)
Di f fere ntia l shading (MacArthur and Connell , 1966 : 154)
Equa l frequency el l ipses (Sakal and Rohlf , 1969: 527)
Regression lines and other curves (Sokal a nd Rohlf, 1969)
Iso 1ines (Grieg-Smith, 1965: 193)
Pictorialized scatte r diagrams (Davis and Heywood, 1963: 477)
Illustrated sca tter diagrams (Smith, 1966: 84 )
Directed networks (Allee et al . , 1949: 208)

Trivariate scatter diagrams:

Bar diagrams (HcKell et a1.. 1969)
Regression and other respon se surfaces (MacArthur and Connel l .
1966, 161)
Triangular graphs (~~cArthur a nd Connell, 1966 : figs . 1- 16)
Perspec t ive drawings (Figure 22 - 2)

Reduced axes:

Reduced character space axes:

Princ ipal components axes (Figure 22-2)
Discriminant functions (DuPraw , 1965)
 470 22

TABLE 22 -3 (Continued )

Canonical axes (Oxnard, 1969)

St e reograms (Rohlf, 1968)

Reduced character s pace axes and ne t wo rks (Figure 22- 2)

Reduced object space axe s (Cravello, 1968c)

Character space a nd reduced character space combined (Davis and

Heywood, 1963: 66)

C. Directed graphs

Open

Branched (denclograms, trees ):

Taxonomic phenograms:
Two-dimens i onal (Figure 22-3)
Three-dimension al (Waterman and Horowitz, 1965: 270)

Cladograms:
Pla in (Dayhoff, 1969; Mor ishima. 1969)
Illustrated (Figure 22- 3)
Wagner trees (Figure 22-3)
Cladochronograms
Typological diag r am (Mayr , 1963 : 599)

Phy l og r ams :
Two-dimensional (Eaton, 1970)
Three-dimensional (Saka l and Snea th: 234)
Il lustrat ed (Davis and Heywood, 1963 : 70)
Cha racte r cladograms (Davis and Heywood: 189)

Closed:

Flow charts ( Sakal and Crove llo , 1970)

Some near es t-neighbor taxonomic graphs (Figure 2 ; Wirth et al . , 1966)

Directed crossing experiments (Figure 22-4)

D. Undire cted graphs

Open: Some nearest-neighbor taxonomic graphs

Clos ed (Gabriel and Saka l, 1969)

22 471

TABLE 22-3 (Continued)

Some nearest - neighbor taxonomic graphs (Prance et al .. 1969)

CrosSing experiment or pollen fertility polygons (Figure 22-4)
Character correlation polygons (Clausen, 1967)
Polygram (Davis and Heywood, 1963 : 331)
Polygonal graphs. Polygraphs (Figures 21- 8 and 21-10)

E. Geographic maps

Profile diagrams :
Plain (HacAr thu r and Connell, 1966 : 76)
IUth frequency distribution (Lewis and Taylor, 1967: 223)
With glyphs (Kendeigh , 1961: 28)

Uniform. discrete symbols by computer (Perring and Walters, 1962)

Uniform symbols of different size (Nayr, 1963 : 263)
Different symbols (Nayr , 1963: 344)
Glyphs (Figure 22- 6)
Actual character values (MacArthur and Connell , 1966: 53)
Isophenes (Adams, 1970)
Dice-grams (Sokal, 1965: 355)
Multiple comparison homogeneous lines (Sokal, 1965: 358)
Undirected networks (Figure 22- 7) ,
Directed networks; cladograms (Figure 22-8)
Bar diagrams (Figure 22-9)
Pie diagrams (Mayr, 1963: 431)
Differential shading (Mayr, 1969: 191)
Character polygons (Figure 22-10)

F. Misce llaneous

Photographs and drawings

Differential shadin g of cells of a matrix (Figure 22 - 1)
Pie diagrams (Odurn, 1956: 380)
Skyline plots and sequence diagrams (Prance et al., 1969; Wirth et al., 1966)
Contour diagrams (Hoss, 1967)

*Parentheses next to each category or item contain references to pub l i-

cations that use the item or explain it. Each reference is separated from
the next by a semicolon . If a colon appears, the information immediately
to its right refers to specific pages or chap t ers.
 472 22

Biva ria te f reque ncy distributions are natural extensions of the

above and are most frequently presented in two dimens ions with changes
in frequencies indicated by 1s011nes or by d if fer ent-s iz ed circles .
With the growing availability of computer graphics software packages
(e . g ., Rohlf, 1969). we may expect per sp ective drawings to be come
more common i n sys t emat i cs . Hult ivariate frequ ency distributions
a r e put in a s a separate categor y on l y to serve as a s timu lus t o
r eaders to consider the shape that such graphs might take--be side s
those in two dimensions that have several d iffe r e nt scales for the
abscissa.
2. Scatt er Dia grams. These are graphs in one or more d imensions on which is
located a set of points , each indicating the position of an ob jec t
(Ca tegory B of Table 22-3; also see Figur e 21-12) . The axes may be
r egular characte r s or they may be some combination of chara cters , such
as a principal components axis . !.J'ithin this oonce pt of scatter dia-
grams , the frequency distributions d i scussed above are just a par-
ticular type of scatter diagram. I separate them on l y because they
are so often used and are such an involved entity in the mse lve s .
Fo r convenience only , I separate the scat ter diagrams listed in
Table 22 - 3 into those involv in g un reduced (r egula r characters) axes
only an d those i nvo lving at leas t one reduced axis . The meaning of
the dif fere nt types and subtypes s hould be c l ear from their names and
from the examples cited . The only very recent type is a figure that
con tain s reduced character (or obj ect) sp ace axe s and networks . For
examp l e, consider a sty rof oam model or a pers pe ctive d rawing tha t d i s-
plays tbe positions of a set of obj ects in the first t hree axes of a
principal components spa ce . Assume the po sition of each object is
represented by a sma ll circle or a sphere . As men tioned ear lie r,
this gives the best (according to the l eas t -squares c ri te rion) three -
dimensiona l view of t he relationships among the ob j ects. But to be
comp l etely accurate, we r ea lly need mo r e than three d imensio ns . One
way t o see what would hap pen to the obj ec t relationships if we could
see furth e r i n to the hyperspace is to draw a dire c t ed l ine on the
drawing from each objec t to its near est ne ighbor in the hyperspace .
This neares t-neighbor info rmation is available from the object by
obje ct similarity or distance matrix cal culated e arli er . Rohl f
( 1970) projected the sho rtes t spann ing tree down to three d i mensions
of a principal components analysis . I feel st rong l y that f or man y
purposes i n systematics , this i s the best po ssibl e summa r y of descrip-
tive relations hips in a mult i dimensional charac t er spac e especially if
the graph ends a re directed (Figure 22 - 2) . Natura lly one could add
a second netwo r k o f directed edges (using dashes ) to inuicate t he
second c l oses t neighbo r, or perhaps the fa r thes t removed. The only
probl em is that as success ive networks are added. the figure becomes
harder to comprehend .
3. Dir ected Graphs . Categories C and D o f Table 22 - 3 more fully conside r
graphs a nd networks. As defined in the disc uss ion of Tab l e 22-2.
graphs can be separat ed into op en or clos ed and direc ted or und irected
g r aphs . Busacker and Saaty (1965) reserve the term network for open ,
directed graphs . but t he use of the term is no t uniform. Unlike Mayr
(1965) I reta in the t e rm dendrogram as a gene ral t e rm to mean a ny
t r eelike diagram . Busacker and Saat y (1965) define a tree as a g raph
t ha t is connected and has no c ircui t s . A tree would i nc l ude our open,
directed and open , undirected graphs . To di scover me tho ds to cons truct
22 473

graphs that require involved cal culations, such as phenograms and cladograms ,
consult the references of Table 22-2, Sections C and D.

Category C presents uses of directed graphs in character variation analy-

sis. The first type of open , directed graphs are dendrograms . Most dendro-
grams li ste d in Table 22-3 do not have exp li cit directed edges . But they are
implied either by direction of descent , as i n cladograms , or by the property
of incl usion , as in phenograms . Undirected cladograms would belong in Cate-
gory D of Table 22-3. hut phylogenetic systematists usually are willing to give
directions. Whether they are a lways warranted is a different matter .

Phenograms are directed graphs that r eflect phenotypic relationships .

The result of c l uster analysis , they may be of two or more dimensions. They
may be directly derived from an object - similarity or distance matrix which
is based on a character by object basic data matrix. Or they may be derived
from a similarity or distance matrix from reduced character or object space .

Cladograms , graphs depicting the branching sequences of evolution, have

been a major systematic tool for years . Cladograms can be drawn in two dimen-
sions, since the only property they r eflect is the branching sequence of evo-
lution. Over the last five ye ars many methods of const ructing them by com-
puter have been introduced . At regular i ntervals Dayhoff (1969) recompu te s
cladograms on the basis of the expanding data bank of known macromolecular
sequences . Cladochronograms are included for completeness to emphasize that
if one also has accura te time estimat es of the branching events , he no longer
has just a c ladogram. A typological diagram is a peculiar cladogram which
shot~ s series of changes over time but without branchin ~.
,
The term phyl ogram is r eserved only for those grap hs that summarize in for -
mation on the cladistics, chronistics, and phenetics of the group under study .
If information on any of these essential parts is missing , t hen the graph is
not a phylogram . but a cladogram or phenogram . Some phylograms are naked , in
the sense that only the extan t objects are known. Others are full , in that
taxa at branch points in the past are known in detail. Paradoxically , naked
phyl ograms are much easier to fit into the formal taxonomic system than full
phyl ograms . l.Ji t h naked phylograms we are not required to make arbitrary deci-
sions to determine with which branch the ancestral types should be placed
when we are delimiting formal taxa . The taxonomic hierarchy is characte rized
by the property of inclusion of s ubsets within sets. Thus , species are in a
genus , genera in a family, etc . But phylograms are characterized by taxa of
the same catego r y at each br anch point as , fo r example, in the wel l-known
horse sequence , where Parahippus supposedly gave rise to Herychippus and it
gave rise to Hippa rion, Nannippus, and so fort h (Eaton, 1970). I cannot under-
stand why systematists wish t.o distort supposed phylogenetic r elations by
making them fit into a taxonomic hierarchy that is defined by the principle
of inclusion . It seems more sensibl e to pre s ent two separa te graphs , one a
taxonomic tree (phenetics) and the other a phylogram or a cladogram. Then
one can see where they disagree and can ask the important evolutionary
question, why do they disagree?

What about reticulate evolution? The definition of cladogr am given

above is based on open branching processes and does not consider fusing
processes which are thought to be common event s at the micro evolutionary
level. Algorithms to const ruct cladograms and other closed , directed graphs
474 22

at the. species level and b e low should be a f r ui tful field of future research .

A charact e r c l adograrn is a graph that depicts successive changes in the

states of a characte r, or of a su i te of characters , over evolutionary time .
Examples include the change of angiosperm ovary and placentation types (Law-
rence, 1951 : 74 , 76) or of amino acid se quences in a prote in . Much can be
learned about the processes of evo lution by the construction of character
cladograms . Since only one or a fe\" characters are cons ider ed in a character
cladogram , the problem of possible reversals in the e volution of the charac-
t er states is more acute. Systematists have neglected allometry and the r ela-
tion of fo rm and function of a set of adaptively correlated characters .

NORTH AMER ICA SOUTH AMERICA

.,,,., ,.' ," -, ....

-- --- -- - - - -
OU" - L . . .. "

-- -- - ---- - ,-I-:~----
~,o:. -I--j ' 0' .. ,

,
- -- ,

,,
II
II
- ',~~ ;,;~""'"
,,
,,
Fi r,u r e 22-3. Hagner "target" diagram int1icatinr, a probahle phylogeny for
Gutierrezia according to a non-computerized method . Habit of plants dravn
to Rcale; hroken lin es indica t e l ess \·!ell established relationships , (From
Solhrig, O. T , 1 970. Th e phy logeny of Gutierrc7.ia . Brittonia 22(3): 217-
229 . Used Nith permission , J
22 475

Hany taxonomic keys used to identi fy unknown specimens are open , directed
graphs.

Flm" charts th at serve as a pr e liminary step i n complex computer p rogram-

ming are common i n sys t ema tic studi es . The f l ow char t devised by Sakal and
Crove llo (1970) to del imit biologica l species i s also a c lo sed, directed graph .

A closed, directed graph some time s represents the nearest neighbor rela-
tionship among a set of objec ts that a systematis t is studying . Undirected
graphs are used by some , but this is l ess info nnative since a neares t-neighbor
line connecting two ob j ec ts is not necessarily true for both directions. One
may be closest to a third obj ect . The results of any directed , crossing exper-
iment c an be pre sented as a c lo sed , directed g raph ( Figure 22-4) . Natu ra lly ,
a l og i c al re finement of s uc h a gr aph wou l d be to use di ffe r ent symbols (solid
line s , dashes, dotted lines) on one graph to ind icate t he s trength of the
c rossing relationships , o r s i mply the orde r of the r e lation ship . Reade r s
s hould reali ze t hat di r ec ted graphs have more informa t i on than undirected
g raphs . So where possible, use a directed graph .
Flo wer ing F1 , 60~ or mor e
stainable pollen
Vir ginianum
Flowering F1.O'7. stain~bl e
pi 10$um
Crou p pollen
n· 39
___ • _. Fl vegetat ive only
, lefluifofium

,
,,
,
, ,,
,,
.- ,
,'

se/osum Fluuo Slim Croup

n. 38 chnQPQdiQides

.. lbeHens In canum Croup

n,I9

,: ,~
loomisii pre n enl hemo ide II

n. 19 n.3 6
."

n' 3 8

Figure 22--4 . Cro ssing diag ram o f artificial hybridizations . ArrOHS point from
po llen parent to pistillate parent . [From Chambers , H. 1. , and K. 1. Chambers .
1971. Art i f icial and natural hyhrids in Pvcnallthemum (La bia tae) . nrittonia
23 : 71-88 . Used \-lith permission.]
 476 22

4. Undirected Graphs. Category D of Table 22-3, undirected graphs , also can

be separated into open and closed graphs. Bot h open and closed , undi-
rected networks are exemplified by some nearest-neighbor diagrams. I n
these and all previous references, the reader should be aware that
successive nearest - neighbor graphs are possible and often are quite
useful. Character correlation polygons ai:e another example in which
sets of lines are added in successive operations . Wirth, Es tabrook and
Rogers (1966) alter this procedure by showing successive clustering
stages .
Polygonal graphs are polygons embedded in a coordinate system from
whose origin extend several radii, each calibrated \-lith a different
character's character states . The polygon is formed by joining the
one state of each character possessed by the object (See Figure 21-10).

ATLA~U

/\0
SPRINGVILLE

POLLEII FERTltlTY
o_ 19~
l'O
RELAnONSH~,,,~~2~~:===~~~~~~~l
_)') ~ ._ ......
ARIZONA
~O_S?~· --
10_1910 _
!O_l00 ~ _

Figure 22-5. Pol l en fertil ity relationships among various populations of

Clarkia rhomboidea. [From Hosquin , T. 1966. Toward a more useful
taxonomy f or chromosomal races . Brit tonia 18! 203-214. Used \-lith
permission . 1
22 477

5. Geog r aphic Maps . Category E, geographic maps , is extensive . For-

tunately, t he meaning of each item is fairly evident and I shall
for ego rev ie~" in g them here . The references given in Table 22- 3
should be adequate for understanding .
6. Niscellaneous. The first three en tries of Category F of Table 22-3
a l s o are se l f - expl anatory . Skyline plots and se que nce diagrams
are interesting , different methods clearly pr esented in the
references cited . Finally, Hoss ' s (1967) contour diag rams are
quite different from all other me thods of depicting relation-
ships among objec t s . Interested readers shou l d consult his
paper for details .

E. Eval uation of Result s and Reanalysis

Evaluation of results is a serious ste p in char acter variation ana l y-

s is. Fi r st . the rese archer ascertains wha t the result s mean to his origi-
na l purpos es and hypotheses . Have they f ul f i lled and s upport e d , or have
they negated, t hem? Sec ond , at this s ta ge the analyst must integrate his
findings with tho se of other s . This is becoming increasingly difficult
because of the expanding number of produc tive worker s . Perhaps a com-
puteri zed data bank will he l p to alleviate this problem .

Reanal ysis, either by the same or by a subsequent invest igator , is

a necessary endeavor that serves as a constant check on our theories and
on the accuracy of our facts. Reana l ysis is justified by an i ntensional
or an extensional expansion of the da t a . Intensiona l expan sion involves
co llection of mo r e of the same, whe the r it be more tempe r at ure r eadings ,
from the same place at the same time of year or more individuals fro m
sites a l ready s tudied . Extensional expansion involves the collec tion of
data f r om new systems , or f rom the f ie ld or from a data bank . The dis-
tinct i on between intensional a nd extensional data is somewhat arbitrar y
since it can be negated with certain examples, but I think it has some
value .

F. The Dvnamic Data Bank

We now turn to the f i nal stage in our multistage decision process

(Table 22-1). Until recently , at the close of a character variation
analysis, information not published was usual ly stored in one ' s office
and then discarded when the space it occupied was needed for a later
project . The outstanding exception has been the maintenance of colle c-
tions of animals and plants i n museums and herbaria throughout the world.
Earlier i n this article I outlined the differen t s ources of information of
actual or potentia l value to syst ematists .

I do not wish to be labo r the obvio us. Given the potential of j us t

the computing equipment avai l ab l e today , what I call the dynamic data
bank could become a real i t y. Consider the experimen t al data from eve r y
character variation analysis that has been irr etrievably f i l e d in exper i-
ment notebooks that are being discard ed r egularly . Today , such data of t en
can be put onto punched cards or tap e to speed up the initial analysis .
As a by-product of this pr ocess , the data can be deposited into a r egiona l
data bank . Also , optical scanners are now commercially available which
" read" any of 2000 different type fonts and put t hem direct l y onto
478 22

Figure 22-6. Distribution of

Clematis columbiana. Closed
circles with arms to right,
extreme form of var . columbiana;
closed circles \."ithout arms,
"repe!ns" fo rm of vaT. c.olum-
biana; half-closed circles,
intermedi.ates between var. col-
umbiana and var. tenuiloba; open
circles Hithout arms. extreme
form o f var. tenuiloba; open cir-
cles \.,rith arms to left , var .
tenuiloba Hith coarsely divided
l eaves ; open circles Hith arms
dOWTIHard, var. tenuiloba \"ith
elongated aerial stems; x's,
probable hybrids betHeen .f.
columbiana and C. occidental is.
Glyphs Hith short arms represent
coll ect ions Hhich ,,,ere variable
Ivith respect to the trait indi-
cated. [ From Pringle, J. S.
1971 . Taxonomy and distribution
of Clemat is, Sect . Atragene (Ra-
nunculaceae), in No r th America .
Brittonia 23: 361-393. Used
\"ith permission .]

Figure 22-7 . Correlation coefficients

of the terpene composition spectrum of
seven populations of Rursera microphylla.
Absence of line between localities indi-
cates no correlation. [ From Hooney , H. A.•
and {oJ. A. Emboden. Jr. 1968. The rela-
tionship of terpene composition, morphology
and distribution o f populations of Bursera
microphvlla (Burseraceae). Brittonia 20:
44 51 . Used with permission . ]
22 479

I , L. UI. LU~

Figure 22-8 . Hypothetical evolu-

tionary relationshi ps among the
~"hite-rayedtaxa of Helampodium.
[From Stuessy , T. D. 1971 .
Systematic relationships in the
Hhite-rayed species of !1elampodium
(Compositae). Brittonia 23 : 177
190. Used with permission .]

..

•
pedicel le ngth
,
I
petiole/blade
pub",y r"n<e
/ //colyx lobe len g th

WlJ
Figure 22-9. [From Horley, T. 1'9'71 . Geographic variation in a Hidespread
neotropica l species , Mouriri myrtil l o i des (He l astomatac eae). Br i ttonia 23 :
413-424. Used with permission . ]
 480 22

A
B

9 C

F • o
E

Figure 22-10. Graphic

r ep resentation from popu-
1ation samples of Ver-
ncoia marginata and ~.
tenuifolia (sensu Gleason)
of Means for : floNers per
head (A); ~"idest bract.
mm. (n): involucre length,
mm . (C); invo l ucre 1Vidth, mm .
(D); leaf margin score (E); bract
tip shape score (F); leaf blade length,
em . (G); and leaf blade width , em . (1-1) .
Geographically the two southernmost polygons
represent~ . tenuifolia . [From Jones, S. B. 1972.
A systematic study of the fascicu l ate group of
Vernonia . Rrittonia 24(1) : 28-45 . Used Idth
permission. ]

a magnetic tape . Not only I"Quld this make published material contained in
floras and faunas avai lable to researcher s for new and valuable analyses. but
also, with increasing publication costs, such a data bank system might gradu-
ally replace formal journals as we know them. Such proced ur es are not without
problems, however (Crovello, 1968b; Keller and Crovell o, 1973). The big prob-
l em in systematics, and in biology in general , has been the lack of a l arge,
driving force actively campaigning to bring about a data bank system . The
full value of such data banks to future work is still unexplored . Clearl y ,
the taxonomic community must organize now. The methods of implementing such
a data bank system are available.

CON1'iON PROPERTIES OF CHARACTER VARIATION ANALYSES

Even in the above brief survey of character variation analysis in syste-

matics certain connnon properties emerge . The purposes of the analysis are the
description of the pattern of variation in nature and its explanation. Most
character var i ation analyses in systematics have the same building blocks:
i.e ., characters, objects , and system.

Another common property that emerges is that any character variation

analysis is a multistage decision process and the decisions made at each stage
are arbitrary to some degree . All use the same general process , and systema-
tists should benefit from the study of methods used in analyses in areas other
 22 481

than th eir own specialty . This also helps us to get out of our though t-rut s .

Op timiza tion i s a propert y of all systematic stud i es . But optimization

can occur at al l stages of a char acter variation analysis a s well as at dif-
f e ren t l e vel s within one s tage . Taxonomists ~... ish t o maximize the val id ity of
a phylogram or of a phenogram to ref l ec t true r elat i ons hips . Evolut ionists
us ually consider mode l s of evolutionary processes as tend i ng towards op timal
l evels , eithe r in i so l at i on or, more reali s tically, with all processes con-
sidered a t once . Of course, eve r yone attempts to optimiz e t he use of his
own t ime.

Hany have observed that most knowledge is or ganized in to hierarchies .

Any taxonomic system i s a hie rarchy. Aside f rom i ts general hierarchia l
arrangement the re a r e more specialized hierarchies in the sys tem. These includ e
a hie rarchy of purposes, or objectives . I n any complex character variation anal-
ys is, deci sions must be made a s to the direction research e ffor ts should t a ke ,
and t his is determined by the re l a tive i mportance of the purposes and of the
hierarchial re l a tion among them. The problem with cha r acte r variat i on analysis
in systematics is th at it cont inually s uggests so many i nte r es t i ng side pro j ec t s
tha t it i s di fficult to keep t he main pu r pose i n mind. Data hierarchies were
disc ussed ear l ier. These a r e formed by partitioning the original cha racte r
by obj ec t data matrix into more compact matrices . Thus, char acte r i nforma tion
on several species in many sites may be compressed i nto a character by spec i es
tab l e . Character hie r archies are possible . Davis and Heywood (1963) refer to
mul tiple c hara cter wh i ch can be divided into severa l sma l ler cha ra c t e rs. For
example , type of l e af pubescence (a charact er ) can be further separated into
characte r s l ike density of pubescence. length of ha i rs, number of ce lls per
hair , and hair color. Sampling procedure hie r archies occ ur when differ ent
samp ling t echniques mus t be comb ined in a succe s sive procedure. To obtain a
f air es timate of the characte r istics of a species , a s tratified sampling desi gn
may be con st ructed throughout equal areas of the spec i es ' range , but then
s amplin g within each area wil l be random . Decis ion hierarchies are foun d i n a
complex systems analysis where de cisions are made in several subs t ages and the
r es ulting to t al outcome i s us ed to make a decision at the maj or stage . Also ,
in any s tudy of a n e volu tionar y sys t em one can think of a con trol s trategy
hierarchy . For exampl e , i f a specie s such as marijuana i s to survi ve in cul-
tivated areas, what are the cont rol s tra t egi es of the species as a whol e , of
each population in different areas o f t he world, a nd of each individua l i n a
gi ve n population? I r ealize that this las t is a nt hropomorphic to say the
l east . Bu t if it pr ovide s a worke r with i nsight s into the systems of inte r e st
t o him, it i s a valuable concep t . Last, and perhaps most impor tant , i s the
conc ept of a hi e r archy of chara c ter variation sys t ems . Tha t is , how can we
group separate c haracter var iat ion a nalyses to pr ovi de maximum value to futur e
work i n systematics?

We also should not be s o conce rned with the analysis of variation that we
f ai l to be aware of the oppos i te si tuat ion, i nvaria nce . I n any sys tem cer-
tain cha racte r s are invar iant over all or part of the set of objects. What
does t his mean in t erms of the action of environmental s e l ec t ive forces , of
control strategies , and other piological event s and evolutionary r esponses?
 482 22

COMMON PROBLENS OF CHARACTER VARIATION ANALYSIS

Nost read e r s are awar e o f t he probl ems of character variation analysis.

Among t hose a l ready mentioned is the effect of a r bitrar y decision-maki ng
(conce rning the weighting of c haracter s , the clu stering of objects , and t h e
choice of the characters and parameters to be measured in a system ) . The
proble m of data compatibility at all l e vels (choice of charact ers , character
states, etc .) also HilS di s c u ss ed . An addi tional prob lem is dete rmining whether
an i nference abou t some specific part of the cha r acte r variation study , or the
t otal study , i s warranted o r unwarran ted . Also , how does one overcome redun-
dancy in data a cc umulation beyond t hat which is necessary fo r verification?
The t ime e l ement is a problem . How can one i nsu r e tha t data co llected se pa-
rat e ly, at different times and pl aces by di ffe r ent people \olit h differ en t
instrume nts , are at all comparable?

ANALYSIS OF CHARACTER VARIATION LITERATURE

Adams, R. P . 1 970 . Contour mappin g and d i fferential sys t ematics of geographi-

cal variat i on . Sys t emat i c Zool ogy 19: 385-390 .
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Philadelphia .
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systems for herbaria. Canadian Journal of Botany 50 : 21 97- 2209 .
Blackit h, R. E. 1960 . A synthesis o f multivariate techniques to distinguish
pattern s o f growth i n grasshoppers . Biometrics 16 : 28- 40 .
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Academic Press . Ne\~ York .
Bossert, H. 1969. Compute r techniques i n sys tema t ics . I n : Systematic Bio-
logy . Pub1. No . 1692 . National Academy of Science , 595-605. \.Jash ington .
D. C.
Burnaby, T . P . 1966 . Grm... th i nvariant descriminant functions a nd ge neralized
distance s . Biometrics 22(1) : 96-110 .
Busacker , R. G. , and T . L. Saaty . 1 965 . Finite Graphs and Networks : An
In troduction and Applications . l-lcGraw- llill Book Company . New Yo rk.
Clau sen , J . 1967 . Biosystematic consequences of ecotyptc and chromosomal
differentiation . Taxon 16 : 271- 79 .
Cochran, \~ . C.• and C. N. Cox . 1 957 . Experimental DC!signs . Hiley . New Yor k .
Col e, A. J . (Ed . ) . 1969 . Numerical Ta xonomy . Academic Press. New York .
Crovello, T . J . 1968a . A numerical taxonomic study o f the genu s Salix. sec-
t ion Si tchenses . Un iversit y of Califo rn i a Pub lication Bot. 44 : l-61.
1968b . The effect of missing dat a a nd of two so ur ces of charact er
values on a ph e n etic study of the willows of Ca li fo rnia . l-!adrono 19 : 301-15 .
1968c . The eff ec t of alte r a tion of techniq ue at two stages in a
numerical ta xonomic s t udy . Un iversity of Kansas Science nulletin 47(12):
761-85 .
1968d . Key communality clus t e r analysis as a taxonomi.c tool . Taxon
17, 241- 58 .
1969 . Effec t s of c h ange of characters and of number of characters
in numerical taxonomy . Ameri can Hidland Naturalis t 81 : 68-86 .
1970. Analysis o f character variation in ec ology and systematics .
Annual Review of Ecolo gy and Systemat ics 1 : 55- 98 .
1972. Computerization of specimen data from the Edwa rd Lee Gr eene
Herbar ium ( ND-G) a t No tre Dame. Brit t onia 24 : 131-141 .
22 483

Cravello, To J .• and R. D. HacDonald . 1970 . Index of EDP-IR projects in

systema t ics . Taxon 19: 63 - 79 .
Davis . P . H., and V. H. He~1ood. 1963 . Princip l es of Angiosperm Taxonomy .
Oliver and Boyd . London .
Dayhoff, ~1. O. 1969 . Atlas of Prote in Sequence and Structure . Vol. 4 .
National Biomedical Research Fo undation . \~ashington, D. C.
Dupra,,,, E . J. 1965 . Non-Linn aean taxonomy and the syst ematics of honeybees .
Systema t ic Zoology 14 : 1-24 .
Eaton, T. H. 1970 . Evo l u t ion . H . hI , Norton . Ne\.,r York .
Fitch , \ol. N . • and E. MargoUash. 1967 . Construction of phylogenetic trees .
Science 15.'): 279 - 84 .
Gabriel , K. R. • and JL R. Sakal. 1 969 . A ne", s t atis tical approach to geo-
graphic variation analysis . Systematic Zoology 18 : 259 -7 8 .
Gomez - Pompa , A., and L . I . Nevling . 1973 . The u se of e l ectronic data process-
ing methods i n the Flora Vera Cruz Program . Con trib. Gra y Herb . No . 203 .
Go odall , D. W. 1968 . Id e ntification by compute r. Bi oscien ce 18 (6) : 485-88.
Goodman , H . H. 1967 . The identification of hybrid plants i n segrega t ing
pop ulat ions . Evolution 21 : 33ll-40 .
Got"er , J . C . 1966 . Some distance properties of late nt root and vector
methods used in multivaria t e analysis . Biometrika 53 : 325- 38 .
1 967 . A compar i son of some methods of cluste r analys i s . Biometrics
23(4) , 62 3- 38 .
Grieg- Smith , P . 1964 . Quantitative Plan t Ecology . Butterworths . London .
Hall, A. V. 1970 . A compute r based sys t em for formin g identification keys .
Taxon 19 : 12-18 .
1972. Computer based data banking for taxonomic collections.
Taxon 21: 1 3-26 .
Hawkst"orth, F . G. , G. F . Estabrook, and D. J . Rogers . 1968 . Application of
an information theory model for character analysis in the genu s Arceutho-
bium (Viscaceae). Taxon 17: 605-19.
Jackson , R . C. , and T. J. Cr ovel l o . 1 971. A compa rison of numerical and
biosystematic studies i n lIaplopappus . Brittonia 23 : 54-70 .
Keller , C. , and T . J . Cr ovello . 1973 . Problems and procedures in the us e of
floristic data i n a computerized data bank . Proceedings Indiana Academy
of Science 82 : ( in pres s) .
Kendeigh , S. C. 1 961. Anima l Ecol ogy. Pren tice lIall . Englewood Cliffs,
New Jersey .
Kluge , A., and J . S . Farris . 1969 . Quantita t ive phyletics and the evolut ion
of anurans . Systematic Zoology 18(1): 1-32 .
Lawrence , G. H . N. 1951. Taxonomy o f Vascular Plants . Hacmillan and Company .
New York .
Lewis , T., and L . R. Taylor. 1967 . Introduction to Experimental Ecology .
Academic Pr ess . New York .
Li.ndsey , A. A. et a1. 1969 . Natural Ar eas in Indiana and Their Preservation .
American Nid 1and Natur alis t Pr ess. Not r e Dame, Indiana . 594 pp .
HacArthur, R. H. , and J. Conne ll. 1966 . The Biology of Populat i ons . Wiley .
Ne\" York .
Hayr , E . 1963 . Animal Species and Thei r Evolution . Harvard Un i versity Press .
Cambridge .
1965 . Numerical pilenetics and taxonomic theory. Sys t ematic
Zoology 14(2) : 73-97 .
1 969 . Princip le s of Systematic Zoology . l'-lcGraw-Hil l. New York .
HcKe11 , C . H., C . Duncan , and C. H. Huller . 1969 . Competitive r elationship s
of annual ryegrass (Lolium mu1tiflorum) . Ecology 50 : 653-57 .
Horishima , H. 1969 . Phenetic simi l arity and phylogen etic r elat i onshipa among
strains of Oryza pe r ennis estimated by methods of numerical taxonomy .
Evol ution 23 : 429 43 .
484 22

Horrison, D. F . 1967 . r-Iultivariate Statistical Nethods. NcGraw-Hill-. New

York.
Norse, L. Eo, J . H. Beaman , and S . G. Shetler . 1968 . A comput.er system for
editing diagnostic keys for Flora North America . Taxon 17 : 479-483 .
Noss, \.J . \oJ. 1967. Some new analytic and graphic approaches to numerical
taxonomy with an examp le from the Dermanyssidae (Acari). Systematic
Zoology 16(3) : 177-2 07 .
Odurn, E. P. 1956. Fundamentals of Ecology. \." B. Saunders . Philadelphia .
Oxnard, C. E. 1969. Nathematics . shape and func tion: a study in primate
anatomy . American Science 56: 75-96 .
Perring, F. H. , and S . H. Walt ers . 1962 . At las of the British Flora . Thomas
Nelson . London.
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Angiosperm family: generic delimitation in the Chrysobalanaceae . New
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Biology . Publication No . 1692 . National Academy of Science , 542- 587 .
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the taxonomy problem . Journal Theoretical Biology 15(1): 103-144 .
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New York .
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Systematic Zoology 14 : 148-149 .
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J ournal of Zoology London 151: 65-122 .
__~~~.' and R. R. Sokal . 1973 . Numerical Taxonomy: The Principl es and
Practice of Classification . H. II. Freeman Company . San Francisco .
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critical evaluation. American Naturalist 104 : 127-153 .
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San Francisco.
--"7.~c.-' and P. H. A. Snea t h. 1963. Principles of Numerical Taxonomy.
H. H. Freeman Company . San Francisco .
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Processing. Prentice Hall. Englewood Cliffs , New Jer sey.
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New York .
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cal Biology . Blaisdell . New York .
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taxonomy. Advanced Botanical Research 2 : 35-68 .
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systematic biology, with an example for the Oncidunae (Orchidaceae) . Sys-
tematic Zoology 15(1) : 59-69.
23 485

Chapter 23. NUHERICAL AIDS IN TAXONOl'fYl '~

The use of quantitative methods in plant taxonomy was pioneered by Edgar

Anderson (see especially his 1949 r eview) in the 1930 ' s. The tec hn i ques us ed
at t hat time wer e simple but eff ec t ive . A few car ef ully chosen c harac ters
were usually s tudied on r elat ivel y smal l numbers of pl ants from at most several
populations . Today computers make similar sorts of compa ri sons possible for
many characters using large population sizes and studying many populations
simul tan eous ly. Since the late 1 950 ' s , Sakal , Sneath . and their co-workers
have actively publicized the use of s uch techniques in a wide variety of
organisms .

The objec tive of numer i cal aids 1n taxonomy i s to clarify and illustrate
degrees of relationship or similarity in an objective and repeatable manner.
While some have seemingly argued i n the past t ha t numerical techniques woul d
replace traditional taxonomy , the well - worn expression "garba ge in -~ garbage
out " perhaps best sununarizes many problems not only those of numerical tax-
onomy .

The problem most of ten encountered in t he use of numeri cal techniques i n

taxonomy is the fact that many techniques exi st , the techniques may result in
differen t conclusion s , and little basis is available fo r choosing the "best"
tec hnique(s). Certain methods are fai rly basic , however , and , if clearly
und erstood , can be used fo r most of the situations that th e taxonomist will
encounter . A comprehensive s urvey of a l most all available t echniq ues has
recently been published by Crovello ( 1970) .

Section A. SAHPLING

While taxonomists wish t o make conclusions about populations, they

actually wo rk with samp l es from such populations. These sampl es may be her-
barium specimen s , l ive plants growing in a garden or gree nhouse , or wi ld
plant s growing in the ir " normal " environmen t. In many case s it is possible
only t o samp l e cer tain s ubpopulations ra ther than the e n t i r e population about
which it is wished to make conclusions. Clearly if one ' s observations are to
be pe rtin-e nt to a populat ion , a sample representative o f the population must
be studied. Random sampling is gener al ly inappropriate since a small random
sample from a lar ge population wi ll not include the extremes of variability
which a r e necessary to prop e rly delimit t axa . An ext r eme example of the
inappropriateness o f random sampling i s in the collect i on (and preservation)
of varieties of cultivated plants _ Random samples of s uch plants woul d con-
sist a l most completely of t he mos t widely us ed commercial types . The botani -
cal ly and historically most int e r esting types would probably not be collected
at all , an d in a partic ular region the sample would mos t likely consi st of
e ssentially the same genotypes collected rep eated l y . Common sense and a
r easonab l e knowle dge of the range of variation possible i n the taxon under

IThis title was suggested by Hall (1969) as an alternative to either nume ri-
cal taxonomy or t aximet rics _

*Contributed by Hajor N. Goodman , North Carolina State Univer sity, Raleigh.

486 23

study are p r obably the major prerequ~s~tes for assembling samp l es of plants
representative of the populations about which inferences are to be made .

Section B. DESCRIPTI VE STATISTICS

Each character, X, in each sample can be described as having an average

value or mean, which may be calculated from class val ues and frequencies as
in Chapter 20 (Section B I ) or dir ect l y as simply the sum of the values of
character X for the N different members of the sample divided by the number
of individuals in the sample .

x
N
The measure of variation used is us ually not the range, which depends solely
upon the two extreme i ndiv i d uals, but the variance, which may be calculated
using frequencies as in Chapter 20 (Section B I) or directly from the raw
data as

N
L I (N-1) ,
i=l

or alternatively the standard deviation, the square root of the variance .

A simple bar graph of the frequen cies of occurrence o f the vario us possi-
ble values of X is often i nformative. The shape of such a graph is often
bell-shaped . In such a case the population is said to be approximately
normal:

r-
r- r-
r-
r-
;- L

o 100
X
Under such circumstances the interval (x - sx) to (x + sx) includes
about 2/3 of the sample . If the population frequencies are not symmetri-
cal about the mean (i.e. , skewed) then the range and mode , the most com-
monly occurring value, should be presented .

lfuen two characters X and Yare being studied , then the c l o seness of
the relati onship between them can be measu r ed by the corre l ation co-efficient

N
L [(Xi-x) (Yi - Y)]
i-I
p
N N 2
[L (Xex)2 ] [' (Ye Y) ]
i=l i=l
23 487

The correlation coefficient can range in value from - 1 to +1. ~fuen p is c los e
to - 1. then high values of X are associated Hith low values of Y and vic.e
~. When the correlation is close to +1 then high values of X are asso-
ciated with high values of Y; conversely low values of X and Y tend to occur
together. Hhen the correlation is close to zero , there is very little linear
relationship bet\veen X and Y. (Under most circumstances non-linear relation-
ships may be safely ignored except in comparisons involving functional rela-
tionships such as that between the diameter and surface area of a tree trunk.)

When several populations are being studied at the same time, several
types of means, variances, and correlations can be considered. First there
are the overall mean and variance for each character and the overall correla-
tion for each pair of characters. Second. each population has its own means,
variances, and correlation coefficients. These latter variances and correla-
tions can be averaged over populations to give "pooled" or "common" within
population variances and correlations . Third. there are t he variances and
correlations which can be obtained from the population means . These are
called the Among Population variances and correlations . (See any introduc-
tory statistical methods textbook. especially the section on nested analysis
of variance , for the computational details.)

EXAMPLE 23- 1

The samples below were obtained from 6 imaginary populat ions representa-
tive of real data . but simplified for arithmetic purposes .

Sample I Sample II •
X Y X Y
10 24 10 35
1 16 8 37
2 17 6 39
8 23 12 33
4 20 14 31

Sample III Sample IV

X Y :{ Y
1 13 1 49
7 17 29 31
3 17 15 40
9 13 3 35
5 15 27 45

Sample V Sample VI
X y X y
62 94 97 100
50 86 93 100
48 100 80 100
60 90 91 95
55 80 89 105

The mean of X for sample I is : (10 + 1 + 2 + 8 + 4)/5 ~ 5 = Xl

For Y for sample I: (24 + 16 + 17 + 23 + 20)/5
488 23

The variance of X for sample I is estimated as :

[(10- 5)2 + (1_5)2 + (2-5)' + (8_5)' + (4-5)']/(5-1)

(25 + 16 + 9 + 9 + 1)/4 = 60/4 15 = 52

X,
For Y in sample I:

[(24- 20)2 + (16-20)2 + (17-'0)2 + ('3-20)2 + (20 - 20)']/(5-1)

= (16 + 16 + 9 + 9 + 0)/4 ~ 50/4", 12.5 '"

The numerator of the correlation coefficient between X and Y for samp l e I is :

5
, (Xi-x)(Yi-Y) = (10- 5)(24-20) + (1- 5)(16- 20)
i=1

+ (2-5)(17-20) + (8-5)(23-20) + (4-5)(20-20) =

(5) (4) + (-4) (- 4) + (-3) (-3) + (3) (3) + (-1) (0) = 20 + 16 + 9 + 9 = 54
5
The denominator we have mostly calculated above. We need E (X i -x) 2 '" 60
5 _ 2 i= 1
from our calculations of and I. (Yi-Y) = 50 from our calculations of
i=1
Thus :
54 54 54
o~ 54 . 8 '" 0.99
/(60) (50) 13000

Section C . CHOICE OF CHARACTERS

Characters \.,rhich va r y greatly among populations in relation to their

variabi l ity wi thin populations are clearly more useful fOT systematic pur-
poses than are those which shm" lesser var i ability among populations in pro-
portion to their variance within populations . Similarly characters which are
less affected by t he environments in which the pl ants are found are usually
more useful than those ~olhich are highly subject to environmental effects.
Analyses of var i ance can be used to sel ect characters tolhich have such desir-
able properties (see Goodman and Paterniani , 1969, for an example) . Standard
computer programs are availab le which perform such anal yses and provide esti-
mates of the proportion of total variation attributable to differences among
populat i ons, differences among collection sites , and di fferences among years
or seasons of the collections . Such programs are found in the UCLA Biomed
(see Dixon, 1968), the TSAR (see Anonymous, 1970), the SAS (see Service , 1972),
and the NTS (see Rohlf et a!., 1971) software packages, at least one of which
should be availabl e at most major U. S . computing centers. In simple cases it
is often unnecessary to do more than carefully examine you r data. For example
for on ly a fe\ol taxa 'olith data collected i n only a few years or locations , t he
data (either raw data or means) might look like:
23 489

Character X
Taxon A Taxon B Taxon C Average
Year (or Season or Location) 1 5 10 15 10
" " " " " 2 10 15 20 15
" " " " " 3 7 13 16 12
" " " " " 4 8 13 18 13
Average 7.5 12.75 17 . 25 l2.5

Character Y
Taxon A Taxon B Taxon D Average
Year (or Season or Location) 1 5 10 15 10
" " " " " 2 12 10 9 10.3
" " " " " 3 4 10 15 9.7
Average 7 10 13 10

Clearly despite the fact that year to year (and/or s e ason to season and/or
location to location) sampling does affect character X, it is still a useful
character effectively separating taxa A, B. and C. However, character Y is
quite unreliable (it does not consistently separate the taxa; a comparison of
taxon A collected in year 2 compared fa taxon D in year 1 is more similar than
taxon A is to itself in years 1 and 2).

Since the amounts of time, effort, and money available for any study are
limited, the number of characters which can be studied is also limited. There
is little reason to study both members of a pair of highly correlated charac-
ters when a third character with lower correlation is available and has an ,
equivalent proportion of its variation attributable to diff erences among popu-
lations. In most studies, the number of available mutually non-(or slightly)
correlated characters with high proportions of their variability attributable
to differences among populations is large in relation to the amount of time
available for measurement. Under such circumstances a choice of Hhich char-
acters to use arises . Common sense suggests that the characters should as
much as possible depict the entire plant, not some part icular f eature.

Section D. HATHEHATICS NEEDED BY TAXONmlISTS

In addition to making possible comparisons among large numbers of indi-

viduals and populations, computers make possible the use of correlated char-
acters, whereas some Andersonian techniques were really appropriate only for
non-correlated characters. The understanding of the procedures used for
correlated characters does require some knowledge of basic undergraduate
mathematics.

The use of mathematical techniques in plant systematics has been limited

by the general lack of mathematical training received by taxonomists. I t
seems reasonable to expect that in the futur e a l most all students expecting
to pursue graduate work in any field of science will complete at least a year
of calculus and a semester of matrix algebra. At present, however, most
biologists possess littie more mathemat i cs than somewhat rusty high school
algebra. Even that is quite sufficient for most currently used procedures in
numerical taxonomy (taximetrics). This rather f ortui tous circumstance results
from the fact that the more complex mathemat i cal procedures usually arise due
to correlations among the characters being utilized . By choosing non-corre-
lated characters, the mathematics becomes much less difficult. Thus a small
amount of preliminary field work designed to de termine t"hich characters are
490 23

essentially un correlated Hith other characters can often make up fo r a gap in

one r 5 knm.,rledge of higher mathematics . Such preliminary studies can o f ten also
be used to sel ect characters wh i ch not only differ from population to popula-
tion but whi ch also yield consistent results from year to year and from reg i on
to r egion .

Section E. GROUPING POPULATIONS

Once the characters have been chosen, the populations sampled, and the
specimens measured , then data are available to use in grouping the populations .
Th i s is us ually a tHo - step process . First some measure of similarity or dis-
simila rity betl.een members of each possible pair of populat i ons studied is
chosen . The number of possible similarity measures is very large indeed (most
are fo und in Sakal and Sneath, 1963), bu t two general k i nds have been widely
used .

I . MATCHING A.L'lD CORRELATION COEFFICIENTS

Instead of the correlation coeffici ent described earlier, other measures

of agreement between characters are sometimes used . These inc l ude match i ng
coeff i cients and various types of non-parametric procedures usually resu l ting
in some form of non-parametr i c correlation coefficient (see Kendall , 1962,
for details) . These usually result in values rang ing from 0 for no agreement
to +1 fo r complete agreement (and sometimes -1 fo r perfect disagreement) . As
an example , consider two characters, say length of a leaf and Hidth of the
same leaf , ea c h divided into three classes: large, medium, and small :

Numbers of Leaves Hith Small, Medium, and Large Lengths and Hidths
Small Medium Large

Length 15 25 13
Width 17 23 13

A total of 53 l eaves Here exami ne d . Suppose , f or each l eaf, He score it as a

one if length and width fall in the same class, say small , and a if they fall
in differen t classes such as small length but medium \"idth . Then the average
score for all 53 leaves would be a matching coefficient t"hose possible range
Houl d be f r om zero (for no matches) to one (for all S3 matches) .

It i s poss i b l e to use matching and correlation coe ff icients to measure

the sim i larity bet\"een pairs of taxa as Hell as betHeen pairs of characters .
The data for the first taxon i s treated as the X variable and the data from
the second taxon i s trea t ed as the Y variable. Then the matches bet..,een X
and Yare ta llied or the correlation coefficient betHee n X and Y is calculated.
See Sokal and Sneath (1963), but i t should be noted that the characters are
usually standardized by subtracting the character mean and subsequently divid-
ing each observation by the standard deviation among population means for that
characte r before cal c u l ating the correlation or matching coefficient .

II . DISTANCE HEASURES

The s i mp l est measure of the square of the distance betHeen tHO taxa i
and j is:
23 491

where Xkj is character k of taxon j. Usually the characters are standardized

to avoid compounding various scales of measurement such as mi l limeters , meters ,
l eaf numbers . etc. The standardization most commonly used i s to divide each
character by its among taxa standard error . Other standard i zations can als o
be employed (see Goodman, 1969) .

Such a s i mp l e measure of d i stance works q uite well f or cha racters \.;rhich

are reas onably non-correlated . However, \"heo the value of one character c an
be reasonably predicted on the basis of the koO\>,ledge of the values of some
of the other charac t ers , the equation above results in distances which differ
mo r e amongst themselves than is justified on the basis of the da t a . To a vo i d
such a situa t ion , it is poss i b l e to adjust or transform the charac t ers so that
the adjusted characters are not correlated before cal culating the dis t ance s.
The resulting distances are cal l ed generalized distances (Hahalanobis, 1936) ;
the procedures involved require a knmlledge of matri x algebra and ,dll not be
discussed at this point .

EXAMPLE 23-2 .

For the six samples examined previously (Example 23 - 11 , we had

Sample I II III IV V VI Nean Vari a n ce

X 5 10 5 15 55 90 30 1220
y 20 35 15 40 90 100 50 1310

To standardize these scores, we can subtract 30 from each value of X and then
di vide each va l ue by 11220 " 34 .9 . For Y we subtract 50 and divide by {13 l 0
" 36 . 2 . Thus our standardi7.ed scores are:
Sample I 1I III IV V VI Hean Variance

Yo s -0 . 72 -0 . 57 -0 . 72 -0.43 0 . 72 1. 72 0 1

Ys -0 . 83 -0.41 -0 . 97 -0.28 1.10 1. 38 0 1

(Act ually due to small sounding errors the means and variances of the stan-
dardized scores do differ triv i ally from 0 and 1, respectively . ) Since the
co r relation hetHeen the sample means of X and Y is large (about 0 . 96) , a
dec i sion should be made as to ,,,hether to use just X, j ust Y, both X and Y,
or even neither X nor Y. In pract ic e we should probably choose eithe r X o r
Y or a new " character " Z wh i ch combines X and Y i n an " optimum" manner to be
discussed l ater . Anyone of the three methods should give about the same
results . For now , we wil l i gnore this problem and cal culate the di st a nces
from hoth characters :

D~ " , - [-. 72 - (- 0 . 571]2 + [-0.83 - (-0 . 411]2 (0 . 15)2 + (0 . 42 ) 2

0 . 0225 + 0 . 1764 Z 0.20

Thus Dr , II :::: /0 . 20 ::: 0.45 .

492 23

The complete distance tab le is :

I II III IV V VI

I 0 . 45 0 . 14 0.62 2.41 3.29

II 0.45 0.58 0.19 1. 99 2 . 91
III 0.14 0.58 0 . 75 2 .52 3 . 39
IV 0 . 62 0 .19 0 . 75 1. 38 2.72
V 2.41 1.99 2.52 1. 38 1. 34
VI 3.29 2.91 3.39 2.72 1.34

Clearly samples I, II , I II, and IV differ little, while V and VI differ from
each other as well as differ greatly from the other samples. The same con-
clusions are evident from either X or Y alone , but the technique can be used
for many characters at once, ,,,here simple inspection of cite data may not be
suffi ci ent for reaching conclusions.

III . CLUSTERING PROCEDURES

Once measures of between taxa similarity or distance have been obta i ned .
several techniques can be used to group the taxa . The most widely used one
first groups those taxa \"hich are mu tually most simi l ar or closest together
to form groups, then proceeds to group the groups together on the same basis,
treating remaining individual taxa as groups . This results in a tree- like
diagram called a dendrogram, where the taxa are located at the branch tip s and
where a pair of branches join together at the average distance (or average
similarity) between all possible pairs of taxa located on the two branches ,
the first member of a pair being from one branch , the second member from the
other . I n Example 23-2, the first step would be to group samples I and III ,
which have the small est dis t ance , 0 .14 . Second , samp l es II and IV would be
grouped with their distance of 0.19. Then samp les V and VI are grouped a t a
distance of 1.34, since they are closer to each other than to any other sam-
ple . The average distance from {I ,II I } to {II , IV} is (0 . 45 + 0 .62 + 0.58 +
0 . 75)14 = 0 . 60, which is much less than the distance of either to {V , VI}, so
{I , III} and {II, IV} are grouped at 0.60. Finally the average distance from
{I , II , III , IV} to {V , VI} is (2.41 + 1.99 + 2.52 + 1.38 + 3 . 29 + 2 . 91 +
3.39 + 2. 72)/8 m 2.58 which completes the dendrogram .

EXAMPLE 23- 3 .

Example of a Dendrogram
I

III

II

IV

VI f--
I I I I
0 1 2 3
Distance
23 493

A quick inspection of the data indicates that either the use of X alone or Y
alone would have resulted in a dendrogram of essentially the same form .

I nstead of average dis tances, smallest or largest distances between the

members of on e group and those of another can be used for clustering. HO\</'-
ever, the agreement between the original table of distances and t he distances
read f r om the dendrogr am obtained is much less than is obtained using average
distances . In Example 23-3 , using smallest distances between members of dif-
fere nt groups wo u ld change the grouping of {T . III} with {II , IV} to 0.45
rather than 0 . 60 , and of {I . II, III , IV} with {v , vI} to 1. 38 rather than
2.58, but would leave the form of the dendrogram unchanged. Using largest
distances , Hould result in a dendrogram of the same form also , but ,.;rith {T , III}
grouping ,<lith {II , IV} at 0 . 75 and Hith {t. II. III, IV} grouping with {v , VI}
at 3.39 .

Rather than averaging distances , the means themselves can be averaged

afte r each clustering and distances recomputed. The latter techniq ue is
ca lled centroid clustering . Results generally differ trivially from distance
averaging .

Several theoretically appealing alternatives to these procedures have

been proposed. One is to form k groups in such a manner that the variation
among the within group distances is as small as possible in relation to the
among group variation. A second is to find the dendrogram for which the den-
drogram distances fit closest t o the actual distances among the taxa . Several
criteri a of closeness of fit could be used , but the usual one is l east squares
,
Nin E (D -cl .) 2
i,j ij i J
which is the minimum sum of the squares of differences between the distances,
Dij ' and the dendrogram distances, dU' Neithe r procedure has proved to be
practical i n the case of more than a lew taxa. Gower (1967), Wishart (1969),
and Cormack (1971) have recently reviewed a number of alternative clustering
procedure s .

Section F . PRINCIPAL COMPONENTS

\-lhen it i s necessary to Hork t-lith sets of highly correlated characte rs,

the simple application of distance equations is often unsatisfactory . Cal-
culating distances from severa l highly correlated characters is essentially
equivalent to calculating them from a single one of the correlated charac-
ters and then multiplying the distances obtained by an unknmffi constant .
lfuen several sets of highly correlat ed characte rs plus a few rel atively un-
cor related ones must be dealt with, then the distance equations are clearly
inappropriate.
To reme dy this situation . the characters can be transformed to a new
se t of uncorr elated charac ters . Principal components are simp l y the neH
charac ters obtained by . transforming the original characters to an uncorre-
lated set . If X and Yare tHO corr elated characte r s .
494 23

,,
H
/
z

y
@
/ '
/ ',
x
then the 111- 2 axes define the principal components. An individual plant or a
taxon mean has values of Hand Z that correspond to its values of X and Y.
Theoretically these values could be read from a graph or calculated arithmeti-
cally , but in practice computer programs provide them directly. t.J and Z are
uncorrelated (know ing the value of H for an individual tells us nothing about
that individual ' s value of Z and vice versa). Usually iol' and Z are also stan-
dardized , by dividing each by its standard error, so that each has about the
same range:
H
I
I
I
I
I
- - -- - t--- -- z z*

where w* and Z* are the standardized forms of 111 and Z.

The computations involved require a knowledge of matrix algebra, but an

understanding and use of the technique does not. Computer programs such as
the SAS , UCLA Biomed, and NTS software packages can first calculat e the cor-
r elations among the characters used and then make the transformation of the
data.

The first principal component (here Z) has the greatest variance and the
last one (her e \.J ) has the least. Thus , sometimes the fi rst several principal
componen ts can be used to summarize most of the variation among the taxa,
rather than the fu ll set of characters initially studied . Since the sum of
the variances of the principal components is the same as the number of charac-
ters studied, if the original character s are " standardized" to mean zero and
variance one (by subtracting the mean from each and then dividing by the stan-
dard deviation) , t he proportion of this sum attributable to each of the prin-
cipal components is easily calculated. If the first two components account
for most o f the among taxa variability, then a plot of the taxon means, pos-
sibly wit h their within taxon standa r d errors, for the first two p r incipal
components should illustrate quite well the inter-taxa relationships. See
Rao (1964) and Goodman (1972) for further discussion . In any case, distances
between taxa can be computed using the principal components which account for
most of the variation among taxon means . Often the last few principal com-
ponents are little more than "noise ," if the characters are highly correlated .
23 495

Section G. CANONICAL VARIATE ANALYSIS

When t he problem at hand is to assign individuals to t heir proper group

or when the gro ups i n a dendrogram cluster t ogether so closely that lit tle
reliance can be placed upon the partic ular arrangement obtained (see Rohl f ,
1968) , canonical variate anal ysis can often be profitably used. It is closely
related to principal components anal ysis . The first canonical variate. YI. is
a linear transformation of the original characte rs

where Xi is character i. The hi are determined in such a manner that YI has

the greatest possible variat ion among taxa in proportion to its variation
within taxa. The second canonical variate has the greatest possible varia-
tion among taxa in proportion to t he within taxa variation of all possible
variates constructed by

where the correlation among and within taxa of Yl and Y is zero . In a simi-
lar manner , Y3 is formed so as to have zero among and wIthin taxa correlation
with both YI and Y2 and to maximize the among/within taxon variation on that
basis. By using the first two canonical variates. the taxon means and within
taxon standard deviations can be plo tted and unidentified individuals assigned
to thei r nearest neighbor. Simi l arly by using the first few canonical var i-
ates as input da t a for the construction of a dendrogr am, the relationships
among gr oups will be emphas ized over t he relationships among individual pairs ,
of taxa .

In the case of only two group s , the first canonical variate is identical
to the discriminant function of Fisher (1936), first used for a case of
hybridization in Iris discovered by Edgar Anderson . These techniques (dis-
criminant functio~canonical analysis) a r e s till widely used in studies of
hybridization among plant, especially forest tree, species. (A recent example
is Smouse. 1972).

Section H. HISCELLANEOUS

There are a number of specialized t opics which will only be men tioned
here. There has been much discussion about the use of transformations of
the characters (see Wright , 1952) and the use of ratios, rather than the
original data (see Sakal , 1965) . Tests of significance are discussed i n all
statistical method s books. but most taxonomic comparisons are among popula-
tions known to be different . In that case , the l evel of significance of a
difference will probably depend mostly on the number of specimens measured .
Significance i s not synonymous ~.,.ith importance. The test for differences
among within population covariance matrices is given in Horrison (1967) .
Such mat r ices do us ually differ significantly, but t his may not be important
for multivariate studies, which are based on samples of approximatel y e qual
size . Regression and covariance analyses can be used to adjust the data to
eliminate (or at least ameliorate) the effects of variation in characte ris-
tics for \.,.hich little or no variation would be preferabl e . These might be
sex or age differences , location or altitudinal differences, size or weight
differences, etc. Standard statistical methods texts give details and exam-
ples.
496 23

Gower (1966. 1972a, h) has shown hOH various analytica l techniques for
measuring similarity may be related and has developed a simple technique for
measurin g the degree of agreement between sets of analyses based upon either
di ffer e nt characters or differ ent measures of relationship or both . Recent
examples of taximetric applications of general interest outside the area of
plant taxonomy are those of Horne (1967). Fitch and Nargoliash (1967, 1970).
and the symposium edited by \.Jeiner and Huizinga (1972) , while Far ris (1970,
1972) has formalized a technique suggested by Wagner and ....idely used by
botanis ts for deriving tentative phy logenies .

Examples of possible p roblems encountered in the application of numeri-

cal techniques to botanical data can be found in Heiser et al. ' s (1965) analy-
sis of Solanum species and hybrids. An informed critique of the philosophy
behind much of the early (and indeed of some of the current!) work in numeri-
cal taxonomy has been Nri tte n by Johnson (1970). He also has contrasted the
mathematical simplicity being applied to the biological complexity under
study, and he has done so in very specific terms . It is a paper well worth
reading. A sho rt review of the whol e field of numerical taxonomy has recent l y
appeared (Gower, 1969) , and edition two of Sokal and Snea th' s (1963) basic
text has just appeared (Sneat h and So kal , 1973). Another comprehensive
review (Blac kith and Reyment, 1971) is a book of applied examples (along with
extensive cO!!lputer programs), but t he amount of explanation is often minimal.
It has an exce llent current reference list and can serve as an introduction
to the vast literature which has accumulated in recent years.

Section I . ARITH}illTIC

To speed up calculations you may wish to code your data by subtraction ,

addition, multiplication, or division. (For example you might prefer to work
with I, 5 , 3, 8, .... rather than IDOl, 1005. 1003. 1008, .. .. etc . ) . After
coding your data in that manner and calculating your means and Variances, the
fol lowin g rules apply to convert t hem to their original units of measurement.

Neans : If you added a constant K, subtract it.

If you subtracted a constant K, add it.
If you multiplied by a constant K, divide by it .
If you divided by a cons tant K, multiply by it .

Variances: If you added or subtracted a constant K, fo rget it .

If you multiplied by a constant K, divide by K2
If you divided by a constant K, multiply by K2.

Finally the following equation is exceedi ngly helpful in the calculation

of variances:
N N
l: (X - x)2 = L
i X2 -
i"'l i=l i

and a corresponding equation is very helpful i n the calcul ation of correlation

coefficient s :
N N N N
E [(X.-X)(y.-y)]
1 1
'" L X.Y. - ( E X.)( E Y.)/N
i=1 1=1 l. l. i~l l. i= l 1
 23 497

NilllERIGAL AIDS LITERATURE

I. General Texts or Reference Books

Anderson , E. 1949 . Introgressive Hybridization . John lJiley and Sons , I nc .

llew York .
Anonymous. 1970. TeleStorage and Retrieval System User's l'lanua l. Duke Uni-
versity Computation Center . Durham, Horth Carolina .
Blackith , R. E., and R. A. Reyment . 1971. Multivariate Horphometrics . Aca-
demic Press . London and NeH York .
Dixon, H. J . (ed.) . 1968 . BHD Biomedical Computer Programs , 2nd ed . Univer-
sity of California Press . Berkeley and Los Angeles.
Kendall, H. G. 1962 . Rank Correlation Nethods . 3rd ed . Charles Griffin and
Company, Ltd . London .
Morrison, D. F . 1967. Hultivariate Statistical Hethods. HcGraw- Hill Book
Company . Nel" York .
Ostle, B. 1963. Statistics in Research . 2nd ed. Im~a State University Press .
Ames, ImV'a .
Rohlf, F. J ., J . Kishpaugh, and D. Ki rk. 1971 . NT-SYS . Numerical Taxonomy
System of Hultivariat e Statistical Programs. State University of New York .
Stony Brook, Ne\V' York .
Service, J . 1972 . A User's Guide to the Statistical Analysis System . Stu-
dent Supply Sto r e s. North Carolina State University . Raleigh . North
Carolina .
Sneath, P . H. A. , and R. R. Sokol . 1973 . Numerical Taxonomy: The Princ iples
and Practice of Numerical Classification. H . H . Freeman. San Francisco .
Snedecor, G. H., and H. G. Cochran. 1967 . Statistical Methods . 6th ed . Io\.<.'8 ,
State University Press . Ames , IO\~a .
Sokol, R. R . • and P. H. A. Sneath . 1963 . Principles of Numerical Taxonomy .
H . H. Freeman. San Francisco .
Heiner, J. S., and J. Huizinga (eds.). 1972. The Assessment of Population
Affinit i es in Nan . Oxford University Press . London.

II. Pe riodicals

Applications of numerical techniques can be found in most botanical, as

well as agricultural and forestry , journals . Periodicals such as The American
Naturalist , Genetics. and Evolution also frequent ly carry articles dealing
with numerical taxonomy . The follmV'ing are more specifically aimed at the
application of numerical techniques to biology:

Biometrics . 1945+ . F . A. Graybill , Editor . Department of Statistics, Color-

ado State University, Fort Collins, Colorado. Publication of the Biometric
Society, \11ash ington, D. C.
The Classi f ication Society Bulletin. 1970+ . P . H. A. Sneath , Editor . H. R. C.
Hicrobial Systematics Unit, University of Leicester , Leicester LEI 7RH,
England. Pub lication of the Classification Society .
Systematic Zoology . 1952+ . A. J . Rm.,rell , Editor . Nuseum of Invertibrate
Paleonto logy , Departm~nt of Geology , University of Kansas, Lawrence , Kansas.
Publication of the Society of Systematic Zoology , Nel. Haven, Connecticut .

III . References

Cormack , R . N. 1971. A revieH of classif ication . J . Royal Statistical

Society A 134 : 321-367 .
498 23

Cravello, T . J . 1970. Analysis of character variation in ecology and system-

atics . Annual RevieH of Ecology and Systematics 1: 55-98 .
Farris, J. S . 1970 . l'!ethods for computing l~agner trees . Systematic ZooloBY
19 : 83-92 .
1972 . Estimating phylogenetic trees from distance matrices . The
American Naturalist 106: 645-668 .
Fisher, R . A. 1936. The use of multiple measurements in taxonomic problems .
Annals o f Eugenics 7 : 1 79-188 .
Fitch, H. M.• and E . Margoliash. 1967 . Construction of phylogenetic trees .
Science 155: 279-284 .
1970. The usefulness of amino acid and nucleotide sequences in
evolut ionary studies . Evolutionary Biology 4 : 67-109 .
Goodman, N, H. 1969 . Heasuring evolutionary divergence . Japanese Journal of
Genetics 44 (Suppl . 1) : 310 - 316 .
1972. Distance analysis i n b i ology . Systematic Zoology 21 : 174-
186.
and E . Paterniani. 1969 . The races of maize : III . Choices o f
appropriate characters for racial classification . Economic Botany 23 : 265-
273 .
Gm.Jer, J. C. 1966. Some distance properties of latent root and vector
methods used in multivariate analysis. Biometrika 53 : 325 - 338 .
1967 . A comparison of some l'lethods of cluster analys is. Biometrics
23 : 623-637.
1969 . A survey of numerical methods useful in taxonomy . Acarologia
11 , 357-375 .
____ 19 71. Statistical methods of comparing different mul tivar iate
analyses of the same data . In : Hodson, F . R., D. G. Kendall , and P . Tautu
(eds . ) . Mathematics in the Archaeological and Historical Sc i ences . Uni-
versi ty Press . Edinburgh .
1972 . Neasures of taxonomic distance and their analysis . In : The
Assessment of Population Affinities in Nan . Oxford University Press .
Oxford .
Hall , A. V. 1969 . Group form i ng and discrimination l.Jith homogeneity functions .
In : Cole, A. J . ( ed .). i'h lmerical Taxonomy . Academic Press , Inc. Ltd .
London .
Heiser, C. B., J . Soria, and D. L . Burton . 1965 . A numerical taxonomic study
of Solanum species and hybrids . American Naturalist 99 : 471-487 .
Horne , S . L . 1967 . Comparisons of primate catalase tryptic peptides and
impl ications for the study of molecular evolution. Evolution 21 : 771-786 .
Johnson, L . A. S . 1970 . RainbohT's end : The quest for an optimal taxonomy .
Sys tematic Zoology 19 : 203- 239 .
Nahalanobis, P . C. 1936 . On the generalized distance in statistics . Pro-
ceedings National Institute of Science, India 2 : 49--55 .
Rao , C. R . 1904 . The use and interpretation of principal component analysis
in applied research. Sankya Series A 26 : 329-358 .
Rohl f, F . J . 1968. Stereograms in numerical taxonomy . Systematic Zoology
17 : 246-255 .
Smouse, P. E . 1972. The canonical analysis of multiple species hybridiza-
tion. Biometrics 28 : 361-371.
Sokal , R. R. 1965. Statistical methods in systemat ics . Biological Review
40 : 337-391.
lJishart , D. 1969 . Node analysis : A generalization of nearest neighbor which
reduces chain ing effects . In : Cole , /I.. J . (ed.) . Numerical Taxonomy .
Academic Press , Inc. Ltd . London .
23 499

Ihight, S . 1952 . The genetics of quant itative variab ility . In : E. C. R.

Reeve a nd C. H. \\Taddington {eds . } . Quantitative Inheritance-:- Pp . 5-41 .
Her ~~jesty ' s St ationer y Office . London .

EXERCISES

23.B . 1. The table below derived from the data presented i n Example 23-1 con-
tains four incorrect means . Fi nd them and corr ect t hem .

X
-
Y 2
5X 52
Y P
Sample I 5 20 15 12 . 5 0 . 99
Sampl e II 15 40 10 -1. 0
Sample III 5 15 10 4 0
Sample IV 10 35 170 53
Sample V 55 90
Sample VI 90 100 12.5 -0 . 11

Z3 . B. Z. Complete the table above .

23 . B. 3. The overall mean in this expe r iment for character X 1s

(10 + 1 + 2 + 8 + 4 + 10 + 8 + 6 + ... + 80 + 91 + 89)/30 ~ 30 .
Wha t is the overall mean for Y?

23 . B. 4 . The overall variance for X is

[(10-30 )2 + (1-30) 2 + ... + (89-30)21/29 = 1090 . 6
\.fhat i s the overall variance for Y? \~hat is the corresponding over-
a ll correla tion be tween X and Y?

23 . B. 5 . The average within sample varian ce fo r X is

(15 + 10 + 10 + 17 0 + ... )/6 c 47.
Hha t i s t he average within samp l e variance for Y?

23 . B. 6 . The variance of the sample means for character X is

[( 5-30) 2 + (15-30)2 + (5-30 )2 + (10- 30) 2 + (55 -30)2 + (90-30)2] /(6-1)
• (625 + 225 + 62 5 + 400 + 625 + 3600 )/5 • 6100/5 • 1220 .
Hot.,! does t h is compare to the average within sample var ian ce for X?
Hake the same comparison fo r Y. Compare these values with the
values of the overall variance of X and Y.

23 . B. 7 . Compute the correlat ion among t he sampie means of X a nd Y. The

numerator i s calculated as [(5-30)( 20- 50) + (15- 30)(40- 50) +
(5 - 30)(15-50) + (10-30)( 35- 50) + (55 - 30)(90- 50 ) + (90- 30)(100-50) 1.
Compare t his t o the overall correlation between X and Y.

23 . B. 8 . Not e that you wer e not asked to ca l culat e t he average correlation

wi thin samples. For-technical r eas ons beyond the scop e of this t ext
the best est i mate of th e ave rage within samp le corr e lation i s not
(0 . 99 + (-1.0) +
0 + . .. + ( - 0.11»)/6 . Instead we a ve rage the six
5
values of t (X -x)(Y 1
i
-y)
and call t h at txy , ave r a ge t h e values
5 1::=1
of t (X . - x) 2 and call that t x 2 , and average the values of
1=1 1.
500 23

5
r (Y Cy) 2 and c a ll that [y2 . Then the average Hithin samp le
1=1

correlatlon coefficien t between X and Y is

Calculat e the latter value.

23 . E.1. From th e dendrogram in Example 23-3 read off t he distances het\-!een

all possible pairs of samples (for example , t he dendrog ram distance
hetween I and VI is 2 . 58), Hake a tahle of these dendrogram distances .

23.E.2. Hake tables for the de nd rog ram distances hased on smallest distances
and largest distances as ,...e ll .

23 . r. . 3 . Camuare these three tables with the original distance t"h I e on whIch
the dendrograms are hased . Hhich table compares mo st closel y?

(These should he requi r ed of al l those ",ha have not previouslY used com-
puters.)
23 . f . 1. Using the table of sBr.:Jple means of characters X and Y Hhich we have
calculated for samples I, II , ... , VI of Exercise 23 . B.l ., punch the
means of sample I on an Iflll car d . Place the decimal point for charac-
te r X in col umn 5 and the decimal point for character Y in column 10 .
(If you have not used a keypunch before, you l.-i ll finq it \wrks very
much like a t yp ewriter. First experiment wi th the " Feed ," "Register , "
"Relea se , " and "Duplicate" keys. You will q u ickl y get the kn ack of
simp l e keypunchin g) . Pla ce the means fo r samples II through VI on
separate cards , being su re to place the decimal points in the proper
columns .
23. F. 2. Locate a program for computing principal compon ents (SAS is suggested) ,
punch the necessary control cards , and run the program .
23 . F . 3. Ca ll t he principal components Wand Z, r espec t ively . Calculate the
me ans of l-l and Z and t hei r var i ances. IJhich is more variabl e? I f
W has no t been standardized , standardize it.
23 . F . 4 . Fr om the standardized values of H, calculate the distances between
the sample means I, II , III , IV , V, and VI. Compare these di stances
to those obtained earlier in Ex ample 23-2.
23.G . I. Punch , as i n Exercise 23.F.I. , values of X and Y for each memher of
samples I. II , . ..• VI of Examp l e 23-1. The val ues of X and Y for
the fir s t individual go on the first card, etc ., as befor e . Be cer-
tain the decimal poi::o.ts are in the co l umns specified in the previous
exercise.
23 . G. 2 . Using the UCLA Bio med program titled "Stepwise Hultiple Dis c riminant
Function Analys i s " or some other program which does canonica l variate
analysis (not the same as canonical correlation analysis) , punch the
necessar y cont r ol ca r ds and run the program . Be sure to get your
output plotted if pos sible . Experiment \.,fith t he vario us options your
progr am permits . How do the plotted distances compare with those
p r eviously ca l culated? (You may have to pl ot the means yourself i f
you u se a program other than t h e one suggested ) .
23 . r. . 3. St udy the entir e outp u t of your program . Note the points which you
cannot understand. Consult a statistician or numerical taxonomist
to c l arify those poin ts.
24 501

Chapter 24 . DESCRIPTION AND DESCRI PT IVE FOR}~T

Plant description shou ld convey an image or imp r ession of plant charac-

ters , attributes and nature . Descrip t ion of plants and their parts and t axa
provide the basic information for all of taxonomy, i . e., characterization ,
identification and classif ication, which form the core of floras , manuals ,
revisions , monographs , and systems of classification . Descriptive terminology
consti tutes the frametwrk fo r determination of relationships between taxa .

No absolutely comprehensive description has ever been written for any

species or higher taxon . In data banking, systems are being devised fo r input
and retrieval of all descriptive information as it is produced by various
spec ialists. Honographic treatments are fairly comprehensive for se le c ted
types of evidence such as morphological, cyto logical, ecological but definitely
not for al l fi elds of evidence (see Chapters 6-19). Nanuals usually con tain
descriptive terminology relevant to diagnosis and accurate identification , with
little else. Simple floras have a more limited descriptive content , frequently
not much more than geographic and ecological data for each species listed .

The major problem in descriptive \~riting is making the description rele-

vant to the type of publication contemplated or in the selection of pert inent
characters with the app r opriate character states for each taxon described .
Another problem is the following of a descriptive sequence I established by
the author or editor, for consistent and comparative treatment. All taxa of
coordinate rank sho uld have the same characters described in the same sequence. ,
Sti ll another problem will be the bridging of the terminology gap due to an
inadequate vocabulary or inadequacy of definition of the terms in the glossary .
t-lany other problems will have to be solved or resolved as one describes parts ,
plants, and taxa accurately and relevantly.

A general descriptive format is included to encourage consistent key and

description writing and for the orderly use of terminology as well as to help
the student understand the descriptive sequences in previously published flor as
and manuals. This example of format is based on that now being used i n t he
writing of t he "Flora of the Southeastern United States" (Radford et a1. , 1967)
but it i s adaptable, with necessary changes in geography and ecology , for use
in writing floras or descrip tions of species for any region (Sections A-F).

This format introduces the student to some of the problems in forma t

design and to some of the difficulties involved in the use of any consis tent
sequence of descriptors . For the understanding of the descriptive language
used in any flora , manual . revision , or monograph , the basic phytograph ic
glossa r y or vocabul ary should be part of the descriptive format or , at a
minimum , the basic s ource of terms should be indicated.

For a flora or manua l covering a large area in which two or more authors
are participating , a Guid ebook with an explanat i on of taxonomic philosophy ,
concepts , and policy as we ll as plans, procedures, form and style of treat-
ment (format) is an absolute requirement. Hany of the major and minor prob-
lems related to descriptions of taxa can be eliminated with appropriate guide-
lines . The most comprehensive set of instructions ever published for a taxo-
nomic project are those in "A Guide for Contributors to Flora North America
502 24

(FNA)" by Shetler et al . (1973) . The taxonomic philosophy . concepts and poli-

cies for Flora North America are present ed in Sect ion G to show the philosophi-
cal framework and organizational detail necessary for the making of descriptive
treatments , according to a Format, of the taxa within a large flora or manual .
DESCRIPTIVE FORI-tAT

Keys and DescriPtions

1. Family number and name (a, b. f). No authority. Common name (c).
2. Family description Cd).
3. Key to Genera (e) .
4. Generic number and name (a , f , i) . Authority. Common name (c).
5. Generic description (d, f) .
6. Key to Species (e ) .
7. Species number and name (1). Authority. Common name (e) .
8. Species description Cd).
9. Infraspecific taxon number and name (j). Authority . Common name (c).
10. Infraspecific taxon description (g , h , j) .
(a) I f a family or genus has a single species with i n our area , the fami ly
and /or generic desc riptions are omitted . The spec i es description is modified
to include the family and/or generic characteristics fo r our material.
(b) Family numbe rs and name s are according to Sec tion F (included in part
only) .
(c) The common name , if available , should be the most commonly used one
(rarely more) for the taxon within our area.
(d) All taxon descrip t ions s hould follow the de tailed arrangement in
Sect ion A, examples in Sec t ion 8, abbreviations in Section C, geography-ecol-
ogy in Section D, and synonymy in Sect ion E.
(e) Keys to all taxa are to be strictly dicho tomous , ind ented , and each
couplet successively numbered (se e Sect ion 8 for examples and Chapter 25 for
key writing) . Each line or heading of a couplet should start with the same
noun or morphological character. All characters used should have the con-
trasting condition in the second couple t entry . Nouns usually come before
adjectives. No more than three characters should be used within a coupl et to
distinguish any two taxa or groupings . The character easiest to determine
should be used first in the couplet . Vegetative , flower and fr ui t characters
should be used whenever possible. Quantitative measurements or ratios should
be used, not qualitative statements. Positive statements should be used when-
ever poss ible, negative only when the positive con trasting characters occupy
too much space. If combined flowe r ing and fruiting or seasonal and sexual
keys become too cumbersome, write keys for flowering material and fruiting
material , for wint er condition and summer condition, fo r male and female
plants and so forth.
(f) All marking, punct uat ion and manuscript spacing should be according
to the samples in Sec tion B.
(g) Briefly no te only the distinctive features of the infraspecific taxon,
the n fol low with habitat, distribution, synonymy , e tc .
(h) Keys should not be written for infraspecific taxa unl ess th e infra -
specific taxon is the only representative in our area .
(i) Genera should be succes sively numbered in each family beginning with
one; species should be s uccessively numbered in each genus beginning with one;
each infraspecific taxon should have the specie s number and each suc cessive
one indicated, a, b. c , (4a , 4b, 4c , etc.) .
(j) Infraspecific taxa should not be i ncluded unless well - defined or
easily delimited. If the a uthor desires, reference can be made to previou s l y
described infraspecific taxa in the remarks section of each species .
24 503

Section A. FORNAT

I. GENERAL COHNENTS

In writing family , gener ic , species a nd infraspecific description s, the

order as given in " B" of Fo r mat Details s hould be followed fo r the characters
cons i dered . In groups where it is not practical t o f ollow this order essen-
tially the author should submit his format to the chairman of the editor i al
board for approval before final writing of the manuscript . Descriptive ter-
minology should conform to that of Lawrenc e , G. H. H., 1951 , " Taxonomy of
Vascu lar Pl ants . " The Hacmilla n Company , New York, or prefe r ab ly that i n
Chapter 6 .

Unless otherwise s tated in descriptions it is assumed tha t the p l ant is

hermaphroditic and a u totrophic ; the woody plan t i s per ennial; stems are erect ;
roots are fibrous; woody plant l eaves are deciduous; leaves are Simple, green,
pe tiolate ; flowers are per fect, regular, pedicellate ; f l oral parts are sepa-
rate for sepals, petals, stamens and fused for carpels; the ovary is super ior
and the other floral parts are inserted hypogynously . Authors should be con-
scious of these de sc riptive assumption s so that exceptions are alway s indi-
cated in t h e des c riptions .

Key characters should be i ncluded in descript i ons and amplified where

necessary . Family characters common t o all genera within the f am ily should be
included in the famil y descript i on and not in each generic description; generic
c haracters common to all species sho uld be i n c luded in t he generic description
and not i n each s pecies descr i ption . In families with t wo or more gene r a a nd
gen era wi th two or more species, struct ures desc r i bed shou l d be compared for ,
the c l osely r elated taxa within t he tribe, section or natura l g ro up to e l imi-
nate uncertainty, e . g., if ge nus "a" or species "a" have opposite leaves and
cordate stipules , then genera " b ," " c," "d" or species " b, " " c ," li d " should
have lea f arrangeme nt and s t ipule shape de scribed .

Each page of manuscript must be numb ered consecutively within the f amily
and genus treatment . Pages for the family description and the keys to the
genera s houl d have two parts , t he f a mil y number and the page numb er separated
by a hyphen, e .g., 1 40- 1 , 140-2 , 140-3. Page numbers for t he treatment of
each gen us and its s pecies wi ll have three parts; the fa mily n umber , the genus
number , and t he page number aga i n all sepa r ated by hyphens . The ~ number
starts with 1 again fo r each genus , e . g . , 140-1-1 , 140- 2-1, 140-2-2, 140-2-3 ,
140- 3- 1 , etc . Family numbers can be obtained from Section F and the generic
numbe r s within a family are assigned by each author for his aIm contribution .
Each fami l y and each gen u s shoul d start at the top of a new pa ge . All pa ge
numbers should be in the upper , right-hand co rner .

J Note the following : Fractions should be written "1/2," "1/4 , " e tc ., not
2 , 4, etc . , except after who l e numb e r s where the decima l system s houl d be use d:
"l.SX" or 1. 2SX, " not 1 l /2X or 1 l/4X . "Times" is to be indicated by cap it al
X; "no . 13" should b e u sed rat her than " U13." When u sing "mor e " or "l ess" in
meas urements follow measurement with appropriate phrase , e . g . , 7- 9 c m or more
long , 3-6 mm or less wide . For width of flat objects use the term "wide, " e . g . ,
l eaves 2-3 nun wide , not 2-3 nun broad; on "3-dimensional" width measurements use
"broad" or " in diam ., " e . g . , nutle t 2 mm broad ; pome 2 cm i n diam . ; for height
use the ter m " ta l l ": 2-3 dm ta ll, not high ; fo r prostrate or spread i ng s t r uc-
tures use " long " for length . In hyph e nation use "3-nerved" and "2-3 nerve d ,"
not 3 ner ved or 2- 3-nerved. Where the number of parts described per plant is
more than one use the plural, i f only one , use singular , e . g ., fo r the l i l y,
" leaves" rather tha n "leaf " and "bulb" rath er than "bulbs " would be desc r ibed.
504 24

II . FORMAT DETAILS

A. Manusc r ipt Or der

Note Punctuation: period (.) always at end of entry descr i ptions ma rked with *
1. General modif iers • ( l arge , small , 10. Fruit s ; or •
coarse, evergreen , parasitic , epi- a. type,
phytic, in sec tivorous, etc.) b . receptacle (hypanthium)
c . pedicel; or
2. Sex of plan t . (monoecious , etc.)
*11. Seeds.
3. Plant duration • (annual , perennial) a. type
b. seed coat • embryo
4. Habit; or . (tree, shrub. vine, endosperm ;
herb) c. placentation .

5. Stem ; or . (nodes, internodes) 12 . (n=) No punctuat i on [o r

(3n= ) or (Sn= ) in
*6 . Roots . the case of triploids
and pentaploids J
*7. Leaves .
a. duration *13 . Flowering data
b. structure (simple , compound)
J

c. arrangement J position ; 14 . Habitat data ; (see Sec-

d. blade; s heath ; ligule ; tion D)
e. petiole;
f . st i pules . *15. Distribution (see Sec-
tion D)
*8 . In florescence . a. province or .
a . type ; or no punctuation b . state .
b. posi tion ; (terminal, a xillary)
c. peduncle, and/or rachis 16 . [Distribution in adjacent
d . bracts ; or . bractlets ; or . sta t es] No period unless
e . pedicel bracket last item in des-
cription.
*9 . Flowers.
a. sex; (perfect, imperf ec t) *17 . Synonymy . (see Section E)
b. symmetry;
c. calyx, sepals ; *18 . Remarks.
d. co rolla , petals
e. stamens , anthe rs, filaments ; *19. Hajor reference (s) .
f. carpels, stigma, style . ovary (Literature cita tion as in
(position first) , ovule ; American Journal of Botany)
g . receptacle; (hypanthium)
h. ped i cel

B. Desc riptive Sequence for Each St ructure Treated

(App l ies most often to leaf blade, calyx and corolla)
l. number of parts 5 . shape or form •8. venation
2. fusion of parts (unequal al so ) 9. vestiture
3. habit (erect , spreading, etc . ) 6. size, (length, width) 10. apex ,
4. col or • odor • 7 . texture and /or surface11. margin •
12 . base; or
General: male before female; chasmogamy before cleistogamy ; combined struc-
ture before single; tube before lobes ; uppe r surface (use " above") . before
lower surface (use "ben e ath" or "be low" ) .
24 505

Section B. SPACING , MARKING , PUNCTUATION, FORMAT EXA}WLES

97-1-1

Period at No punctuation
end of name if authority is
entry only written in full
if authority
is abbreviated ~
97 . brZABALACEAE 1. lines between
headings in order to

M
~---,,-"' allow room to mark
1. AKEBIA Dcne . / for type face
fVVVVV~/
1. A, quinata (Houttuyn) Dcne. --Honoecious, woody , twining vine.
~

Leaves deciduous or semi-evergreen, palmately compound, alternate ,

exstipulate; leaflets 5, elliptic or \"eakly obovate, 2-8 cm long , 1-4 cm

wide, retuse, entire , terminal leaflet usually largest; pet iolules 4- 15 mm ,

long. Inflorescence axil lary, racemose. FlO\07ers frag rant, functionally

unisexual, 3-merous, petaloid sepals and petals light purple ; staminate

flowers proximal, smaller , stamens 6; the pistillate distal, 1 cm or more

broad, carpels 3-10, separate . Follicles purple or violet, fleshy , to 8 cm

long. (n=16) La te spring. Estab l ished in low woodlands and on roadbanks;

mts. and pied . of NC. [Tex, Pa l A rare escape f rom cult .

No period
a f ter ( )
or [ 1
unless last
entry; ~
spaces
to start of
next entry
506 24

0pa~
78.' FUMARIACEAE...y
5 spaces

~~nnual' biennial or perennial, usually glabrous and glaucous

erect or scandent herbs . Leaves basal or cauline, pinnately or

ternately parted or divided , alternate, exstipulate . FloHe r s per-

feet ; sepals 2, bractlike , often caducous; corolla bilaterally

symmetrical or irregular, petals 4 , I or 2 of the outer petals

basally spurred or saccate, the 2 inner petals usually connate

apically over the stigma; stamens 6, diadelphous; stigma lobed,

style persistant, ovary bi carpe l late, I- l ocular .

~ ~ period after
~ Corolla 2- spurred.~~ number and name

2. Vines ; petals persistent ........ .... .... .. ... 1. Adlumia .

2. Acaulescen t herbs ; petals dec i duous ..... . ... . 2. Dicentra .

l.

I
Corolla I-spurred.

3. Ovary and fruit elongate; fruit dehiscent;

~
seed crested .... .... ............ ..... .. 3. Corydalis.

8
3 spaces
'il~~:"
6 spaces
': : ,: : :'"',,~' :"::':::~: e
)

do not run keys

, '"~'"
past right ha:1d
column of numbers

o By C. Ritchie Hell, University of North Carolina

24 507

78-2-1

c::!:)
YT
2.
spaces_

DICENTRA Bernh.
period at end of
entry only when genus
entry ends in abbrev.
authority .

Perennial, scapose, glabrous, often glaucous herbs from scaly

rhizomes or bulblet-bearing rootstocks . Leaves basal , long-petiolate,

pinnately or ternately decompound, ultimate segments minutely

apiculate. Inflorescence a raceme or panicle . Sepals 2, reduced ,

often caducous; corolla zygomorphic, petals 4, the outer 2 spurred

or saccate at base, the inner 2 spatulate, crested dorsally ,

distally connate; stamens 6, in 2 groups of 3, each group fused

and opposite an outer petal. Capsules valvate. Seeds lustrous

black, reniform to suborbicular, ca . 2 mm broad, crested. Stern, ,

K. R. 1961. Revision of Dicentra. Brittonia 1 3 : 1-57 .

1. Inflores cence a panicle ; flowers more than 18 mm

long , dark pink .................................. 1 . D. eximia .

1. I nflorescence a raceme; flowers less than 18 rom

long, white or light pink .

2. Spurs 1/3 or less of total corolla length;

corolla base cordate .... .... . ......... ..... .. 2 . D. canadensis .

2. Spurs 1/2 or more of total corolla l ength;

corolla base sagittate ....................... 3 . D. cucu11aria .

508 24

Section C. ABBREVIATIONS

A. Nomenclature
sp., spp . species (sing . ), species (pl.)
ssp. , sspp. subspecies (sing.) , subspecies (pl.)
var. , vars. variety (sing . ), varieties (pl.)
f. form (sing . ) , forms (pl . )

Authority abbreviations- -follow those in the "Hanual of the Vascular Flora of

the Carolinas." In case of doubt, write out in full.

Cornmon name for species which dup l icates that of genus, use capital first
letter for generic common name; e.g., OAK for Quercus, WHITE O. for Q. alba .

B. Description
Without period With period

mm millimeter (s) ca . circa, about

em centimeter(s) cult. cultivated
dm decimeter(s) diam. diameter
m meter(s) freq . freq uent

Flowering times - -No abbreviations; do not use months . Spell out spring, sum-
mer, f all, all year; late or earl y can be used as modifiers where appropriate.

C. Distributions

Compass directions (small letters, no periods): n , ne, e , se, s, w, nw, sw

SE (caps, no period) = general distribution throughout all states of the

southeastern United States

States (standard abbreviations without periods or internal spacing): Ala,

Ark, Del, Fla , Ga , Ky, La , Hd, Miss , NC, SC, Tenn, Va , WVa, Tex, Okla , Mo,
Ill, Ind , Ohio, Pa, NJ

Physiographic provinces (caps, no period):

SA - Southern Appalachian Highlands pp - Piedmont Plateau

BR - Blue Ridge IP - Interior Lm" Plateau
RV - Ridge and Valley Cli - Cumberland Plateau
CP - Coastal Plain (both Atlantic and AP - Alleghany Plateau
Gulf and Mississippi Embayment) CA - Cumberland - Alleghany Plateau
AC - Atlantic Coastal Plain OH - Ozark - Ouachita Highlands
GC - Gulf Coastal Plain OZ - Ozarks
ME - Hississippi Embayment OU - Ouachitas
D.
Synonyms from Manuals
S - Small, 1933 G - Gleason & Cronquist, 1963
F - Fernald , 1950 R - Radford et al . , 1968
incl. If name used includes synonym ; e . g . , "incl. A. discolor Pursh-- F t S"
under Aesculus pavia.
Journal Abbreviations: Follow those in Schwarten , L. & H. W. Rickett. 1958 .
Abbreviations of titles of Serials cited by Botanists. Bulletin of the Torrey
Club 85: 277-300.
24 509

Sec tion D. GEoGRAP HY--ECOLOGY

I. GEOGRAPHY

Distribution sheets , one pe r species, for pl ott i ng data can be obtained

by writing Albert E. Radford , Department of Botany, Un iversity of North
Carolina , Chapel Hi ll , N. C. 27514 . See Sample. Dis t ribution is to be by
major phys i ographic provinces , unless restricted (see below), wit hin the south-
eastern states , and by state within the adjacent states . Each county in the
list on the sample is keyed, by type face , to one of the three major provinces :
Mountains (Bo l d Face) , Piedmont or I nterior Lot., Plat eau s (Italics) , and Coa stal
Plain (Regular) . The symbols f or t he major provinces are : mts . for moun tains
which includes all subprovinces under SA and oH (see back of-s3mple sheet) ;
pied . for t he piedmont and int e rior lot. plateaus; and cp o for the coastal
plain wh i ch includes the sub provinces AC , GC , and ME (see Section C for
abbreviations) .
Province data will follow semicolon after habita t ; e . g ., Rich woods; mts .
If a species is found in two pr ovinces the provinces I.,i ll be connected by "and,"
e . g . , Rich woods ; mts . and pied . If a species is found in a l l major provinces
in the sou theastern states, "all pr ov ." is used , e . g . Rich woods; all prov o If
a species is typically foun d i n one or two provinces a nd occasionally in one or
two others, province data should be preceded by " chi e fly "; e . g . , Rich woods;
chiefly mt s . and pied. This means that the species is occasionally fo und i n
the cp ., but primarily in mts . and pied . If "chiefly mts. " had been i ndicated,
then the species also might be f ound occasionally in the pied . and/or cp o
Dis t r i bution by states within the southeast will be a separate ent r y with
the s t a t es a l phabetized; e.g ., Ri ch woods; mts . Ala, Ark, NC, SC. If distri-
bution is i n each state of the southeastern United States , the symbol " SE" is
used . If distribution is thr oughout the southeastern states except one, tHO,
,
or three s t ates , then SE is fo l lowed by "except" and the one , two , or three
states lis t ed alphabetically.
Fields ; all pr ov o SE except Fla and HVa .
Fie l ds ; al l prov o SE except De l , Fl a , WVa .
Distr i bution in adjacent s t ates will be a separ ate ent r y to be shown by
abbreviat i ons , in brackets [], from southwest to northeast , e . g ., Tex, Okla ,
No , Ill , Ind, Ohio , Pa , NJ . If the species is present in all eight adjacent
states , use the symbol [ALL ].
Rich woods; mts . Ky , Va, INa . [Ind . OhiO, Pa , NJ]
Fie l ds ; al l provo SE [ALL}
If di st r ibution is restric t ed , then the author should give precise data
using the appropriate symbols f r om Section C.
JunCuS tri f i dus--Rocky ledges ; BR . NC .
Heuchera rucigei- -Rocky woods ; I P. Ky and Tenn . [Ohio ]
Avicenn ia ni tida - -}1angrove sl...amp s ; Keys. Fla .
Sedum pusil lum -Granite outcrops; PP. Ala, Ga , SC .
If a species is found in tl"O or three major provinces within the so uth-
eastern Uni t ed States but not in the two or th r ee ma j or provinces fo r each
state li sted , then t he pr ovincc(s) in which i t is f ound within a state is
given af t er the state symbol.
Plan ta herba- -Rich woods; mts . and pied . Ala mts ., Ga, NC, SC .
This means that the species is found in the mts . and pied . provinces of the
three state area , but in Ala only in the mts .
Planta herba--Fiel ds ; a l l provo Ala , Ark cp ., Del , Fla, Ga , Ky .
This means that the species is fou nd in all thr ee ma jor provo of the south-
easte r n st ates listed but in Ark on l y in the cp o
Distribution information i s to be based on specimens examined by the
510 24

Table 24- l. Countie s and Ph ysio graphic Provinces of the Southeastern United
States .
Species
ALA. o Wins o Soar [l Osee Cl Douy n P olk o CaTT o Mull! o More o Car r
o o Soba [] P13ea o Doug o Pub o Carl o Nc!.< O Nate o Chic
Auta ARK. o Sc"i o P"s~ o Ea r l o Pmr! o en,e o N ich [J Orlc o Choe
C Ba ld
o o Shar [J Pine [J Echo o Q u it o CI,r; o OIlio o Ouae o C la i
13nrb [] Ar ka n Sion r-] Pol k n Em [] Rabu' o Ciar o Oldh rJ o Clar
o Bibb o Ash! o Voio o o Elbc
Putn o j{rmdo Clay D Owen o PCou
Plag
n
o Blcll o Baxl o VBllr [l SJoh r I Eman o Hieh o eli" o Owsi [] Rapi
Clay
o Bull o Benl o W "sh n SLue o Evan o Rock [J [] Coah
oo n Copi
Crit [l RR;"
o But! o Boon o Whil o SRos n F ann' o Sehl [J Cumb Pc"d
Perr [ j Rich o
o Calh o Brad o Wood o Sn'-;l [J Faye [J Sere D Oll!); o Pike o Sabi
Covi
(J LlSot
o Chum o C" lh o Yell [1 Semi o Flay o Semi o Edmo [] Powl' [] SBer o Forr
[] Chef n Carr n Sumt o Fors' o Spa! [] Elli [] Pula [] SCha Il Fran
[] Chi l
o
[] Chic
o
DEL. [] Suwa rJ Fran D Step o Esti [] Robe r:J Sl le l n Geor
Choe
n
CiaI'
o
[l
n
T~yl D
o
Full o Stew oFaye r) Rock o SJam o G ,.f'C
[] CiaI' C la y
n
Kent Unio Gilm [] Sumt oFlem !] Rowl! n SJBa n G"en
[] Clay
o
[] Clob NC<ls [] Volu r l Gbs [J Ta/b oFlay n Russ IJ SLa n [] Hane
Cleb [J Clev
o
o S u ss rl Wa ku o Gl y n [] TI,li [] Fran [] Scot rl SIIln r o Harr
["] ColT
o
Col u [l Wa lt CJ Gord [J Ta t! oFu ll o She! n 5 ]\:lay fl Hi nd
[] Colb
o
Conw FLA . D Wash ~l G rad o Tay! oGal! o S imp
[] STarn n Holm
O Cone
o o
Crai
o
rJ Cree [] Te lf oGarr n I-I
Spc n Tang o Hu mp
o
Coos Craw Alae
n Bake
GA. 11
n
Cwin n Terr n Gral! o o
To"l Tens n Issa
Covi [] Crit
o Habe '
o
[] Th orn n G ra v r) o
Todd Terr n Ilaw
[] Cren
o
Cros [J Bay [] Apr l rJ HaW Tift o
Gray n fl
Trig Unio n Jaek
[] Cull
o
Dnll
o
[] Brnd [] Alb
o
11 Hune o Toom n
Grell n [J
Trim Verm [l .htsp
[] Dale
o
Desh Brev Baco
n Bnke
o Hum [J Town· [] Grep o UlliO n Vern n Jeff
[]
o
Dall
o
Dr ew
o
[1 Brow
o {i Harr o T,.eu I I Halle o War.- rJ \Va.,h n JDav
o
DK al
o
Faul
o
C~lh Bald n Han n '/'ro" :l Hard o Wo s l, D Webs n Jone
o
Elm o Fran Char [] Ban!; n /lear o Turn o Harl n W".'!" o WnRo rJ Kemp
o
Esca [] Full o Citr OBa rr r I Hene [J Twig n Harn nWe bs o WCar n Lafa
Elow o Garl n Cla y [] Barl n Hous [] Unio· [-J Harl o Whit [] W~'cJ o
o F~ye o Gran [] Coli n llHil Il Trw i [J Upso [l Hend n Wolf o Winn n
Lama
[J Fran o Gree o Colu n Berr :1 Jilek Ll Wal k Ij llenT o W ood n
Lnud
O G en e o Hemp [] Dade: n Bibb n Jasp [J Wall f"J Hick MD.
Lawr
r l Leak
[] Gree [] HSpr o DSot n Blec n .JDav o Ware rJ Hop" LA. o
o Hale [] Howa [] Dixi n Br~n n JeIT o Warr [1 J ack
Lee
o Hen r o Inde o Duva [] Broo [] Jenk rJ Wa sh n Jeff n Aead
D Aile
o AAru
[] Lell
n L ine
[] Hous o har o Esca r B ~' ya n .Ioh n o Wuyn rl Je.<.< n Ail e [] Bait n
o
o
Jack o Jnck n Flag n Bull n Jone [J Webs [J John n Asce o BCi!
Lown
n I\Iadi
Jefl [] JelT o Fran rJ Burk n Lamll o Whee n Kent n o n
o Lama o J oh n o Gads o BUI! n L:l.l1i o Whil' [j Kno!
Assu
[] Avoy o
Cal" Mari
n ll;lars
o Laud o Lafa n Gilc lJ Calh o La ur Ll Whid n Knox n B~au o
Caro
Carr D !'Ilonr
o Lawr o Law r o Glad n Camd n Lee [] Wi!c [] La ru n Bien [] Ceei o Mon t
oo Line
"",
[J Lee
o Lime
[] Gu lf
n Hami n
[l C,md Il Libe [] Wilk
n r.;fle [J Wi In
[] Lau f ro Boss o Chnr [1 Nes h
o LR iv o Hard
Carr
rr Calo n Long o
[] Lawr n C~dd [1 Dorc n Ne wt
[] Lown
o Wor t n Leo Il Calc o Fred n Nox u
Ma co n Loga n Hend n Ch3r [l Lown [l Lesl n Cald
o Ma di o Lono n Hem n Chat 11 Lump' KY. n Lelt n C~mc
[] Ga rf
o Hurf
:I Okti
: Pano
[] Mare o Madi
o Mar; o High [] Chac n ;"leDu o Lewi 'I Cala o Hawa n Penr
L! Mari
o Mars o Hill II Ch~a fl 1\1cJn n Ariai r-I Liu(' n Cbi o Kent r--; Perr
n Mi ll o Holm n Chcr ' . I M~co n Aile n l.ivi n Cone :l MOll ! n Pike
[] I1'lobi o illiss [J lRiv r:J Clar ci Mild; n Jlude o Loga n DSo t CJ PGeo [] Pont
o Monr o MOll!" n J~('k n Clay n Mal'i rl Ball 11 Lyon n EB Ro [l QAnn [j Pren
[] Mont o Mon t n JeIT Il Clal f"J t1/(' ri [) Barr [] MeCn [) ECar n
o Morg
o Pe r r
o Neva o Lafa i i Clin n "-li ll n
Bath o McCy n EFel
SMar
fl Some
[l Quit
o R~ nk
o Nowl [] Lake [J Cobb r J Mite n Bell [J McLe n E\'an r '] Ta lb I.J Scot
[] Pick
o Pike o Ouae n Lee IJ ColT 'I Mm,r [] Rooll Il M",li [J Fra n [] Wa5 h 1 Shnr
o Perr o Leon !l Colq .1 :Vlont n [j"lIr o Mago [J Gr~n n W,co '1 Simp
O Rand o Phil o Levy IJ Col" n Morg o
Boyd rl Mad II I ben l"' Wore [] Smit
[] Russ o Piko IJ Libe [] Cook n Murr [] Boye n ilIars n Ibee n Ston
o SCla o P a in o M~di [J Cowe n Muse n Brae [] Mart n Jilek MISS. 1"1 S un f
O Shel [] Polk o Mana n Crnw -! Newt [] Brea o Ma so r.l JeIT II Ta ll
[J Sum! [J Pope n )l.lari I ! Cris n O('on n Bree n Mead r) .ID,,,, n Adam o
o Tald o Prai n M~rt o Dade rTOgle [] B,,/I n Me ni n Lnfn rl Ako
Tate
[- I Ti pp
o
o
Talp o P ula o I"onr o Daws ' n Palll [] Bll!! [] lIlere [] LFou o Amit n Tish
Tusc r::! Rand [J N~ss II Deca 'l Pe~e [] Cald [] lIlete n LS~1 'l Tu ni
o Walk [] SFra o Obi o DKal [] Pick Calln [J !IIOTiT r ) Line n
[] Alta
Be n t [j Un io
o Wa sh o Sali n Okke 0 Dodg [] Pier Cnmpo [] Mom n Livi [l Boli [] Wa lt
[] Wile o Seol [] Oran 0 0001 o Pil;e C~rJo o Morg o i'l-lndi o Ca lh . l Warr
24 511

o o Cumb o o o Ke rs o Clay o Laud o Amcl o Jl.la th o

Gr an
o
Wash
o Curr o
M U,;'
o
Wayn
Wilk o Lane o Coek o Lawr o AmJle o Meek o
Gree
o
o
Wayn
o Dare
Mon t
o Moo r o o Lallr o CofJ o Lewi o Appo o Midd Hamp
o
Webs
o Nash o
\Viis
o o C roc o Line o Atfi o Monl fJ Hane
o
Wilk [l Davit
o Dave o NHan
Y adk
D Yane ' o
Lee
Lexi o Cmnb o ~o\! d o Aug\! o Nans o
Hard
o
Wins
o Du pJ o No r t o MeCo o Davi o MeMi o Ba lh oNels o
H a rr
o
Ya la
o DUTil o Onsl S.::. o Mari o Dcca o McNa o Bcd! O NKen o
Jack
Yaw
o o Marl o DKal o Maco o B lan oNorn o
Jeff
Edge D Oran
o Paml o Abbe o Newb o Dick n Modi o B olo oNord o
Kana
N.C. :J Pors
o Pasq o n o o o o
Lewi
o
ooo AAnd Deon Dye r Mad Bnm NOH
o o
Fran
o Pend
ike
D Oran [J r'ayc o Mars o Buch oDran o
Line
o
Atam Ga~(
o Gate o Perq
Al1c
e o Pick o Fenl o MaliT o Bli ck oPago o
Loga
Alex
o Griib' o Pers o 8amb o o Frau o Moig o Camp [J P a t r o
MeDo
D Aile'
o Gran o Pitt CJ Barn o
Rich
Sal" o Gibs o Monr o Caro oPitt o
Mari
D
o
Ansa
o Cree o t'olk' o o Spa r o Gile o /'.'lont o Carr oPowh o
Mars
Ash o'
o GlIil o
Beau
o Sumt o G ra i o Moor o CCit U P Edw o
Maso
D Av er'
o Bali
O Rand
o mch o
Berk
o Unio o Gr oe o Morg o Char oPGeo o
Mere
D Beau
o Berl oo HHarn o Robe
Calh
fJ Char o o G ruD Ll Obio o Ches oPWil o
Mine
o Blad o Rock o
Will
York o Ham b [l Over o Cl3 r oPula o
Ming
o Emn
ayw'
o H ond' o
O Cher
o o H am; o PerT o Crai oRapp o
Mono
o Bunc' o Hert o
Rowa
o
Ciles
TENN. o H ane o Pick o Culp oRich o
Monr
o o Hoke
Rutli
o Samp o
Ched
Clar o Harm D P olk ' o Clim b oRoan o
Morg
o
BUTk
o H yde o Scot oo D And o n Harn o l'ufII [1 Dick oR ock o
Nich
o
Cabn
o Ired o SIan
Coll
Darl o Bcdj o H a wk o Rhea o Dinw oRoe m o
Ohio
o
Cold
o o Siok o o Bent o Hayw D Roan o f:sse ' oRuss D Pend
o
o
Camd
o
J ack '
o o
Dill
Dore o Bled o Hend o Robe o Fa ir oScol Plea
o
Ca rt
o Casw o
John
o
SUrT
o Edge o Blo\! n He m o Rlllh o FlI uq oShen P oea
o
o Cara o
J ane
o
Swai'
o Fa ir o Brad o H ick o Seol o fl ay oSmyl Pres
o Chat o
Lee
o
T ra n '
o Flor o Ca m p o o Soqu o Fhw oSout o
P utn
o Chllr" o
Lena
o
Tyrr
o Geor o Cann rJ
H OllS
Hump o Sovi ' o Fra n oSpot O Rale
D Chow o
Line
o
Unio
o Gre v [J Carr o Ja ck i J She\ o Fred oStaf O Rand
o
o Clay' o
ilfcDo
o
Vane
Wilke o Gre w o Carl· o Joff o Smi! o G ile oSurr Rile
o
o elev o Mildi '
Maco'
o o o Cll ca o Jo~ o Stew 0 0 101.: oSuss Roan
o
Warr Ham p
o o
o o o oo
Summ
o Colu oo M:lrt o Wash Horr
o
Ches :<::..e~
o
.~ ".lll Gooc TUB
oo o
r ayl
o Crav Meek o W a la ' o Jas p Clai [1 La!~~
o
Swr.n
Tipt o
G ray·
O ree
W arr
Wash o
Tuck
o T rOll o Ore\' oWest o
Tyie ,
o !,blc' o Hali oW ise o
Upsh
o -'::lio o Ha:1o oWylh o
W ayn
o V 91:~ o H c nr oYork o
Webs
o '!Iarr o Hcny o
Weh
o o n W ash o High W.VA. o
Wirt
o WOlin o IoWi o
Wood
o WCllk o JCit oBarb o
Wyom
[] Whit o KaQu [J Bark
o
o
o oo
Will KCeo Boon ILL.
o Wiis I(Wil oBrax [l IN D .
o Lane o Broo o
MO.
VA. o L oo o Cabe o
N. J.
o Loud o CaIh n
OH IO
o
o Aceo o L oui o C lay OKLA
o Albe o Lune o Dodd o
PENN.
D Aile o Madi o F aye o
TE X.
o G Hm o
L_ I.

Standard = Coaslal Plain: CP (including AC. CC. ME).

Italic = Picdmonl : PP. and Inte r ior Low Plateau : IP
Boldf ace = Mounlains: RV; CA (including CU. AP); OH (includ ing OZ. eu).
Boldfaee' = Blue Ridge: Bn.
 512 24

author in several major herbaria and from publications by competent authori-

ties. Do not base distribution on general ranges given in manuals.
These geographic instructions do not cover all distribution possibili-
ties so each author is enjoined to be consistent in desc r ibing his r ange
exceptions .
II . ECOLOGY--HABITATS

A. Notes

Designation of fr eq uency or density should not be used unless author is

entirely confident of statements such as rare, infrequent, common, abundant .
Endemism should be stated \~here certain , e . g . , Shortia ga l acifolia. en-
demic to BR of Ga , SC, NC .
Specific habitats should be used f or highly localized species when defi-
nitely knot-m : i.e . , Sand dunes ( Uniola paniculata) . Shale ba r rens (Oenothera
argillicola), Limestone or dolomite outcrops (Asplenium resiliens), Hammocks
(Clusia f 1ava) , et c.

B. Habitat Designations
(See Chapter 17)
Aquatic Balds

river lake heath balds

stream pond or pool grassy ba l ds
springhead marsh
ditches bog Disturbed habitats

Us e modifiers \"here certain : t~asteplaces gardens

roadsides pastures
brackish brackish marsh railroads meadows
marine shores peat bog fence rows, hedge rows
acid (pH 5 & belO\~) shrub bog
rapid cypress swamp Savannahs (use modifiers when needed)

\"et
dry
Fields (use modifiers when needed ) pine

\"et clay Outcrops (use modifiers when needed)

dry loam
low rocky granite
upland old sandstone
sandy cult . limestone

l-,l oods (use modi fie rs to describe cond i tion and also cha r acteristic species
where needed) .
evergreen pine, spruce-fir, longl eaf pine
deciduous oak woods, mixed hardwoods , oak- pine
low (alluvial or swamp) Taxodium, Nyssa
rich oak woods , mixed me sophy tic
dry oak-hickory, oak-pine
dry sandy turkey oak
dry clayey blackjack-post oak
open pine
dense cove hardlwods
 513

Sect ion E. SYNONYMY

Synonyms are to be included only ~."hen the names employed in the Flora of
the SE differs from those used i n the recent manuals of the Eastern United
States . These recent manual s and their abbreviations are in chronological
order by author:

S J . K. Small . Manual of the Southeas t ern Flora. 1933 .

F M. L . Fer nald . Gray ' s Manual of Botany , 8th edition . 1950 .

G H. A. Gl eason a nd A. Cr onquist . Nanual of the Vascu l ar Plants of

the Nor t heastern United States and Ad jacent Canada. 1963 .

R A. E . Radford, H. E . Ahles and C. R. Bell. Nanua! of the Vascul ar

Flora of the Carolinas . 1968.

A) Previous segregates accepted in the above recent manuals but not by

the contributor should be listed as follows:

(Under Cassia nictitans L . )

"Incl . Chamaecrista mul tip i nna ta (Pollard) Greene--S ; Cassia n . var .

hebecarpa Fernald , f . !!.. var. l e iocarpa Fernald--F, G. "

B) Taxa previously i nc luded as synonyms in the above recent manual s but

regar ded as distinct by the contributor shoul d be listed in synonymy as
follows :

(Un der Desmodium n u ttallii (Schindler) Schubert )

" . . . Heibomia viridiflora (Nuttall) Kuntze- - S, in part; D. viridif l orum

(L . ) DC.--G, i n part."

(Under Cassia obtusifolia L . )

. . Emelista!.£!2. (L .) Britton & Rose--S ; f. ~ L . - -F , G. "

"
C) Taxa previous l y passing under the name of other taxa should be indi-
cated as fo l lows :

(Under Amorpha georgiana Hilbur)

"A. cyanostachya Curtis-- sensu S, not Curt i s"

(Under Aeschynomene indica L . )

"A. vir ginica (L.) BSP--sen su 5, in lar ge part ."

(Under Sabatia s t ellaris Pursh)

"5. campan ulata (L . ) Tor r ey- -sensu S . i n part ."

514 24

D) Obvious synonyms f rom the above recent manuals should be listed :

(Under Diamorpha smallii Britton)

" inc1. D. cymosa (Nuttall) Britton ex Sma11--S, Sedum sma l1ii (Britton)
Ahles--R."

(Under Psoral ea lupinellus Michaux)

" Rhytidomene lupinellus (Hichaux) Rydberg-- S . "

(Under Lotus helled (Britton)

" Acmi spon helleri (Britton) Small--S . "

Note : Synonymy is not as mechanically categorized as the above might indicate .

The above is meant merely as a tentative guide and the contri butor is encour-
aged to convey the facts accurately and concisely . Constributors are strongly
urged to publish their nomenclatural research in full elsewhere . Scholarship
demands more than the legal minimum of the basionym .

Section F . VASCULAR FAMILIES OF SOUTHEASTER..N UNITED STATES

(Selec ted)

1. Acanthaceae - Long 15 . Aristolochiaceae - Bell

2. Aceraceae - James 16 . Asc1epiadaceae - Bell
3. Aizoaceae - \.,Tard 17. Aspidiaceae - Evans
4. A1ismatac eae - DePoe 18. Aspleniaceae - Evans
5. Amaranthaceae - Massey 19. Asteraceae - Shinners
6. Amaryllidaceae - Bro~me 20 . Azol laceae - Evans
7. Anacardiaceae - Long 21 . Balsaminaceae - Hard
8. Annonaceae - Kral 22 . Bataceae - Bell
9. Apiaceae - Bel l 23 . Begoniaceae - Smith
10. Apocynaceae - Bell 24 . Berberidaceae - Bell
11 . Aquifol i aceae - Clark 25 . Betulaceae - Hardin
12 . Araceae - Hard 26 . Bignoniacea e - Bell
13 . Ara1 i aceae - Be l l 27 . B1echnaceae - Evans
14. Arecaceae - Read 28 . Boraginaceae - Honnnersand

Section C. TAXONOHI C PHILOSOPHY, CONCEPTS, POLICIES

(From Shetler et a1. [1 973J A Guide fo r Contribu t ors to Flora North
America [FNA] [Provisional EditionJ)

I . TAXONOHIC PHILOSOPHY

The Editorial Connnittee will encourage in principle a broad species con-

cept and taxonomic practices that promote stability in the taxonomy and nomen-
clature of North American plants . To this end, it is expected that the authors
Hill keep changes in rank to a minimum and make them only after adequate taxo-
nomic invest igat i on and publication of the results in an appropriate journal
or monog r aph . h'here current authorities offer a choice of rank for a particu-
lar taxon , the author's decision should be made in the light of the taxon's
variation throughout i ts Hho1e geographic range . Radical treatments involving
excessive speCies-splitt ing or species-lumping relative to past practice Hill
be given the closest scrutiny by the Editorial Committee, and the authors of
such treatments must be prepare d to defend them on objective gro un ds . In no
24 515

case \dll the pages of the Flora be open for publication of nely taxa or new
combinations. Such publication should be done in the recognized journals.
where appropriate evidence and documentation can be presented .

II. TAXONOmC ~ONCEPTS AND POLICIES

The contributor will define the limits of all taxa within the group he
1s treating, while the Editorial Committee , in consultation with the. sp e cial-
ists . wil l be responsible fo r delimiting taxa of coordinate or higher rank.
In practice, this means that in most cases the contributor Iyill be responsib le
for the limits of infrageneric taxa , mainly the species, and the Editorial Com-
mittee wil l be responsible for setting the limits of genera and higher taxa .
Contributors dealing \·lith whole families naturally viII be r.esponsible for
delimiting all in cluded genera. All decisions by contributors will be subject
to the usual editorial reviev .

The contributors and Editors should take into account all available evi-
dence--mo rphological , geographical , ecological, cytological, chemical--in
delimiting genera and species, but the taxa so recognized must be definable
ultimately on a practical morphological basis .

A. Genera . The Editorial Committee will strive to achieve a reasollably

uniform generic concept from group to group , but especially \~ithin fami li es .
Although there can be no fixed criteria for delimiting gener a, due considera-
tion should be given to historical precedent and to the importance of main-
taining nomenclatural stability . Familiar genera should not be merged or
split on the basis of personal preferenc e or other t r ivial grounds, but only
on t he basis of carefully we i ghed new evi dence that is documented fully in
the literature. Furthermore , generic a lignments shoul d be made in the light
of treatments in Flora SSSR and Flora Europaea . For the sake of advanci ng
common understanding, every effort should be made to eliminate artificia l
gener ic distinctions bet\~een the FNA and comparable European or Russian treat-
men t s of the same or c l osely related species .

lL SpeCies. The FNA Hanual above all is to be a practical Hork , and th e

primary units of the Flora , the species , must be practical entities Hith
cons i stent , if small, morpho l ogica l distinctions . At the same time i t is
understood that no tlW species, nottdthstanding their equivalent rank, ever
are equivalent in t heir origin , nature, and ex t ent of evolutionary differen-
tiation . Even wit hin genera the species, though definablE' by mor phological
cri teria, \~ill illustrate different theoretical or evolutionary situations .
All availab l e biosystematic e vidence should be used in delineating spec i es in
the FNA treatments . Some examples of different kinds of species , viewed
t heoret ically, that may be recognized in the FNA Nanual, as in Flora Europaea ,
a r e discussed in the Flora Europaea guidebook (Heywood 1958 , Appendix E). At
least the folloHing kinds of species may be recognized whenever t he mo rphologi-
cal requirements can he met :

- High l y distinct, t..ddely distribu t ed plants ,... ith little varia tion .
- Species with different ploidy levels in which the levels are not
recognizable morphologically or in which the variation is so nearly
continuous that infraspecific t axa cannot be recognized.
- Species in wh ich aneuploidy occurs , but in which the different chro-
mosome numbers are not correlated with morphological features of
the plant s .
516 24

Species distinguished by minor morphological characters , but which are

distinct ecologically from related species.
Po lymorphic species with morphologically definable subspecies or varie-
ties.
Polymorphic species in which clina! varia tion occurs and in which infra-
specific taxa cannot be distinguished satisfactorily.
- Species that are widely separated and morphologically different from
related species with which they are highly interfertile.
- Asexually reproducing plants that are morphologically distinctive and
which occupy a significant geographical range.

All recognized specie s are to receive full and comparable treatment in

accordance with the FNA guideline s, and the Editors will not include any spe-
cies in the Data Bank that are not recognized clearly by the contributor him-
self on a coordinate basis with all other species recogniz ed by him . The
recognition and treatment of microspecies is discouraged , although microspe-
cies may be grouped as complexes into broader species. In such cases, ortho-
dox binomials must be used for the broad species (e . g., Campanula rotundifolia
L. ~ . !., not Campanula rotundifolia complex), and the micro species must be
treated as taxonomic synonyms, not as a l is t of included microspecies. To
summari ze , there can be only one level of species . Species may be aggregated ,
however, to simplify keying or to show close simi larit ie s, provided that the
"aggregate ," "group," or " complex" can be keyed out and diagnosed as a unit
on a practical morphological basis and can serve. therefore, as a useful
though informal supraspecific category .

C. Infraspecific Taxa . Only major infraspecific taxa that are morphologi-

cally definable entities \.;rill be included in the FNA Manual. Either the cate-
gory subspecies or the category variety may be used , depending on the history
and knO\.;rledge of the group, hut both may not be used in the same species .
Furthermore, each contributor should use one category or the other consistently
throughout the genus or other group that he is treating, except that individual
authors treating medium-sized or large families may be permitted to vary usage
from genus to genus if, in the Family Editor's judgment, such inconsistency is
justified. The us e of subsp ecie s is preferred, because i n the Editorial Com-
mittee 's view this is the more meaningful evolutionary category , but histori-
cally the category variety has often been used in about the same sense . To
exclude varieties from the Flora would encourage \.;rholesale publication of new
combinations, a result that in itself would not contribute to our knowledge
of the given taxa and \wuld be contrary to the Committee's goal of promoting
nomenclatural and taxonomic stability . The Committee is attempting, therefore,
to strike a balance in the use of subspecies and varieties between total incon-
sistency and total but artificial uniformity , realizing that the guidelines
set forth here will cause a certain amount of r ecombination to bring about the
required consistency \ilithin groups . In general, the contributor should be
guided insofar as possible by existing knowledge and nomenclatural precedent .
In groups where subspecies are recognized , traditional varieties should be
dropped altogether if they cannot be defended on good evolutionary grounds
as subspecies .

The following kinds of taxonomic groups may be treated as subspecies or

varieties in the Flora :

- geographically iso lated populations that differ from one another by

feHer characters than do species of the same genus and can be
24 517

demonstrated to have some degree of interfer t ility .

- ecolog ically specialized populations , particularly with respect to
certain edaphic or climatic conditions, when there also is a mor-
pho logical mean s of distinction .
- cyto logical variants, if they have characteristic distribution patt erns
and can be recognized by featu res other than chromosome number alone.
- chemical varian ts, with concomitan t morphological a nd geographical or
ecological attributes .

A key to inEra specifi c taxa must be provided when either subspec ies or
varieties are recognized. All formally recognized infraspecific taxa must be
treated in the standard way, includ ing ful l description . Taxa that are not
accepted by the author himself may not be included.

Any variation not meritin g treatment at the subspec i fic or varietal l e ve l

(e . g . , clinal or local eco typ ic variation, minor variation in color or pubes -
cence), is no t t o receive formal recognition . Specifically , the forma l use
of the category form will not be permitted . Informal comment on infraspecific
variation must be brief , germane , and without the haphazard use of such terms
as " culti var," " form," " type ," and "race"; otherwise, it \"ill be pruned rigor-
ous ly or e liminated .

D. Other Categories . Infrafamilial and infrageneric categories sanc tioned

by the International Code of Botan ical Nomenclatur e (IeBN ; Stafleu and Voss
1972) may be intercalated between family and genus or bett"een gen us and spe-
cies , respectively, provided they are formally described and serve a useful
purpose in the treatment. The judicious use of s uch ca tegories is encouraged ,
as a means of subdividing large families and genera into more natura l units,
thereby making the t r eatmen t of such a family or genus mo r e useful as a sys-
tematic consp ectus. Their use also a llows for more· concise treatment , be-
cause characters that are common to a whole group of species or genera can
be described under the appropriate inf rageneric or infra f amilial category
and can be omitted from the descriptions of the individual species or genera.
Excessive s ubdivid i ng is discouraged , however, because the treatments must
remain practical. General ly, a singl e rank be t ween family and genus and
between gen us and species sho uld be suff icient. Ordinarily, this will be
subfamily or tribe between family and genus , and subgenus or section
between genus and species , depending on traditional usage t"ithin the group .
The ranks s ubtribe, subse ction , ser ies , and, particularly , subseries should
be used sparingly if at all . Usually, the infrafamilial and infrageneric
taxa \~ill not be included in the keys .

Experimental categories, e . g . , coenospecies , ecospecies, ecotype. deme,

may not be used in the formal nomenclature . Likewise, the t erm "cultivar"
and other special t erms in use for cultivated plants wil l not be permitted
as d esignations of formal categories. If essential to an understanding of
the contributor's treatment , the current bios ystemat1c concept of the group
may be summari zed tersely in pertinent experimental terminology under the
section " Observations ." Expansive hypo t he tical discussions in rel a t ionships
will be condensed to th ei r essence or eliminat ed by the Editors.

E. Hybrids. Intergeneric and intersp ecific hybrids may be include d in the

Flora only when they are common , self-propagating taxa twrthy of treatment as
ordinary genera or species . Such hybrids must be given regular generic or
binomial names prefixed by the multiplication sign (x), i n accordance with
518 24

Appendix I of the IeBN . Fo rmul a s to expr ess hybrid paren tage, as provided by
t he IeBN, may not be used in t he formal nomenc lature , but may be used spar ingly
i n an explanat ion of the parentage und er "Observa tions " whe n s uch explanation
is abso l utel y necessary.

The i ndiscriminate naming of hybrids is to be a~oided. and in general

the Editor ial Committee wi ll pe rmit the inclusion of designated hybrid taxa
only when t heir hybrid origin has been thoro ughly establis he d and wide l y
accep ted and vlhen s uch taxa c l early represent e nti ties in the No rth Amer i can
flora t ha t must be r ecognized. Long l ists of putative hybrids wi ll be e limi-
nated . If i t i s essen ti al to an unders t anding of the g r oup . t hen the signi -
f icance of hybridization may be explained by simpl e comment under "Obs e rva-
tions ."

F . Apomicts . The t a xonomy of groups in which apomixis occurs must be

approached with specia l caution . To attempt t o describ e or even mention all
apomicts in such critical genera a s Rubus and Tar axacum is out of t he ques-
tion , and the Editors will not accept treatments in which apomictic plants
have been named and described indisc riminate l y. Insofar as i s possible .
a pomic t s s hould be refe rre d to their closest sexually-reproducing species.
Howeve r, when an apomic tic plan t is widely dis tr ibu ted and mo r pho l ogically
dis tinct on a constant basis , i t may be t r ea t e d as a species. If it is
essential t o an understanding of t he group , then the significance o f apomixis
may be discussed brief l y und er "Ob servations ," but without giving a l ong li s t
of named apomicts .

G. Alien Species . The Editorial Committee will be guided by a l iberal

policy on th e inclusion of al i en (introduced) species. Contribu to rs wi ll be
given wide latitude t o decide which species to include, on the con dition t hat
a ll i ncluded a li ens be inco rporat ed in t o the key and described in full.
Al i ens tha t a r e t horo ughl y na t uralized, as judge d by the contrib u to r, are to
be includ ed a u tomatically . These i nclude no t only weeds a nd accidental i ntro-
ductions bu t also ga r den pl ants t hat have escaped and are propaga ting them-
selves well beyond the immedia te vicinity of where t hey are cu lt iva ted . In
addition , impe r fectly naturalized or casual aliens (adventives , waifs , escap es ;
i .e. , a l iens that a pp arent l y can propagate t hemselves in the wild, but which
are not known to be spreading nor to be l ong pers isting without constant rein-
troduction) should be included when in t he contributor ' s opinion they are of
special intere s t or are likely to be encountered frequently enough in the flora
to pose pr oblems in identification . Even c ultivated perennials (espec i a l ly
trees and shr ubs) may be in cluded if they ar e species that often are found per-
sisting un tended in the wild in st ands o r as i ndivid ua l s . Finally , there are
taxonomic groups i n which it may be desirable to trea t certain of t he purely
c ultivated species because they are trees or c r op plant s that a r e planted or
cultivated in con tinuous stands or on an extensive scale . Cultivated plants ,
i.e . , species that will not reproduce in t he Hild or per sist without tending
or managemen t, s hould be i nclud ed sparingl y . however , and onl y Hhen the i r
i nclus i on s e rves a r eal purpose . Any cul tivated species that a r e i nc luded
must be keyed and described in the usual manner .

H. Ra r e and End angered Speci es. A special effort will be made to i dentify
those taxa native to the FNA territo ry which, on the basis of their ent ire
wo rldwide range, are known or believed to be r are or enda ngered or to have
become e x t inct i n the wild r e ce n tly. I n their trea t ments , contributors should
i nc lude recen t l y extinc t species (e .g .• Frankl inia alatamaha) , desc r ib ing and
keying them , and also should attempt to designate the rare and endangered spe-
cies . Criteria for recognizing and classifying rare and endangered species
are given in a later section of the Guidebook . All introduced , naturalized ,
and cultivated plants are to be excluded from cons ideration. A species that
is rare o r endangered in one or two states. but is common elsewher e in its
geographic range, does not qualify for recognition i n the FNA rar e and endan-
gered l ist. Primary a t tention is to be given to spec i es tha t a r e endemic to
the FNA terr itory. especially if they are locally endemic or relict species.

A l is t of rare and endangered s pecies is urgently needed fo r North

America . but none has ever been compiled on a continental basis . The need
for at least a national list for the United States has been emphasi zed
repeated l y by those who have prepared environmental i mpact statements a nd
studies on the location and preserva tion of natural areas and ecological
preserves . and by conce rn ed citizens and conservation organizations . Pre-
serving ra r e and endangered species has become a matter of policy a t national
and international leve ls . The first s tep in doing s o is to provide scientifi-
cally accurate information . The designation of ra r e and endangered species
is the subject of recent bills before t he U. S. Congress. The Inte rnational
Union fo r Conservation of Nat ure a nd Natural Resources is concerned abo ut
endanger ed plants on an i n ternational leve l. Certain rare and endangered
p l ants a r e being considered for international protect i on to prevent t heir
export and import . FNA is the appropriate body to compile a scien ti fically
a uthorit at ive list of endangered plant s for the Un i ted States (less Hawaii)
and Can ada . Pe r sons concerned about endangered species on the s t a te or local
l evel are encouraged to use the same c r iteria and classification scheme set
fo r t h fo r t he FNA Da t a Bank. There is a definite need for a hierarchy of
lists of endangered species - - local. state. national. and i n terna t ional .

I . Na tu re and Scope of Descriptions . All descriptions must be comparative

as ~e ll as diagnostic . The primary object of a f l ora is to provide a rapid
mea ns of correct l y iden t ifying the pla n ts of some s pecified region . For
this reason descriptions in Floras usually include on l y the most diagnostic
cha r acter s of the pl an t s treated . Because these characters differ greatly
from group to group, t he descriptions i n a Flora r arel y are compar ab l e from
one family o r genus to another or even among species of the same genus .

The FNA Hanual is i ntended to serve the usua l purpose of a Fl or a . and .

accord i ng ly , it will i nclude only di a gnostic desc riptions . ~... hich must be as
concise a s is consistent with clarity and utility . The FNA Data Bank is
intended to serve the additional. critically impo r tant , new f unction of
corre l ation (comparison). made poss ible by the computer -based Infor mation
Sys t em i n which the Data Bank is maintained . Through t he query l anguage of
the FNA System . it is possible to retrieve the data in any of a v i rtually
limitless array of permuta t ions . Thus . simple to highly complex correlations
of morpholog i cal . eco lo gical , geographical . and other parameters a r e possible
within an d among taxonomic groups at t he same or a t different level s .

Rel i ab le. meaningfu l correla t ion is possible, however , on l y when the

descriptions are al l comparable in content, forma t, and termino l ogy. Other-
wise. t he pr oblems of missing data and unstandardized formats and terms
largel y s ubvert retrieva l strategy and make generali zation from query respon-
ses hazardou s , if not i mpossible . The r efore . the Editorial Conunit t ee has
defined a min imum set of characters that are common to most vasc ular plants
for which data must be supplied consistently and comparatively in every
520 24

desc r iption by every contributor over the full range of taxa treated , except
where the characters are not applicable . This common requirement is intended
to guarantee some measure of comparab ili ty and comp l eteness in the FNA Data
Bank from the QutS E't .

Each contributor will be expected to provide data not only for all required
comparative char acters , but also for those characters that he considers to be
the most diagnostic for the taxon in question, which o f ten may not be among the
required set . The guidelines for description are to be followed for the diag-
nostic characters as \.,re l 1 as for the comparative characters. The fu l l descrip-
tions will be entered into the Data Bank and included with the treatments when
they are published individually, but only the diagnostic characters will be
printed in the concise Manua l.

Al l descriptions are to be based on the diversity and variation of the

taxa as they are represented in the FNA territory-. An exception is to be
made only when this procedure would lead to a distorted description of the
taxon , be cause its FNA representatives are highly unrepresentative of the
taxon as a t... hole . If the diversity or variability f ound outside the FNA area
is taken into consideration. this should be i ndicated clearly , character by
character . I t is assumed that the need for taking exception to the above
rule wil l be minimal at the species level and will increase upward through
the hierarchy of taxonomic categories .

Chromosome Number is an optional character. to be given at the end of the

morphologica l description when reliable information can be supplied . Contri-
butors are encouraged to include chromosome number in their descriptions when-
ever possible , especially if number is taxonomically important or critical .
However, all counts must be verified in the primary sources and must be based
on material of known wild origin within the FNA ter ritory . It is in t ended
that in due course the FNA Data Bank will become the central repository for
authentic , critically evaluated information on the chromosome numbers of North
American plants. In the interest of producing the FNA Manual rapid l y , hm...-
ever , available information on chromosome number is not being requested on a
mandatory basi s at this stage , because of the extra effort that would be
required of the contributors and Editors to search out and verify the data .
An unverified FNA compilat ion would not add s i gnificantly to the secondary
compilations of chromosome numbers already available to botanists . It is the
hope of the Editorial Committee that a separate effort to compile a complete
file of documented chromosome numbers can be undertaken once the FNA Manua l
is published. and sooner if possible.

The FNA Data Bank also is designed so that in the long run it can include
comprehensive ecological character iz ations of the taxa. Fo r the immediate
purposes of the FNA Manual . however, the ecological data will be restricted
for practical reasons to characterizations of habitat preference and flowering
time. The contributor sho uld provide a brie f description of habitat and
flow ering time for each taxon .

As a f i nal no te on descrip t i on, it f;bould be emphasized that these require-

ments apply to all taxa at the level of fami.ly or belm.... Orders and higher
taxa Hill not be described. The treatmen ts are to be c omparable betHeen taxa
of different ranks as l ... ell as bett...een taxa o f the same r ank. Furthermore , each
taxon shoul d be described so as to encompass all of i ts subo rd ina te taxa . To
the maximum degree poss ibl e, the computer will be used to check whether
24 521

descriptions take into account the characters and character states of all
subordinate taxa.

MANUAL FORMAT EXERCISE

Write a dichotomous key to ten species of angiosperms . The species can be

from one to ten genera in one to ten families. Write complete descriptions
for two species and a diagnostic description for one of the two species .
Consult the format instructions .

DESCRIPTIVE FORMAT LITERATURE

Heywood, V. H. 1958 . The Presentation of Taxonomic Information-- A Short

Guide for Contributors to Flora " Europaea. Leicester University Press.
Leicester .
Radford, A. E., C. R. Bell, J. W. Hardin, and R. L. Wilbur. 1967. Contri-
butors' Guide for the Vascular Flora of the Southeastern United States.
Privately published . Chapel Hill .
Shetler. S. et al. 1973. Guide for Contributors to Flora North America (FNA)
(Provisional Edition). Smithsonian I nstitution. Washington, D. C.
522 25

Chapter 25. PLANT IDENTI FICATION

Ident ification i s a basic activity , and one of th e p rimary object ives

of systematics . Al though identif ication is a separate activity or process ,
in pract ice i t invo lves both classification and nomenc lature . I dent ification
is simply the determination of the si milarities or differe nces betwee n t\.JO
ele ment s . i.e . • two e l ements are the same or they are different. The compari-
son of a n unkn Olffi plan t with a named specimen and t he de termination that t he
t wo element s are the s ame a l so involves classification, i.e .• Hhen one co r-
rect l y decides that an unknown belongs to the same gro up (s peci es , gen us ,
f a mily, etc.) as a known specimen, the informati on stored in c l assif i cat ion
systems becomes availab l e and applicable to t he material at han d . Ro th proc-
esses--identificatic,n and c.Lassitication--inv olve cOliIDaris on and judgme nt and
require a defini tion o f cr ite ria of similariLies . Identification is , there-
f ore, a basic proce ss i n classification with n omenclature pl a ying an ess e ntjal
role in t he retrieva l of informa tion a nd as a means of conunun ication . Acc o r d-
ing to Blackwelder ( 1 967) "identification enable s us to ret r ieve the app r o-
priate facts from the system (c lassification) to be associa t ed with some
specime n at hand" an d is " . . . bett er def>cribed as the recovery side of taxon-
omy." In pract i ce on e common l y identifies a plant bv direct comparison or
the use of keys and arriv es a t a name . The practi ca l aspects and methods of
plant identification and iden ti fica t ion systems arc discu ssed in this chapter .
For f ur ther i nfo rmation see Harrin gton and Dur re ll' s book Hm" to Ident i fy
Pl ants .

Section A. TRADI TIONAL IlJENTIFICAT ION I'!ETHOIJS

The traditiona l netho ds of identific ation include (1) expert determina-

t i on , ( 2) r ecognition , (3) compa rison , and (4 ) the use of keys and similar
devices . For a thorough Bnd technical discussion of specimen identification
see Sneath and Sokal (1973).

In terms of rel iability or accuracy the best method of id entificat ion is

expert determination. In general the expert will have prepared treatments
(monographs, revisions, synop ses) of the gro up in question , and it is probable
that the more recen t flo r as or manuals include t he exper t' s concep t s of taxa .
Although of g reat r eliabili t y , t hi s me thod present s problems b y requiring the
valuable time of exper t s a nd c reatin~ delays for identification . Recogn ition ,
according to Ho rs e (1971) app r oaches expe rt determination in reliability .
This is based on extensive , past experience of the identif i e r with t he plant
group in question . In some groups this is virtually impOSSible . A third
method is by compar ison of an unknot.m. with named specimen s, photographs,
illustration s or descriptions. Even though this is a reliable method , it may
be very time consuming or vir tually impossible due to the l ack of suit able
materials for comparison . The reliability is , of course, dependent on t he
accuracy and authenticity of the specimens , illustrations , or descr i pt i ons
used in th e comparison. The use of keys or similar devices (synopses, out-
lines, etc.) is by fa r the most widely used method and does not req uire the
time , materia ls, or experience involved in comparison a nd reco gnition.

Keys in t he tradiLional sense are a type of taxonomic lit era ture (see
Figures 25-1 & 25-2) . Keys a re devices consisting of a se r ies of con trasting
o r contradictory statements o r propositions requirin g the identi fie r t o make
compar i sons and dec isions based on sta tements in the key as related to the
25 523

material to be identified. The first modern type keys (dichotomous) clearly

designed fo r identification ,,rere those of Lamarck in his Flore Fran;aise in
1778 (see Voss , 1952. for an interesting account of the history of keys and
phylogenetic trees in systema tic biology) . The comments of A. P . de Candolle
in his dedication to Lamarck in the third edition of this flora concerning
keys are equal l y appropriate toda y :

"As to the artif icial method , I have without hesitation , given

preference to the one vhich you have contrived, and \.;hi ch consists
of leading the student to the name of the plant by all.aY8 forci ng
him to choose betlveen two contradictory characters : in this ana-
lytic me thod I have permitted mysel f only the slight changes neces-
sitated by the increase in the number of plants described . There ,
after your example. I have sough t to distinguish the plants by the
easiest and most apparent characters; and \"hen these characters
were not constant, I. have tried to foresee their aberrations and
to arrive at the same name by differen t routes ; but this ease in
the distinguishin g of plants is very different- in different fam i-
lies: in some, such as t he c rucifers, it is impossible to distin-
g uish the genera wit-hou t examination of the f ruit. . . tJhen
be ginners undergo these d i fficulties in the use of the analyt-ic
method , I beg th em , before blaming it, to r ef l e ct that the most accom-
plished botanists meet ~.;ith th e same embarras s ment , and that 110
method can make the ;.Jork easiE'.r to studt::nts than it is to the
masters . . . . But \.;rhen the pupil kllOHS the name, let him take
care not to think he knm.;s the thing ! Reterred bv a number in
the anal y tic method to the desc ription, he wil1 find in this ,
second part th e details which p u t together constitu te the whole
science ." (Ouoted f rom trans la tion by Vcss. 1952)

Section B III of this chapter includes a discu ssion of the const ruction
of identification keys and the appli ca tion of computers to this process .

The followinj! J ists of suggestions are for the use and constr uct i on of
tr adi tional dichotomous keys .

1. Select appropriate keys for the materials to he identified . The keys may
be in a flora, manual, g uid e , handbook , monograph, or revis10n (see Chap-
ter 30) . If the locality of a n unknown olant is known. select a f l ora ,
guide , or manual treating the plants of that geog raphic area (see Gu i des
to Floras in Chapter 30) . If the family o r genus is recogni zed , one may
choose to use a monograph or revision . If locality i s unknown, select a
general work . If materials to be identified were cultivated, select one
of the manuals treating such plants since most floras do not include
cultivated plants unless naturalized.

2. Read the introd uctory comments on format details, abbreviations , etc .

before using the key .

3. Read both leads of a couplet before making a choice . r.ven though the Eirst
l ead may seem to describe the unknown rna terial , the second lead may be
even more appropri ate .
 524 25

4. Use a glossary to check the meaning of terms you do not understand .

5. Measure several similar structures \"hen measurements are used in the key,
e . g . , measure several l eaves not a single leaf. Do not base your deci-
sions on a single observation. It is often desirable to examine several
specimens .

6. Try both choices when dichotomies are not clear or when information is
insufficient, and make a decision as to which of the two answers best
fits the descript ions.

7. Verify your resu l ts by reading a description , comparing the specimen with

an illustration or an authentically named herbarium specimen.

Suggestions for Construction of Keys

1. Identify all groups to be included in a key .

2. Prepare a description of each taxon (see Chapter 24 for details for

description and descriptive format) .

3. Select "key characters" ,"ith contrasting character states . Use macro-

scopic, morphological characters and cons t ant character states when
possible. Avoid characteristics that can only be seen in the fie ld or
on specially prepared specimens, i.e ., use those characteristics that
are generally available to the user .

4. Prepare a Comparison Chart (see Figure 25-3).

5. Construct strictly dichotomous keys .

6. Use paral l el construction and comparative terminology in each lead of a

couplet.

7. Use at least two characters per lead when possible .

8. Follow key format (indented or bracketed; see Figures 25- 1 and 25 - 2).

9. Start both leads of a couplet with the same word if at all possible and
successive leads with different words.

10 . Mention the name of the plant part before descriptive phrases , e . g .,

leaves alternate not alternate leaves or flowers blue not blue flowe r s.

11. Place those groups with numero us variable character states in a key several
times when necessary .

12. Construct separate keys for dioec i ous plants, for flowering or fruiting
materials and fo r vegetative materials when pertinent.
25 525

A DICJ-IOTONOUS KEY TO SELECTED GENERA OF SAXIFRAGACEAE

Shrub or woody vine .

Haady vine; petals 7 o r more . . .... . . . . . .... ... .. .... ... 3. Decumar ia
Shrub; pe t als 4 o r S .
Leaves alternate or on short s pur branches .
Leaves pinnately veine d; ovary superior;
fruit a capsule ........................ ••• • .. . l. Itea
Leaves pa l mately veined ; ovary inferior;
fruit a berry .........................• ... .... 2. Ribes
Leaves opposite .
Petals us ually 4; stamen s 20-40; fruit
longitudinal l y deh i scent. not r ibbed . .. ..• • . . . 4. Philade l phus
Petal s usually 5; stamens 8-10 ; fruit pori-
cidal l y dehiscen t , 10- to I S-ribbed . .. .... • .. . 5. Hyd r angea
Herbs .
Staminodia pres ent ; petals more than 10 mm l ong . . . . •... 6. Parnassia
Stamino dia absent; petals less than 10 mm long .
Leaves t ernately decompound .... .... .............. •... 7. Astilbe
Leaves simp l e .
Flm"ers solitary in l eaf axils, or in shor t ,
leafy cymes .
Sepals 4; carpel s 2 . ..... .... .... ............•. .. 8. Chrysosplenium
Sepa l s 5; carpe l s 3 ...... . .. . ..... ...........•.. . 9. Lepuropetalon
Flowers in racemes or pan i c l es .
Pet als pinnatifid or f r inged; s tem leaves
opposite ....... ....................... ..... . 10. Hitella
Petals not pinna tifid or fringed ; stem leaves
alternate or absent.
Ovary I-celled .
Inflorescence paniculate ; stamens 5 ....••. . . . 11. Heuchera
Inflor escence racemose; stamens 10 .......... . 12. Tiarella
Ovary 2-celled .
Stamens 5; leaves pa l mately l obed .. .. ...... .. 1 3. Boykinia
Stamens 10 ; leaves not palmately lobed ... .... 14 . Saxifraga

Figu re 25-1. Examp le of an indent ed (yoked) key . [From Radford , A. E.,

II . E. Ahles , and C. R. Bell . 1968 . Hanual of the Vascula r Flora of the
Caro linas. University of Nor t h Carolina Press . Chapel Hill , North Carolina .
Used with permission . J
 526 25

A DICHOTOHOUS KEY TO SELECTED GENERA OF SAXIFRAGACEAE

1. Shrub or woody vine ..................•••• • ••• •• .•••••••....•......... 2 .

1. Herbs . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . •• • •• •• •• •..................... 6.
2. Hoady vine; petals 7 or more ........ ... ... . .. ... .......... Decumaria.
2. Shrub ; petals 4 or 5 ................................. . ......... . . 3 .
3. Leaves alternat e or on short spur branches .. ..................... . ... 4.
3. Leaves opposite .. . ....... .... .. ....... .......... . .................... 5.
4. Leaves pinnately veined; ovary superior ; f ruit a capsule ...... I tea .
4. Leaves palmately veined ; ovary infe rior; fruit a berry ....... Rihes .
5. Pe tals usually 4 ; stamens 20-40; fruit longitudinally -----
dehiscent , not ribbed .... .......... ................... Philadelphus .
5. Petals usually 5 ; stamens 8-10 ; fru it poricidally dehis-
cent, 10-15 ribbed ....................................... Hydrangea .
6. Staminodia present ; petals more than 10 mm long .......... Parnassia.
6. Staminodia absent; petals less than 10 mm long ................... 7.
7. Leaves ternately decompound ..... ..... . ...... ................... Astilbe .
7. Leaves simple ........ . ...................... . .. . ...... ... .. . ......... 8 .
8. Flowers solitary in l eafaxils , or in short, leafy cymes ......... 9 .
8. Flol<Jers in racemes or panicles ................................ . . 10.
9. Sepals 4; carpels 2 ... .......... . ......... . ............. Chrysosplenium .
9. Sepals 5; carpels 3 ...................................... Lep uropetalon.
10. Petals pinnatifid or fringed; stem leaves opposite ........ Hitella .
10. Petals not pinnat i f id or fringed ; stem leaves alternate
or absen t ................................................. . 11.
11 . Ovary I-celled ..... .. ............................................... 12.
11. Ovary 2-c e l led .. . ................................................... 13.
12 . I n flores cence pan ic u1ate; stamens 5 ... ...... . . ... . . . . .. .. Heuchera.
1 2 . Inflor es cence racemose; stamens 10 ..... .. ............ . ... Tiarella .
13. Stamens 5; leaves palmately lobed .. . . .. . . . .. ......... ......... Boykinia.
13 . Stame ns 10; leaves not palmate l y lobed ....... ... ...... .. .... . Saxif raga.

Figure 25-2. Example of a b r acketed key . [Hodified from Radford , A. E.,

H. E. Ahles, and C. R. Bell. 1968 . Hanual of the Vascular Flora of the
Carolinas. University of North Carolina Pre ss. Chapel Hill , North Carolina.
Used with permission.]

PLANT IDENTIFICATION EXERCISE

1. Identification o f an unknmffi. Select an unknmffi specimen and identify it

by keying in an appropriate manual, flo r a , or monograph . Verify your
results by readin g a description, by comparing with an illustration or by
checking \"ith your instructor .

2. Preparation of a comparison chart . Select 5 or more specimens from t he

group provided by your instructor. Identify each by keying . Veri fy your
results . Prepare a de scription of each simil ar to those in a flora or
manual. Be sure characters and character states are in the same order.
Select contrasting character states and prepa re a comparison chart (see
Figure 25-3) .

3. Construction of keys . Construct a dichotomous key to these specimens using

the information in the comparison chart .
 N
~

COMP ARISON CHART

Decumaria Itea Ribes Parnassia Heuchera Sax if raga

Habit Haady vine Shrub Shrub Herb Herb Herb

Leaf
arrangement Opposite Alternate Alternate Basal Basal Basal
or on spur (Rosulate) (Rosulate) (Rosulate)
shoots

Petal
Number 7-10 5 5 5 5 5

Locule
Number 7-10 2 1 1 1 2

Stamen
Number 7+ 5 5 5 5 10
(staminodia 5)

Fruit Type Capsule Capsule Berry Capsule Capsule Capsule

Figure 25-3. A comparison chart used in the construction of key s (for six of the genera in Figures 25-1
and 25- 2) .

~
N
~

,
528 25

Section B. RECENT AND NE1~ IDENTIFICATION METHODS*

1. POLYCLAVES

Po ly c l aves of various kinds allow one to select the characterist ics for
use in identifying each spec imen , taking his choices from some charac ter set
and repeating an elimination process until a t e ntative identification is made.
A printed data table, chart , or matrix giving the stat us of various taxa for
useful characteristics is readily used as a poly clave by listing the possible
taxa on sc r atch paper and crossing out those which do not agree with the speci-
men ' s characters . Such data tables appear irregularly in t he taxonomic lit-
erature, often for only the more difficult groups involved but occasionally
for all the treated taxa , as done for medical bacteria by Cowan and Steel
(196 5) . For large groups , the diagnostic tables are not only more powerful
than the equivalent key, but also take less space to print. Lists of taxa
having various characters were among the first non tabular poly clav es . These
resemble the inverted fi l es common in compu t erized i nformation systems, where
entries are li sted according to their cha r acteristics , rather than charac t er-
istics by entr ie s . Lists of taxa lacking specified feat ures have also been
produced; this modification expedites use as one may then jot down the possi-
ble taxa and rapidly cross off those dif fer ing from the specimen. Polyclaves
are readily mechanized, as shown by the familiar edge-punched cards and the
less familiar t.,rindow keys, as well as various mechanical devices. The possi-
bility of a computerized po l yc la ve t..as noted by Sokal and Sneath (1966) and
by I.Ji U iams (1967); implementation is st raight foI'\.,rard once suitable data
formats have b een devised. The computerized polyclave system we developed
at Hichigan State University (Norse, in press) uses a General Electric Hark II
timesharing system, hut has also been tested on several other computers.
Other computerized polyclaves inc lude those of Boughey, Bridges, and Ikeda
(1968), Dybowski and Franklin (1968), Goodall (1968), and li'alker et a 1. (1968).
As yet no comparison of these many parallel approaches has been made.

If an identif ication method requires, in general . use of all the charac-

ters in some list, it i~ no longe r a polyclave, for no user options remain in
selecting characters. Several such charact er- set methods have been devised ,
mostly statistical . Some computerized "keys" are also based on this model
(e.g . , Gyllenberg, 1965; Bogdanescu and Racotta, 1967; and Rypka , 1971) .
Although we ll intended, characte r- set algori thms seem to offer no advantages
over polyclaves . However, pattern recognition methods , using no formal char-
acters, might be employed effectively fo r identification in some cases. Such
techniques linked Hith optical scanners , spectroscopy, or chromatography could
even offer ful ly automated identification.

II . TAXA , CHARACTERS, AND DATA t-1ATRICES

An objective identification system requires ~ priori information of t hree

kinds: the pertinent taxa , the useful differentiating cha racte r s of these
taxa, and the taxon/character data themselves .

*Adapted from " Specimen Identification and Key Construction t.,ith Time-Sharing
Computers" by Larry E. Morse (Harvard University, Camhridge , Nassachusetts).
in Taxon 29: 269-282 (1971) , with extensive revisions by Hr . Horse. Used
td th permission.
25 529

The hierarchy of natural taxa is the foundation of biological informa-

tion retrieval as well as systematic synthesis, since data can be stored,
retrieved, and studied at any level of generalization (Horse, 1974) . In data
processing, \ve may usually regard a taxon 3S a group of one or more individ-
uals or lower taxa judged sufficiently similar to each other to be treated
together f ormally as a single evolutionary or informationa l un it at a parti-
cular level in the taxonomic hierarchy, and sufficiently d i fferent from other
such groups of the same rank to be treated separately from them . Tradition-
ally each taxon (except the highest) belongs to one an d only one taxon of the
next higher rank, implying each individual belongs to exactly one species (and
has one name) in any particular taxonomic treatment of its group . Taxa are
either monothetic or polvthetic . For monothetic taxa. possession of a certain
set of diagnostic characters is both necessary and sufficient for membership.
Pclythetic taxa are more loosely circumscribed, since presence of only "a
large number" (rather than all) of a l ist of characters is required for mem-
bership. I n other Hords, the members of .1l polythetic taxon exhibit overall
similarity. Hodern systematists employ the polythetic taxon concept in most
of their Hark, as has b een \"idely recognized in the past decade. However,
dichotomous keys and other monothetic methods of identification are still
used. Theoretically, identification schemes for modern biology ought to
assure that no single-character difference, either in population variability
or observer error, can result in a misidentification . Polythetic polyclaves
offer a solution; here , no possibility is e liminated until several dif fe rences
have accumulated be tt"een the taxon description and the unknown specimen . The
appropriate threshold depends on the variability of the taxa, but toleration
of just one or two differences often gives a marked improvement in identifi-
cation success. ,
Our character model was originally developed for a list - structure repre-
sentation of dichotomous keys (Norse, Beaman, and Shetler, 1968). It centers
on the concept of character couplets (or two - state characters), and also
allows coding of "dependent characters" (Hilliams, 1969). In our model, each
character couplet represents two contrasting alternatives about possible
features of an individual population . (For convenience we often label the
possibilities A and B, or true and false.) In other words, the character
couplets are equivalent to the pairs (couplets) of contrasting leads in
dichotomous keys, specifying a part of the organism and asking Hhich of two
alternative modifications of i t is present. Such a question implies the
specified part itself is present--the couplet "petals red" versus "petals
yellO\."." is mean i ngless for apetalous plants as well as those having petals
of different colors. A couplet can thus be inapplicable or noncomparable,
often coded NC in numerical-taxonomy data . An additional code is needed
Hhen a specimen is variable in its expression of the character states of a
couplet; more precisely. variable means sometimes A and sometimes B. At
other times the state of a couplet is not known for a specimen, perhaps
because the character is difficult to determine or not recorded, or the
specimen is incomp let e, poorly preserved, or at the wrong stage of develop -
ment; these cases are coded unknown. All binary-couplet characters can be
readily encoded with these five cha rac ter states , listed belot" with our
code numbers for them :

o unknown 3 false, or B
1 true, or A 4 = inapplicable
2 variable
530 25

Each character couplet in our model is a hierarchy of two binary (tHO-

state ) characters, one implicit and one explicit. Expression of either state
of the explicit char.acter. depends on the presence of one particular state of
the implied more general character--petal color depends on petal presence.
As \,'ell illustrated in a diagram by Dale (1968), larger character hierarchies
can be built by combination of a number of such couplets .

Extensions to multi-state and quantitative characters are, of course ,

necessary before our methods can be seriously considered fo r handling taxo -
nomic information in general. For this He implemented a f ile system involving
three kinds of taxon/character data : two-state characters, as described
a bove; mu lti-state characters, and quantitative (numeric) characters (Horse,
Peters, and Hamel , 1971). Hulti-state and quantitative characters may also
be coded as couplets for use with the current programs . The former can ahlays
be expressed as a list of yes/no binary characters, while the ranges of quan-
titative characters may be segmented as appropriate . Details of the !'!SU pro-
grams are available e lseHhe r e (Horse , in press).

For identification and allied procedures, the preparation of data matrices

as taxon/character rather than specimen/character entries is preferable . In
compiling effective taxon/character descriptions it is of cours e important to
st u dy an adequate sampling of specimens and other information, including the
relevant literature, as otherwis e the data cannot ind icate the nature of the
taxon as a "'hole . Yet if enough examples of a taxon I.ere examined, virtually
all the characters studi ed ,wuld eventually be marked variable, rendering the
whole description useless . Hmlever, \~ith polythetic identification methods
the occasional oddity can be safely ignored in preparing a taxon description.
For polythetic taxa, tbe code variable means noticeably variable, Hhile A
(or B) means usually A (or B), perhaps 80% (CO\"an and Steel, 1965) or 85%
(Lapage et a1., 1973), or a number calculated from the data (Ho11er, 1962) .
In our present system, taxon/character data are presented in the format
illustrated e1set~here (Hor se , in press; She tler ct a1., 1971). Rriefly, such
files consist of a taxon list, a cha racter list, and a coded taxon/character
data matrix, \~ith additional fields for various parameters used by the pro-
grams . AllOlvance is also made for extensive documentation .

Hierarchical matrices aid in processing of taxonomic data since different

characters may be used fo r different subgroups of taxa. In a hierarchical
taxonomic matrix system, each taxon-description line may refer to another com-
plete matrix differentiating subordinate taxa, permitting programs to work
dOl"n through the hierarchy much as one uses a key to families, then a key to
genera , and then a key to species. This method also a llOl']s the user to query
the data bank beginning at any level he desires--if he knOl"S his specimen is
a Rhododendron , he need not first verify that it is an angiosperm !

In preparing identification data, one should include only characters of

potential value in making identifications . A number of computer algorithms
are available for character-set selection, but subjective screening appears
ordinarily adequate for choosing a manageable set of characters for further
study . The traditional key characters of the group offer an initial list,
but net-! diagnostic characters should be sought, especially in difficult groups .
Guidel i nes for character selection , offered in a multitude of ,"orks, include
the following : presence in the usual mate rial to be identified, ease of
observation and interpretation, distinctness hetHeen the variolls states ,
independence from other characters used , and tolerance to environmental
25 531

influences . Identification characters must not only be ,,,ell expressed in

nature but shou l d also be clearly expressible verbally or pictorially . Occa-
sionally one should describe \-fhat the user \~ill think he is seeing, ,,,hether
o r not this be the actual case morphological l y . The previous experience of
the intended users should also be conSidered , and especially in educational
I-Jorks the characters might be easily leanled and remembered in as s ociation '"1th
t hei r tD.xa.

III. CONSTRUCTION OF IDENTIFICATION KEYS

The presentation of dichotomous keys is usually considered a mandato r y

part of scholarly taxonomic publication , yet until recent years few advances
,,,er e made in key-constructing theory over the methods of Grew· or Lamarck ,
or even Aristotle and Th cophrastus (Voss , 1952) . I n his study of the theory
of keys. Osborne (1963) conc l udes that the most efficient key is the one in
\"hich each dichotomy divides the remaining taxa symmetrically , or as nearly
so as possible , assuming the characters have equal probabilities of mis i nter-
pr etation and the taxa are eq ually abundan t. However , i n r eality some c harac-
ters are far easier to use than others, and some taxa are keyed out more often .

Th e relative conveniences or ease of use of the var i ous characte r s should

be a major consideration in key construction , as one wants the easiest and
fastest ke y \"hich is sufficiently accurate . Ledley and Lusted (1960) suggest
the characters (disease symptoms in their paper) be grouped into numbered sets
or blocks of similar difficulty . In making a diagnosis they employ a l l a vail-
able characters from one block before using any from the next. life a1101" nine
such blocks, \-lith the characters i n block one hardest and those in block nine
easiest; any cha r acters labelled zero are comple tely ignored by some routines .
In our programs \"e treat the bl ock codes, or character convenience values, as
na t ural logarithms of a convenience- of- use number . Thus the nine symbols
r e pr esent a range of about three to eight thousand. IJhcn rating characters .
on e should assign codes such that the intended users would prefer to st udy
any three characters at some level than any s i ngle character at the next
lOt"er level . seven to one for a two- level difference , t\~en t y to one for three
levels. and so forth . The conveniences may , of course , be revi s ed with exper-
i ence or a change in user groups . Hal l (1970) cons iders conveniences ,,,hich
var y from taxon to taxon, but finds it easier to rate each character only
once , instead of once per taxon .

If the re l ative frequencies of the various taxa can be estimated for a

sitUation, t h en the taxa can be considered a ccordingly i n determining the
evenness of division at a dicho t omy , in effect taking each expected occur-
rence as a sepa r a t e object to be identified . Determining such frequencies
is not at all easy , since \~e a re interested in how often a taxon will be
collected and keyed out, a quite different value than how often it occurs
in t he field .

The intra-taxon variabi l ity of the characters is also an important con-

sideration in keys . In our methods , characters are coded as either variable
or as (nearly) constant, leavin g selection of a variability threshol d to the
researcher preparing t he data . An alternative approach often used involves
stating the variability of each taxon for each character as a quantitative
e nt ry in the data matrix , us ually as a percent age . This percentage mat rix
woul d be valuable in a general t axonomic information sys t e m, especially
532 25

whil e data were being collected and variability patterns were developing .
However, the true-false-variable matrices present their data more clearly ,
and take far le ss effort to prepare . Also , we know no programs which utilize
the percentages directly in constructing keys .

Hany texts reconunend the use of data tables in \olriting keys , but other
mechanica l aid s are rarely described . }letcalf (1954) presents an index-card
technique, and Peter s (1969) used a computer to help incorporate additional
taxa in keys . t!hi le develop i ng our key-constructing techn iques we prepared
a key-editing program (}Iorse , Beaman , and She tler. 1968). but work on the
"advanced system" outlined there was suspended in favor of our current
research on k ey cons truction and identif ication procedures. Key editing is
important in lar ge projects such as Flora Europaea and the planned Flora
North America , where nume rous minor improvements are made in the keys during
editing. Hm"ever , if taxa are added or deleted , it is often better to de-
velop an entirely new key .

The possibility of comp uter ized key const ru c tio n is often mentioned , yet
\.'e know of only four programs for cons tru ction of biological keys, name l y
those of Holler (1962) , Hall (1970), and Pankhurst (1970, 1971) , as well as
our own (Norse , 1971 and in press). Pankhurst (1974) p r ovides a comparison
o f some of these programs . Noller 's method requires complete b i nary data,
and has attracted litt le attention . Hall ' s program utilizes quantitative
data , printing a numeric version of the key which mus t then be rewritten be-
fore use. Pankhurst's algor ithm differs from ours primarily in his use of
rigi d charac t er- convenien ce blocks and his employment of the attribute-
value rather than hierarchical-couplet character concept . His program, like
ours , prints the key directly . The production version of our HSU programs
is described below; these a llaH mixed -data key construction with our new
data matrices . Several matr ix-reduct ion and monothetic-devisive algorithms
i n the l iterature resemble key const ruction : the KEYCALC program by Niemala,
Hopkins , and Quadli ng (1968) is typica l. Also , some aspects of decision- tree
and game-tree research in computer science could contribute to the theory of
keys.

In preparing a key , one usuall y divides the initial group of taxa by a

ch aract er couplet into two subgroups , each of Hhich is independently divided
into further subgroups , and so forth, until every taxon is distinguished from
all o t hers . Indeed , the further subdivision of a subgroup can be consi dered
as construction of a full key to that l ocal group , s ugges ting a concise recur-
s ive algorithm for computerization: construct the key E.Y. div iding the original
taxa into tHO subgro up s E.Y. the best possible character couplet , then consider
each of these subg r oups separately, dividing them simila rly. It is readily
established that this procedure Idll produce a k ey if one can be made at all,
but the optimality of such a key appa rently remains uninvestigated . The four
key - construction programs cited above all opera te on thi s principle .

For use of this algorithm one needs a means for selecting the "bes t"
character couplet for a given dichotomy, given the appropriate taxon/charact e r
data for a set of potentially u sef ul characters. In our program , a pre liminary
screening of the characte rs is followed by a detailed evaluation of the poten-
tia lly u5eful ones . In the preliminary scan , a cha racter is eliminated from
further consideration if it is coded as unknown or as inapplicable for any
taxon in the loc al group, or i f it is coded in only one Hay Idthin that group .
In other words, the remaining characters a r e all coded partly ~ and partly
false; true and variable; fa lse and variable; or~ , false, and va riable .
For keys these are obvious l y the only characters of immediate interest. Next ,
a combined measure of ease of use and dichotomizing power is determined for
each character. A tal ly is made for each state (true , false, and variab l e ),
givin g the sums of the taxon frequencies of the taxa so coded for a character.
In effect. these three tallies give the total number of encounters expected
unde r each state of a character. The tally for var i able is then multipli ed
by the sum of the other two tallies, and half this produce is added to tt,lice
the product of the ~ a n d false tallies, giving a "d ichotomizing value . "

whe re qT , qF. and rtv are the respective tallies . High values of DV indicate
well-divided dichotomies . In considering the character conveniences , these
ratings are treated in exponential (values 3 through 8103) rather than con-
densed (1-9) form . The "b est " couplet is found by maximizing the produc t of
the character- convenience expon ential and the dichotomizing value . The char-
acter couple t having this product greatest is then used for that dichotomy of
the key , and the taxa in t he local group are sorted according to their recorded
status for that character, placing variable taxa in both subgroups . Note the
charac ter conveniences have a powerful but not exclusive effect , as a 5-5
split at conve n ience 6 (7254) as well as a 6-4 (or worse) split at 5 (7004) .

If the indented style is desired, the program prints the key as it selects
the dicho t omies. However, for the bracketed style the key must be stored (in
our list-structure representation) until completed , since division of variable
taxa af fe cts the numbering . It is then printed in one operation . The l is t -
structure condensation can also be printed if desired , perhaps for late r us e
in editing . ,

Ordinarily a batch-processing computer is adequate for key construct ion

since no user interaction during processing is necessary. HOI,lever. on-line
key construction allows the effects of changes in the data to be seen quickly .
An on- line key-editing sys tem Hould also be possible , b u t most changes can be
made with the text - editing softl,lare usually supplied with time-sharing sys-
tems . For small groups , these methods may also be used manually .

IV . DYNANIC POLYCLAVES

Polyclaves n eed not be automated , but might be computerized when a lar ge,
rapidly developing information base is involved; otherwise , imaginative pub-
lication methods suffice (e . g . , Ogden , 1953 ; Leenhout s , 1966, Archbald , 1967;
Duke , 1969 ; Han sen and Rahn , 1969; Shultz , 1973). Here on-line (usually time-
sharing ) computers offe r a clear advantage over batch-p r ocessing systems, sin ce
the on -lin e system allows the us er to submit additional data during the execu-
tion of a program. This permits a dialogue or conversati on between the us er
and t he computer, l1ith the machine pri nting questions and a ....'aiting responses
before continuing the processing . Such conversations can be about as terse
or ver bose as desired; we have t aken a middle course, Hriting out most questions
and taxon names in full. but sometimes usin g nume r ical codes for charac t er
states and coup l ets , as well as for several program op t ions . Numeric coding of
lengthy ansl"ers saves prin ting or typing t i me and r educes the chances fo r
errors , assuming the user can copy a short number more accurately than a long
descriptive phrase. For regular us ers of the system a faster abbrev i a ted
terminology is planned . As yet, the idea l of free - form language input is not
prac tical for this or almost any information-retrieval system (Simmons . 1970 ) .
 534 25

Following selection of a group of possible taxa by t he choice of the

in itial da t a f i le. our polyclave algorithm consists of th r ee steps , rep eated
as necessary: ( 1 ) request the user to give one or more charac teristics of
his specimen; (2) e limin ate all pos s ibilities incon sistent with this partial
description; and (3) print the res ults of the elimination , either an i denti-
f i cation or some oth er action , and recycle to the first step if n eces sary .
Several a l ternatives are available, including production of a useful-charac-
t e r s lis t and deletion of the effects of the last charac ter set submitted .
Si n ce a user may start with a d ata matrix at any l evel i n the taxonomic h ier-
a r chy , continuation to a subsequent matrix may be possible after a family,
genus , or other higher-level taxon is identified. After each identification ,
sever al diagnos t ic or peculiar characters of t he taxon are listed as an
immediate check of the suggested identification . Particularly with higher-
l evel matrices , " false posi t ives" may occasionally occur when no single spe-
cies has a cha r acter combination suggested by the generali zed description of
a taxon .

Poly the tic identification , \,Ihere one difference no l onger implies elimi-
na t ion , is avai l able as a n opt i on on our computer sys tem . Here the program
tallies the number of differences and eliminates a taxon only when its tally
e xceeds a use r -determined val ue , commonly one , two, or three . Although it
allows for greater taxon va riability or user error, the polythetic po l yclave
is slower than the monothe t ic one since more characters must be submitted to
assure comple te elimination of all the other taxa .

When a l is t of use fu l characteristics is desired part way th rough a poly-

clave procedure , a s ubrou t ine selects the best characters to continue divi ding
the set of remaining possibilities . Actual l y, a portion of the key-construct-
, ing algorithm is used to determine the first such character , and repetition
of the procedur e (ignorin g this character) giv es the next- best character , and
s o forth . l.,rith the polyclave , of course . such r ecommended characters are
merely suggestions which need not be employed . No matter which state the
specimen shows , the recommended c haracter will eliminate about half the pos-
sibilities . Any other cha racter, on the average , wi ll delete fewe r taxa
because it el i minates a larger number on l y \,Ihen i n its rarer s tate . However,
if the user happens to notice his specimen displays a rare character , he can
e l iminate a large number of possibilities at on ce and identify hi s s p ecimen
much more rap i dly . Since t he ability to recogniz e rare characters and realize
their powe r comes only through training and exp e rience , the expert delights in
the efficiency and power of a polyclave. while the neophyte is lost in its
multitude of choices and prefers th e supervision a nd security of the t r adi-
tional dichotomous key .

PLANT IDENTIFICATI ON LITERATURE

Archbald , D. 1967. Quick- Key Guide to Trees: Trees of Northeas tern and
Centra! North America. Doubleday . Ga r den Ci ty, New York.
Bl ackwelder, R. E. 1967 . Taxon omy , A Text and Reference Book. John l-'fley &
Sons , Inc. New York .
Bogdanescu , V., & R. Racotta . 1967. Identification of mycobacteria by overall
similarity analysis . Journal of General Microbiology 48 : 1!1-l26 .
Bossert, H. 19 69 . Computer t echniques in systematics . I n: Systematic Biol-
ogy . National Academy of Science Publication 1692 . Hashington , D. C.
Bou ghey , A. S . , 1( . W. Bridges. and A. G. Ikeda . 1968 . An automated biological
identification key. Nuseum of Systema t i c Biology . University of California
(Irvine) .
25 535

Co\... an, S . T ., and K. J. Steel. 1965 . Nanual for the Identification of Hedi-
cal Bacteria . Cambridge University Press . Cambridge, England.
Dale, M. B. 1968. On property structure , numerical taxonomy , and data hand-
ling . In : HeY"lOod, V. H. Hadern Hethods in Plant Taxonomy . Academi c
Press . London and New York .
Duke, J. A. 1969 . On t r opical tree seedlings . I . Seeds , seedlings , systems ,
an d systematics . Annal s of t he Hissouri Bo t anical Ga r den 56: 125-16l.
Dybowski, I.J., and O. A. Franklin . 1968. Conditional probab ility and t he
identification of bacteria : a pilot study . Journal of General N1crobiology
54: 215- 229 .
Goodall, D. H. 1968. Identif i cation by computer . BioScience 18 : 485-488.
Gyllenberg . II. 1965 . A model f or computer identification of micro-organisms.
Journal of General Microbiology 39: 401-405 .
Hal l , A. V. 1970 . A comp uter-based system for forming identification keys .
Taxon 19: 12-18 .
Hans en, B., and K. Rahn . 1969 . Determination of angiosperm families by means
of a punched-card system . Dansk Botanisk Arkiv 26 : 1- 46 + 172 punched
cards .
Harrington, H. D. , and L. W. Durrell. 1957. How to Ident i fy Plants . Swa ll ow
Press. Chi cago .
Lapage . S . P . , S. Bascomb , \L R. Hillcox. and M. A. Curtis . 1973. Iden t ifi-
cation of bacteria by computer: genera l aspects and perspectives . Journal
of General Microbiology 77 : 273- 290 .
Ledley, R. S., and L. B. Luste d. 1960 . The use of electronic compute rs in med-
ical data processing: aids in diagnosis , current info rmat ion retrieval , and
medical re co rd keeping. Institute of Radio Engineers Trans actions on Medi-
cal El ectronics 7 : 31-47 .
Leenho ut s. P . W. 1966 . Keys in biology : a s urvey and a proposal of a new ,
kind . Pr oceedings KOllinklijke Neder l andse Akademie Van I-Ietenschappen 69
(ser. C) : 571- 596.
Metcalf, Z. P. 1954. rhe construction of keys. Systematic Zoology 3 : 38-45 .
Moller , F . 1962. Quantita tive methods i n the systematics of Actinomycetales.
IV . The theory and applicat ion of a probabilistic ident ification key.
Giornale di Micr obiologia 10: 29-47 .
Morse, L. E. 1974 . Comp ute r-assisted storage and retrieva l of the data of
t axonomy and systematics. Taxon 23. in press.
(in press) . Computer programs f or specimen identification, key
construction, and descrip tion printing using taxonomic data matrices . Pub-
lic ations of the Museum, Hichigan State Un iversity (Biological Series) .
East Lansing, Mi chigan .
_-,;-:-:::-" .' J. H. Beaman , and S . G. Shetler . 1968 . A computer system fo r edit-
ing diagnost ic keys for Flora North Ame rica. Taxon 17 : 479-483 .
_-,==:-.' J. A. Peters, and P . B. Hamel. 1971. A general data format for
summarizing taxonomic information. BioSc i ence 21: 174-180 , 186.
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binary test battery for a set of microbial cultures . Canadian Journal of
Microbiology 14: 271- 279 .
Ogden , E. C. 1953. Key to the North American species of Potamogeton . New
York State Huseum Ci r cul ar 31.
Osborne . D. V. 1963 . Some aspects of the theory of dichotomous keys . New
Phytologist 62 : 144-160 .
Pankhurst, R. J . 1970 . A compu ter program for generating diagnostic keys .
Comp uter Journal 13: 145-151 .
1971. Botanical keys generated by computer. Hatsonia 8 : 357- 368.
 536 25

Pankhurst, R. J. 1974 . Automated identi fication in systematics . Taxon 23,

in press .
Peters, J. A. 1969 . Discussion . In: Systematic Biology. National Academy
of Science Publication 1692 . l~ashington, D. C.
and B. B. Collette. 1968 . The ro le of time-sharing computers in
museum research. Curator 11: 65-75 .
Rypka , E. loJ. 1971. Truth table classification and ident ification . Space
Life Sciences 3 : 135-156 .
Shetler, S . G., J . Il . Beaman, M. E. Hale, L . E. Morse, J . J . Crockett , and
R. A. Creighton . 1971. Pilot data processing systems for flor istic
information . In : J . L . Cutbill . Advances in Data Processing for Biology
and Geology. Academic Press. London and New York.
Shultz , L. M. 1973 . Random-access key to the genera of Colorado wildflowers .
University of Colorado Museum . Boulder, Colorado .
Simmons, R. F. 1970 . Natural language question-answering systems : 1969 .
Communications of the Assoc iation for Computing Hachinery 13: 15-30 .
Sneath , P. H. A., and R. R. Sokal. 1973 . Numeri cal Tax onomy : The Princi-
pl es and Practice o f Numerical Classification . H. H. Freeman and Company.
San Francisco.
Sokal, R. R. , and P . H. A. Sneath . 1966. Efficiency in taxonomy . Taxon 15 :
1-21 .
Voss, E. G. 1952 . The history of keys and phylogenetic trees in systematic
biology . Journal of the Scientific Labo ratories of Denison Unive rs ity 43 :
1-25.
Halker , D. , P. :t·.filne , J. Guppy, and J. t.Jilliams . 19 68. The computer- assisted
st orage and retrieval of po l len morphological data . Pollen et Spores 10 :
251-262 .
Hilliams . H. T. 1967 . The computer botanist . Australian Journal of Science
, 29 : 266-271.
1969. The problem of attribute-weighting in numerical c l assifica-
tion . Taxon 18: 369-374.

POLYCLAVE EXERCISE

Preparation of a s imple polyclave . Select a family of ten or more genera.

Construct a master sheet listing an abbreviation or number for each genus and
print in red on a 5 x 8 card. This list is the taxa master list underlay . By
examining herbarium specimens or using an appropriate manual or monograph
select and compile a list of 50 or more characters and character states for
each genus . Duplicate (in black) the taxa master list under lay to equal the
number of characters selected. These will become character state overlay
cards . Label ea ch card idth a specific character state from th-= character
state list . Search each specimen or description for the character state in
question and punch out the generiC name or number listed on the card for each
genus having the character state in question. Punch a card for each character
state, e.g. , if one character state selected is leaves opposite , punch out the
names of all genera known to have opposite leaves. To use the polyclave select
a specimen identified as belonging to the family in question . Notice charac-
ter states and begin sorting through overlay cards until you find a card ~"ith
a characteristic that matches one of the cards you have prepared. Place the
character overlay card over the taxa master li st. Those names or numbers
appearing in red through the holes will represent all genera possessing the
character in question . Continue sorting and overlaying until all cards have
been used or until a single genus name or number appears .
26 537

Chapter 26. VARIATION A.IIlD EVOLUTIONARY RELATIONSHIPS

Evolutionary studies provide the essen tial concepts and core of informa-
tion necessary fo r the realistic evaluation of experimental and descriptive
data concerned with species orig i ns and relationships and , by extrapolation ,
the origin s and r elationships of higher ranks . Nore specifically , organic or
biological evolution i s respon sible for the var i ation and change in organ isms
through time . The operation of the basic evolutionary mechanisms--mutation ,
r ecombination , and selection--in different environments today accounts f or th e
cu r r e nt patterns of variation o f i n terest and import anc e to taxonomi sts .

Indivi dual organisms do not exist apart from their environment and their
ge netic history but are the resu lt of present and past inter actions bet\"een an
array of gen etic backgroun ds and a long seq uence of e nvironments . Thus dif-
fe r ent kinds of organisms are not, as once thought , the result of special crea-
tion but are knmm to be the product of different evolutionary lines which
themselves result from the basic interactions between heredity and e nvironment.
These interact ions have prod uced , through time , all of the vartations in bio-
logical form and fun ction found in the fossi l record and all o f the continuous
and di scontinuous variation pa ttern s which make possible the c urrent concepts
of taxonomic categories as Hell a s the ecological concep t of plant communi ties .

An awa r eness of biologica l variat ion and also an awareness of differen t

ki nds of organisms dates back i nto man ' s ear ly hi s tor y . Attempt s to account
for the differ ent kinds of o r gan isms , bi.olog ica lly or Scientif ically , cover ed
a century or mor e before the ans\"er--natural selection--Has independently dis-
covered but join tly p r oposed by DanJin and Hallace in 1 859 . Even though the ,
fact of universal variation was a key point in t he Darwin-lvallace hypothesis,
t h e genetic mechanisms of variation - -mutation and r ecomh ination--\"ere no t dis-
cover ed and understood for several more decades. It \"as mo r e than half a
century after Darwin that cyto logical and genetic data began to be generally
applied to taxonomic problems . The result \.as " eyto-taxonomy ." As the
mechanisms of selection became better understood through experimental work ,
as more was l ear ned about the evolu tionar y role of the population breeding
s t ruct ure , an d as more was learned as to the causes of na t ura l variation ,
taxon omy in much of t he ,,,or ld became concerned \"ith " spec i ation" a nd thus
became primarily a study of evolution--or "Biosystematics ." The r esults of
similar evolutionary studies by t he plant eco lo gis t s interest ed in variation ,
adaptations, and community interact ions , oft en found ready application in
taxonomy. Ind eed , the fina l use of application of informa tion from hiosys-
tematic studies ( i n either taxonomy or eco logy) may of ten be secondary in
importance t o t he further demons tration or el ucidation of evolutionary proc-
esses e it her in general or in the specific o rganism studied.

The broad taxonomic consequ ences of mor e detailed e voluti onary s tudies ,
and especially tho se which have used comp uter - ass isted comparisons of the dat a ,
have been a clear e r under standing of the kinds of rel ationship s that ex ist
be t ween and amona or ganisms and a mo r e p r eCise , and useful, set of definitions
associated with such terms as "relat ionship ," "a ffinity , " and " simi l ar ity . "
As pointed out by Sn eath and Sakal (1973) the unqualified \"ord "r elat ionship"
may mean , in our language , ei t her r elat i onshi p by descent (a truly phylogenetic
relationship) or any of one or more types of r elationships based on a common
chara c t er or s imila rity (a phenetic relationship) . A phenetiC relationship may
exist quite apart from any genetic relationship or , on the other hand, it may
538 26

be the result of genetic relationship . and thus also be cladistic- - the term
applied to genetic relationship s that express or r eflect recent commo n ances -
try . Cladistic re lationships , if plotted against a re lat ive time scale, are
also chronis tic relationships and the resu lting dendrogral'l that \"ould illu s-
trate su c h a situation is called a cla dogra m (s ee Table 22-3-C, p. 470) . The
relationship s usua l ly sought by evolutionary st udies are phyl et ic or phylo-
genetic , but the relationship s usually shOlm are , at b est, phene tic . 1I00"ever,
even \-then no genet i c relationship is implied , phenetic similarities (or
"aff i nit ies" or "re la tions hip s" ) may be the res ult of s i milar evolutionary
processes . For example, the evolutionary process o f convergence migh t pro-
duce strong phyletic similarities between genet ically unrelated organisms .

The experimental and analytical approaches necessa r y to sound evolution-

a r y r esearch involve variation studies in morphology , anatomy, phytochemistry ,
embryology and physiolo gy ; studies o f evolutionary me c hamisms involve genetics ,
cytology, and e cology . Because of this holistic nature of biosystematics,
facility \·Iith many of the met h ods , and an understanding of much o f the vocab-
ulary, treated in previo us chapters l.Jill be essenti al to the demonstration and
under s tand i ng of evolut i onary (or other) relatio nships between and Idthin
plant species . Special note should be made o f the material in Chapters 10 ,
1 1 and 12 (cy tology , geneti cs , and r e pro ductive h iology) , the section on adap-
tive fe atures i n Chapter 1 5 (ecology), and the material on data accumulation
and analysis i n Chapters 20 , 21 and 22.

A very high degree of specialization, a n d subsequent inte rre lationship ,

also exists within the broad field of evolu tio nary studies. Each of Darwin ' s
five generali z ed f eatures of organic evo lu ti on are today mtljo r specialized,
but h i ghly r eticulated , lines of evolutionary rese arch. The fo llow ing o ut line
attempts to re la te some of the p resen t evolutiona r y concepts , mechanisms , and
term i nol ogy to the five primary aspects of classi c al evolution . Fo r d efini-
tion of the terms see the Evo l ut ion Glossary at the e nd of this chapter and
also the glossaries for Chap ters II , 12 , and 15, all o f Hhich are indexed
under the ~eneral s ubject and terms index beginning on page 881. Fo r examples,
see pertinent items from the lis t s o f g e n eral references for t his , and
related , chapters. For the more f requently used methods of data accumu l at ion ,
analys is of variation, and methods of presenting the da ta , see Chapters 20 ,
21, 22 , and 23 .

I. VARIATION

Variation stud ies , and a careful a n alysis of t he information gathe r ed

(s ee Chapters 20-22) , are the first steps in any evolu t i onary ,..,r biosystematic
study . Once the type and magnitude o f var iation a r e knOloffi it is fa r more
likely that further specif i c Hork I·rill enabl e one to go beyond t he descriptive
phase and account for t he var. iation observed and thus learn something of th e
actual hiologic al relationsh ips involved .

A. Types of variation a n d variationy-attern~ .

1. By primar.y con t ro l.
a . genetic (an ess e ntial aspect of t he life c y cle , e . g . , reproductive
characters ; usually has a relatively 1 011 amount o f variab i l ity ,
e . g ., a 1011 coefficient of variability) .
h . envir.onmental ( a less essen t i al character in the life cycle , e . g . ,
l e a f lengths; usually has a relatively high amount of varia-
bility , e . g . , a high coefficient of va ria b ilit y ) .
26 539

2. Ry uniformity .
a. continuous or clinal (the cline might be altitudinal due to tem-
perature, or other e nviro nmental f actors but 'h'ould not permit a
clear separation of the series of populations into discrete
units or subcategories).
b. discontinuous (isolat ion and adaptations to different env i ronments
can produce discrete variation patterns Hith fe\" or no i nter-
mediate forms and thus easily separable, on one or mor e charac-
ters, into categories) .

3. By form .
a . monomorphic c. polymorph i c
b . dimorphic d. heteromorphic .

B. Sources of variation in natural populat ions (also see Chapter 11, Genetics) .
1. Mutation 2 . Recombi nation
a . gene mutation a . crossing over at meios i s
h. chromosomal mu tation b . random assortment of homo logs
deletion at meiosis
duplication c . out-crossinp, (limited or pan-
inversion mictic)
aneuploidy d. hybridization
polyploidy e. introgression
3 . Genetic interact ion
a . epistasis c . euheteros is
b. transgressive variat ion
;

c. L imi tat i on s on va r i a t i on in 11 il.\t,uu"a"lc. .JP"o"ppuill.l~a"t.li"Q"tl"'s •

1. Self-fertilization . 4. Fixation .
2 . Asexual reproduction (apomixis) . 5 . Genetic homeostasis.
3 . Stabilizing selection .

II. REPRODUCTION

Evolutionary survival depends more on the reproductive methods and repro-

ductive potential of the population than on individua l " fitness . " Further-
more, since different rep roductive methods have a direct influence on the
amount of genetic recomhination (and thus often on the total amount of popu-
lation or spe.cies variation) a thorough understanding of the reproductive
biologv of a species , or series of congeners, is essential to an understand-
ing of both the variation patterns and the relationships involved in any b i o-
systematic study (see Chapter 12) .

A. Sexual reproduction (high evolutionary potential) .

1. }lechanisms that favor or insure outcrossing or allogamy and thus maxi -
mum heterozygosity and variability (either actual or potential) .
a . plants monoecious e . heterostyly
b . plants dioecious f . self sterility
c. protandry g . chasmo gamy
d . protogyny

2. Nechanisms that favor or insure selfing or autogamy and thus maximum

homozygosity and genetic uniformity .
 540 26

a. self fercility c. flowers perfect

h. cleistogamy d. autogamy

B. Apomixis or asexual rep r oduction (lov evolutionary potential) ; he re there

is no recombination and maximum phenotypic unifo rmit y . variability is
pheno typic and under environmental control .
1. Vegetative . 2. Parthenogenetic .
a. r hizomes, stol ons , runners a. apospory
b . bulbUs b . apogamy
c. fragmentation ( " cuttings") c . advent itious eIilb r yony
d . layering d. agamospermy
( Note that very few plants are completel y rest ricted to a single reproductive
method: many violets, for examp le , have chasmo gamous flower s that may he out -
crossed, clei stogamous fl mvers that self pollinate. and rhizomes which provide
effective asexual reproduction . Also , even monoecious and dioecious plants
often produce a few perfect flowers tha t may be self-po llinated and many
" inbreeding species" sho'II" some degree of outcrossing . The more reproductive
pathways an or ganism has t he greater its l ong r ange evo l u t ionary potential
and also its short range probabi lity of survival.)

C. Isolating mechanisms (interspecific) .

1. Prezygo tic (isolat i on which prevents zygote fo rmation) .
a. geographical (the r esult of geographic sepa ration of the entit ies)
b . ecological (the r es ult of eco l ogical separa tion of the entities,
even if sympatr ic)
c . seasonal (phenological separation of the entities)
, d . indirect etholog ical (isolation caused by the animal pollinator
preference)
e. mechanical (structural diff erences , such as corolla tube size or
anther placement , that prevent crossing)
(Note: Excep t for geographical isolation the above forms of iso l at i on may
not be comp l ete and occasiona l hybrids may be formed ; it i s not uncommon for
t,...o species of plants to be isolated by a combination of two (or more) mech-
anisms.)
2 . Postzygotic (zygote form ed , but effect ive gene exchange between enti-
ties still preven ted) .
a . failure of embryo to develop
b. hybrid inviabi li ty or sterility
c. reduced fitness and reproductive potential in F2 and la t e r genera-
tions .

III. Co:-tPETITIQ}l k\lO SURVIVAL

Competition is a test of preadaptation and illustrates the often i ndirect

effect of natur al selection . Seed size , viability , a nd dispersal mechani sms ,
for example , can in no way be of any select ive advantage ~ the plan t t ha t
produced the seed ! But seed viability per se is only o ne factor in survival
as is shovn by the intricate physiological adaptations of the relatively l ong-
lived seeds of desert annuals that enable them to "meas ure" the rainfall and
germinate only after a given amount of rain has fallen in a short enough time
period to ins ure adequate soil moisture for a reproductive life span.

The seed dormancy in cockl ebur (Xanthium sp . ; A~te raceae) sho'II"s another
germination pattern presumab l y of adaptive value . Here the seeds of a single
annual plant contain different amounts of dormancy producing auxin wi th the
26 541

result that a £e\" seeds from this one plant germinate each year over a period
of years rather than all at the first oppor tunity . Every organism i.e; to some
degree in compe ti tion with every other organism in the area . It is in compe-
tition with other organisms of its DIm kind as \"e11 as with organisms b elong-
ing to other species . These organisms a r e in competition for e nergy (light
or food), water , raw mate rials, space , and all of the other elements of the
environment nec essary to su rvival and r ep roduction .

Some Evolutionary Results of Competition

l"Iithin a population Between species

Genetic coadapta tion Organism coadaptation

Self-thinning Competitive exc l usion
Decrease in var iabi l ity (stabili zing Incr ease in variability (niche sub-
selection) divis ion)
Himicry
Suc c ession
Su rvival of t he fittest Survival of the fi ttest

For an ou t line of the aspects of adaptation related to the biotic select ive
force s (competition) see V- D. below) .

IV . NATURAL SELECTI ON

Nat ural selection is t he "steering mechan i sm" of evolution . It may vary

i n type , kind . and i n tensity and may have varied r esul ts--from adap tation and
su r vival to ex tinction . Ar tif i cial selection . or the se l ection of spec ific
charact ers or sets of characters by man , is the basic tool of plant and anima l
breeding programs essential to modern agricul t ure , and is an excellent example
of " applied evo lut ion " from l"h1ch He can draw many valid conclusions conc ern -
ing evolution in nature .

Nat ura l selection, the evo lut ionary p rocess that results in v a r iation and
speciation. may be general l y c lassified either by resu lts . by se l ect ive agent
o r by i ntensity. as fo lIous :

1- By resu lt . 2. By agent . 3. By i nt ensi ty.

a. stabi lizing a . abiotic a. DaDJinian selection
b. directional b . hiotic b. catastrophic
c. disruptive

A. Types of Select i on Compared

1. Cent r ipet al. stabilizing or normalizing selection--the selection that occurs

in every normal popu lat ion eliminat ing extr eme variations. maintaining
the adaptation of the group somewhere near its optimum for the environ-
ment in which it lives .
2. Balancing se l ection--a type of stabiliz ing selection in Hhich t \"O or more
d iscon tinuous forms are maintained in t he same interbreeding popul ation
(balanced polymorphism) because each fo r m is of advantage to the popula-
tion part of the time . or in part of th e environmenta l circumstances
under I"hich the populat i on lives . s u ch as seasonal flu ctuations in
temperature .
542 26

3. Centrifugal. disruptive o r divers ify ing selec tion--sel ec t i on 8t..ay f rom the
original mode of a pop ulat ion in t\o/O or more direc tions either as a
respon se to a more or les s s udden change i n the original e nvironment
(short dura t ion) or as a f orm of balancing selection \.;hich produces a
comple x polymorphism.
4. Li near or directional sel ection- -con tinued select ion in a given direction
producing an evo lu t i onary trend. a sequence of a dapt i ve changes in
resp onse to an env i ronment al gradient .

B. Comparative Features of Abiotic and Biotic Selec t ion

Abiotic Biotic

1 . Evolution direction fixe d 1 . Evolution direction not fixed

(dependent on unal ter ab l e selec- (dependent upon interaction with
tive fo rce , s uch as c l ima t e ). ano th er evolving or ganism) .
2 . Hay often result i n parallel 2 . P~rely res ults in parallel evolu-
evolution . t ion.
3. Po pulation si ze not importan t . 3 . Population size important.
4. Bio logical interac tions general . 4 . Biolog ical inte ractions very specific.
5 . Does not produc e niche subdivision . 5 . Produces n ich e s ubdivi sion .
6. Pr o duce s low s pecies densit y . 6 . Prod uces high species density .

C. Comparative Features of Darwinian and Catastrophic Selection

Darwinian Catas trophic

1 . Operates on all individua l s in a l l 1 . Operates on ma r ginal i nd ivi dual s i n

popula tions . marginal populations .
2. Involve s gradual ch ange i n th e 2 . Involves r ela t i vel y sudden changes
selective force rar e l y exceed- in the e x t remes of the selective
ing range of tol erance of most fo rce that may exceed the tol-
individuals . e rance s of most i ndivi dual s .
3. Sel dom produces drastic decreas e 3 . Often produces drastic decrease in
in population density . population dens i ty .
4 . Large gene pool maintained ; fixa - 4 . Gene pool often dr ast ical ly de-
tion rare . c reased ; fixatio n l ikely .
5 . Selection is homeostatic (multi- 5. Selection 1s often unidirect ional.
directional) .
6 . Ra t e of evolution slow , long cyc l e . 6 . Rate of evolution fast er . shorter
cyc le .

V. ADAPTATION

Adapt at i on . which produces diversity and specia tion , is the r es ult of

n atur al select ion . Howeve r , organ isms can only adapt I"ithin the genetic
l imits of their existing gene pool . Ther efore, different organi sms . some-
times even when close l y related , may adapt in different I~ ays to a part icular
selec tive force . However , a h igh degree of adap t at i on (e.g . • overspecializa-
tion ) can l ead to a decrease in evolu tiona r y potential and ultima t e ext inc-
tion . The adaptation, when possibl e , of di fferen t popula t ion s of a n organism
to different select i ve factor s of their envi ro nme nts produces polymorphi sm
which may, in t urn , lead to speciation .
26 543

A. Results of Adaptation at the leve l of the i ndividua l.

1. Developmental homeostasis (canalization).
2. Correlation pl eiades .
3. Euheterosis .

B. Results of adaptation at the population level.

1. Heterozygosi t y .
2. Genetic coadaptat ion .
3. Genet ic homeostasis .

C. Some possible results o f adaptation to (us ually) abi otic selective f orces.
1. Eeotypes.
2 . Ecoclines .
3 . Eco logical equival ents .
4 . Paral lel evolution (convergence) .
5 . Periodicity .

D. Some possible results of adaptati on to (usually) biotic se lect ive forces.

1. Allopatry (competitive exclusion) . 6. Parasitism .
2. Succession (competitive exclusion) , a . facu ltative
3. Character displ acement . b . obligate
4. Coadaptation or coevolu tion (species) 7. Mimicry .
5. Symbiosis . a . Batesian
a . mut ualism b . Nueller i an
b . commensa lism c . Hertensian
d . au t omimi cry
,
For a summary of evolut ionary mechanisms we can paraphrase the neoDarwin-
ians. I t is clear f r om the above out l ine that there is indeed " unive r sal var-
iation ," both genet i c and pheno typic ; through various sexual and asexual means
of r eproduc tion there is an " overproduction of offspring " and the potential of
a geometric increase i n the numbers of organisms ; however, there is " competi-
tion " for space and raw materials, thus the numbers of organisms of each kind
generally tend to remain constant; those organisms wh ich have t he proper com-
binations of characters (Le ., are best " adapted ") tend to survive in higher
p ro portions than the "weaker" or unadapted organisms ; "adaptation" can take
many forms a nd may lead to gr eater uniformity o r greater diversity, depending
upon the organism and the circumstances; if adaptation r esul ts i n direct i onal,
here ditary variation we have evolution . Since biologica l variation , at dif-
fe ren t levels , is a t once the alpha and omega of organic evolution i t is
impor tant tha t its causes , and res ults, be understood.

EVOLUTION GLOSSARY

Abiotic Sele ction . Selection pri- Adapt i ve Radiat i on. Evolutionary

marily by the physica l e nviron- diver sificat ion wi thin a gr oup of
ment , usually unidirectional . related species o r other taxa
(See parallel evolution .) brought about by adaptations to
Adapt ation . The result of natural different selective forces through
selection ; t h e evo l ution of a time or migration.
character, or g r oup of charac- Adaptive Zone . The "place" or "niche"
ters that favor s u rvival under of an organism in the e volutionary
certain selective forces . reticulum of its community ; the set
of conditions to wh ich t he offspring
of an organi sm wi ll be preadapted .
544 26

Advanced . An or ganism , character or Batesian Nimicry . The superficial

cond ition presumed to be more resemblance of a harmless organ-
specialized or evolved than ism (the mimic) to a sympatric
another to which it is compared . and more plentiful organism (the
Adventitious Emb r yony . The production model) .
of an embryonic spor ophyte by mito- Biosystematics. The systematic
tic divisions from the tissues of study, usually includ ing repro-
the parental sporophyte without an ductive biology and chromosome
intervening gametophytic generation . studies , of the variation and
Agamospermy . The formation of seeds evolution of a species or species
without ferti li zation; pollen or complex .
male gametes may be necessary to Bio tic Potentia l. The total repro-
stimulate division of the zygote . ductive potential of an individual
Allogamy . Cross-fertilization . or a population; it is directly
Allometry . Different growth rates in re lated to the number of func-
different parts of the same organism . tional gametes , and seeds , pro-
Al lop atric . Populations or species duced.
with separate and nonoverlapping Biotic Selection . Selection pri-
geographic ranges. marily by the biotic environmen t.
Allopatric Speciation . TIle differen- See comp etitive exc lu sion and
tiation of geographi cally isolated character displacement.
populations i nto separate species. Bradytelic Evo l ution . Evolution at
Allopatry . The occurrence or origin a relatively slow rate as compared
of species or populations in differ - with most other knmm or presumed
ent geographical regions . evolutionary r ates .
Apogamy . The production in plants of Canalisation . The property , possessed
a sporophyte directly from a game- by developmental pathways, of
tophyte cell withou t the production achieving a s tandard phenotype
of a zygote derived by f us ion of despite gene tic or environmental
gametes . disturbance .
Apomict. An organism that reproduces Catastrophic Selec tion . A harsh
by asexual means; it may be facul - selective for ce (e . g., fire ,
tative or an obligate apomict. severe drought) acting on an eco-
Apomixis. Asexual repr oduction, includ- logically marginal population so
ing vegetative propagation . there are only a fe\~ survivors and
Apospory . The condition found in some a small gene pool remaining; results
higher plants in which a diploid in operation of the Founder Princi-
embryo-sac is formed directly from ple.
a somat i c cell of the nucellus or Character Displacement . The increase
chalaza and an embryo is then in the differential of a charac-
formed \"itllout fertilization . ter of two competing spec ie s in
Autodeme . A group of individuals of a an area of sympatry .
t axon composed of pr edominantly Cleistogamy . The flower self poll i -
self-fertilizing individuals . nates , Hithout open ing in the bud .
Autogamy . Self- fertilization; persis- Cl ine. A character gradient , often
tent autogamy results in an increase re lated to geographic distribution;
in homozygosity and the division of a more or less uniform series of
the popula tion in to a number of variants from one extreme to the
"pure lines . " other in a population or series of
Automimicry. Nimicry ~"ithin a species . populations; a continuous varia-
Allaesthetic Selection . Disruptive tion pattern .
selection via differential polli- Clone . A group of individual organ-
nation. isms produced by vegetative asex-
ual reproduction (rhizomes, stolons ,
budding, cuttings, etc.) and thus
all of the same genotype .
26 545

Coadaptation . The adaptive response Dimorph ism . Th e occurrence of two

o f two organisms to each other; morpho l ogical , or other , fo rms of
e . g . , flowe r structur e anu pol lina- a given organism ; e . g . , sexual
t or . dimorphism.
Coe nogamodeme . A unit composed of all Dipl ospory. The phenomenon in some
the hologamodemes which a r e consi- highe r plants in '~hich a dip l oid
dered t o be capable of exchan ging embryo- sac is formed directly from
genes to some extent, but not with a megaspore mo th er- cell ; an embryo
freedom , under a specified set of is then form ed withou t fer tiliza-
condit-ions. tion .
Coenospecies . An ecological category Dire ctional Selection . Selec tion
approximating a Lin naean genus . occu rr ing ,.hen the environment is
Commensalism . A r elationship b etween changing in a systematic fash ion ,
individuals o f t ,,,"o species in '''hieh l eading to a regular directional
one species be nefits but does not change of the adaptive charac-
harm the s econd, or host , species . teristics of a gamod eme .
Comparium. A genet ic term to des ig- Disj unc t . A geographically isolated
nate a group of coenospec ies which popula tion or species outsid e of
c an hybridize either direct l y or the range of other simi l ar popu-
through intermediates. lations o r species .
Competitive Exclusion . The ef f ect of Dis rup tive Selec t ion. Sel ection
stron g bio tic selection vhich (usually bio tic ) \"hich b reaks up
e liminates one of two compe ting a ",eakly polymorphic popula tion
sympatric organi s ms ; environmen- into a number of different l y -
tal modification by one organism adap ted variants .
t hat resul ts in the exc lus ion of Ecoc line . A varia tional trend corre-
another; e . g . , plant suc c ession . lated ,~ith an ecological g radient.
Convergence . A type of paral le l evolu- Ecodeme. A group of i ndividuals of
tion r esulting in similar f orm or a taxon occurring in a speci fied
stru c ture in different or ganisms; kind of habitat .
e . g ., ecolog ical eq uivalen ts; the Ecospec ies . An ecological category
morphologi cal similarity of de sert- approximating a Linnaean species.
dwelling p l ants of the Eupho r h ia- Ecotype. An ecologi cal catego r y
ceae and Cactace a e . indicating an infra-specif ic
Corre l a t ion Pleiades . The t endency of group of p l an ts a dapted to a
t wo or more characters of an in - par t icular habitat .
dividua l that are under the same Endemic . A pop ulation or spec ies
selec t i ve fo r ce to have corre- ,~ ith narrow phys iological or othe r
lated patterns of variation . re strictions I.hich limit it to a
Cross Pol l ination . Pollinat ion between special habitat or a very re -
two differen t p l ants of the same str icted geogr aphic range . or both .
species; ou t cross. Environmental Var iance. That portion
Cytodeme . A group of individual s of of the phenotypic var iance caused
a taxon differing from others cyto- by differences i n the envi ronment s
l og ically , e .g ., in chromosome to "'hich the indiv iduals in a popu-
numb er . lation have been exposed .
-deme. A biologically and taxonomi- Epigenotype . The series of interre-
cally neutra l suffix , whi ch should lated developmen t a l pathways
not be used alone , that deno t es a through which the adu l t fo r m is
gro up of individuals of a s pe c i - realized .
fied taxon ; the prefix indica tes Ep istasis . The interaction of non-
the context of th e relat ionship or allelic gen es .
similarity between the ind i viduals . Euheterosis . The condition in ",hich
(See topodeme , gamodeme , cyto- the heterozygotes in a popu la tion
deme , etc .; also Chapter 11 ,111) . are at an adapt i ve or sel ec tive
advantage ove r the homozygo tes.
546 26

Evolution . A directional change Geneti c Assimilat ion . The incorpor-

in gene frequency , over time , in ation in to the genotype , by a
response to selec tive factors of selective process, of charac t er-
the environmen t ; the con tinuous i st ic s appearing in ontogeny as
genetic adaptation of organisms a respons e t o the envi ronmen t.
to their environmen t . Geneti c Coadaptation. Adap tation
Expr essitivity . A mea sure o f the of blocks or combinations o f
uniformit y of th e phe notypic gen es to eac h other tn a n inte r-
expres sion of a gene i n a parti- breed ing popul at i on .
cular env i ronment . Genet ic Dri ft . Random fl uc tuation
Feedback . The in flue nce of th e of gene f requen cy (us ually due
result of 0. proc ess upon the to sampl i ng error inheren t in
fun c t ion ing of the pr ocess ; th e genet ic mechanism) ; p r esen t
success i n producing viable and in al l population s , its effects
fertile offspring . a r e mos t eviden t i n very small
Fi t ness. The s ur vival value and populations .
reproductive capability of a Genet ic Homeos tasis . The tendency
given genot ype relat iv e to other of pop ulations under selec tion
genotypes in a popul ation . to main t ain , or regress tOl. . ard
Fixation . \.fhen the frequency of a th e adaptive mean rather than
par ticular al l ele rea ches zero or an a dap tive peak .
100% i n a (usual l y small) pop ula- Genet ic Load . The average number
tion . of po tent ial deaths per indi -
Founder Principle . The establish- vidual due to gen e t ic caus es ,
ment of a new pop ulation in iso- such as lethal s and semilethals,
la tion so that its gene pool is in a populo. tion .
not i denti ca l \·, ith tha t of the Genetic System . All the fac to rs
parent population; these di ffe r- that affect genetic recombina-
ences ar e e nhan ced as different tion in an organism .
evolutiona r y press ures in th e Genot ype . Refers either to the
area s occupied by the two popu- total genetic component of an
lations must th e r e for e oper ate or gan ism or , in a di ffe rent
in cJiff e r ent genetic e nviron - sen se , to a specific allele , in
ments; th e result is increa sed a par ticular ind ivi dual .
divergence . Genot ypic Variat i on . Genet ic varia-
Gamodeme . A group of ind ividuals o f tion; di ffe ren ces in geno t ypes
a spe ci fied taxon I .... hich , within within a pop ula tion (or species)
the limits of the breeding sys - ar e a re sult of mutation and
tem, can inte rb reed . r ecomb ination .
Gap . A d i scontin ui ty in variation . Gynogenesis . Reprodu c t ion by pa r-
Genecology . Th e st udy of in tra spe - thenogenesis in wh ich stimu la -
cifi c var ia tion in plan ts in tion by a spe rm is n ecessa r y fo r
r elation to environment . the development of the egg .
Gene FIOl..•. The d is tribu tional Haplo phase . That p.'lrt of the life
"move ment " of ge nes wi th in and c ycle in I~hich t h e game tic c hrom-
among pop ulations . osome number i s fo und in r epro-
Gene Frequency . The percen ta ge of ductive ce lls .
an allele in a population; the Ilardy-l.Jeinberg Lm.. . . Th e lat. . stat ing
number o f loci at whi ch a given that , in a panmictic population ,
allele is fo und Id thin a popula- in the a bsence of selection , muta-
tion, divided by the total num- t ion , mi gr ation a nd genet ic d rift,
b e r of loci at I.;hich i t could t he f r eq uency of au t osoma l genes
occur. at a given l ocu s r emains constant
Gene Pool. The tot a l gen e tic infor- and that , af ter one generation ,
ma tion posse ss ed by a population . the freq u ency of genotypes at the
locus reaches eq u il i b r ium .
26 547

Heritability . The Belletic variance Imprinting . The i mposition of a

divided by the phenotypic var iance; stable behavi or pattern by expo-
an estimation of the degree of sure, during a part:l.cular period
resemblance between offsp ring and in development , to one of a
paren t . restricted set of stimuli .
He r maphroditic . Possessing the pheno- Inb r eede r . A plant that is normall y
type of both sexes in organ isms self-fertilized .
\~ith male -female different ia tion . Induction . Dete r mination of the
Het eroblast ic Change. The tran sition developmental fate of one cell
fro m a juvenile to an adult form mass hy anot her .
accompanied by a more or less Introgression . lncorporation of ge-
abrupt change in morphology . netic material f rom one pop ulation
Heterogametic . Produ cing game t es Id th into that of ano th er through hy-
differ i ng chromosome complements . bridization fo l lowed by repeated
Heteromorphic . Hith many forms ; backcrossings ; usually r est rict ed
polymorphic . t o populations regarded as dis-
Het e ro troph ic . Requiring organ ic tin ct by taxonomists .
carbon for nutrition. Introgressive Hybridization . Genet ic
Higher Categories or Ranks . Taxonomic modification of one species by
categories or r an ks above the level another by gen e flow through fer-
of genus in the established hier- tile hybrids and backcrosses ;
archy of our taxonomic system . introgression .
Hologamodeme. A group of in dividuals I so l ating Hechanism . Any mechanism
of a taxon Hhich arc believed t o or condition wbich prevents gene
interbreed Hith a high level of exchange betlJeen the individuals
frecdom under a specified set of of two or morc popula cion s ; i so-
conditions . This term is preferred lation may be e ither prezygo ti c
to ' biologi cal species,' Hhich or postzygotic and may be of
r elates to a ta xonomic category . several types .
Homomorphic . 11ith a single form; r.!acroevolution . Evolutionary event s
monomorphic . vicIJed through the perspectiv e o f
Horotelic Evol ution . Rates of evolu- geologic time, such as the evolu-
tion falling Hithin the distribution tion of different and distinctive
(asymmetrica l Hith a mode nearer the organisms .
upper than the lower end) most cor.t- Matroclinous (r.laternal) Inheritance .
monly found ,,,hen evolut i onary rates The condit ion when a hybrid is
are plotte d in a frequency distri- closely similar to its seed
bution . par ent.
Hybrid . An individual resulting from Heiotic Drive . A higher p robab i lity
cross fe r tilization bet,.reen or- of one allel e at a lo cus being in-
ganisms belonging to different cluded in the gametes t han t ha t of
taxa, usually different species . others .
Hybrid Zone. A zone of sympa try and Hen del1an Pop ulation . A r ep roductive
hybridization bet,"ecn tl'lO species group sha ring 11 common gene pool.
I"hich may , desp ite hybridizat ion Neristic Variation . Variation in
Hithin the zon e , be a barr ier to number s of parts or of organs .
extensive gen e flow or int ro- Her tensian Himic . Hhen one deadly
gression between the spec ies in- and one harmless organism both
volved. m1m1C a moderately poisonous or
Hybridization. The forma tion of a distasteful model .
zygote th rough the fusion of gametes Nesopolyploid . A polyploid o f i nter-
from organisms belonging to tHO mediate biological Or geo logical
different taxa . ag e . (S ee paleopolyploid , neo-
polyploid) .
548 26

Hicroevolution . Evolutionary events Panmixis . Completely random mating

usually viewed ove r a short period within a population .
of time, such as changes in gene Parallel Evolution. The r es ult of
freq uency within a population ove r strong direct i onal selection pro-
a fe\o,' generat i ons . ducing similar adaptations i n
Nigratlon. The dispersal and estab- related organisms .
lishment of organisms beyond Parallelism. Evolu tionary convergence
their place of origin; a periodic along similar pathways among close l y
movement of individuals; both related organ isms.
result in the transfer of genetic Parasitism . "'lhen one organism obtains
information among populations . f ood or raw materials , or both from
Himicry . The superficial resemblance another living organism (the host)
of one organism by another . pre- ,·lith no apparent benef i t and often
sumably affecting the actions of harm to the host; some organisms
predators. are obligate parasites and cannot
I'lonomorphic. One form. l ive without their host, other
Morphogenesis . The process leading organisms, facultativ e parasites ,
to the development of the charac- may be auto tropi c under some condi-
teristic ma ture form of an organism. tions .
Hullerian l>Iimicry . The superficial Parthenogenesis . The development of
resemblance. and thus reinforcement an individual from an unfertilized
of the pattern on predators, of tt.)'Q gamete .
or more harmf ul organisms . Penetrance . A measure of the propor-
l'\utualism. A type of symbiotic re l a- tion of ind ividuals homozygous for
tionship between t\,JO organisms of a gene that shows its phenotypic
different species which bene fits effect.
both of the organisms invol ved. Periodicity. The periodic occu rrence
Natural Selection. Nonrandom (differ- of a given biological phenomenon,
ential) reproduction and sur vival pre sumab ly as a result of natural
of genotypes without the conscious select i on.
intervention of man . Phenetic . Presumed natural (but not
Neopolyploid. A relatively recent phyletic) rela tionsh ip based on
polyploid, the parent(s) of which current t otal similarity ; relation-
are still ex tant . ships indicated by the use of un-
Niche . The status (way of life) of ' . . eighted characters, as in n umeri-
an organism in a community; e . g ., cal taxonomy .
its position in the fo od chain or Phenocopy. An environmentally pro-
relationships Hith the physical duced specia l i zed phenotype that
substrate . r esembles the phenotype of a dif-
Ontogeny . The development of an ferent, more specialized, genotype;
individual. fa cultative dimorphism .
Organizer . A portion of an emhryo Phenology . Biological processes that
that determines the developmental are correlated with the seas ons ,
fate of the cell masses wi th which e . g . , flOl. . ering , mat i ng , dormancy.
it comes into con tact. Phenotype . The actual expressed
Outbreeder . A plant that is normally characters or observable form or
cross fertil i zed ; outcross . physio l ogy of an organism .
Outcross . Pollination bet'. . een t,. . o Phenotypic Variance. Th e total vari-
different plants of the same spe- ance observed in a character.
cies ; cross pollination . Phenotypic Variation. Morphological
Overdominance. The res ult of the or physiological variation; the
heterozygote being mo r e extreme result of the action of different
than either homozygo te. environments on one or more geno-
Paleopolyploid . An ancient poly- types.
ploid the parent (s) of wh ich are
extinct .
26 549

Phyle tic Evolution. Any change occur- Primitive. An organism or character

ring sequentially in a single line judged to be less changed from a
of descent . presumed common ancestral state
Phylogenetic . Pertaining to phylogeny, than another with \olhich it is
the evolutionary history of an compa r ed .
organism ; relationship through Ps eudocopula tion. A mode of polli-
lineal descent from a common nation in some orchids in which
ancestral form . st ructures of the flowe r resemble
Plast i city . The deg ree to which a a female insect and male insects
given genotype may vary pheno- attempting copulation transfer
typically under different environ- pollen from one flO\oler to another.
ments; also, the total variation Pseudogamy. The phenomenon found in
(both genotypic and phenotypic) of some apomictic plants , whereby
a species or population. pollination is necessar y for seed
Polymorphism . The occurrence at a development .
given stage of development, of tHO Pure Line . A lineage of ind ividuals
or more distinct variants of a originating from a single homo-
species or population . zygous ancestor .
Polyploid . An individual with 3 or Quantum Evolution . Rapid evolution-
more (haploid) se t s of chromosomes. ary change resulting in what a
(See additional ploidy and chromo- taxonomist would r egard as a new
some number terms in Chapter 10 . higher taxon .
Section A VII , p . 246 . ) Ramet. An individual belonging to a
Polyploid Complex. A series of inter- c lone.
related polypl oid s , often with mor- Random Sample . A subset of a popula-
phological s imilarities , involving tion selected or obtaine d in such
one or more level s of polyploidy . a \-lay that a l l variants in the
Poly topic. The occurrence of a spe- pop ulation ar e equal ly likely to
cies , or other taxonomic group , in be included in the samp le.
t\olO or more separate areas . Ratio-Cline. Clinal variation oc cur-
Population . A group of individuals ring in polymorphic species , in
of one kind in a given area at a which successive populations show
given time; often used as a syno- progressive change in the propor-
nym for mendelian population . tion of the variant s .
Population Structure . The s um of all Recapitulation . The idea that in the
the factors that govern the pattern course of development an individual
in which gametes from various passes through stages similar in
individuals unite with each other. form to adults of it s presump tive
Postzygotic Isolation . The prevention ancestors; often stated as "ontogeny
of gene flow bet\·leen hybridizing recapitulates phylogeny . "
populat ions (or spec i es) caused by Recombination . The recombining of
the fai lure of the hybrids, or genetic material through c rossing
their derivatives, to be viable over and random as so rtment at
or reproductively competitive. meiosis and the fusion of geneti-
Preadaptation. The historical (evo- cally different gametes .
lutionary) or fortuitous ability Recombination Index . A measure of
of an organism to surv ive in a the number of new gene combinations
particul ar environment in which it tha t can be produced in a given
finds itself . time ; of Darlington : the average
Pre zygotic Isolat i on. The prevention number of chiasmata plu s the
of gene flo w between populations genetic chromosome n umber .
(o r species) caused by geographi- Recombination System . The interac -
cal, ecological, seasonal , mechani- tion of a series of genetically
ca l, e tholo gica l or other di ffe r- controlled processes which more
ences which variou s ly act to prevent or less determines the amount of
cross fertilization . recombination possible in a
 550 26

species ; the system may be Spontaneous Generation . Abioge nesis .

"open , " "restricted," or Sporophyte . The diploid spore-pro-
"closed . "
ducing phase o f the life-cycle of
Replicate. To duplicate repeatedly . a plant .
Reproductive Cells . The gametes and Stabil izing Sel ec tion . Selection in
their immediate predecessors from which genotypes closer to the mean
Hhich they are produced by divi- for a character have an advantage
s i on . over those at the extremes .
Restitution Nucleus. A nuc leus Structural Hybridity . Heterozygosity
f ormed by the fusion of tlW for a chromosomal rearrangement .
daughter nuclei, usually in a Subspecies . A geographic subdivision
cell which d i d not divide after of a speCies deemed Iwrthy of for -
its nucleus divide d . mal recognition by a taxonomis t .
Reticulate Evolution . Evolution in Succession . The sequence of transient
which phyletic lines merge as wel l communities occurring in an area
as diverge . before the climax community is
Sample . A subset of a population; a es tablished.
group of items picked by some pro- Symbiosis . Living togeth er ; see
cedure and f rom '''hi eh one hopes to mutualism. commensalism and para-
learn certain things about the sitism.
population . Sympatric. Occurring i n the same
Sampling Error. Varia tion due to geographic area.
random elements in the sampling Sympatric Speciation . Speciation
process . without geographic isolation ; see
Se l ection . The action of environ- divergent selection .
mental factors on each member of Sympatry . Populations (or species)
a population that results in the occupying the same geographical
'. differential reproduction of area .
genotypes .
Syngamodeme. A unit composed of all
Selection Coefficient. The measure coenogamodemes which are connected
of the disadvantage of a given by the ability of some of their
genotype relat ive to other geno- members to form viable but sterile
types in the populat i on . hybrids under a spec if ied set o f
Self- fe rtile . Capab l e of self conditions .
fertilization. Systematic Pressure. One of the non-
SeXual Reproduction. Reproduction random evolutionary pressures :
involving the fusion of gametes s e lection. mutation , or migration .
produced by a prior meiotic Tachytelic Evo l ution. Evolution at a
process . relatively rapid rate, e . g ., at a
SpeCialized . Apply ing to an organ- much faste r rate than horotelic
ism or character judged to be evolution.
more changed f rom a presumed Taxon. A unit of taxonomic classifi-
common ancestral state than cation such as family, genus, or
another ,dth t.,hich it is com- species .
pared . Territoriality. The defense of an
Speciation. The splitting process area against organisms of the same
of evolution that results in the or similar kind .
existence of different kinds of Topocline . A geographical variational
organisms that are classified as trend I.hich is not necessarily
species by taxonomists . correlated with an ecological
Species. A category in the taxonomic gradient .
system; a group of organisms judged Topodeme. A group of individuals of
by taxonomists (b y diverse cri- a taxon occurring in a specified
ter i a) to be Horthy of formal recog- geographical area.
nition as a distinct kind. cate-
gory . or rank .
 26 551

Transgressive Variation . Hhen the Viability. The capability for living

heterozygote (or individual segre- or continuing to develop; often,
gating) is more phenotypically but incorrectly, used as synony-
extreme t han eit her homozygote (or mous with fitness.
pa rental form); ove rdominance . Vicariads . Close l y allied but al10-
Variant . A neutral term applied to pat rie taxa.
any definable individual or group Vivipary . In plants, vegetative re-
of individuals. production in which propagules
Variation Pattern. The type , form and replace flowers in the inflores-
extent of variation in one popula- cence; in animals, the production
tion or taxon as compared with that of living young rather than eggs .
in another population or taxon . Xenogamy . Outcrossing ; outbreeding .

EVOLUTION LITERATURE

I. Periodicals.
American Journal of Botany. 1914+. Published by the Botanical Society
of America.
Brittonia. 1949+ . Published for the American Society of Plant Taxono-
mists by the Ne\~ York Botanical Garden.
Ecology . 1920+. Published by the Ecologic al Society of America .
Evolution . 1946+. Published by the Society for the Study of Evolution .
Index to Plant Chromosome Numbers . 1956 . Published annua lly 'by the
International Association of Pl an t Taxonomist s as a volume of Regnum
Vegetabile.
Journal of Eco l ogy. 1913+. Published by the British Ecological Society . ,
The American Naturalist. 1867+. Published for the American Society of
Naturalists by the University of Chicago Press.

II. General References.

Bell , C. R. 1967. Plant Variation and Classification. Wadsworth Publish-
ing Company. Belmont, California .
Briggs, D., and S. M. Walters . 1969. Plant Variation and Evolution .
HcGraw-lIilL New York .
Brosseau, G. E. (ed .). 1967. Evolution : A Book of Readings . }Iilliam C.
Brown Company . Dubuque. Iowa. [A collec tion of basic reprints. 1
Dressler, R. L. 1968 . Pollination by Euglossine bees. Evolution 22:
202-210.
Ehrlich, P. R., and R. W. Holm . 1963 . The Process of Evolution . NcGraw-
Hill. New York .
-~C7o--.' and P. H. Raven . 1969. Different iation of populations . Science
165, 1228-1232.
Evolutionary Biology . 1967. A book of review articles published annually
by Appleton-Century Crofts. New York. Edited by T . Dobzhansky , H. K.
Hecht, and W. C. Steere .
Faegri, K., and L. van der Pij l . 1971. The Principles of Pollination
Ecology. Pergamon Pres s . New York.
Glass, B. , et al. (eds .). 1959. Forerunners of Darwin . The Johns Hopkins
Press. Baltimore .
Grant, V. 1968 . The Origins of Adaptations. Columbia University Press .
New York.
1971 . Plant Speciation. Columbia University Press . New York.
Hadley, E. B., and D. A. Levin . 1967 . Habitat differences of three Liatri s
species and their hybrid derivatives in an interbre eding pop ulation.
American Journal of Botany 54: 550-559 .
552 26

Harper , J. L . 1961. Approaches to the Study of Plant Competition . Symposia

of the Society for Experimental Biology X:V. Nechan isms in Biological Com-
petition.
Levin, D. A.• and H. 1". Kerster. 1967. An analysis of interspecific pollen
exchange in Phlox . American Naturalist 101: 387-400.
Levins, R. 1968 . Evolution in Changing Enviro nments. Princeton Univ ersity
Press. Princeton, New Jersey.
Lewis, H. 1962. Catastrophic selection as a factor in speciation. Evolution
16 : 257-27l.
Nelson, A. P . 196 7 . Racial diversity in Californi an Prunella vulgaris. The
New Phytologist 66 : 707-746 .
Ornduff, R. (ed . ) . 1969 . Papers on Plant Systematics . Little, Brown and
Company. Boston.
Solbrig, O. T. 1970 . Principles and Hethods of Plant Biosystematics. The
Mad-lilIan Company . NeH York .
Sneath, P ., and R. R. Sokal. 1973. Numerical Taxonomy . \~. H. Freeman and
Company. San Francisco.
Stebbins, G. L. 1950. Variation and Evo lution in Plants. Columbia Unive r-
sity Press. Net" York .
1966. Processes of Organic Evolution. Prentice Hall , Inc . Engle-
wood Cliffs, New Je rsey.
1969. The Basis of Progressive Evolution . Un iversity of North
Carolina Press. Chapel Hill.
Tax , S . (ed.). 1960. Evolution After Danlin . Vol. I : The Evolution of Life.
The University of Chicago Press. Chicago.
Wicker, W. 1968. (translated by R. D. Martin). Mimicry in Plants and Ani-
mals . McGraw-Hill Book Company. Net... York .
Williams, G. C. 1966. Adaptation and Natural Selection . Princeton Un iver-
sity Press. Princeton , New Jersey.
27 553

Chap te r 27 . STRUCTURAL EVOLUTION AND PHYLOGENY

The word phylogeny refers to the origin and evo lutionary history of or-
ganiSms in time. Phylogene tic char acter s. accordingly , are c haracte r s asso-
ciated with taxa over long period s of time and which a r e interpre table evo-
lutionarily . Some botanists have argued that attempts to construct phyloge-
netic relationships among angio s perms are pure speculation in view of the
incomplete fossil record of flow ering plants, the con flict betl. een phyloge-
nies based on diff e r ent kinds of evidence. the varying r ates of e volution in
both st ructures and groups of pla nts , and the a pparen t inabi li t y of botanists
to decide what characters are primitive. As no ted by Solbrig (19 70) , however ,
the term phylogeny can be used to r epr esent both t he actua l evolu tionar y
history of a taxon and the in ferred histo ry as deduced from available data.
Davis and Heywood (1963) suggest that most phylogene tic studies involve inter-
pre tations of evolutionary trend s in organs or tissues and, a l though this may
r esult in an i ndication of the relative advancement of a taxon rel ative to
other pl a n ts . it t e lls no thi ng about the deriva t ions of t he organisms concerned .
It is important, there f ore , to make the distinction between a natural (phenetic)
classification and a phylogenetic classification (see a lso Chapte r 28) . A
phenetic classification is one in which plant s are grouped together on the
basi s of their ove rall s imilarity whereas a phylo genetic system attempts to
interpret plant structure and function in terms of time and evo lutionary change .
Phylogenetic ist s are concerned with how par ticular morphologi es have been
attained . We believe it is essential that the student of systematics have
thorough training in vascular plant morphology, anatomy and evo lutiona r y prin-
cipl es not only to evalua te effec tively conflicting concept s regarding the
nature and evolution of angiosperm organs , structures , and taxa , but also to
help answer the four basic questions of systematic biology as outlined by
A. C. Smith (1969) -- (1) \ITbat is it? (i.e ., a circumscription of t he taxon and
an indication of how it differs f rom other taxa) ; (2) When did it arise? ( 3)
~~er e did it originate? and (4) How di d the taxon acqu ire its pr esent distri-
bution?

Sect ion A. ORIGIN OF THE ANGIOSPERHS

(Darwin ' s Abominable Hystery )

I. PROBLEMS O~ I NTERPRETATION

Prior to a consideration of possib l e ancestors of the angiosperms and the ir

time and place of origin , the following questions conf ronting t he botanist i n
his search fo r the or i gins of angiosperms must be answered: (1) What is an
angiospe rm? (2) Would all featur es considere d unique to flm. ering plants be
fossilized? (3) Since structures evolve at diffe rent rates when does a plant
become an "angiospe rm" ? and (4) loler e t he angio sperms monophyletic or polyphy-
leti c? The majori ty of botanis t s now favo r a monophyletic or igin a nd use the
following arguments to s upport their pos it ion- - the common possession by angio-
sperms of a similar type of embryo sac ; possession of endospe rm fo rmed by union
of sperm and two polar (female) nuclei; similar pollen types in primitive di -
cotyledons and mono cotyl edons; and the common possessio n of sieve tube elements
and companion ce lls as well as stamens and carp e l s . The statistical probabil-
ity of all of these c haracters ariSing together more than one time is extreme l y
s ma ll.
 554 27

II . HAJOR THEORIES ON THE ANCESTORS OF ANGIOSP ERMS

The re is no general agreement as to t~hich group of p lan ts represents the

inunediate ancestors o f the angiosperms. A \o.'ide range of putativ e ancestors ,
or close relatives, has been suggested at one time or anothe r and include :
algae , Filicales, Coniferales, Gnctales, Bennetti tales , Cycada les, P terido-
sperms, Pentoxylales, and Glossopteridaceae. As noted above, studies on the
origin o f the angiosperms are fu rther complicated by the sugp,est i on that
flovlering plants may be polyphyletic . The more promin ent theories on the
ancest ors of anRiospe r ms are br iefly disc ussed beIOl~ .

A. Coniferale s - Amcntife r ac Theory . Compar iso ns Nere made in the early

literat ure between the strob ili of the Coniferales and the naked , "'ind polli-
nated flowers of the Amentiferae I'i hich I'lere regarded as the most primitive
angiosperms. Acco rding to this interpretat ion, the stamen with its paired
microsp oran g ia Has regarded as homologous to the I'ri.crosporophyll of many con i-
fe rs with its tlVO micros porangia . Anatomical data hav e shown the Amentiferae
to be advanced, hOh'ever, a n d no support i s fo un d for this idea today .

B. Gnetalean theory . The combina tion of vessel-bearing \vood, broad

leaves I"ith reticulat e venatio n in Gne t um , and angiosper-m-Ilke ceprod uctive
morphology , particularl y in _Ep hedra, has prompted suggestions that this g mup
Has the ancestor of floHerin g plants . This theory Ho uld derive the catkin -
bearing angiosperms f rom a n Ephedra-like ancestor Hith th e magnolial ian typ e
flOl"er resulting from f urth er condensation of the catkin . The vessel elements
of the Gneta le s \-lith p e r for ations derived fro m circ ular-b o r de r ed pits rather
than scalariform pits ne ga tes th is idea according to most botanists .
'.
C. Be nnett i t alean theo S[. This theory is based up on the suppose d resem-
blance of the Meso zoic cycad e oid cone to a primitive flotver of the Nagnolia
type . The bisexual fruct i fication of Cycadeoidea had a basal I"horl o f peri-
anth-like bracts and a I~horl of pinnately compound microsporophylls beloH an
ovuliferous r egion . Thus, there ex i sted a superficial similarity to a flOl"er .
Along Hith other di ff icult i e s , it has been shown that the cone of Cycad eo idea
never possesse d expanded microsporophylls but rathe r had a compound mi c ro sporan-
giate synangi um Ivhi c h seems , to some, to present insurmoun table objections to
this theo ry.

D. Pteridoso erm theories . The discovery of the Caytoniales , a g rou p o f

Mesozoic seed ferns, caused considerable interest I"hen comparisons I·Jere mad e
between their fema le reproductive stru ctures and the a n giosperm carpel. llam-
$ha''''' Thomas (19 34) initially concluded that the basal lip of the Cayton i a
" fru it" fun ctioned like a stigma , and as a result the primit ive carpel might
also have had a gy nobasic style . A follicle i n this interpretation ",ould
rep r esen t a more advanced fruit type . Subsequent study of Caytonia demonstrated
that the pollen grains came i n to direct contact Hith the micropyle , Hhich r efuted
any connection be tll/een this extinct gro up and modern angiosperms . Although
there are points o f comparison , the la t e Paleozoic seed ferns are often regarded
as possible angiosp erm ancestors primarily on the basis tha t all other gro ups
can be eliminated. There are , hOI'l ever , objections to this idea such a s the
natur.e of the microsporangiate f ructification and tracheid pittin g that have
not been satisfa ctor ily resolved .
27 555

E. Cycadales theory . Recent evidence from fossil cycads indicates that a

few primitiv e c.ycads bore 2 rm.,s of special ized ovules that were part l y
covered by a lamina, thus represen ting an early stage in carpel evolution .
These clata would lend support to the opinion that the carpel of Tasmannia
and Drimys represents the primitive condition in angiosperms , although the
idea that the Cyendales Here angiosperm ancestors has not been Hidely accepted .

III . TIHE AND PLACE OF ORIGIN

I"ith regard to the time of origin of flm;ering plants there are basically
two opposing schools of thought. One group advocates a Cretaceous origin ( or
late Jurassic) \o,fhereas others strongly support a pre-Cretaceous origin of
flowering plants. Scott , Barghoorn and Leopold (1960) reviewed the fossil
r ecord and concluded there we r e no valid angiospermous fossil remains p r ior
to th e Cretaceous. The pollen record is generally in agreement with this
conclusion . Hany have argued, however, that since the angiosperm fossils of
the Cretaceous seem , to some, rather advanced in morphology and since these
fossils appear rather diversified, the flm.,rering plants must have had a long
pre-Cretaceous history, perhaps evolving in situations \.;hich were not condu-
cive to fossiliza t ion . Various Hesozoic angiosperm fossils have been reported,
however, these materials have always come under serious interpretational doubt
or otherwise have been shown to be incor r ectly dated .

Najar theories on the place of origin and subsequen t migration of angio-

sperms include : (1) Angiosperms had a holarctic origin with s ubsequent south-
\,rard m:igration; (2) Th~ flm.;ering plants attained much o f their present dis-
tribution by long-distanc e dispersal; (3) Angiosperms arose in the general
area of Southeastern ASia-Indomalaysian tropics and reached their present di s -
tribution by Antarctic land connections; i.e., migration \-las over land sur-
faces not much different from those existing today; and ( 4 ) Angiosperms
evolved in pre-Cretaceo us times and the process of continental drift was the
major factor in establishing present distributional patterns .

The geological and biological evidence supporting sea-floor spreading

(continental drift) is now so impressive that few disp ut e it, however , as
related to the early evolution of a.ngi osperms the important consideration is
what were the relative positions of the continents in geologica l time . The
term GondHanaland refers to the southern land mass formed by Antarctica, Aus-
tralia , southern Africa, India, and South America prior to their separation .
The northern group of continents prior to their separation is knOlvu as Laurasia .
Axelrod (1970) is of the opinion that the flowering plants evo lved in mild up-
l ands at 1m.; latitud es ,dth the breakup of Go nd\~ana 1and r esulting in a trend
tm.;ard widesp r ead , more equitable clima te \-.'hich favore d the invasion of the low-
l ands by the early angiosperms . Smith (1 970) argues fo r an origin of angio-
sperms in the general area of southeastern Asia, adjacent Ha1aysia, at a time
the Gondwanaland and Laurasia land masses Here undergo ing initial fragmenta-
tion by seafloor spreading . Nigration was subsequently over three principal
routes: (1) Northward overland to northern Asia and Europe, some elements
crossing the Bering S trait and then moving south,.,rard , (2) \.Jestward along the
coast of the Indian Ocean . This took place during moist periods or along
insular chains skirting the coast of Hadagascar and Africa , and (3) South-
ward to i.,restern and Central Austra.lia . Subsequent migration into the "I-lest
Antarctic Archipelago resulted in angiosperms reaching South America by means
of insular stepping stones . In a recent discussion of the evolution and
relationship of the floras of Africa and South America, Smith (1973) suggested
 556 27

that continental drift cannot be used in exp l aining the plant distributions
of these land areas .

IV . ORIGI N OF THE NONOCOTYLEDONS

The origin of the monocotyledons is a subject which has long been debated.
At one time or another the mono cots have been regarded as ancestral to dicoty-
ledons and polyphyletic in origin . The current widely held view is that the
monocots were derived from dicotyledons in a monophyletic fashion , although
there has not been uniform agreement on \olhich group of monocots represents
the ancestral stock.

As in dicots. the primary xy l em vessels of monocotyl edons probably orLgL-

nated more than once from scalariformly pitted tracheids (Cheadle , 1953) . How-
ever , the phylogenetic sequence is different; vessels first arose in the late
metaxylem of roots, as evidenced from the fact that the most specialized ves-
sel elements always occur in the root metaxylem , and only subsequently extended
into the stem and leaf . \.Jithin anyone organ , vessel element origin and
specialization occur first in the late metaxylem and then progressively in the
early metaxylem and protoxylem. The distribution of vessel elements in mono-
cotyledonous families shows they may be restricted within the plant body to the
root , to the root and stem, or occur in all part s . In the opinion of Bailey
(1944) and Cheadle (1953) the independent or i gins and specializations of vessel
e l ements in monocots and dicots indicate that if the angiosperms are considered
to be monophyletic , the monocotyledons must have arisen from a vesselless woody
dicotyledon. Cronquist (1969), hm...ever , does not accept the position that ves-
sels evolved independently in monocots and dicots and, therefore , that monocots
arose from a primitively vesselless ancestor. In his view , monocots had an
, aquatic origin (which profoundly influenced their evolutionary history) from
dicotyledonous ancestors resembling the modern Nymphaeales . That is, they . had
apocarpous flowers without a specialized per i anth, monocolpate pollen, herba-
ceous habit; i . e . , did not have a functional cambium, had laminar olacentation,
lacked a functional vessel system either through being lost or never having
had one , and were aquatic . The aquatic habit resulted in the loss of a cam-
bium that subsequently resulted in the reduction and loss of vessels . In
keeping with this viewpoint, the Alismatidae are considered a near-basal side-
branch of the Liliatae. Kosakai , Hoseley , and Cheadle (1970) reported the
occurrence of primitive vessel elements in the primary xylem of roots of
Nelumbo and seriously questioned the idea that monocots were derived from the
Nymphaeaceae . These authors emphasized that the Butomaceae and Alismataceae
have advanced vessel elements only in the root metaxylem . Accordingly , it is
difficult for them to believe, in viet... of t ... hat is knmm of vessel evolution,
that the putatively primi tive A1ismatidae evolved advanced vessels in an
aquatic environment yet gave rise to terrestrial monocots that had more primi-
tive vessel elements in the metaxylem of their roots . The vessel story ,
according to them, favors the origin of the A1ismatidae from terrestrial forms
and does not support an aquatic "rana1ian" ancestry of the monocotyledons .
Suggestions have been made from time to time , e . g ., Tomlinson (1970), that
the \.JOody . arborescent and perhaps branched habit represents the primitive
form in the monocotyledons and not the herbaceous condition .

V. PRI HITIVE ANGIOSPER}l FAHI LIES

The families most videly believed by evolutionary botanists to possess

the greatest number of primitive characters are listed below . It must be
27 557

remembered that evolution proceeds at different rates in different tissues

and thus these families exhibit a mosaic evolutionary pattern in which both
primitive and advanced characters occur together. At one time these families
were nearly all relegated to the large order "Ranales." however, contemporary
systems of classification have created additional orders to accommodate them .
The approximate size and distribution of each of the 39 families is presented
(after Smith , 1967).

Number of Number of
genera species Di stribution

1. i.;rinteraceae 7 95 Indomalaysia , eastern Australia.

Helanesia, New Zealand; Madagascar,
South and Central America
2. Magnoliaceae 12 250 Tropical and temperate S . E. Asia and
Indomalaysia; S. E. North America
through ~.;rest Indies and Central
America to ep..ste:::-r. Erazil
3. Degeneriaceae 1 1 Fiji
4 . Eupomatiaceae 1 2 Southeastern Australia to New Guinea
5. Himantandraceae 1 2 New Guinea; Holuccas , eastern Aus-
tralia
6. Illiciaceae 1 42 Japan , China , and Himalayas, Philip-
pines, Borneo, Malaya, Sumatra;
S. E. United States, eastern Mexico,
Cuba, Haiti
7. Schisandraceae 2 47 Sakhalin and S. E. Siberia to India
and into Nalaysia to Amboina and .,
Java; S . E. United States
8 . Annonaceae 122 2,100 Predominantly pantropical, extending
into subtropical and temperate
regions
9 . Myristicaceae 16 380 Pantropical
10. Canellaceae 6 20 Tropics and subtropics but lacking in
Asia and Indomalaysia
1l. Aristolochiaceae 7 600 Tropics and subtropics , extending
into temperate areas
12 . Austrobaileyaceae 1 2 Northeastern Australia
13 . Chloranthaceae 5 70 Tropics and subtropics but not in
African region
14. Amborellaceae 1 1 New Caledonia
15. Trimeniaceae 2 7 New Guinea and eastern Australia to
New Caledonia and Fiji
16 . Monimiaceae 33 450 TropiCS and sub tropics , predominantly
of Southern Hemisphere , with cen-
ters in South America , Malaysia,
and Hadagascar-Hascarene regions
17 . Calycanthaceae 2 9 North America, eastern Asia and
eastern Australia
18. Lactoridaceae 1 1 Juan Fernandez Island
19 . Gomortegaceae 1 1 Chile
20 . Lauraceae 47 2,250 Pan tropical , abundant in Central and
South America and in Malaysia; also
subtropical with extensions into
t emperate regions
558 27

Number of Number of
genera sEecies Distribution
2l. Hernandiaceae 2 52 Pantropical
22. Gyrocarpac e ae 2 17 Pantropical
23. Piperaceae 12 2,000 Tropical, abundant in tropical
America and Halays i a
24. Saururaceae 4 5 Himalayas to East As i a and Philip-
pines; Atlantic North America
and Southl.estern United States
and Hexico
25 . Lardizabalaceae 8 30 Indian to eastern and S. E. Asia;
also in Chile
26 . Sargentodoxaceae 1 1 China and southeastern Asia
27 . Menispermaceae 70 425 Tropical , with slight northern and
southern extensions into extra-
tropical areas
28. Ranunculaceae 47 2 , 000 Cosmopolitan; pre dominantly Northern
Hemisphe re, frequently in moun-
tains , ascending to 5,700 ffi . in
Himalayas
29 . Circaeasteraceae 1 1 Himalayas to northwestern China
30. Berberidaceae 14 650 Northern Hemisphere. ~lith only Ber-
be ris and Mahonia extending into
Southern Hemisphere
31 . Papaveraceae 28 250 Mostly subtropical and temperate
regions of Northern Hemisphere ,
Hith centers in western North
America and eastern Asia; rare
in Southern Hemisphere
32. Fumariaceae 19 425 Host1y subtropical and temperate
regions of Northern Hemi sphere ,
center i ng in Eurasia; also in
southe r n Africa
33 . Nymphaeaceae 8 80 Cosmopolitan
34 . Ceratophy11aceae 1 6 Cosmopolitan
35 . Trochodendraceae 1 1 Japan and Formosa
36 . Tetracentraceae 1 1 China and Burma
37 . Eupteleaceae 1 2 China to Assam
38. Cericidiphy11aceae 1 2 Japan and China
39 . Eucommiaceae 1 1 China

LITERATURE ON ORIGI N OF A.IIJGIOSP ERHS

AndreHs , H. N. 1963 . Early seed plants . Science 142 : 925-931.

Arber , E. A. N., and J. Parkin. 1908. Studie s in the evolution of the angio-
sperms . The re l ationship of th e ang i osperms to the Gneta1es . Annals of
Botany 22 : 489 - 516 .
and 1907 . The origin of the angiosperms . Journal of
Linnean Society (Botany) 38 : 29-80 .
Axelrod, D. I . 1952. A theory of angiosperm evolution. Evolu tion 6 : 29-60 .
1959. Po1eHard migration of early angiosperm flora. Sc i ence 130 :
203-207.
1970 . Hesozoic paleogeography and early angiosperm history . Botan-
ical Review 36 : 277 - 319 .
Bailey , 1. H. 1944. The development of vesse l s in angiosperms and its signif-
icance in morphological research . American Journal of Botany 31: 421- 428 .
1949 . Origin o f the angiosperms : Need for a broadened outlook .
Journal of the Arnold Arboretum 30: 64 - 70 .
27 559

Bancroft, H. 1930 . The arbo re scent habit in angiosperms . The New Phylolo-
gist 29: 153-169 ; 227-275 .
Benson, M. 1904 . The origin of flm,fering p l ants . The New Phytol ogist 3 : 49 - 51 .
Bel1s. J . \·1 . 1927 . St udies in the ecological evolution of angiosperms . The New
Phytologist 26: 1-21; 65-84; 129-14 8; 209-248; 273-2911 .
Camp , \\' , H. , and 1>1 . N. Hubbard. 1963 . On the origi ns of the ovule and the
cupule in t he lyginopterid pteridosperms . American Journal of Botany 50 :
235 -24 3 .
Cheadle, V. I . 19 53 . Independent o rigin of vesse ls in the monocotyledons and
dicoty l edons . Phy t omorpho l ogy 3 : 23-44 .
Cronquist , A. J . 1969 . Broad features of the system of angiosperms . Taxon
18: 188-193 .
Doyle, J . A. 1969 . Cretaceous angiosperm pollen of the Atlantic coastal plain
and its evo l utionary sign i ficance . Journal of the Arnold Arboret um 50 : 1 - 35 .
Er dtman , G. 1948. Did dicotyledonous plants exist in ear l y Jurassic times?
Geologiska Foreningens I Stockholm Forhandlingar 70: 265 - 271.
Greguss, P. 1966. The polyphyletic origin of angiospermae . Advan cing Fr on-
tie r s Plant Science 15: 37 - 49 .
Harr is, T. 1960 . The origin of the angiosperms . Advancement of Science 67 :
207-213 .
Hu ghes , N. F . 1961. Fos s il evidence and angiosperm ancestry. Science
Progress 49 : 84-102 .
Just , T. 1948 . Gymnosperms and the origin of angiosperms . Botanical Gazette
110, 91-103 .
Karsten , G. 1918 . Zur phylogenie der angiospermen. Zeitschrift fur Botanik
10 : 369-388 .
Kosakai , n ., H. F. Noseley , Jr., and V. L . Cheadle . 1970 . Horphological
stud i es of the Nymphaeaceae . V. Does Nelumbo have vessels? American
Journal of Botany 57 : 487-494 .
Li, H. L . 1960 . A the ory of the a nce stry of angi osp erms . Acta Biotheoretica
13 : 185-202 .
Haguire, B. 1970 . On the flora of the Guayana Highland . Biot r opica 2 : 85-100 .
Hamay. S . I], 1969 . Cycads: fossil evidence of Paleozoic origin . Science 164 :
29 5-296.
He l vi lle, R. 1966 . Con tinen t al drift, Hesozoic continen ts and the migrations
of the angiosperms . Nature 211: 116-120 .
1969 . Leaf venation patterns and the origin of the Angiosperms .
Na ture 224 : 1 21-125 .
1970 . Links betwee n the Glossop teridae and the angiosperms . In :
Second Gond~,'an a Sympo sium . Council for Scientific and Ind ustrial Res ear ch ,
Scien tia . Pretoria , South Africa .
1971 . Some general principles of leaf evolution . Sou t h African
Journal of Science 67: 310- 316.
Meeuse , A. D. J . 1963. The multiple origin of the angiosperms . Advancing
Front i ers of Plant Science 1: 105-127 .
Puri , V. l Q67 . The origin a n d evo lu tion of angiosperms . Jou rnal Indian
Botanical Soc i ety 46 : 1-14 .
Schopf , J . M. 1 970 . Relation of floras of the south ern h emisphe r e to conti-
n ental drift . Taxon 19 : 657-824 .
Schuste r, R. H. 1 972 . Continental movemen t s, "I.,ra l lace ' s lin e " and Indomalayan
Austral asian dispersal of land plants: some eclectic concepts . The Botani-
cal Review 38 : 3- 86 .
Sco tt, R. A., E. S . Ba r ghoorn , and E. B. Leopold . 1960 . HOt" old are the
angiosperms? American Journal of Science 258 : 284 - 299 .
_-;;--;;:_' P . L . 1.Ji1liams , L. C. Craig , E. S . Barghoorn, L . J . Hickey, a nd
H. D. MacGin ite . 19 72. "Pre-cretaceous" angiosperms from Utah : Evidence
560 27

for tertiary age of the palm woods and roots . American Journal of Botany
59 , 886-896 .
Smith, A. C. 1967 . The presence of primitive angiosperms in the Amazon Basin
and i ts significance in indicating migrational routes. Atas do Simposio
s obre a Riota Amazonica 4 (Botanica) : 37-59 .
1969 . Systematics and appreciation of reality . Taxon 18: 5-13 .
1970 . The Pacific as a key to flm.;ering plant history. Haro ld L.
Lyon Arboretum Lecture Number 1 .
1973. Angiosperm evolution and the relat ionship of the floras of
Africa and A~erica . In: Tropical Forest Ecosystems in Africa and South
America : A Comparative Revie'IV , B. F. Hegge rs, E. S . Ayens u, H. D. Duck-
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Sporne, K. R. 197 1. The mysterious origin of flowering plants . Oxford
Biology Readers 3, F. F . Head and O. E. Lowenstein (eds . ) . Oxf ord Uni-
versity Press. Oxford .
Thomas, H. II . 1934. The nature and origin of the stigma . A contribution
to\.. ards a new morpho lo gical interpretation o f the angiosperm flO\~er . The
New Phytologist 33 : 173-198 .
1936 . Paleobotany and the origin of the angiosperms . Botanical
Reviet~ 2 : 397-418 .
1958 . Pal aeobotany and the evolution o f the flower ing p l ants .
Proceedings of the Linnean Society of London 169: 134-143 .
Tidwell, hT • D. , S . R. Rushforth , J. L. Reveal , and II . Behun i n . 1970. Palmoxy-
Ion simperi and Pa1moxv10n pristina : THO pre-Cretaceous angiosperms from
Utah . Science 168 : 835 840 .
----C7--- ' and A. D. Simper . 1970. Pre-Cretaceous flowerine p l ants :
furthe r evidence from Utah . Science 170: 547 -5 48 .
--,c--""",' D. A. Hedlyn, and G. F . Thayn . 1972. Fossil palm materials from
the Tertiary Dipping Vat Formation of central Ut ah. Great Basin Natur-
alist 32 : 1-15 .
Tomlinson, P . B. 1970. l>lonocotyl edons--towards an understand ing o f their
morphology and anatomy . In : R. D. Preston (ed.) . Advances in Botanica l
Research . Academic Press . NeH York .
HolEe , J . A.• and E . S . Ba rghoorn. 1960 . Generic change in tertiary f loras
in relation to age . American Journal of Science 258 : 388-399.

Se ction Fl. PHYLOGENETIC DICTA, PRINCIPLES AND GENERAL EVOLUTIONARY

TRENDS IN FLOh'ERING PLANTS

For the most part, the basic dicta and princip le s presented in this sec-
tion form the basis of modern phylogenetic thought and classification . In a
discussion of phylogeny the terms primitive and advanced with respect to
different character states cannot be avoided. Sporne (1948) used the term
"primitive character" to mean one which is possessed by a present day taxon
and was also possessed by its ancestors . An "advanced character ," on the
other hand , is one possessed by a present day taxon and not possessed by its
ancestors , that is, it replaced an ancestral character during evolution . A
fundamental problem that confronts an evolutionary botanist involved in phylo-
genet i c pursuits is to determine the direction and rate in which structural
modifications have occurred or are occurring , i.e., to identify primitive and
advanced characters within the taxonomic units under study , and t o determine
,,'hether this direct i on has changed . Becaus e the errat i c fo ssi l record is of
comparatively little assistance in this regard , other approaches must be
27 561

employed (see Sparne, 1956) . The most successful methods in determi ning
advancement or primitiveness, as related to comparative morphology, are
termed by Sparne (1956) the doctrines of association, correlation . and
common ground plan.
That vessel elements e volved from trac.heids appears certain . Advantage-
ously , the sequence involves characters that are measurable and, therefore,
can be dealt \-,lith statistically . Bailey and Tupper (1918) provided the ear-
liest systematic eviden ce for this trend in a comparison of length of tracheary
elements of vascular cryptogams , gymnosperms , and angiosperms. Frost (1930a,
b; 1931) extended this \wrk and initiated a detailed statistical analysis of
specialization of the secondary xylem in dicotyledons. Frost made two basic
assumptions that were proved by subsequent tests of association and correla-
tion . The first I.as that the primitive state of the derived structure l.,rill
resemble the ancestral structure if it can be determined from which structure
the other has evolved . Si nce it l.,ras assumed that vesse l elements evolved
from tracheids, Frost deduced that the most primitive vessel elements l.,rere
those which most resembled tracheids (doctrine of association). If this
were not true, then the assumption of a direct relationship is incorrect, or
the structures are so widely separated in the evolutionary scale that the
tracheid-like condition of the vessel elemen t has been lost. The doctrine of
association has been criticized because of assumptions that are made prior to
its application . Frost's second assumption, the doctrine of correlation ,
takes advantage of the fact that primitive characters are frequently asso-
ciated l.,rith other primitive characters. Thus, if one character can be demon-
strated to be primitive (e.g .• scalariform perforation plates on the vessel
end wall) then characters co rrelated with this feature are probahly also primi-
tive. The doctri ne of the common ground plan assumes that any character
present in all or most individuals of a taxon, or the most common character
,
state among a group of related plants , has probably been inherited unmodi f ied
from a common ancestor and must be the ancestral or primitive state .

Phylogenetic speculation invoking the concepts of tissue conservatism,

recapitulation of phylogeny during ontogeny, and teratology generally have
met t.,rith criticism and limited success. To date, only the secondary xylem
of fo ssil angiosperms has revealed significant phylo genet ic information .

~.Jagner (1962) has pointed out that in the detennination of the most
probable pathways of phylogeny t.,rithin a group of plants there is one overrid-
ing evolutionary phenomenon which must be taken into account. This is that
common ancestry gives rise to divergent derivatives . Some derivatives will
closely resemble their ancestor whereas other derivatives l.,rill become
highly modified and not resemble their ancestor. If this pattern is valid
three basic assumptions are required : ( 1) plants l.,rhich have a large number
of similar characteristics in common have the sallie common ancestor (simi-
larity corresponds to recency of common ancestry); (2) evolution proceeds in
various directions . Different lines will change in different characters and
different character-complexes (evolutionary divergence) ; and (3) evolution
takes place at different rates at various times and in different lines . Some
forms will remain fairly stereotyped, resembling the common ancestor ; others
will change radically in the same period from the common ancestor (inequality
of evolutionary rates).

It must be emphasized that evolutionary i nterpretations of this type must

be made and evaluated in light of certain basic principles and pitfalls in
phylogenetic study . For example , i t is often necessary to determine l.,rhether
562 27

simplicity of structure ref lects a primitive condition or is the result of

secondary specialization . Similarity of structure may lIot imply or demon-
strate close genetic relationship because similar structures may have arisen
independently and more than one time in Hidely unrela ted taxa . Horphological
similarity between indi vidual organs and ent ire organisms has frequently re-
sul ted from the convergence het\.Jeen two distinct phyletic lines in response
to ada p tations to similar. habitats . or morphological similarity has sometimes
result e d from para llelism . Similarity r esult in g from common ancestry is termed
patri stic similarity whereas the actual pathHay or relative position of diver-
gences is spoken of as cladistic relationship . Both patristics and cladistics
are measurements of phylogeny involving time as opposed to strictly phenetic
cons i de rations . Thus, morphological resemblances have been attained patristi-
cally or by convergent evolution . From a taxonomic standpoint it is the degree
of mo rpho l ogical divergence that is significant and not the time at which the
divergence occurred. Anoth er f undamental basis of phylogenetic interpretat i on
i s homologous and analogous structures . Homologous structures refer to struc-
tures similar i n evolutionary and developmental origin regardless of function
whereas superficially similar structures not the result of corranon ancestry are
analogous . An organ or structure is homologous with another because of \"hat
it is , not becaus e of what it does (de Beer, 1971). Understanding which
structures and organs are homologous is a basic goal of comparative morphologi -
cal study and provides evidence of affinity between organisms that have evolved
with structural modification from a common anc estry . Deciding \.olhether a par-
ticular taxonomic group is monophyletic , having had a single origin, or poly-
phylet i c , having had multiple origins, is a complex problem not easily resolved
in the absence of a complete fossil record. If true phylogenetic relation-
ships are to be expressed, it is widely believed that higher taxonomic categor-
i es should be monophylet ic .

The u l t i mat e goal of phylogenetic study Is to determine , with the highest

degree of probability. the pathways of r e lat ionship . Hagner (196l , 1962) has
of f ered one method (The Groundplan/Divergence Nethod) of reconstructing the
best estimate of phylogeny utilizing all characters . This approach involves
three basic steps (quoted from Hagner, 1961): (a) systematic or comparative
analysis of the plants in question to find and understand thei r contrasting
characters; (b) determination of ground plans to f ind the character states
common to a l l or most o f the plants in order to deduce the most probable
an cest r al or primitive st a tes ; and (c) phylogenetic synthesis to assemble the
taxa according to thei r respective deviations from the basic ground plan and
from each other . Specific steps are as f ollows : (l) to compare and study
all the variable charac ters among the taxa; (2) to determine the gene r alized
or primi tive conditions on the principle that characters fo und in most or all
of a number of related taxa are inherited essentia lly unchanged f rom the com-
mon ancestor, using data also f rom related taxonomic groups of the same l e ve l .
( If no obvious trend can be determined in a given character that character may
be used only for grouping purposes . ); (3) to assign for each character the value
o for the generalized or primitive condition, and 1 f or the specialized or
secondary c ondition (the inte rmediate states being assigned the value 0 . 5);
(4) to list in tabular form the taxa and for ea ch give the divergence values
from the ground p lan , both for individual characters and in total; and (5) to
dete r mine the mutual character groupings between taxa and then arrange them in
sequence according to the se groupings on a concentric chart or graph , the radii
and branchings to be determined by the mutual character complexes , and the dis-
tances by the divergence indices . I t should be noted that this procedure
27 563

attempts to understand evolutionary trends in individual structures and to

correlate these in cladograms that illustrate the degree of divergence f rom
ancestors . those characters ,~hich have undergone modification, and the direc-
tion in which the evolut ion has progressed .

Cronqu ist (1968) has stated, " It thus appears that a Iwrkahle taxonomic
syst em cannot provide a perfect reflection of evolut ion, no matter hm., abun-
dant the evidence on which it is hased . Indeed , the more abundant the evi-
dence , the better understanding of phylogeny, the clearer i t hecomes that the
correlation betHeen phylogeny and taxonomy must be general rather than exact.
However, the phylogenetic concept still provides the underlying rationale for
the natural system . Taxonomy can provide only a somewhat muddy reflect ion of
evolution, but a r eflect ion al l the same . "

I . DICTA (After Bessey , 1915)

A. Gen eral Dicta

1. Evolution is not a hmys upI"ard, but often it involves degradation and
degeneration .
2. In general , homogen eous s tructures (\-li th many and similar parts) are
lowe r, and het ero geneous structures (with Eel,'er and dissimilar
parts) are higher .
3. Evolution does not n ecessarily involve all organs of the plant equally
in any particular period , and on e organ may be advancin g \olhile
another is retrograd i ng.
4 . Up\olard d evelopment is sometimes through an increase in complexity,
and sometimes by a simplification of an organ or a set of organs .
5 . Evolution has generally been consistent and I~hen a par t icular pro-
gression of retrogression has set in , it is persisted in to the
end of the phylum .
6 . In a phylum the holophytic (chlorophyll- green) pla n ts precede the
col or less (hyst erophyt ic ) plants, and the la tte r are d erived from
the former .
7. Plant relationships are ~ and down th e genet i c lines . a nd these must
constitute the framelwrk of phylogenetic taxonomy .

B. Dicta having special reference to the general structure o f the flOlvering

p l ants
8. The stem structure Ivith col l ateral vascular bundles arran g ed in a
cylinder i s more primitive than that I"ith scattered bund les , and
the latter are to be regarded as derived from the former .
9 . Hoody stems (as of trees) are more primitive than herbaceous stems,
and herbs are h el d to have been derived from t r ees.
10 . The simple, unbranched stem is an earlier type, from \>.' h ich branching
stems have been derived .
11. Historically the arrangement of leaves in pairs on the stem i s held
to have preceded the spiral arrange ment i n I~hich the l eaves are
so li t ary a t the nodes .
12. Historically simple leaves preceded br<lllched ( " compound") leaves .
13 . Historically l e ave s \"ere f irst pers i stent ("evergreen ") and later
deciduous .
14 . The reticulate venation of leaves i s the normal structure . and the
para llel v enation of some leaves is a special modification derived
fro m it .
 564 27

C. Dicta having reference to the flm.;rers of fioHering plants

15 . The polymerous flower structure precedes, and the oligomerous str uc-
ture f01101"5 f rom it, and this is accompanied by a progressive steri-
lization of sporophylls .
16 . Petaly is the normal perianth structure , and apetaly is the result of
perianth reduction (aphanisis).
17 . The apochlamydeous perianth is earlier and the gamochlamydeous peri-
aoth is derived from it by a symphysis of the members of perianth
whorls .
18. Actinomorphy is an earlier structure than zygomorphy, and the latter
resulted from a change from a similar to a dissimilar growth of the
members of the perianth \"horls .
19 . Hypogyny is the more primitive structure , and from it epigyny was
d e rived later .
20. Apocarpy is the primitive structure , and from it syncarpy was derived
later .
21 . Polycarpy is the earlier condition, and oligocarpy was derived from
it later .
22. The endospermous seed is primitive and lower, while the seed t.;tithout
endosperm is derived and higher .
23 . Consequently , the seed with a small embryo (in endosperm) is more
primitive than the seed t-.tith a large embryo (in scanty or no endo-
sperm).
24. In earlier (primitive) flm-lers there are many stamens (polystemon-
ous) while in later flOHers there are fewer stamens (oligostemonous).
25 . The stamens of primitive flowers are separate (apostemonous) , while
those of derived flowers are often united (synstemonous) .
, 26 . The condition of powdery pollen is more primit i ve than that with
coherent or massed pollen .
27. Flo\.,rers with both stamens and carpel s (monoclinous) precede those in
\"hich these occur on separate flowers (diclinous) .
28 . In diclinous plants the monoecious condition is the earlier, and
the dioecious later .

II . PRINCIPLES (After Thorne, 1958)

1. Exi sting species have descended with change from pre-existing species, and
are therefore the products of evolutionary forces .
2. Ancestral conditions and trends of specialization are often recognizable
in the organs, tissues, and cells of living and fossil angiosperms .
3. The primitive , ancestral condition of any given characteristic can be no
more special ized than its condition in a derived, existing species most
primitive for that characteristic.
4. The presence of vestigi al rudiments of organs , or sometimes the presence
of vestigi al vascular s upply to greatly modified or miSSing o r gans, often
f urnishes evidence of evolutionary reduction , loss, fusion , or other
major modification o f structures.
5. The prevalence of parallel and convergent evolution in habit, function ,
and structure is predictable consequence of the relatively limited means
angiosperms have for effective reproduction and for adaptation to avail-
able environmental niches .
6. All parts of plants at all stages of their development may produce evidence
that is valuable in establishing relationships .
7. Evolution may tend toward elaboration and diversity or toward reduction and
simplicity .
 27 565

8. The rate and direction of evolution may vary in the different organs and
tissues of plants.
9. Most existing angiosperms are highly specialized and greatly modified
from their primitive, generalized ancestors .
10. Evolutionary trends are sometimes reversible under the influence of
changes in environmental factors .
11. Once lost, organs usually are not regained.
12. New angiospermous structures have arisen as modifications of or as out-
growths from pre-existing structures.
13 . The sporadic or restricted occurrence of unusual or uncommon characteris-
tics lacking apparent evolutionary significance is often an indication
of relationship when correlated with other characteristics .
14. The occasional attainment of certain characteristics of certain levels
of evolutionary development is frequently valuable in determining the
affinities of families and orders.
15. Embryos and seedlings of related though dissimilar plants often resemble
each other more than do the adult plants because of their apparent reten-
t i on of primitive characteristics .

III . EVOLUTIONARY TRENDS IN Al~GIOSPERMS

A. General trends (Expanded from Smith, 1967)

Primitive Advanced

1. Tropical plants Temperate plants

2. Hoody pl ants Climbing or herbaceous plants ,
3. Wood homoxylous (vesselless) \\!ood heteroxylous (vesselled)
4. Perennial habit Biennial or annual habit
5. Terrestrial habit Aquatic, epiphytic, saprophytic
or parasitic habit
6. Vascular bundles in cylinder Vascular bundles scattered
(Dicotyl edons) (Monocotyledons)
7. Chlorophyll present Chlorophyll absent
8. Evergreen plants Deciduous-leaved plants
9. Stipules present Stipules absent
10 . Node unilacunar Hith two traces Node variously altered
11 . Leaves spiralled Leaves opposite or whorled
12. Leaves simple Leaves compound
13. Flowers hermaphrodite Flowers unisexual
14. Fl owers solitary Flowers in inflorescences
15 . Flm..rers entomophilous FloHers anemophilous
16 . Floral parts spirally imbricate Floral parts whorled or valvate
17. Flowers polymerous Flowers oligomerous
18 . Perianth undifferentiated Perianth differentiated into calyx
and corolla , or further reduced
19. Flowers with petals Fl m..rers \..rithout petals
20 . Petals free Petals connate
2l. Actinomorphy Zygomorphy
22 . Hypogyny Perigyny, epigyny
23 . Stamens many Stamens few
24 . Stamens separate Stamens connate
25 . Stamens 3- veined microsporophyl1s Stamens various l y modified
Hith linear embedded sporangia
566 27

26. Pollen monocolpate (or derived) Pollen trico l pate (or derived)
27. Carpels many Carpels few
28. Carpels free Carpels connate
29. Carpels conduplicate . unsealed . Carpels variously modified
styleless
30. Placentation laminar Placentat ion submarginal to axile
to parietal, etc.
31. Fruits single Fruits aggregate
32. Fruit a fo llicle Fruit a capsule. berry , drupe,
etc.
33. Seeds large; embryo small ; endo- Seeds small ; embryo ~"ell devel-
sperm ahundan t oped; endosperm little or none
34. Cotyledons 2 Cotyledons 1 or 3 (or more)
35 . Anatropous ovules Other types
36. Two integuments One integument
37 . Nuclear endosperm Cellular or helobial endosperm
38. Crassinucellar ovule Tenuinucellar ovule
39. Long, slender sieve cells with Shorter sieve tube elements Hith
scattered sieve areas specialized sieve plates
40. Chromosome number low; n = 7 Higher numb ers

B. Sporne (1954, 1971) has discussed the statistical correlation between

floral and vegetative characters in the dicotyledons . According to him
the most reliable indicators of primitiveness are the folloHing :

1. Woody habit 12. Nuclear endosperm

2. Presence of secretory cells 13 . Carpels free
3. Leaves alternate 14. Axile placentation
4. Stipules present IS. Unisexual flower
S. Flowers actinomorphic 16 . Leuco- anthocyanins present
6. Petals free 17 . Small number of seeds
7. Stamens pleiomerous lB . Scalariform perforation plates
B. Carpels pleiomerous 19 . Scalariform intervascular
9. Seeds arillate pitting
10. Two integuments 20 . Apotracheal wood parenchyma
11. Integumentary vascular bundles 21. Unstoded ~'lOod
22 . Pollen binucleate

LITERATURE ON PHYLOGENETIC DICTA

Bailey , 1. H., and H. H. Tupper. 1918. Size variations in tracheary cells.

I. A comparison between the secondary xylems of vascular cryptogams , gym-
nosperms , and angiosperms. Proceedings American Academy of Arts and
Sciences 54 : 149-204 .
Bessey, C. E. 1915. The phylogenetic taxonomy of flm.;rering plants . Annals
of Missouri Botanical Garden 2 : 109-164 .
Frost , F. H. 1930a. Specialization in secondary xylem in dicotyledons . I .
Origin of vessels. Botanical Gazette 89 : 67-94 .
1930b. Specialization in secondary xylem in dicotyledons. II.
Evolution of end wall of vessel segment. Botanical Gazette 90 : 198-212 .
1931. Specialization in secondary xylem i n dicotyledons . III.
Specialization of lateral wall of vessel segment. Botanical Gazette 91:
88-96 .
Smith , A. C. 1967 . The presence of pr i mitive angiosperms in the Amazon
Basin and it s significance in indicating mi grational routes . Atas do
Simposio sobre a Biota amazonica 4 (Botanica) : 37-59 .
Sporne, K. R. 1948. Correlation and classification in dicotyledons . Pro-
ceedi ngs of Linnaean Society of London 160 : 40-47 .
1954 . Statistics and the evolution of dicotyledons . Evolution 8 :
55 64 .
1956 . The phylogenetic c l assification o f th e angiosperms . Bio-
logical Reviews Cambridge Philosophical Society 31 : 1-29 .
Thorne, R. F . 1958 . Some guiding principles of angiosperm phy l ogeny.
Brittonia 10: 72-77.
1963. Some problems and guiding principles of angiosperm phylogeny.
The American Naturalist 97 : 287-305 .
\~agner , H. H. 196 1. Problems in the classification of ferns . In : Re c ent
Advances in Botany 1, sect . 9 : 841-844.
1962 . The synthesis and expression of phylogenetic data . I n :
Benson , L ., 1962 , Plant Taxonomy, Methods and Pr inciples . The Ronald Press
Company . NeH York.

Section C. STRUCTURAL EVOLUTION

Since the fossi l record is incomplete, the evolutionary history o f angio-

sperms, for the most part , can only be inferred from data existing in extant
plants. The objectives of this section are to reconstruct a hypothetical pr im-
itive flol",ering plant from available info rmation and determine in which direc-
tion or directions structural modif ica tions have occurred .

I. VEGETATIVE MORPHOLOGY AND ANATOHY

A. Hab i t . The ancestral dicotyledonous angiosperm i s widely believed to

have been perennial, t",oody, and arborescent. The evidences for the primitive-
nes s of the woody habit has been summari zed by Constance (1955) : (1) the
prevalence of the woody hab i t in gymnosperms; (2) its correlation with anatomi-
cal a nd floral characters regarded as primitive and, conversely , the associa-
t i on of advanced anatomica l and floral characters tdth the herhaceous habit ;
(3) the predominance of twody plants in moist tropical and subtropical areas ,
habitats which are widel y bel ieved to be modern equivalents of the climatic
conditions widespread at the period of angiosperm inception; (4) the revela-
tion of the persistence of cambium in seedling and herbaceous stems; (5) the
s upposed correlation between f l eshy fruits and woody habits; and (6) the lack
of undoubted fossil herbaceo us angiosperms in older geologic strata. The
viny habit along ,,,,ith epiphytism , saprophytism , and parasitism are derived
conditions.

B. Secondary Xylem . During the past fifty years the value o f \..'ood
anatomy , and vessel element evolution in particul ar , in the study of the
phylogeny and classi f ication of angiosper ms has been c l early established .
In no other vegetative tissues o f the plant are t he trends of evolution as
clearly defined. These trends were recognized entirely without reference to
existing taxonomic systems and thus the relative primitiveness or advancement
o f the plants in which they occur . The trends of evolution of the ves s el ele-
ment have been firmly established thr ough extensive and intens i ve comparative
anatomical studies on both fossil and living material, statistical anal yses
(i . e ., measurements and correlations) , and ontogenetic studies . Therefore,
the evolutiona r y trend from tracheids to vessel elements is the most reliable
 568 27

..

"

...
HIt.lAN TANDRA

Figure 27-1. Stamens of Representative Hagnolialian Genera . (Canrigh t, J . E. 1952)

2
3

Figure 27 - 2. The Conduplicate Carpel of Tasmannia (Hinteraceae) . 1. Si de

view . 2 . Ventral view, sho\~ing the paired stigmatic crests . 3 . Tran sverse
section , showi ng penetration of pollen tube as three major carpellary bun-
dles. 4 . Cleared, unfolded lamina , showi ng vasculature. 5 . Unfo l ded
l amina , shm."ing placentation, distribution of glan dular hairs (stippled),
at the course of pollen tubes . (Bailey , I . W. , & B. G. L. Swamy. 1951)
27 569

too l knOI-.TIl in the study of phylogeny . This is true because this trend is both
largely unidirectional and irreversible .

It has been sholoJl1 that the angiosperm vessel element (in secondary l'lOod)
has been phylogene t ic ally derived from a scalari f orm-pitted tracheid by the
loss of pit membranes in the regions o f tracheid overlap. The refore , vessel
elements Hhich are most tracheid-l ike , viz ., e longate , narrOH , many scalari-
form perforations i n the e nd walls , etc. , are the most primitive type . Other
knmm major trends of vessel element specialization are : (1 ) change in per-
forati on pl ates from scalariform , many barred, and ful l y bordered to few
barred and borders ahsent to more advanced porous oblique and transverse por-
ous plates ; (2) angle of end 1·1811 highly oblique and tapering (much ove rlap)
to transverse ; (3) reduction in vessel e l ement length; (l!) change in appear-
ance on the transverse section from angular to circular ; (5) increase in pore
diameter ; (6) change in intervascular pitting from scalariform and bordered
to opposite and alternate multiseriate; and (7) change in vessel distribution
from solitary to extensive aggregate groupings o f vessels . Concomitant with
these changes in the vessel element are modifications in the f usi f orm cells
of the vascular cambium from long initials Idth overlapping ends that divide
in a pseudo-transverse manner to advanced initials I"hich undergo radio-
longitudinal divi sion and are reduced in length .

Although t he above trends of vessel evolution are believed to be firmly

estab l ished , Beck (1970) has pointed out that circular bordered pits had
evolved by Middle Devonian time and occurred in either the primary or secon-
dary xylem tracheids of coniferophytic progymnosperms and primitive pterido-
sperms . In his opinion the acceptance of the scal ariform bordered pit as
primitive in seed plants shou l d be reconsidered . To quote Beck, "In view o f
the early appearance of the circular bordered pit in the progymnosperms and
the preponderant occurrence of this type in the secondary xylem of cycadophy-
tic and coniferophytic gymnosperms , it seems not unreasonable to suggest that
the circular bordered pit might be the basic, primitive type in the gymnosperm-
ang i osperm line of evolution . This is not to say, necessarily, that the cir-
cular bordered p i ts of some angiosperms might not have evolved f rom the scalar i-
form pits of their ancestors, I-.'hether gymnosperm or ang iosp erm. In these
groups this mi ght be interpreted to have been accomplished through neoteny .
The circular bordered pits of other groups of angiosperms might simply reflect
their ances try from gymnosperms Hith circular bordered pits. "

Other I"ell known major phylogenet i c trends in the secondary xylem , as

r evealed by statistically significant correlations using fe atures of vessel
elements , include the evolution from tracheids to f ibe r - tracheids to advanced
libriform fibers throu gh a progressive increase in cell length (as compared
to vessel elements) \"ith a concomitant reduction in pit size and numbe r and
eventual elimination of the tracheid pit border . Primitive woods have both
uniseriate and multiseriate rays that are heterocellular and high-celled; the
multiseriate rays have long uniseriate wings. Specialization results in the
reduction a nd l oss of either the multiseriate or uniseriate rays (or both) ,
the rays becoming homocellular and reduced in height with the unise r iate
wings of multi seriate rays reduced to a single cell . The condition of strati-
fied rays formed by a storied cambium is an extreme advancement. Apotracheal
diff use axial parenchyma is regarded as the primitive state f rom I"hich more
advance d paratracheal types (vasicentr ic , aliform, confluent) were derived ,
hOHever . Bailey (1957) points out that the available data do " . . . not provi de
a single linear series of increasing structural specialization . " The phylo-
genetic significance of the absence of Iwod parenchyma is unclear.
570 27

Additional features of wood structure are believed to represent sporadi-

cally occurring divergent trends of specialization whose presence in Hoods
does not imply a close genetic r e lationship . These include helical thicken-
ings on the walls of tracheary elements and fibers , impe r forate tracheary
elements t~hich become septate or retain a living protoplas t, formation of
vascular tracheids and vasice ntric tracheids, excessive thickening of I-Jalls
and widening of vessel element diameter (as i n lianas), various reticulate
perforation plates or other modifications of perforation plates , and exten-
sive aggregations of vessels in diversified patterns (Ba i ley , 1957) . Further-
more , Carlquist (1962) has shown that primitive features of the primary xylem
may occasionally be extended i nto the secondary xy lem (paedomorphosis or juve-
nil ism) in plants that are othGnlise specialized .

The great value o f \.,rood anatomy as a guide to understanding angiosperm

evolution l ies not only in the f act that t ren ds to\,'a rd specialization are Hell
defined, but that all stages in evolutionar y specialization are observab le in
extant angiosperms . Therefore , there is nO L the slightest doubt that the
absence of vessels in eleven genera of ,.,rood y di cotyl e dons represents the
retention of a primitive feature . The la r g e ::;t number of vesselless genera
(7) occur in the \~interaceae, \,'herea::; the Trochodendrace ae, Tetrace ntraceae,
Amborellaceae , and Ch lorantllaceae contain a s ingle vesselless genu s each .
However , based on the sale evid e nce o f lac k o f vessels, these genera are not
necessarily the most primitive angiosperms , ';ince evolut ion proceeds indepen-
dently in all organs and tissues of plants. The absence of vessels in plants
belonging to families , as Cactaceae , clear ly the rp.sult of extreme special-
i zation (Bailey , 1966 ). The importance o f hOI.' plant habit and the factors o f
the environment affect Hood structure and the trends of xylem evolution have
been emphasized recently by Carlquist (1966, 1969a&b) . The singl e most impor-
tant guiding principle of \wod anatomy and phylogeny (Bailey, 1944) emphasizes
the impossibility (unlikelihood) that a taxon ,.,oith sp eci alized xylem gave ris e
direct l y to a taxon wi th less specialized x y l em . Accord i ngly , Hood anatomy is
more useful in negat i ons than assertions of positive relationship .

Other features of the secondary xyle m are diagnostic in so far as they

are restric t ed to ce rt ain families or genera; 1l0\",ever, thei r phylogenetic Sig-
nificance remains dubious. Distinctive tile ray cells are confined t o certain
genera of the Bombacaceae, Sterculiaceae, and Tiliaceae ",hereas upright sheath
cells located on the ffi<'Hgins o f multiseriate rays are o f comparatively I",ide-
spread occurrence . Vestured pits are reported for t",enty- four families of
dicotyledons (Bailey , 1933) h'ith a concentration in the Hyrtales. Solitary
prismatic crystals are relatively "'idespread, although such forms as raphides,
druses , and crystal sand have a more restri cted distribution , A.nd, therefore,
are of greater taxonomic interest (Hetcal fe and Chalk, 1950) .

C. Secondary Phloem . Unlike the xylem, information on the compara tive

anatomy and evolutionary trends in phloem tissue is meager (Esau et a!., 1953) .
For example, I",itness the absence of phloem data in Hetcalfe and Chalk (1950) .
The subject of phloem is thoroughly reviewed in Esan (1969) . Suggested trends
of specialization relate mostly to sieve element s in secondary phloem of
dicotyledons and metaphloem elements of mono co tyledons, and have by no means
been demonstrated to be unidirectional or irreversible . Composing the secon -
dary phloem are the principal conducting cells, the sieve elements, in com-
bination "'ith various amounts of phloem parenchyma , sclerenchyma , and ray
parenchyma . Hith regard to sieve elements. Cheadle and Hhitford (1941) recog-
nized t",o basic types , the sieve cell and sieve tube element . The sieve cell
27 571

is interpreted as a le ss s pecialized element 1"1th r e l atively undi ffere ntiated

sieve areas (i. e . • narrOH pores and thin connec ting strands), lacking highly
developed sie ve pIa tes and companion cells . t-lore adva nced sieve tube clemen ts
have specialized sieve plates on the end \·,a11s, have accompanyin g companion
cells . and are superimposed on one another in a longitudina l series forming a
sieve tube. Sieve plates may be compound, that is , composed of several highly
differentiated sieve areas, o r simple and composed of a single s i eve area .
Also , end \.;a11s ran ge in orientation from highly oblique to transverse.
Although the distinction between thes e cell types is not always sharp , gym-
nosperms and Im·,e r va sc ular plan ts possess sieve cel l s t.;he rea s aneiosperms
are characterized by sieve tube ele ments and asso ciated companion cel ls .

The sieve cell i s generally r e garded as the primitive type of conducting

e lemen t from \-fhich the si e ve tube el e ment a r ose . Najor trend s of sieve tube
element special ization (Esau et al. , 1953) are believed to involve (1) locali-
zation of specialized sieve areas on the end wa ll s , (2) change in orientation
of end walls from v ery oblique to transverse , (3) progressive change from com-
pound to simple sieve plates , and (4) progressive decrease in conspicuousness
of the sieve areas on the s ide walls . althoug h the r e lation betl_·een specializa-
tion of lateral sieve a r eas and the ::;ieve tube element is not consistent
(Zahur, 1959). Esau (1969) has s t "(essed that compa rat i ve data of lateral sieve
areas sho uld come only frOr.l Halls he tween con ti guous sieve elements . The same
general trends have also been ag rf'e d upon for monocotyledons (Cheadle , 1948 ;
Cheadle and lfuitford , 1941) . NeaB llrements o f cell len g th, width, and si ze of
sieve pore area has yet to demon st rate the s ame phylogenetic significance pro-
posed for tracheary tissue. HOh!f'V@ r , Zahu r (19 59) i s of the opinion that there
has been a phy logenetic decrease i n sieve tube el(!;ment l e ngth (realizing sec- ,
on dary septation can occur) accomp anied by decrease in the length of the end
Hal l s and increase in size of si eve plate po r es . Follol.. ing ea rlier workers .
Zahur classif i e d sieve tube element s into three categories of increaSing
specialization : I, elements long (l+00 )Jm or more), with very oblique sieve
plates containing 10 or more sieve areas ; II . e l ements of medium l e ng th, ~.'ith
oblique sieve plat es co ntainin g 2 to 10 sieve areas; a nd III, element s short
Hith slight l y ob lique to t r ansvers e , s imple sieve plates . It is clear that
there is ra t he r uni form agreement th a t the mos t primitive sieve tube element s
are ones tha t most resemble sieve cells . As far as is known , unspecialized
sieve cells occur in only one angiosperm, Austrobaileya scandens ( Aus t ro-
baileya ceae ) (Bailey and S"amy , 1949 ; Srivastava , 1 970) .

Zahur (1959) also classified companion cells lnto types refle cting ascend-
in g evo lut ionary advancement. Type A are cel ls that are much shorter than the
sieve tube elements and occ ur singly, usually at t h e corner s o f sieve type ele-
ments. Type B are cells as long as the sieve tube e l emen t. and Type Care
cells that are septate in add ition to being as long as the sieve t ub e ele-
ment. Zahur found companion cell type to be con s t ant within families and ,
occasional ly, orders \.;ith the greilt+?st variability occurring in families \-lho se
"naturalness " has been questioned .

Al tho ugh indecis ive, Zah ur regarded abundant , extrem(!; l y variab l e and
irregular pa renchyma to be advanced ,.,..hereas scanty pa r enchyma showin g litt l e
variation in size probably represented a primiti.ve cond ition because of its
association \-li t h polypetalous famili e s . Zahur could fi nd no bases for decid-
ing I~heth e r the pres e nce or absence of phloem sclerenchyma \-las structurally
primitive or advanced . lIowever, Eames (1961) regarded the prp.s ence of phloem
fibers to be an advanced state .
 572 27

D. Nodal Anatomy. Since the first comprehensive study of nodal struc-

ture and evo lu t i on by Sinnot t (1914), nodal anatomy has assumed much impor-
tan ce in discu ssions r el ating t o a n giosperm phyl ogeny . l-Thech e r the p rimitive
nodal pattern was trilac unar (Sinnott, 1914) , multilacunar (Ozend a , 1949) ,
unilacunar, two-tra ce (~~rsd en and Bailey , 1955 ; Canrigh t . 1955), or tri-
lacunar or multilacunar \·lith two traces diverging f r om the median gap
(Takhtajan . 1 969) has yet to be aosHercd . The primary evidence used to
support the primitive nature o f the 2 : 1 nod e are its occurrence (or modifi ca-
tions thereof) in members of the ~mg n oliidae (Calycanthaceae , Lactoridaceae ,
Ill iciaceae , Amhorellaceae , Laur aceae , Chloranthaceae , Nonimiaceae , Austro -
baileyaceae. Schisandraceae, a n d Trimeniaceae) , t he distribution of this type
i n vascular plan ts other than angiosperms , and its presence i n t he cotyledon-
ary nod e o f many dicots even though the mature foliar nodes of the s ame plant
may be t r i - or mult i1acunar ( i . e ., cotyle donary nodes are conservative and
thu s reflect ance stra l conditions) .

Recen t papers by Conde and Stone (1970) a nd Stone (197 0 ) have des cr ibed
an amazin g d i ver sity of cotyledonary nodal patt e rns in the Jugl a nd aceae in
which nodal evo lu tion appears correlated wi th functional demands of the seed-
ling . These authors \o,Iere able to confirm the 2 :1 cotyledonary node as primi-
tive in Jug1andaceae and s uggested the diversification to mor e comp l e x pat-
terns as func tionally related to independent shifts "'!thin the family from
epigeal to hypogea1 seedl ings . Furthermo r e , a sugges tion was made that the
apparent conservatism of cotyledonary node s i n dicotyle dons as a who le may be
directly related to the prevalence i n many g r oups of primi tive , ep igeal germi-
nation , and thus , co tyle donary nodal ana t omy should be considered independent l y
of the mature fo l iar nodes .
,
Although mo st fami l ies tend to have a uniform nodal anatomy, some fami l ies
and even gene r a have variable nodal struc ture . Tri l acunar n odes occur in the
majority of dicotyledon s (Sinnott, 191 4) wher eas mul t ilacunar nodes are r e l a -
tively uncommon in dicotyledonous fami lies ( Howard , 1970), but are fo und in
members of primitive orders such as Hagnolial es, Piperal es , and Trochoden-
drales , and advanced orders such as Umbellales and Aste rale s . The uni lacunar
node has an interesting dist r ib u tion , namel y the Laurales ( sensu Takhtajan) ,
Caryophyllales. Ericales , Diapensiales. Ebenales , Primulales , Myrtales , and
a majority of the families in the Asteridae . Some orders show transit i ons in
nodal stru ct ure (e. g ., the Theales) where Schofield (1968) described t wo pos-
s i ble lines of evolution or i g in ating f r om the a n cestral tri lacunar conditio n
in the Dilleniaceae. One l i ne maintained the t r i l acuna r node , and event ual
amplificat i on culminated in the multilacunar node of the Car yocaraceae . The
second line involved reduction to the unil acunar nodes of the Harcgraviaceae
and Theaceae .

Namboodiri a n d Beck (1968) have recent l y proposed t ha t the euste le of

gymnosperms evo lved direc t ly from a protos t ele by g radual medullation and
l ongitudinal dissection followed by formation of a cylin de r of longitud i n al
sympodia l bundles from which l eaf traces diver ge radially . In this inter-
pretation the primary vascular system o f seed plants is not directly compar-
able to that of ferns so that the classic filicinean-type l eaf gaps may not
occur in e usteles. This concept has a l so been extended to angiosperms ( Slad e ,
1971; Devadas and Beck, 1972). Benzing (1967) concl uded that t h e unilacunar ,
one-trace o r t r i l ac unar. three-trace noda l patt e rn was probably primitive .
27 573

E . Leaves. Although anatomical leaf characters are generally useful in

the delimitation of taxa , the majority of these characters cannot be readily
interpreted ecologically or evolutionarily. Suggested trends of foliar evolu-
tion have generally not been supported by convinc ing evi dence . The ancestral
angiosperm is envisioned as having alternate, s imp l e , entire, stipulate , and
pinnately net-veined leaves . Thi s is one modern d i ctum which is markedly
opposed to Bessey ' 5 original opinion s .

Trends of evolution of stomata have long been unclear. Baranova (1972)

has recently advocated para c ytic typ es as primitive \,,1th1n the angiosperms on
the basis of the nearly uniform occurrence of this pattern in the Magnoliales
(sensu Takhtajan). The on l y taxa of the order that do not have this stomatal
pattern are Liriodendron tulipifera (Hagnoliaceae) and Bubbia penied (Inn-
teraceae) , both advanced in their respective fami lies. Corre l a ted with para-
cytic stomata in the Hagnoliales are thickened l amel l ae on the outer walls of
t he guard ce ll s a nd t hick- walled epidermal ce lls wi th pores i n the outer wall.
both characters are regarded by Baranova (19 72) as primitive in angiosperms
as a whole . Since i t is I... ell known that simi lar mature stomatal patterns can
arise by differen t developmental pathways , an ex tension of this work ut i liz-
ing the ontogenetic clas sification of stoma tal types by Fryns-Claessens and
Van Cot t hem (1973) t.,rould be desirable. It should also be noted that paracytic
s tomata are found in a large number of very unrelated and often advanced
families .

Preliminar y data by Hickey (1971) show that families otherwise regarded

as primitive tend to have venation patterns i n which the secondary veins have
an irregu l ar course i n th e lamina, and intercostal areas lack uniform size and
s hape, l.Jhereas more advanced fami l ies tend to have more re gular venation pat-
terns I ... hich fo llm... cons istent courses . The phylogenetic s i gnificance of the
open dichotomous venation in the only two angiosperms which possess this fea-
ture--Kingdonia and Circaeas t er - -remains un clear (Foster , 1961). Cronquist
(1968) regarded this as a modification that simulates a primitive pattern .
Although there is conside rable anatomical diversity in petiolar vasculation ,
t rends of evolution in this region of the plant are not r eadily discernible
in angiosperms as a l.Jhole. Also , phylogen e t ic trends in t richomes for angio-
sperms as a I~hole have been sugges ted (see Carlqui st , 1961) but they have not
had significant influence on phylogenetic thought.

II. FLORAL HORPHOLOGY AND ANATOMY

Al though the flOl.Jer has histori ca lly been a princ ipa l source of taxonomic
data , a nd the literature dealing uith floral morphology and anatomy is volumi-
nous , there has not been agreement on a I~orld-wide basis r egar ding interpre-
tations of floral morphology , anatomy (Which is often us ed as synonymous with
floral vascular a natomy) and evolution of floral st r uetures . The following
treatment presents some of the bas ic tenets and eoneepts of floral morphologi-
cal study .

A. Taxonomie and evolutionary val ue and interpretation s of floral mor-

phology . Among the c haracters of floral morphology of systematic importance are
the vascular supp ly, i.e ., number of traces and nodes, elaborateness of vas-
cularization, position of microsporangia and ovules in r elation to the vena-
tion, a n d sta ges of morphological alterations that result in syncarpy , eohe-
sio n, etc . (Carlqui st , 1961) . The classic interpretation of the basic nature
of the vascular suppl y to the floral parts is (a) s epals are 3-trace structures;
574 27

(b) petals are I-trace structures ; (c) stamens are l-trace structures; and (d)
car pels are 3-trace structures. Two fundamental premise s regarding the nature
of the floral vascular system that have long been a cornerstone o f anatomical
investigation have been outlined by Hoseley (1967) . First , the vascula r sys-
tem is nearly always more conservative than the organs it supplies. Changes
in the vascular patterns tend to lag behind morphological alterations ; hence ,
the nature of the changes, often not discernib l e by gross morphological stud-
ies , may be revealed by an examination of the internal characters of the vas -
cular anatomy, such as: nodal lacunae, bundle orientation , and vestig ial
traces. Secondly , the pattern of the vascular system may reveal the former
boundaries, relative positions , numbers, and categories of organs , or the ir
parts , which may nm" be obscured by reduction, cohesion , and adnation. I n
addition , there is wide spread belief that a condition of fused vascular bun-
dles, in general , is more advanced than a condition of un f used bundles . Eyde
(1971) has stated that the value of flora l morphological study to taxonomy and
phylogeny l ies in the fact a consideration of flora l vasculature may engender
an extra measure of confidence when external features are difficult to inter-
pret , and in the fact that there i s a general evolut ionary tendency among the
principal floral vascular bundles to unite in various ways. Also, this author
emphasizes that similarities and differences in floral vascular pat terns can
be taxonomically significant characters even though they offer no plausible
evolutionary scheme. Car l quist (1970) and Schmid (1972) have recent l y
reviewed various concepts of floral morphological study.

B. Morphological nature of the flovler . The classi cal theory of the

f l ower can be traced back to the ideas of Goethe in the latter part of the
18th century . This theory states the floHer is a determi nate branch or shoot
whose parts (bracts, sepals, petals, stamens , and carpels) are modified leaves .
That is , floral parts and leaves are homologous. According to this idea the
general trends in floral organography include :

Primitive Advanced

1. Simple bisexual flm"er Compound un i sexual modified f l ower

(Numerous arguments have occurred in the literature as to whether the
primitive f lol"er was unisexual or bisexual. The generally accepted
vie\-! today is that the primitive flower was bisexual.)

2. Rad i al symmet ry (actinomorphy) Bi l ateral symmetry (zygomorphy)

3. Numerous parts spirally arranged Fewer parts with whorled arrange-

ment
(Stebbins [1967] has pointed out that evo l u tion of floral parts has
proceeded both upI,rard and downward in number. Thus, the general
t r end tmmrd fe\-!er fl oral parts is not always valid . )

4. Floral parts free Flor al parts fused (connation and

adnation)
5. Undifferentiated pe rianth Differentiated perianth or complete
absence of perianth
6. Ovary superior (hypogynous) Ovary inferior (ep igynous)
(The general trend from hypogyny to epigyny is well established in
flm.;ering plants. The first documented reverse change--from epigyny
to hypogyny--has been reported in Tetraplasandra gymnocarpa (Aralia-
ceae) by Eyde and Tseng [1969].)
27 575

Two major theories have been suggested to exp lain the inferior ovaries of
angiosperms. It is apparent that the inferior ovary has evolved several times
among different groups and available evidence suggests that this has oc curred
by two opposing methods . The appendicular theory essentially states tha t
there has occurred an extensive fus ion (both connation and adnation) of the
out er floral I.horis to one another and to the ovary wall. Evidence is derived
from the course of the vascular bundles supplying the ovule s . The ovular vas -
cular s upply o riginates in acropetal fashion moving up t he ovary and does not
descend to supply the ovules . Indications are this type o f inferior ovary is
the prevalent typ e in angiosperms. The receptacular theory, in contrast ,
assumes the carpels have "sunk" into the tissue of the floral receptacle.
Evidence is seen in the descending course of the ovular vascular supply.

Nectaries almost certainly have arisen and evolved independently in dif -

ferent groups of angiosperms. Fahn (19 53) has suggested tha t the re has occurred
a general trend among angiosperms for the floral nec t aries to migrate in an
acrocent ripetal manner : in the most primitive cases the nectaries are borne
on the perianth parts . As evolutionary advancemen t occurs the nectaries migrate
to the inner floral parts and in the pistil from the base of the ovary to the
base of the style .

C. Inflorescence. The evolution of inflorescences is poorly understood.

A few authors have traced lines of specia liza tion within ind ividua l t axa but
the general trends fo r angiosperms as a whole are only speculative. At one
time or anothe r solitary flowers , dichasial cymes , and panicles have been
regarded as primitive types. The widely accepted current idea is that the
solitary flower represents the primitive si tua tion in angiosperms as evid enced
by the presence of this character i n various "ranalean" genera .

D. Perianth . The currently widely accep ted interpre t ation is that t he

primitive dicotyledonous flm"e r possessed an undifferentiated perianth whose
component parts (tepals) were thick and foliaceous in nature. Differentiation
of the perianth into calyx and corolla probably took place at many diffe ren t
times.

Anatomical evidence (predominance of three vascular strands) indicates

that sepal s are most likely the equiva l ent of sterile bracts. Sub tending
floral bracts are of foliar origin whereas petals have app arently arisen in
two ways. In some flowers petals have arisen from modification from bracts
or tepal s , in which case sepal s a nd petals are morphologically eq uivalent.
In other plants petals have been derived f rom sterile stamens (staminodes)
and thus are not morphologically equivalent to sepals and bracts.

E. Carpels . Najar opposing theories on the morpho l ogical natur e of the

carpel fall i nto three major categories : (a) appendicular (foliar) theories
based on the sporophyll concept ; (b) axial theories, which are based on th e
concept that ovules were originally,and in some instances s t ill are. axis-
borne (cauli ne) ; and (c) sui generis theories, \"hich state the flo r al parts
do not find homologies in ei ther leaves Or stems but are entirely new enti-
ties.

1. Classical theory. The carpel is considered fundamentally foli ar in

nature. The primitive carpel is envisioned as a 3- veined (dor sal and t\"O
ventra l s) megasporophyll with a lobed stigmatic crest and numerous ovules
borne along its in turned margins. The ovules are vascularized by traces
derived from the ventral bundles .
576 27

2. Conduplicate theory . This conc ept. which resulted from stud ies on the
primitive dicotyledons (genus Tasmannia), is the most widely accepted among
American botanists. The primitive carpel i s described as cladified , stipi-
tate, condupl i cate, open, 3-veined and I.,.ith many ovules borne on its adaxial
surface . Advancement produces a carpel that is unc l adi fi ed. closed , with a
definite raised stigma, and with few (1- 2) ovules adaxi ally or basally attached.

3 . The "phyllospory-s t achyospory" theory . This interpretation states that

the appendages commonly called carpels are of two basic na tures : fol iar and
cauline . Of primary consideration is t he relationship between ovul es, placen-
tas and other carpel pa rts. A fundament al d i stin ct i on is made between the
classic idea in \~hich margi nal ovules and placentas are par ts of l eaves (phy 1-
losporous condition) and the recognition of situations where the ovule and
placenta are axillary cauline structures sub tended by a sterile appendage
(stachyosporou s condition ) . The term s tegophyll is used for steril e struc-
tures that may surround and thus protect stachyosporous ovules .

4. The Peltate Carpel. This concept has been widel y accepted among non-
English speakin g botanists but has received litt l e attention in Ame rica.
According to Eames (1961) , " The pel tate theory presents a morphological ori-
gin for the carpel more comp lex than the simple folding or inrolling of the
margins of the lamina. Pe lt ate form in the carpel is assumed to have arisen
by the turn ing upward (ventrally) of the basal lobes of the lamina and their
fUSion, margin to margin, as in t he forma t ion of pe l ta te l eaves . Whe r e the
t\\/O marginal meristems mee t , they unite, forming a t r ansver se meristem, t h e
cross zone ." The peltate carpel is thus considered homo l ogous with leaves ,
and ovules are borne on the I.al l formed by the cross zone. Vasculat i on i s
by a dorsal - median bundl e and a ventral-median one . The latter is the result
of fusion of two ventral bundles of the classic carpel.

5 . The Gonophyll theor y . According to this theor y the basic component of

the ovary is a s t erile leaf with an e piphyll ous (adna te fe r til e sporangium-
bearin g) branch . This e n tire comp l ex is termed a gonophyl!. The s terile leaf
is a tegophyll and its fertile branch system an androphyll (if male), gyno-
phyll (if female) or androgynophyll (if bisexua l ) . Repetition of the bas ic
gonophyll unit gives an " i n florescence ." Diverse floral struct.ure is explained
by "modifications " of t he basic gonophyll plan. Belville (1962), for example ,
states , "All superior types of ovary examined could be f ormed by a s imple
i nfolding, or dOlffi-folding , of t he gonophyll blade to effectively enc l ose the
ovuliferou s branches . 11 This idea has f aile d t o gain wide s upport .

6 . Meeuse ' s t.heory . This theory has as its basic premise the idea th at
angiosperm pistil s are the homologues of ovuliferous cupules . The cup ule is
r egarded as an organ of independent origi n . The spo rophyll or foliar na ture
of the carpe l is adamantly rejected .

7. Theory of Carpel Pol ymorphism. The unspecia l ized carpel has t hree
functions -- it is receptive , reproductive and protec t ive . Evolut i on has l ed to
the specializ ation of different forms of carpels for different funct ions .
Three forms of carpels may be therefore recognized : (1 ) valve carpel , ( 2)
solid ca r pe l, and (3) semi-solid or pseudovalve carpe l. This division of
labor means that there are usually at least two types of carpels pres ent in
a n yone ova r y . The theory of carpe l polymorphism , as proposed by Saunde r s
(1925) , has been widely criticized and is not gen era lly accepted as valid.
 S77

F . Placentation . Altho ugh the taxonomic value of placentation type is

clear, int e rpretations of the phylogenetic trends within the various pat terns
has been considerably less so . In part , this confusion has been the result
of inconsistent use of terms . Correlated Hith the belief that the condupli-
cate carpel is primitive, ther e is wide - shared belief that the laminar condi-
tion of the ovules is the most primitive pattern in apoCarpOll$ gynoecia . tn
more advanced syncarpous gynoecia , axile placentation appa rently gave rise to
al l other types.

C. Stamens . The classical (ranalean) th eory regards the stamen as fol ia r

in nature . The primi t ive s tamen of the classical school is exemplified by the
stamens of Degeneria (Degn er ia ceae) . Hagnol ia (Magnoliaceae), IHmantandra
(llimantandraceae) and others . The primitive microsporophyll is characterized
as cJ.adified, 3- veined, Idth linear sporangia embedd ed between the veins on
the abaxial or adaxia l s urface. The advanced stamen is uncladified. I-veined ,
with clear differentiation into anther and s l ende r filament.

The telome theory envisions the origin of the stamen from a much reduced
bran ch system. Evidence is found in the androecial vasculation of some flow-
ers in whi ch a strong vascular bundle (stamen trunk) constitutes the central
axis of a telome system which supplies seve r al stamens by branches . This
th eory maintains the flattened stamens of some "rana lean" genera can be
explained through a reduc t ion and webbing of a branch system .

The sequence of stamen differentiation has in recent years been in terpreted

as having phylogenetic significance. Although the observation of " cen trifugal
sramens" or the "centrifugal androecium" has been reported for over one hundred
years. E . J . H. Corner (1946) was the first to ful l y deve l op its potential taxo-
nomic significance . It I"as his idea that those plants with cen tr ifugal stamen
i nitiation represented an evolutionary line quite di stinc t from those with the
more common centripetal pattern. In Cronquist ' s system, th e Dilleniidae are
characterized by cent rifugal stamens whereas the Rosidae have centripetal sta-
mens.

H. Pollen . It is I~idely believed tha t the mono colp ate pollen grain with
a long di stal aperture is the primitive condition in angiosperms . Evidences
a r e seen in the occurrence of this type of grain among gymnosperms , its distri-
bution in the fossil re cord , and its persistence in such other\"ise comparatively
primitive genera as Nagnolia and Degeneria . Outside thp. " ranalean" complex this
type of grain is not found in dicoty l edons. although it is the character istic
pollen form in th e monocoty ledons . The general trend in pollen evolution is
from lar ge , relatively unornamented, monocolpat e polle n I"ith a long , distal
aperture to smaller, ornamented, tricolpate and derived types. A persistent
problem with thi s trend was hOI" to explain the origin of the trico!pate condi-
tion from a monocolpate ancestor . Evidence now favors at least two origins of
the tricolpate condition-- (1) the furrows of the tricolpate grain arose de novo
from inaper turat e precursors; and (2) the trico lpate grain arose f r om a mono -
colpate grain through a trichotomosulcate intermediary .

I . Embryology . Interpretations of the angiosperm ovule have been numerous

and often widely divergent . As of yet no one interpretat i on seems to satis-
factorily account for the morphology of this st ructure. The primitive angio-
sperm ovule is describ ed as bitegmic , anatropous , and cr ass inuce llate primarily
as a result of th e prevalence of this type among plants otherwise regarded as
primitive . The uniformity of the mature emb r yo sac is certainly one of the
 578 27

most distinguish ing featu r es of fl Ol-Jering plants . The monosporic, eight-

nucleate (Polygonum type) is regarded as the primitive embryo sa c condit ion
because : (1) it is the most common pattern among angiosperms ; and (2) the
largest number of divisions are involved in its formation, i .e., there is a
general trend toward reduction in gametophyte size and development. There
is no general agreement as to whether the primitive endosperm type l"a8 Ce llu-
lar, Nuclear , or Helobial .

J . Fruit and Seed . Ideas regarding the evolutionary trends in fruits

and seeds are not \.:0211 supported by morphological data . Various types of
seeds and fru its have evolved many times in many different evolutionary lines .
The current general assumption is that the primitive seed Has large and had
an undi ff erentiated small embryo embedded in abundant endosperm . Takhtajan
(1969) also envisioned the primitive seed as possessing a seed coat of the
"Magnol i an-type, II i . e . , with a strongly developed peripheral layer of paren-
chymatous cells . Some botanists , including Corne r (1949) in his famous Durian
Theory, considered the ancestral seed to possess a bright l y colored red or
yel lm. . aril (this \. . as associated \. . ith a fleshy fru it and short s eed viability) .
Martin (1946) regarded the time of inception of dormancy as significant with
primitive seeds cha r acterized by early dormancy or at least prior to extensive
embryo development . Eames (1961) belie ved the absence of dormancy was primi-
tive . The primitive f ruit i s usually descr ibed as a many-seeded f ollicle that
opens at maturity.

K. Relationship of plants and insects . Many botanists believe that the

ancestral angiosperms \. . e re pollinated by beetles (cantharoph ily ). Evidence
for this is seen in the existence of this relationship among primitive fami-
lies (e. g ., 1'1 agnoliaceae, \~interaceae , Annonaceae, Eupoma tiaceae , Calycantha-
c:eae , etc.) and in the long geologic his t ory of b e etles. From the cantharo-
philous condition more advanced insect pollination relationships and Hind
po ll ination arose. E . E . Leppik (1957) has stressed the importance of con-
Sidering the parallel evo lut ionary development of flol-'ers and insects , i . e . ,
pollinators. This author recognizes six evolutionary levels of floral deve l-
opment . At the primitive level flOlvers do not possess any special symmetry
or particular colors. Numerous stamens and carpels are often su r rounded by
a large number of bracts . Insects do not require any special sensory abili-
ties to distinguish these flow ers . At the simple level flm"ers are slightly
more advanced and poss e ss simple colors such as yelloH, ,. . hite, or greenish .
There is no definite symmetry or number of parts . Thes e floHers are visited
primarily by beetles and flies . The third level is distinguished by floHers
that are act i nomorphic and oE definite size . Floral parts are in definite
numbers and leve ls . Colors are frequently pure , like ,. . hite , blue, red , and
yellOlL The pollinators oE these floHers presumably have the ability to dis-
tinguish colors other than the simple ones . On the numerate level flowers
show distinctive figure numerals. Former radiate flm. . ers are characterized
by a reduction in number of petals . "Protected" level floHers have fea tures
of the simple and third levels but contain protec ted nectaries . Pollinators
of these floNers mus t have the ability to d is tinguish three-d imensional pat-
terns in order to locat e the nectar. Flo,. . ers on the highest level are typi-
cally zygomorphic with various combinations of odor and color .
27 579

LITERATURE ON STRUCTURAL EVOLUTION

lhdley, 1. H. 1933. Structure, diHtributi,on and diagnostic signific ance of

vestured pits in dicotyledons . Journal Arnold Arboretum 14 : 259-273 .
1944 . The development of vessels in angiosperms and its signi fi-
cance in morphological research . American Journal of Bota ny 31 : 421-4 28 .
1953 . Evolution of tracheary t i ssue of la nd plants . American
Jou rnal of Botany 40 : 4-8 .
1957 . The potentialities and limitations of ~lOod anatomy in the
phylogeny and classification of angiosperms . Journal Arnold Arboretum 38 :
243-254 .
1966 . The significance of the reduction of vessels in the Cacta-
ceae . Journal Arnold Arboretum 47 : 288-292 .
and B. G. L. S\~amy . 1949. The morphology and relationships of
Aust r oba i l eya . Journal Arnold Arboretum 30 : 211-226 .
_--,;-;:::-,' and 1951. The condup1 i cate carpel of dicotyledons and
its initial trends of specialization . American Journal of gotany 38 : 373-
379 .
Baranova , M. 1972. Systematic anatomy of the leaf epi dermis in the r-lagnolia-
ceae and some related families . Taxon 21 : 447-469 .
Barghoorn , E. S . 1940. The ontogenetic development and phylogenetic specializ-
zatioll of rays in the xylem of dicotyledons. 1. The primitive ray s t ructure.
American Journal of Botany 27 : 918-928 .
Beck , C. 1970 . The appearance of: gymnospermou s strllcture . Biological Reviews ,
Cambridge Philosophical Society 45: 379-400 .
Benzing, D. 1967 . Devel opmental patterns in stem primary xylem of woody
Ranales . American Journal of Botany 54 : 805-820.
Brown , \~ . H. 1938. The bearing of nectar ies on the phylogeny of flowering
plants . Proceedings of the American Philosophical Society 79: 549 - 595 .
Canright, J . E . 1952. The comparative morphology and relationships of the
Hagnoliaceae . 1. Trends of r,;pecialization in the s tamens . American Jour-
nal of Bo tany 39 : 484-497 .
1953 . The compar ative morphology and relationships of the Hagno-
liaceae - II. Significance of the pollen . Phytomorpho10gy 3 : 355-365.
1955 . The comparative morphology and relationships of the Nagno-
1iaceae : IV . \·Jood and nodal anatomy. Journal of the Arnold Arboretum 36:
119-140.
Cariquist , S . 1962 . 1'1 theory of paedomorphosis i n dico tyl edonous Hoods .
Phytomorphology 12 : 30-45 .
1966 . 1·lood anatomy of Compos itae : a SUllunary , ,·,ith comments on
fac tors controlling wood evolution . Aliso 6 : 25-44 .
1969a . \olood anatomy of Lobe1ioideae (Campanu1aceae). Biotropica
1, 47-72 .
1969b. I~ood anatomy of GoodeniaceaC! and the problem of insul ar
\·lOodiness . Annals ~!issouri Botanical Garden 56 : 358-390.
1970 . Tm.Jard acceptable evolutionary inte rpretations of flo ral
anatomy . Phytomorphology 19 : 332-362 .
Cheadle , V. I . 1948 . Observations on the phloem in the Honocoty1edoneae I I .
Additional data on the occurrence and phy l ogenetic specialization i n s truc-
ture of the sieve tubes in the metaph1oem . American Journal of Bo t any 35 :
129-131-
___,--~ ' and N. B. Hhitford . 1941. Observations on the phloem in the Hono-
cotyledoneae . 1. The occurrence an d phylogenetic specialization in stru c-
ture of the sieve tubes in the metaph1oem . American Journal of Botany 28:
623-627 .
580 27

Conde, L. F., and D. E. Stone. 1970. Seedling morphology in the Juglanda-

ceae , the cotyledonary node. Journal Arnold Arboretum 51 : 463-477.
Corner, E. J . H. 1946 . Centrifugal stamens. Journal Arnold Arboretum 27:
423-437 .
1949 . The durian theory of the origin of the modern tree. Annals
of Botany 13 : 367-414 .
Devadas, Co , and C. B. Beck . 1972 . Comparative morphology of the p r imary
vascular system in some species of Rosaceae and Leguminosae. American
Journal of Botany 59: 557 - 567.
Doyle, J . A. 1969. Cretaceous angiosperm pollen of the Atlantic coastal
plain and its evolutionary significance . Journal of the Arnold Arboretum
50: 1-35 .
Eames , A. J. 1931. The vascular anatomy of the flm.,rer vith refutation of the
theory of carpel polymorphism . Ame r ican Journal of Botany 18 : 147-188.
Ehr e ndor fer , F ., F. Krendl , F. Habe1er, and H. Sauer. 1968 . Chromosome num-
bers and evolution in primitive angiosperms. Taxon 17 : 337-353 .
Esau , K. 1969 . The Phloem. Encyclopedia of Plant Anatomy. V. (2) . Gebrlider
Bo rntraeger. Berlin .
_ _~~' V. 1. Cheadle and E. H. Gifford , Jr. 1953 . Comparative structure
and possible trends of specialization of the phloem. American Journa l of
Botany 40: 9-19
Eyde, R. H. 1971 . Evolutionary morphology : Distinguishing ancestral struc-
ture from derived structure in flm"ering plants . Taxon 20 : 63 - 73 .
_---,,-__ ' and C. C. Tseng . 1 969. Flover of Tetraplasandra gymnocarpa :
Hypogyny I.JHh epigynous ancestry. Science 166: 506 508.
Faha, A. 1953 . The topography of the nectary in the £Imler and its phylo-
genetic trend . Phytomorphology 3 : 424-426.
Foste r, A. S . 1961. The phylogenetic significance of dichotomous venation
in angiosperms . Recent Advances in Botany 2 : 971-975 .
Frost, F . H. 1930 . Specialization i n secondary xylem in d i cotyledons . I.
Origin of vessels . Botanical Gazette 89 : 67-94 .
1930 . Specialization in secondary xyle m in dicotyledons. II .
Evolution of end ~.Ja11 of vessel segment . Botanical Gazette 90: 198-212 .
1931. Specialization in secondary xylem in dicotyledons . III .
Specialization of lateral I"all of vessel segment . Botan ical GRzette 91 :
88-96 .
Fryns-Claessens, E., and W. Van Catthem . 1973. A nev classification of the
ontogenetic types of stomata . The Botanical Reviel.J 39: 71 - 1 38 .
Guedes , M. 1971 . Carpel pe1tation and syncarpy in Coriaria r usicifolia L.
The New Phy tologist 70: 213-2 27 .
Hickey , L. J. 1 971. Evolutionary significance of lea f architectural fea -
tur es in the Hoody dicots. American Journal of Botany 58: 469 . [Abstract]
Hiepko, P . 1965 . Das zentrifuga1e androecium der Paeoniaceae. Berrichte
der Deutschen Botan i schen Gesellschaft 77: 427-435 .
Howard, R. A. 1970 . Some observations on the nodes of voody plants wi th
special reference to the problem of the "split-lateral" versus the " conunon
gap ." In : Nel\' Research in Plant Anatomy. N. K. B. Robson , D. F . Cutler ,
and M. Gregory (eds . ). Supplement 1 to the Botanical Journal of the
Linnaean Society 63 . Academic Press. London.
Kaplan , D. R . 1967 . Floral morphology, organogenesis and interpretat ion of
the inferior ovary in Downingia baciga l upii. American Journal of Botany
54 : 1274-1290 .
Kribs , D. A. 1.935 . Salient lines of structural specialization in the "lOod
rays of dicotyledons . Bo t anical Ga zette 96: 547-557 .
27 581

Kr ib s . D . A.1937. Salient lines of st ructural sp ecialization in the wood

parenchyma of dicotyledons . Bulletin of the Torrey Bot anical Club 64 :
177-186.
Lam , H. J . 1948 . A new system of the Co rmophyta . Biumea 6 : 282-289 .
Leppik , E. E. 1957. Evolutionary relationship het\.;een entomophilous plants
and an thoph i l ous insects . Evo lu tion 11: 466-481 .
Haci or , L. \J. 19 71. Co-evolution of plants and an imals --systematic insights
f rom plant- in sect interac tions . Taxon 20 : 17-28 .
Ha r sden , M. P ., and I. \J. Ba iley. 1955. A fo urth type of nodal anatomy in
dicotyledons, ill ust rated by Cleroclendron trichotomum Thunb . Journa l of
the Ar nold Ar boret um 36 : 1-49 .
Mar tin, A. C. 1946 . The compara t ive internal morphol ogy of seeds. The
American Hid l and Naturalist 36: 513- 660 .
Heew,e, A. D. J . 1971 . Interpretative gynoecial morphology of t he Lactorida -
ceae and the Hi n te r aceae--a re-assessment . Acta Botanica Neerlandica 20:
221-238.
Melville, R. 1962 . A new theory of the angiosperm fl ower : 1. The gynoecium.
Kew Bul l etin 16: 1-50.
1963 . A new theory of the angiosperm fl ower. II. Th e and ro ecium .
Kew Bulletin 17: 1-63.
Hoseley, H. F. , Jr. 1967 . The value of the vascu l a r sys t em i n the s tudy of
the f lower . Phytomorphology 17 : 159-164 .
Hul l e r, J . 1970 . Palynological evidence on early diffe renti at ion of angio-
sperms . Biologica l Reviews 45 : 417 - 450.
Namb oodir i, K. K., and C. B. Beck . 1968 . A comparative study of the primary
va scular system of conifers. III . Ste1ar evolution in gymnosperms . Ameri-
can Journal of Botany 55 : 464-472 .
Ozenda , P . 1949 . Re cherches su r les dicotyledones apoca rp iques . Pub lications
des Laboratoires l ' Eco le Normal Superieure , Serie Biologie Fasc . II . Par is.
Puri , V. 1952 . Pl acenta t ion in angiosperms . Bo t a nical Review 18 : 603- 651.
Raven, P . H., and D. W. Kyhos . 1965 . New evi dence concerning the original
basic chromosome number of angiosperms . Evolution 19: 244-248 .
--7;-C~ ' , and 1'1. S. Cave . 1971. Chromosome numbers and relation-
ships in Annoniflorae. Taxon 20 : 479-483 .
Rickett , II. W. 1944 . The classification of inflorescences . Botanical Revi ew
10 , 187-231-
Saunde rs, E. R. 1925 . On carpel polymorphism . 1. Annals of Bo t any 39 : 123-
167 .
Schmid. R. 1972 . Fl oral bund le fu sion and vascular conser vatism . Taxon 21:
429- 446.
Schofield. E. K. 1968 . Petiolar anatomy of th e Gutti fera e and r elated fami -
lies . Nemo irs New York Botanical Ga r den 18: 1-55 .
Sinn ott , E. \.,1. 1914 . Investigations on the phy logeny of the angiosper ms .
1 . The anatomy of the node as an aid in the classificat ion of angiosperms.
American Jo urnal of Botany 1 : 303 - 322 .
Slad e . B. F. 1971 . Stelar evolution in vascul ar pl ants . The New Phytologist
70: 879-884 .
Srivastava , L. N. 1970 . The secondary phl oem of Austrobaileya scandens.
Canadian Jour nal of Botany 48 : 341-359 .
Stebbins . G. L . 1966. Chromosomal varia t ion and evo lution. Science 152:
1463-1469 .
196 7. Adaptive r adiation and t r ends of evolu t i on in h igher plants.
In : T. Dob zh ansky , N. Hecht, and W. C. Steere (eds). Evolu tionary Biology
1: 101-14 1. Appleton-Cent ury Crof t s . New York .
582 27

Stone , D. E. 1970 . Evolution of cotyledonary and nodal vasculature in the

Juglandaceae . American Journal of Botany 57 : 1219-1225 .
Wilson, C. L . 1942. The telome theory and the origin of the stamen . Ameri-
can Journal of Botany 29 : 759 - 764.
\.,rilson . '1' . K. 1964. Comparative morphology of the Canellaceae . III. Pollen .
Botanical Gazette 125: 192-197 .
Wodehouse , R. P. 1936. Evolution of pollen grains. Botanical Review 2 : 67-
84.
Zahur. N. S . 1959 . Comparative study of secondary phloem of 423 species of
woody dicotyledons belonging to 85 families . Cornell University Experiment
Station Nemoir 358 .

GENERAL STRUCTURAL REFERENCES

Bierhorst. D. W. 1971. Norphology of Vascular Plants. The Nacmillan Com-

pany. New York.
Carlquist, S. 1961. Comparative Plant Anatomy. Holt, Rinehart and l-linston .
New York .
Constance L. 1955 . The systematics of the angiosperms . In : A Century of
Progress in the Natural Sciences 1853-1953 . California Academy of Science .
San Francisco .
Cronquist, A. 1968. The Evolution and Classification of Flowering Plants .
Houghton-Mif flin Company. Boston.
Davis , P . H.• and V. H. HeYHood . 1963. Principles of Angiosperm Taxonomy.
D. Van Nostrand, Inc. Princeton .
de Beer . G. 1971 . Homology, An Unsolved Problem . Oxford Bio l ogy Readers
II . (F . F . Head and O. E . Lowenstein , ed s.). Oxford University Press .
Oxford.
Eames , A. 1961. Morpho logy of the Angiosperms. McGraw-Hill, Inc. New
York .
Meeuse, A. D. J . 1966. Fundamentals of Phytomorphology . Ronal d Press Com-
pany . NevI York .
Metcalfe, C. R., and L. Chalk. 1950. Anatomy of the Dicotyle dons. Clarendon
Press. Oxford.
Smith, A. C. 1969. Systematics and appreciation of reality . Taxon 18: 5-13 .
Solbrig, O. T. 1970. The phylogeny of Gutierrezia : An eclectic approach .
Brittonia 22 : 217-2 29.
Takhtajan, A. 1969. Flmvering P1ants--Origin and Dispersal. (Engl. transl.
C. Jeffrey). Smithsonian Inst itution Pre ss. l.Jashington, D. C.
28 583

Chapter 28. SYSTENS OF CLASSIFICATIONi'

The philosophical bases of plant classification are reviewed and criteria

used by botanists in devising their systems are discussed in this chapter . We
consider here only the classification systems developed in \-Iestern Civiliza-
tion, realizing, of course, that other civilizations (and "uncivilized socie-
ties") have produced classification syst ems independently .

DEFINITION, GENERAL PRINCIPLES, TRENDS OF CLASSIFICATION

The term classification is used for both : (1) the pro~ of classifica-
tion and (2) the product of this process, Le . , the system of classification.
Bio l ogical classification is the process of grouping together like organisms,
and the subsequent placing together of these groups into larger groups . Due
to discontinuities (of greater or lesser magnitude and greater or lesser de-
grees of completeness) in the pattern o f variation in nature , discrete units
in the biological I-Jorld can be recognized . In order to refer to these units,
they must be named , and an accepted system of nomenclature must be established.
The system of classification contains categories ( species , genus, family,
order, class . etc . ) into which the units, groups of units, and groups of groups
are placed. The system of classification thus needs a hierarchy of categories
into which the named units and groups are p l aced .

The recognition of similarities and dif fe r e nces betHeen organisms is a

basic human activity . and , in combination with a system of nomenclature, is ,
essential for accurate conununica tion (see discussion of this topic in Chapter
2). Modern systems of classification attempt to s e rve two needs: (l) to pre-
sent our current state of knm.,rledge regarding the relationships (totality of
similarities and differenc e s, sometimes Hith phylogenetic interpre tation) of
organisms, and (2) to provide, in combination with a system of nomenclature,
a retrieval system fo r information about organisms . In this context, the ad-
vantages and disadvantages of a linear vs. a t\W- or three-dimensional presen-
tation of the classification should be recognized . Also, a general system of
classification should be devised l .. ith practicality in mind . The classification
is made for ~ and must serve not only taxonomists and other scientists but
laymen as well .

Two principles of classification which taxonomists have learned by hard

experience should be noted here: ( 1 ) in the r>rocess o f classi ficatio n, the
taxonomist should assess the totality of similarities and differences of the
organisms being studied , and (2) after groups have been recognized, characters
may be weighted with respect to their value in defining the group and differ-
entiating the group from other groups (~posteriori weighting).

Considerable controversy exists today between the "pheneticists," i. e . ,

those who would generally rej ect our ability to make correct "phylogenetic
interpretations " \.then devising c lassification schemes (especially at the
higher levels) , and the "phylogenists , " Le . , those whose system-making com-
bines use of purely phenetic information with phylogene tic interpretation .

*Contributed by Kenneth H. Becker, The Nel.,r York Botanical Garden.

584 28

Davis and Heywood 's Pr inciples of Angiosperm Taxonomy (1963) is an example of

a work with a strong phenetic point of vieH, \"rhile Cronquist's Evolution and
Classification of Flowering Plants (1968) is an example of a \"rork with a
strong phylogenetic point of view.

The following trends in the development of systems of plant classifica-

tion through the centuries (which can be fo1lm"red in Section A of this chanter)
should be noted: (1 ) the gradual development of the hie r archy of categories
as we knOlv it today; (2) the gradual recognition of the major groups of
plants--especia11y with regard to vascular vs. non-vascular plants , cryptogams
vs. phanerogams, gynmosperms vs. angiosperms, monocots vs . dicots, and the
gradual recognition of more and more angiosperm families as natural groups; (3)
the gradual use of an i ncreasing number of characters in classification and
(4) cilanges in philos o phy in the transitions between the various periods in
the history of plant classification systems .

TAXA

TAXONOHIC CHARACTERS

Use of habit and Use of one or Use of as many Use of as many

importance to man a fe\"r selected characters as charac ters as
as characters characters to possible to possible plus
rxa group taxa

I
group phylogenetic
(evo l utionary)
inter pretation
t
Artificial }lechanical Natural Phylogenetic
Classi ficat ions Classif i cations Classifications Classifications

Figure 28 - 1 . From taxa to system in the development of various kinds of

general biol ogical c l ass i fications.

Section A. BASES OF CLASSIFICATION

Contemporary plant classification systems are the res ult of the labors
and insights of many \"rorkers throughou t the centuries . Although there are,
of course, no sharp lines dividing the var i ous "Periods ," the history of
plant classification may be broken down as follml s :
28 585

Period Horkers Philosophical Base , Criteria Used , Etc .

Ancients Theophra s tus Hrote to transmit what was generally known

Dioscorides ahout plants. Cl assification based on habit .
Pliny A few natural groups recogn i zed .
to ca . 1500 Alb ertus Nagnus

Herbalists Brunfels Hotivated by medical and commercial considera-

Bock tions--the systems E££ ~ were not the primary
Fuchs concern. Classi ficatio n based primarily on
Cordus habit (see discussion of l'Obe l' s system) .
Nattioli Advent of p r inting stir red new interest in
Dodoens pl ants , and 'mystica l respect' for wr itings
l' Ecluse of the ancien t s gave way to a period of
l'Obel original res earches into plant st ru ct ure.
Bankes
Turner
to ca . 1580 Gerard

Hechanical Caesalpino Primary concern was informa t ion retrieva l--

Systems Bauhin. J . t he system i tself was considered i mportan t .
Bauhin , G. Classification based mainly on form, but in-
Jung cludes also a number of other kinds of criter ia .
Horison ~ priori Iveightin g of values of individual fea -
Rivinus tures for classification purpOSeS was the rule
Ray (see discussion of Tournefort ' s system) . Hier -
Hagnol a rch y of ca tegories imp roved; more and more
Tournefort natural group s recognized.
Camerarius

l .. innaeus Artificial sexual system, based on numerical

relations of floral parts , was a great improve -
ment over earlier systems for purely practical
to ca . 1760 r easons .

Natural Adaosen Attempted to put seemingly re l ated plan ts

Systems Lamarck (plants that "l ook alike ") together , using
deJu ssieu, B. as many charact ers as possible (note change
deJussieu . A . in attit ude from mechanical systems). System
deCandolle . A. of A. deJussieu a turning point in the history
Brown of plant c l assification .
Lind ley
Endlicher
Brongniart
Braun
to ea . 1880 Ben tham & Hooker

1859 Darwin Produced no system of his own , but (with A. R.

lo/al la ce) adopted, recast . expanded. and fos -
tered the Theory of Evolution previously de-
veloped , to greater or lesser degrees . by
Lamarck , Lyell . et a 1. \Jhereas pre-Darwinian
natural system-makers wished their systems to
reflect the Divine Plan which was thought to
underlie al l of na ture , most post - Danvinian
 586 28

"ph ylogenetic" system-makers thought that ,

sin ce plants and plant-groups ar e the products
of evolution. it Has the i r task to determine
the course of this evolution , and have their
systems refle ct it .

" Phylo genetic" Eichler Attempt to integ rate what is koo,,'11 about the
Syst ems Bai11an evolutionary history of plant groups into the
Engler-Prantl sy s tem . Host are similar to natural systems ,
Harming but attempt to ar r ange plant groups according
Hettstein to their routes of descent. It became obvious
van Tieghem to several t.Jorkers ill th e ea rly days of the
Hernham phylogene tic p eriod that the natural system-
Bess ey make rs t.Jere r ecognizing evolutionarily-related
Hallie r groups of plants . 1\10 main schools of thought
Rend le developed as to origin of angiosperms and as
l>lez to \~hich character-states Here considered pr ir:d-
Rayata tive :
Hurtt-Davy Englerian Bessey an/Hall ierian
Fulle primitive flower primit ive flower
Skottsberg apetalous, unisexual, \.Jith perianth of
Tippo anemophilous . Ances- many, f ree, e qual
Gundersen tors o f angiosperms parts, bisexual ,
Hutchinson were coniferoid or insect-po llinated .
Benson gnetoid gymnosperms Ancestors of angio-
S06 \<lith unis exual stro- sperms were cycado -
Novak bili. phytes .
'. Emberger
Deyl See Chapter 27, Structural Evolution and
Kimura Phylogeny .
Ne lchior
Takhtajan Some contemporary systems take into account ,
Cronquist in addition to morphological characters, data
Thorne from the fields of anatomy, embryology, chemo-
Banks taxonomy, palynology , cytology , etc .
Bierhorst
Comprehensive revisions o f the classification
of vascular plants as a \.,r hole have appeared in
recent years .

1. SYNOPSIS OF DF.VELOPMENT OF PLANT CLASSIFICATION

A. Period of the Ancients ca . 300 Be to 1500 AD

1. Theophrastus Classification based on habit . Plants divided

JOO BC into trees , shrubs, undershrubs, herbs, then
into cultivated and I"ild kinds . Recognized :
annuals/biennials/perennials, determinate/inde-
t erminat e inflorescences , hypogyny /perir,yny /
epigyny , ape taly /polypetaly / gamopetaly , mono-
cots/dicots (s eed , stem, and leaf differences) ,
roots vs . rhizomes . Described and classified
ca . 480 kinds of plants. Considered trees to
be the most highly developed plants. Groups
strictly artificial . Used no categories as
28 587

we komv them today . Used speculative phi lo sophy

and methods of Plato and Aristotle (especially
the rule of the Excluded Hiddle : any given
object is either A or not A) .

2. Dioscorides 60 AD Deal t \"ith medicinal plants , and recognized ca .

600 species . Some group s eq ual more o r less
natural families (r e cognized a ser i es of lab-
iates and a series of umb els ) .

3. Pliny 70 Na tu ral History \"as an encyclopedic compi l ation .

4. Albertus i-lagnus 1250 Accepted Theophrastus r classif ic ation for the

mos t part . Recogn ized diffe rence i n stem struc-
ture between monocot s and dicots . Criteria used
for major cat egories : lea f y vs . non-leafy ; leafy
plan ts divided into monocots and dicots; dicots
divided into herba ceous and \wody kinds .

5. "Gart der
Gesundheit" 1485 Compilations with crud e descriptions and wood-
Hortus Sanitatis 1491 cuts. No definite natural arrangement evident
(more or less a lphabet ical ) .

B. Period of the Herbalists - -1500 t o 15 80

1. llrunfels, Otto 15JO Rec o gni z ed Pe r fec ti lf10wers visib l e \~hen held
at arm's l ength) allJ Impe r f ecti (flowers not
visible) .

2. Bock, Jerome 1539 Divided plants i nto t rees, shrubs, and herbs .
( Hieronymus Tragu s) Reco gnized 567 species , and gave concise
descriptions .

3. Fuchs, Leonhart 1542 Fine engrav ings . Described New Horld plants .

4. Hattioli, 1544 Excellent ill ustradons .

Pi e randrea

5. Cordus, 1 56 1 502 species, e x ce llent des cr iptions .

Valerius

6. Dodoens , 1583 Excellent descriptions and i llustrat i ons .

Rembert

7. l'Ecluse, 1601 Treat ed plants of America , Spain , Portugal ,

Charles de Austria, Hungary .

8. l ' 0hel , 1581 Classification based upon general form , growth

Nathias de habi t , ec onomic uses, and width of leaf (sepa-
rated monocots and dicots) .
grasslike plants b roader leaved dicot
with na rrow leaves --- b u l bous and - -- •• herbs _ shrubs ____ trees
rhizomatous
monocots
(supposed " simpler " forms ) ("most per fect " )
588 28

9. Bankes, 1525 Dealt with medicinal plants.

Ry charde

10. Turner, 1551 Alphabetical arrangement by Latin name.

t-lilliam 1562
1568

11. Gerard, 1597 Adopted 1 ' Obel ' s classification (based upon super-
John f icial resemblances and uses to man) .

The quality of de scription and illustration improved greatly during the

period of the Herbalists. There was a large increase in the number of plants
known; the discovery o f the New World had an important impact during this
period. There was some attempt to group closely related plants together.
Some larger natural famili e s were recognized, such as composites , umbels,
grasses , rushes, etc. See Arber (1938), and also Chapter 2 (History) .

C. Period of Mechanical Systems --1580 to ca. 1760

1. Caesalp i no, 1583 Sought a classif ication based upon reasoning , not
Andrea upon utilitarian considerations . Used bas ically
an Aristotelean approach (~priori weighting of
characters). Organs of fructi f ication considered more important than
habit. Classification based upon woodiness vs. herbaceousness , then on
characters of f ru i ts, s eeds, and embryos. Fruits classified on basis of
position , l ocul e number, and seed number. Classification stresses number,
position , and form of parts. Recognized : inferior/superior ovary , bulbs
present/absent, sap milky/\"atery , activities of plants. Gave some tech-
nical descriptions . Had a good concept of genera, and described ca . 1520
plants . Recognized Fagales, legumes , umbels, crucifers, composites, bor-
ages as natural groups .

2. Bauhin, 1650 Descriptions \"ere good diagnoses; described ca .

Jean 5000 plants . Recognized crucifers, labiates .

3. Bauhin, 1623 General arrangement as in l'Obel and Gerard (based

Gaspard on texture and form) . Neglect of floral and fruit
characters in arrangement of larger groups . Recog-
nized distinction between genera and species , and described ca . 6000 species
in his Pinax. Often used binomial nomenclature. Placed together Umbelli-
f erae , some composites, crucif e rs, solanads. poppies, labiates, cucur bits,
but out duckweeds among cryptogams and water ferns \"ith mosses .

4. Jung , ca . 1650 Argued against fundamental division into trees/

Joachin shrubs/herbs .

5. Horison, 1680 System s i milar to that of Caesalpino; woody/herb a -

Robert ceous; divided into groups based on fruit and seed
characters .

6. Rivinus. 1690 System with emphasis on corolla .

Augustus Q.
28 589

7. Ray, 1703 lIerbae

John Imperfectae (basically cryptogams)
Perfectae
Dieots
HanDcats
Ar bo r ae
Manocats
Diects
Hanocats and clieots classif ied on fruit type, then l eaf charac ters, £l Ol.,ler
type . Believed all parts of the plant should be used for classification.
Emphasiz ed importance of cotyledon number . Said: "Nature refuses to be
fo r ced into the fette r s of a p r ecise sys t em. 11 Sys tem direct antecedent of
that of deJussieu; classification was aimed at prac t icality . Gave c lear keys
to genera , and recognized ca . 1 8 , 000 species . Dist i nguished between inheri-
ted and environmentally caused difference s , and noted the sign i ficance of the
f ormer for taxonomy . Used anatomical i nforma t ion gathered by Grew , Malpigh i ,
et al . Recognized composites, umbels , mints, crucifers, le gume s , etc . Com-
positae divided into four classes based on rel atively unimportant characters ,
and his Bacciferae includes many unrelated genera having fruits with fleshy
pericarp .

8. Hagnol, 1689 Used conspicuous characters of roots , st ems,

Pierre flowers , seeds. First concept of mode rn families
(listed 76 in 1689).

9. Tournefort , 1694 Classification based on tree/herb , t hen ap etalou s/

Jo seph Pitton 1716 peta lous/gamopeta l ous , then flowers r egul ar/irre-
gula r. System widely used unti l ca . 1760, when
it was superseded by that of L:tnnaeus (used unt il 1780 i n France , whence super-
seded by that of deJussieu) . Author of modern genus concept , and is considered
to have had a good con cept of genera . Gave generic descriptions , and r ecog-
nized 698 genera, 10 , 146 species . Perhaps gave too much emphasis to vegetative
characters in delimi ting ge nera . His important system is largely artificial ,
and is generally con sid ered inferior to that of Ray .

10 . Camerarius , 1694 Demonstrated sexuality in pl ants .

Rudolf

11. Linnaeus , 1735 Produced a "Sexual Sys tem" based upon the numeri-
Carl 1737 cal relat i ons of floral parts . Recognized 24
1753 Classes , based on number , union, and length of
stamens . Classes subdivided into Orders on basis
of number of styles ; unisexuali ty also considered. T.. i nn aeus' ca t egor y Cl assis
more or l ess corresponds to what we would now call an order, his category Ordo
corresponds to families in the moder n sense . Linnaeus' system was later modi-
fied by other workers (Willdenow, Gmelin, Sc hreber , et al.) in many other,
later ed it ions of his works , up t o about 1805. The Sexual Sys tem was of great
utility in identifica tion, but was deliberately and admittedly artificial. It
was widely adopted unti l superseded by system of deJussieu . Influenced in pro-
ducing hi s Sexual System by Aristotelean reasoning , but not by Aristo t elean
reliance upon habit to delimit major groups. Used (1) the principles of Logi-
ca l Division (as did Theophrastus) and (2) ~ pr iori reasoni ng as to t he most
importan t characters (i.e ., those of the flower) .
 590 28

In his Philosophia Botanica (1751). Linnaeus enumerated 67 "natural

orders ." Some represent natural groups, such as palms, orchids, grasses,
coni f ers, composites , borages , etc., but some "natural orders" are quite
mixed , with monocots and clieots appearing together. Linnaeus was in f lu-
enced here by the empirical method of natural science, i.e. , he developed
his natural system based upon years of observational experience , rather than
on an ~ priori basis.

LINNAEUS' SEXUAL SYSTEN (Systema Naturae, 1735; Genera Plantarum , 1737;

Species Plantarum, 1753)

Class Subclass (Orda) Examples of Species

I . Monandria Honogyn i a Canna indica

(Stamen one) Digynia Cinna arundinacea
II. Diandria Nonogynia Ligustrum vulgare, Chionanthus virginica
(Stamens two) Digynia Anthoxanthum odoratum
Trigynia Piper nigrum
III. Triandria Honogynia Va1eriana officinalis, Helothria pendula
(Stamens three )Digynia Phalaris canarienSiS, Triticum aestivum
Trigynia Hollugo verticillata
IV. Tetrandria Honogynia Dipsacus f ullonum , Plantago major
(Stamens four) Digynia Cuscuta americana
Tetragynia Ilex cassine
V. Pentandria Monogynia Heli otropium ind i cum, Lysimachia guadrifolia
(Stamens five) Digynia Asclepias syriaca, Carum carvi
Trigynia Rhus vernix, Alsine media
Tetragynia Parnassia pa1ustris
Pentagynia Aralia spinosa , Suriana maritima
VI. Hexandria Monogynia Tillandsia utriculata, Pontederia cordata
(Stamens six) Digynia Oryza sativa
Trigynia Rumex vertici1latus , Medeo1a virginiana
Tetragynia Petiveria alliacea
Polygynia A1isma cordi f olia
VII. Heptandria (7) Monogynia Aesculus pavia
VIII. Octandria Honogynia Tropaeolum major, Gaura biennis
(Stamens 8) Digynia Galenia africana
Trigynia Polygonum persicaria
Tetragynia Paris quadrifolia
IX . Enneandria Monogynia Laurus benzoin
(Stamens 9) Trigynia Rheum rhaponticum
Hexagynia Butomus umbel latus
X. Decandria Monogynia Cercis canadensis, Nelia azedarach
(Stamens 10) Digynia Tiarel1a cordifo1ia
Trigynia Silene virginica
Pentagynia Oxalis stricta, Agrostemma githago
Decagynia Phytolacca americana
XI . Dodecandria Honogynia Asarum canadense, Rhizophora mangle
(Stamens Di gyn i a Agrimonia eupator i a
12-19) Trigynia Euphorbia chamaesyce
Pentagynia Glinus lotoides
Dodecagynia Sempervivum arboreum
XI I . Icosandria Monogynia Cactus opuntia, Prunus virginian a
(>19, attached Di gynia Crataegus crus galli
to calyx) Trigynia Sorbus domestica
28 591

Pentagynia Pyrus communis

Polygynia Rosa canina
XIII. Polyandr1a Monogynia Capparis spi nosa, Podophy l lum
(Stamens >19. peltatum
attached to Digyoia Paeonia offic inal is
r eceptacle) Trigynia Delphinium a j acis
Tet ragynia Tet racera voluh uli s
Pentagynia Aguilegia canadensis
Hexagynia Hagnolia virginiana
Polygynia Annona glabra, Anemone virginiana
XIV . Didynamia Gymnospermia Ajuga reptans
(Stamens Angiospermia Scrophularia marilandica
didynamous)
xv. Tetradynamia Siliculosa Draba verna }
(Stamens tetra- Si l iquosa Erisymum 0 icina 1 e
----------ff Cruciferae
dynamo u s)
XVI. Nonadelphia Pentandria tolalthe ria ind ica
(Stamens in Decandria Geranium mac ula tum
one bundle) Polyandria Sida rhombifolia
XVII . Diadelphia Hexandria Fumaria officinale
(Stamens in Octandria Polygala incarnata
two bundles) Decandria Erythrina herbacea
XVIII • Polyadelphia Pentandria Theobroma cacao
(Stamens in Icosandria Citrus aurant i um
several bundles)Polyandria Hypericum perforat um
XIX. 5yngenesia Polygamia Aequalis Tragopogon virginicum
(Stamen s wi t h Polygami a Superflua Artemisia annua
united a nthers) Polygamia Frustranea Helianthu s annuus
Polygamia Neces saria Chrysogonum virginianum
Monogamia Lobelia cardinalis, Viola canadensis
xx. Gynandria Diandria Orchis spectabilis
(Stamens adnate Triandria Sisyrinchiutn bermudiana
to pistil) Tetrandria Nepenthes distillatoria
Pentandria Passiflora incarnata
Hexandria Aristo lochia serpentaria
Decand ria Helicteres a ngus tifolia
Polyandria Arum triphyllum
XXI. Honoecia Honandria Zannichellia palustris
(Plants monoe- Diandria Lemna mi nor
----- -----
ciaus) Triandria ~ latifolia, Zea mays
Tetrandria Urtica dioica
Pent and ria Ambrosia trifid a
Hexandria Zizania aquatica
Heptandria Guettarda speciosa
Polyandria Hyriophyllum spicatum , Quercus
phellos
Honadelphia Pinus taeda, Ricinus communis
Syngenesia Homorrlica cylindrica
Gynandria Andrachne fruti cosa
XXII. Dioecia Monandria Na j as marina
(Plants dioe- Diandria Salix babylonica
dous) Triandria §.mpetrum nigrum
Tetrandria Viscum album , Hyrica cerifera
Hexand r ia Smilax lanceolata
Octandria Populus alba
 592 28

Ennaeandria Nercurialis procumbens

Decandria Carica papaya
Polyandria Cliffortia trifoliata
Honadelphia Juniperus virginiana
Syngenesia Ruscus aculeatus
Gynandria Gluria retusa
XXIII. Polygamia Monoecia Celtis occidentalis , Veratrum nigrum
(Plants polyga- Dioecia Nyssa aquatica
mous) Polyoecia Ficus carica
XXIV. Cryptogamia Filices Equisetum arvense , Adiantum pedatum
(Flowers con- Musei Lycopodium inundatum, Hypnum cusp ida tum
cealed) Algae Harchantia polymorpha , Ulva confervoides
Fungi Ilydnum imbricatum

D. Period of Natural Systems 1760 to ca. 1880

1. Adanson, 1763 Rejected artificial classifications. Emphasized

Nichel equal weighting of characters. and believed that
all organs should be taken into account. Rejected a priori reasoning.
Described groups more or less equivalent to modern families and orders .
Produced 65 single-character systems from \"hich he hoped to produce a
natural system (this \"as largely a failure --criticized by deJuss i eu and
deCandolle) . "Grand f ather of Numerical Taxonomy."

2. deJ ussieu, 1759 Rearranged the plants in a garden at LaTrianon ,

Bernard Versailles, according to his own system , which
\"as not based on habit and was not purely arti-
, ficial.

3. deJussieu , 1789 Adopted and expanded the system of B. deJussieu.

Antoine- "First complete system Hhich c an claim to be a
Laurent n atural one"--Rendle. Weighted characters in
the fol lowing order : embryo> sexual parts> fruit = perianth > vegeta-
ti ve characters. Reintroduced concepts of hypogyny /pedgyny / epigyny.
Differentiated, described, and named 100 "natural orders"--corresponding
to most major families. Some related families (such as palms, lilies,
amary1lids, irises) grouped together.

SYSTEM OF A . deJUSS I EU (Genera P1antarum, 1789)

Class Order (Family) Examples
I . Acotyledones I Fungi , Algae , Hepaticae
{ Stamina hypogyna II Aroideae , Typhae
II . Honocotyledones perigyna III Palmae, Junci
ep igyna IV Musae , Orchides
 28 593

Class Order (Family) Examples

epigyna V Aristolochiae
Apetalae perigyna VI Elaeagni , Lauri
hypogyna VII Amaranthi

Corolla hypogyna VIII Acanthi, Vitices

perigyna IX Guaiacanae, Ericae
anthe riS
Monopeta l ae epigyna ( connatus X Cichoraceae
III. Dicoty- . Jan theris XI Dipsaceae
ledoIlcs ep ~ gynaldistinctis
Stamina ep1gyna XII Arali ae, Urnbelliferae
Polypetalae hypogyna XIII Cruciferae , Acera
{ perigyna XI V Cacti, Hyrti
Diclines irregulares XV Euphorbiae, Urticae

4. deCandolle, 1813 An expansion and elaboration of deJussieu ' s system .

Augus tin P . , 1873 Arrangement of clieots improved (united Apetalae and
Alphonse , Diclines) . Downgraded use of physio l ogical charac-
and Casimir ters, emphasized instead the importance of morpho-
logical and anatomical characters . Used ~ priori
weighting. Ranunculaceae at base , arrangement of system based on reduction
and fusion of floral parts . Linear sequence recognized as necessarily arti-
ficial. Alteration of "original symmetry" by abortion, degeneration, and
cohesion. System looks evo l utionary but was pre-Darwinian, and it is gener-
ally believed that deCandol le did not accept the Lamarckian concept of modi-
fication over time . By 1840 had replaced Linnaeus ' Sexual System.

SYSTEM OF AUGUSTIN PYRAHUS AND ALPHONSE deCANDOLLE . 1B19 and lB73 . Prodro-
~ Systematis Naturalis Regni Vegetabilis . (1819 is modified below with
the treatment of the dicots based on actual publication in succeeding vol-
umes and 1873 summary) .

1. Vasculares (Vascular plants with cotyledons)

Class I Dicotyledoneae (includes Conifers)
Subclass I Tha1amiflorae (Polypetalous and hypogynous)
Orders (Families) 1-54 Ranunculaceae. Dilleniaceae. Magnoliaceae ,
Rutaceae. Simarubeae. Ochnaceae , etc.
Subclass I I Calyciflorae (Perigynous or epigynous and polypetalous
or sympetalous)
Orders (Familie s) 55 -lIB Celastrineae. Rhamnaceae. Bruniaceae ,
Samudeae, Homalineae. Pyrolaceae , Monotropeae. etc .
Subclass III Corollif l orae (Gamopetalous and hypogynous)
Orders (Fami lies) 119-155 Lentibulariaceae , Primulaceae , Aegi-
ceraceae , Solanaceae , Plantaginaceae, etc .
Subclass I V 1'1onochlamydeae (Calyx only present)
Orders (Families) 156-207 Phytola c caceae. Salsolaceae. Basella-
ceae , Ama r antaceae, Euphorbiaceae, etc.
Class II Honocotyledoneae (includes Cycads)
Subclass I Phanerogamae (Liliaceae , Iridaceae, Orchidaceae, etc .)
Subclass I I Cryptogamae (Ferns , etc.)
 594 28

II . Cellulares (plants Idthout cotyledons)

A. }losses. Liverworts
B. Lichens , Fungi, Algae
(Ferns placed in Cellulares in last arrangement of deCandolle system) .

5. Brown, 1 827 Proposed no system of his m"l1 . Pointed out nature

Robert of gymnosperms (naked seeds) and their distinctness
from angiosperms . Opposed Linnaean system. Intro-
duced many new "natural orders," and proposed the grouping of "natural orde rs"
(families) into what He l.muld now call orders.

6. Lindley, 1830 Introduction to the Natural System of Botany is

John first comprehensive natural system published in
English . Recognized distinction between gyrrmo-
sperms and angiosperms . Diects divided into Polypetalae, Apetalne, Achlamy-
deae, and Monopetalae--these groups Here considered artificial. Treatment of
monocots is precursor of Hutchinson's system on basis of perianth . Keys to,
and discussions of, bmilies are considered very good . In his Vegetable King-
dom (1846) , Lind ley overemphasized the importance of the parasitic habit of
some dicots and the importance of net vs . parallel venation among the monocots .

7. Endlicher , 1840Cryptogams clearly separated f r om seed plants .

Stephan Cymnosperms separated from, but placed close to,
dicotyledons . Thallophyta: no stem and root vs .
Cormophyta : stem and root present . LOHer dicots \.,rere apetalous groups, a
practice follm.,red in later ("Englerian " ) systems (as opposed to beginning
with Ranalian groups as in the deCandolle-13essey/llallierian sys tems) . Dicots
divide d into Apetalae , Gamopetalae, Dialypetalae . Some parasitic angiosperms
placed with the cryptogaITl<;. Treatment of mono cots differed from that of
Lindley, similar to that of Bentham and Hooker (see belm!).

8. Brongniart. 1843
Basic division into cryptogams and phanerogams
Adolphe (seed plants) . In the monocots, the presence or
Theodore absence of endosperm in the seed considered impor-
tant. His "dicots" included both gymnosperms and
angiosperms . Angi ospermous dicots divided into Gamopetalae and Dialypetalae .
His attempt to distribute apetalous forms among the petaliferous forms to
which they appeared related is considered a maj or advance .

9. Braun , 1853Influenced by Endlicher and 13rongniart. Placed

Alexander monocots and dicots together and separated from
gymnosperms . The monocots Here arranged to shoH
a progress ion from the simple to the complex--began l.,rith Lemnaceae, ended
wi th Orchidaceae. Dicots : apetalae , sympetalae , polypetalae. Influenced
Eichler, Engler, \~ettstein, et al.

10. Bentham, George

1862- System largely deCandollean, and pre-evolutionary
& Hooker, 1883 in conception . Groupings above family level differ
Joseph D. conSiderably f:rom Prodromus of deCandolle . Both
monocots and dicots start \.,rith polypetalous famil-
ies . Fusion and reduction of parts l.,rere basis of organization. Note : (1)
maintenance of all apetalous groups as a unit, and separated from the petali-
ferous groups to which many are probably more related. (2) the Series in the
Honochlamydeae and Nonocotyledones o f ten contain quite unrelated elements ,
28 595

and (3) placement of gymnosperms between dicots and monoco t s . Accepted 200
families; all genera studied aneH for Genera Plantarum . System became very
popular in English-speaking countries, and used fo r he r barium arrangement in
Great Britain .

SYSTEH OF BENTHMI AND HOOKER (Genera Plantarum, 1862-1883 , with some represen-
tative examples; some spellings modernized) .

Dicotyledones Polypetalae
Series 1. Thalamiflora e
Cohort 1. Ranales - Ranunculaceae . Hagnoliaceae , Annonaceae, etc .
(Order) 2 . Parietales - Sarraceniaceae , Papaveraceae, Violaceae , etc .
3 . Po ly ga linae - Pittosporaceae. Polygalaceae, etc .
4 . Ca rvophyll inae - Caryophyl laceae , Portulacaceae. etc .
5 . Guttiferales - Hypericac eae , Dipterocarpaceae, etc .
6 . Nalvales - Halvaceae , Sterculiaceae, Tiliaceae.
Series II . Discif l orae (hypogynous di sc )
7 . Geraniales - 1.inaceae , Ger an i aceae, Rutaceae, etc .
8. Olacales - Olacineae, Ilicineae , etc .
9 . Celastrales - Ce 1astraceae , Rhamnaceae, etc .
10 . Sapindal es - Sapindaceae , Anacardiaceae , et c .
Series III . Ca l yciflo rae (floral cup)
11 . Rosales - Rosaceae, Leguminosae , Saxifrageae, Droseraceae. etc.
12 . Nyrtales - Rhizophoraceae, Lythraceae, Ona graceae , etc .
13 . Passiflorales - Loasaceae , Cucurbitaceae, Pass i florace ae, etc .
14 . Ficoidales - Cactaceae. Ficoideae .
15 . Umbella l es - Umbelliferae, Araliaceae, Cornaceae .
Dicotyledones Garnopetalae
Series I . Inferae (infe r ior ovary)
Cohort 1 . Rubiales - Caprifoliaceae , Rubiaceae .
(Or der) 2 . Asterale s - Valerianae, Dipsaceae, Calyceraceae, Compositae
3. Carnpanales - Stylideae, Campanulaceae , etc .
Series II. Ileteromerae (carpels >2)
4 . Ericales - Va ccinieae, Ericaceae , Epacrideae , etc .
5 . Primulales - Primulaceae , Pl umbaginaceae , etc .
6 . Ebenales - Sapotaceae, Ebenaceae, Styracaceae
Se ries III . Bicarpellatae (carpel s = 2 , rare l y 1 or 3) .
7 . Gentianales - Oleaceae. Apocynaceae, Gentianaceae , etc .
8 . Polemon i ales - Polemoniaceae , I-Iydrophyllaceae. Boraginaceae , etc .
9 . Personales - Sc rophularin eae , Orobanchaceae , Acan thace ae, ecc .
10. Lamiales - Verbenaceae. Labiatae, Selagineae , etc .
Dicotyledones Monochlamydeae (Apetalous)
Series I . Curvembryeae - Orders (Families) Nyctagineae. Batideae , etc .
Series II . Multiovulatae Aquaticae - Order (Family) Podos temace ae
Series III . Hultiovulatae Terrestres - Orders (Families) Nepenthaceae. etc .
Series IV . Micrembryeae - Orders (Families) Pi pe raceae , Ch loran t haceae .
Monimiaceae , e tc.
Series V. Daphnales - Orders (Families) Laurineae, Proteaceae . El aeagna-
ceae, etc .
Series VI. Achlamydospo r eae - Orders (Families) Loran thaceae , Santalaceae,
et c .
Series VII. Unisexuales - Order s ( Fami li es) Euphorbiaceae, Urtica ceae,
Cupuliferae. etc .
Se ries VIII. Ordines anomali - Orders (Families) Ernpet raceae , Ceratophylla-
ceae , etc .
596 28

Gymnosperrnae

Monocotyledones
Series I. Hicrospermae - Orders (Families) Orchideae , Burmanniaceae , e tc.
Series II. Epigynae - Orders (Families) Scitamineae , Bromeliaceae, etc .
Series III. Coronarieae - Orders (Families) Liliaceae. Ponteder iaceae, etc.
Series IV. Calycinae - Orders (Families) Juncaceae , Palmae , etc .
Series V. Nudif l orae - Orders (Famil ies ) Typhaceae , Cyclanthaceae , etc.
Series VI. Apocarpae - Orders (Families) Alismaceae , Naiadaceae , etc.
Series VII. Glumaceae - Orders (Families) Eriocauleae, Gramineae, etc .

11. Ho f meister . IS51 Embryological researches hel ped supply the basis
"\o1ilhelm f or the division of the plant kingdom into the
Thallophyta, Bryophyta, Pteridophyta , Gymnosper-
mae, and Angiospermae .

Asa Gray, during the formative period of Ameri-

can botany , used a combination of the systems
of deCandolle and Bentham & Hooker for use in
floras of North Amer i ca .

E. Period of Phy l ogenetic Systems - 1880 to present

1. Eichler , 1890
Cryptogams : Thallophyta, Bryophyta , Pteridophyta
August W. Phanerogams : Gymnosperms , Angiosperms
Honocotyleae
Dicotyleae
Choripetala e
Sympetalae
Began dicots with Amentiferae (as did Endlicher) . United apetalous and poly-
petalous dicots . Accepted theory of evolution, and attempted to ar r ange
plants in a series from the supposedly primitive to the mo r e advanced ; equated
simp l ic i ty with primitiveness. Classification i n his Bilithendiagrannne (1878)
largely artificial .

2. Baillon , 1867- Took a broader viet., of some families than most

Henri 1895 autho r s (as Ericaceae , etc.) .

3. Engler, Adolf
& Karl Prantl 1892 Classification based on that of Eichler.
Bacteria; Algae ; Fungi ; Lichenes ; Bryophyta;
4. Eng l er & E. Gilg 1924 Pteridophyta; Embryophyta Siphonogama (seed
plants)
5. Engler &
L. Diels 1936 Gymnospermae
Angiospermae
6. Helchior, H. 1964 Class Monocotyledoneae
Cl ass Di cotyledoneae
Subclass Archichlamydeae
Apetalae (no petals)
Choripetalae (separate petals)
Subclass Hetachlamydeae (Sympetalae) (united
petals)
28 597

Phylogenetic principles : the most primitive flowering plants, in both the

Monocotyledoneae and Dicotyledoneae, have apetalous , simple flowers in
strobilus-like inflorescences

monocots clieots
I
Pandanales
I
Amentiferae
primitive primitive

~ polyphy letic /
from unknown
gymnospermous stocks

Engler was we ll aware of the widespread occurrence of pa ral lelism and con-
vergence i n the angiosperms, and did not produce a "phylogenetic tree"
showing line s of descent (it was only later that Engl er claimed his sys-
tem to be phylogenetic).

Progressive morphological (not necessarily phylogenetic) series :

no sepaloid sepals sepals

perianth bracts and and
only free petals unit ed petals

This series was thought to have taken p l ace in several phyletic lines in
both the monocots and dicots. The system emphasizes apetaly, choripetaly,
and sympetaly , with hypogyny, perigyny, and epigyny considered as secondary
criteria. Associated some apetalous groups with related petaliferous
groups. Gymnosperms clearly set off from angiosperms . Engler ian evo lu-
tionary principles challenged by Bessey (1897), Hallier (1905), Arber and
Parkin (1907) , and othe r authors. System has been very widely used in
Europe and America , and many herbaria and floras are arranged according
to one or another of the various editions of the Syllabus . System used
in 20 volume Die natilrlichen Pflanzenfamilien. The latest (12th) edition
of the Syllabus der Pflanzenfamili en proposes , but does not execute, many
changes in sequence and position of groups based on modern knowledge
gathered fro m the fields of anatomy , chemistry , emhryo logy , etc. The
monocots are considerab ly revised. Some of t he evolutionary princ i ples
upon which the Englerian systems rest are , in the opinion of most con-
tempo r ary workers , obsolete .

7. Warmi ng , 1895 Similar to Engler ' s system, but with some changes
Eugenius in positioning of groups .

8. \.j'ettstein, 1911 Similar to Engler's system , but: (1) monocots con-

Richard von 1935 sidered to be derived from Ranales , (2) dicots thought
to consist of several independent l i nes , (3) admitted
more cases of per ianth reduction than did Engler. Supposed primitive
characters of the Ament ife rae: Hoody , flowers unisexual, Hind pollinated,
apetalous , vascular bundl es occasionally present in ovule, gametophytes
may have a larger numbe r of cells than is typical of angiosperms (Casuarina)--
all gymnospermous characteristics. Believed fossil evidence supported
amentiferous ancestry. Casuarinales considered primitive, Ephedroid or i -
gin for Casuarina flower theorized .
598 28

9. van Tieghem, 1898 i. Liorhizeae Monocotyleae (monocot s max . pars.)

Philippe ii. Liorhizeae Dicotyleae
i i i . Climacorhizeae (cli eo ts max. pars.)
These c lasses are based on the ontogeny of the root apex . Class i i contains
the Nymphaeaceae and Gramineae. Class i i i is subdivided on the basis of the
presence of a "per fect seed" at maturity (Seminees--most dieots) or its ab-
sence (Inseminees--mostly parasitic groups) . Overreliance on characters of
root, ovules , and integumen t s. System basically a n histori cal c uriosity, but
provided useful data .

10 . Wernham. 1911 At least seven l i nes of pol ypetalous d ieots thought

T. J. to give rise to Sympe talae.

11. Bessey , 1897Definitely conceived his system to be phylogenetic .

Charles Edwin 1915Consid ered angiosperms to be monophyletic f rom a
Bennettita lean (cycadeoid) type ancestor with bi-
sexual st robili (see "Bessey ' s Dicta"--Chapter 27, p . 563) . Produced a dia-
gram purportedly showing lines of descen t in the angios perms (see "Bessey's
Cactus" -- p. 602). Ranales sensu lato ( flower with many, free , equal parts )
considered primitive. Emphasized ova ry position (hypogyny vs . perigyny or
epigyny) over petal fus ion. Recognized 32 orders and 300 famili es . Accepted
tribes of Compositae at family level. Bessey ' s contribution probably lie s
more in his set of " phylogenetic dicta" than in his system per~. His system
\.fas original in many respe cts, whi l e also dra\.fing f r om bo th the Bentham &
Hooker and Engler & Prantl systems. The elimination of the Polypetalae . Gamo-
petalae, and Monochlamydeae as major groups of dicotyledons s ho uld be noted.
Clements. Schaffner. and Pool supported and elaborated upon the Bessey system
in the early part of the century . The phylogenetic dicta which Bessey pro-
pounded have , with modificat i ons, provided the theoretical base and guiding
principles for most contemporary systems of angiosperm cl assification (in
this regard credit must also be given to Hallier , see be l ow).

Bessey may be cri ticized for assuming tha t perigynous and epigynous forms
constitute a single evolutionary line, and in the impl ication of direct an-
cestry of contemporary orders from one another in his "Cactus ." Also , many
suppos e d family relationsh ips in his system ar e cont rary to current evidence.
The descent of most hypogynous sympetalous forms from a caryophyl lalean-type
ancestor , among other "problems ," is particularly difficult to accept today.

SYST~f OF BESSEY (Phylogene tic Taxonomy of Flowe ring Plants. ~nnals of the
Missouri Botanical Garden 2: 109 . 1915).

Class: ALTERNIFOLIAE (MONOCOTYLEDONEAE)

Subclass : Alternifoliae- Strobiloideae
Order : Alismatales: Alismataceae, Butomaceae , Triur idaceae, Scheuch-
zeriaceae. Typhaceae, Sparganiaceae . Pandanaceae,
Aponogetonaceae, Potamogetonaceae
Order: Liliales : Liliaceae, Stemonaceae, Pontederia ceae . Cyanast ra-
ceae , Philydraceae , Commelinaceae , Xyridaceae, Maya-
caceae, Juncaceae , Eriocaulonaceae, Thurniaceae,
Rapateaceae. Naiadaceae
Order : Arales : Cyclanthaceae, Araceae . Lemnaceae
Order : Palmal es: Palmaceae
Or der : Graminales: Restionaceae, Centrolepidaceae, Flagellariaceae ,
Cyperaceae, Poaceae
28 599

Subclass : Alternifo l iae-Cotyloideae

Order: Hydral es: Vallisneriaceae
Order : lridales : Amaryllidaceae , Haemodoraceae, Iridaceae, Vellozia-
ceae, Tacc8ceae , Dioscoreaceae. Bromeliaceae, Husa-
ceae , Zingiberaceae , Cannaceae, Haran taeeae
0rder : Orch ida l es : Burmanniaceae, Orchidaceae

Class: OPPOSITIFOLIAE (D I COTYLEDONEAE)

Subclass: Oppositifoliae-Strobiloideae
Superorder : Strobiloideae-Apopetalae-Polycarpellatae
Or der: Ranales : Magnoliaceae, Calycanthaceae , Honimiaceae, Cerc idi-
phyllaceae, Trochodendraceae, Leitneriaceae, Anona-
ceae , Lactoridaceae , Gomortegaceae, Hyristicaceae, Saururaceae, Piper-
aeeae, La cistemaceae , Chloranthaceae, Ranunculaceae , Lardizabalaceae,
Berheri daceae, Henispermaceae, Lauraceae , Nelumbaceae, Cabombaceae ,
Ceratophyllaceae , Dilleniaceae, \-Jinteranaceae
Order : Halvales : Sterculiaceae , }\alvaceae, Bombacaceae, Scytopetala -
ceae, Chlaenaceae, Gonystylaceae, Til iac eae,
Elaeocarpaceae , Balanopsidaceae, Ulmaceae , }\oraceae, Urticaceae
Order : Sarracenia l es : Sarraceniaceae , Nepenthaceae
Order : Geraniales: Geraniaceae , Oxal idaceae, Tropaeolaceae, Balsami-
naceae , Linmanthaceae, Linaceae. Humi riaceae,
Erythroxylaceae , Zygophyllaceae, Cneoraceae, Rutaceae, Simarubaceae,
Burseraceae , Meliaceae. Malpigh i aceae, Trigon iaceae , Vochysiaceae ,
Polygalaceae, Tremandraceae, Dichapetalaceae, Euphorbiaceae, Calli-
trichaceae
Order : Guttiferales : Theaceae . Cistaceae , Guttiferaceae. Eucryphia-
c eae . Ochnaceae, Dipterocarpaceae , Caryocara-
ceae, Quiinaceae , Marcgraviaceae, Flacourtiaceae , Bixac eae , Cochlo-
spermaceae, Violaceae, Halesherbiaceae , Turneraceae , Passifloraceae ,
Achariaceae , Caricaceae . Stachyuraceae , Koeberliniaceae
Order : Rhoeadales: Papaveraceae , Tovar iac eae , Nymphaeaceae, Horin-
gaceae. Resedaceae, Capparidaceae. Brass i caceae
Order: Caryophyllales : Caryophyllaceae . El atinaceae , Portulacaceae .
Ai zoaceae , Frankeniaceae , Tamar i caceae, Sali-
caceae , Podostemonaceae , Hydrostachydaceae, Phytolaccaceae , Basella-
ceae , Amaranthaceae . Chenopodiaceae, Polygonaceae, Nyctaginaceae ,
Cynocrambaceae , Batidaceae
Superorder: Strobiloideae-Sympetalae-Polycarpellatae
Order : Ebenales : Sapotaceae , Ebenaceae. Symplocaceae, Sty r acaceae ,
Fouquieriaceae
Order : Ericales : Clethraceae, Erica ceae , Epacr idaceae, Diapensia-
ceae , Pirolaceae, Lennoaceae
Order: Primulales : Primulaceae, Plan taginaceae, Plumbaginaceae.
Myr sinaceae , Theophrastaceae
Superorder: Strobiloideae-Sympetalae-Dicarpellatae
Order : Gentianales : Oleaceae , Salvadoraceae , Loganiaceae, Gentiana-
ceae, Apocynaceae , Asclepiadaceae
Order : Polemoniales: Polemoniaceae, Convolvulaceae , Hydrophylla-
ceae . Borraginaceae , Nolanaceae, So l anaceae
Order : Scrophulariales : Scrophulariaceae , Bignoniaceae . Pedaliaceae,
}1artyniaceae, Or obanchaceae. Gesneraceae ,
Columelliaceae , Lentibulariaceae, Globulariaceae , Acanthaceae
Or der : Lamiale s : Hyoporaceae , Phrymaceae, Verbenaceae, Larniaceae
 600 28

Subclass : Oppositifo1iae- Cotyloideae

Superorder: Cotyloideae-Apopeta1ae
Order: Rosales : Rosaceae. Nalaceae . Prunaceae . Crossosomataceae,
Connaraceae . Nimosaceae. Cassiaceae, Fabaceae.
Saxifragaceae. Hydrangeaceae . Grossulariaceae , Crassulaceae, Drosera-
ceae , Cephalotaeeae, Pittosporaeeae , Brunelliaceae, Cunoniaceae ,
Nyrothamnaceae. Bruniaeeae, Hamamelidaceae, Casuarinaceae, Eucommi-
aeeae , Platanaceae
Order : Myrtales: Lythraceae , Sonneratiaceae , Punicaceae. Lecythida-
ceae, Nelas toma taceae, Myrtaceae, Combretaceae,
Rhizophoraceae. Oeno theraceae, Ha l orrhagidaceae, Hippuridaceae,
Cynomoriaceae , Ari sto lochiaceae. Raffle s iaceae , Hydnoraceae
Order : Loa sa les : Loasaceae , Cucurhitaceae. Begoniaceae , Datisca-
ceae, Ancistroc1adaceae
Order : Cactales : Cactaceae
Orde r: Celastrales : Rhamnaceae , Vitaceae, Celastraceae , Buxaceae,
Aquifoliaceae. Cyrillaceae, Pentaphylacaceae,
Corynocarpaceae, Hippocrateaceae , Stackhollsiaceae. Staphyleaceae ,
Geisso l omataceae, Penaeaceae, Oliniaceae , Thymelaeaceae , Hernandia-
ceae, Elaeagnaceae, Myzodendraceae, Santalaceae . Opiliaceae , Grubbia-
ceae , Olacaceae. Loranthaceae, Balanophoraceae
Orde r: Sapinda1es : Sap1ndaceae, Hippocastanaceae , Aceraceae , Sabia-
ceae, lca cinaceae. Helianthaceae, Empetraceae,
Coriariaceae, Anacardiaceae , Juglandaceae, Betu1aceae, Fagaceae,
Nyricaceae, J ulianaceae, Proteaceae
Order : Umbellales: Araliaceae, Apiaceae . Cornaceae
Superorder: Cotyloideae-Sympetalae
~ Order : Ruhiales: Rubiaceae, Caprifoliaceae, Adoxaceae, Valeriana-
ceae, Dipsacac eae
Order : Campanulales : Campanulaceae, Goodeniac eae. Stylidiaceae ,
Calyceraceae
Order : Asterales: Helianthaceae, Ambrosiaceae, Hel eniaceae . Arcto-
tidaceae . Ca l endulaceae . Inulaceae . Asteraceae ,
Vernoniaceae , Eupatoriaceae , Anthemidaceae , Senecionidaceae , Cardua-
ceae , Mutisiaceae , Lactucaceae

12 . Hallier, 1905
Independently developed ideas similar to those of
Hans 1912
Bessey. Sys tem was designed to be phylogene t ic.
Saw angiosperms as monophyletic from a Bennetti-
talean-type ancestor with some Narattialean affinities . Regarded dico ts as
older and more primitive than monocots . Monocots thought to have been derived
from stocks ancestral to the Lardizaba1aceae . Po1ycarpy and srirality of part s
considered primitive, syncarpy and cyclicity as advanced. Placed large signi-
ficance on ovule morphology and position. Defended use of many kinds of char-
acters for classification . His realignment of families displaye d many insi gh t s,
such as the alliance of Sali caceae with Flacourtiaceae , Cactaceae with Aizoa-
ceae. Cucurbitaceae wit h Passiflor aceae, etc. Four pr i mary divisions of dicots:
Proterogenes (5 orders), Anonophylales (5 orders), Rhodophyles (7 orders),
Ochnigenes (12 orders ). His 1905 and 1912 treatments differ quite a bit; recog-
nized 213 families in 1912 .

13. Rendle , 1930Basically an Englerian - type system (held to amen-

Alfred Barton tiferous primi t iveness) , but wi th some borrowings
from Bentham and Hooker. Did not claim his
arrangemen t to be strict l y phylogenetic. Treated monocots as more primitive
28 601

than dieots . Grouped clieots into three grades corresponding to level s of dif -
ferentiation of the perianth (compare with use of grade concept by Benson) :
Monochlamydeae (simple type of flower, no petals, occasionally with calyx;
some thought to be primitivel y, others secondarily, simp l e; cons i dered
by Rendle to be polyphyletic)
Dialypetalae (Ranales. with free petals (tepals), many spirally arranged
parts .-+ Umbelliflorae . \dth cyclic, reduced parts; considered by Rendle
to be po l yphyletic)
Sympetalae (higher grades of floral development; pentacyclic ~ tetracyclic +
tetracyclic and epigynous).
Phyletic criteria : wind pollination and woodiness considered primitive.
Naoy taxa were tentatively assigned positions awaiting f urther data.

14 . }fez , Karl C. 1926 Produced a "phylogenetic tree " based (in part)
on serological data (see Chapter 13, p . 288) .

15 . Hayata, 1921 Produced a "dynamic sys tem" of angiosperm classi-

Bunzo fication, \,Ihich was basically a rearrangement of
Engler's system . Hayata ' s arrangement points out
the reticulate nature o f character variation in
the angiosperms.

16. Burtt-Davy, 1937 Cooperated with the ,wod anatomist Chalk to pro-
Joseph duce the following classification of dicots :
Amentiferae/Polystemonae (Divs. Apocarpicae,
Syncarpicae)/Oligostemonae (Divs. Pentacyclicae,
Tetracyclicae) .

17. Pulle, A modified Englerian classification. Dicots

1938
August A. patterned after Wettstein; divided into 8 series.
1950
Considered the Sympetalae to be polyphyletic and
divided them up among the various series . Did not consider the Nonochlamydeae
to be a natural group . Revised his classification in 1950, incorporating such
changes as merging Cae tales \,lith Centrospermae .

18 . Skottsberg, 1940
A modified Englerian classification, with Wett-
Carl steinian influence. Monocots considered derived
f rom an unknown primi tive dicot. Considered
apocarpy and polycarpy as primitive, syncarpy and monocarpy as advanced in
both monocots and dicots. Apetalous forms considered polyphyletic . Amenti-
ferae placed after Rosales, but not redistributed among petali ferous families
to which they appear to be more closely related (i.e., case of Sal icaceae) .
Began dicots with Casuarinaceae, but did not cons i der them to be ancestral
to other dicots .

19 . Gundersen, 1950 System only for dicots . Divided into 10 infor-

Alfred mal, more or less tentative groups: Magnoliflorae,
Cistiflorae, Thea Group , Rosaeflorae , Ulmus Group,
~illiva Group , Geranium Group , Dianthiflorae , Jasminiflorae, Rubiflorae . Con-
sidered "Hagnoliflorae" basal to other groups. Took much modern data into
account (Hork of Sprague, et al . ).

20. Hutchinson, 1926 Basically in the Bentham- Bessey tradition.

John 1934 Angiosperms considered monophyletiC, possibly
1959 from an ancestral group of gymnosperms rel ated
1969 to the cycadeoids . Divides dicots into those
602 Figure 28-2

RELATIONSHIPS of the ORDERS

(a ft er Bessey, 1915)

OPPOSITIFOLIAE

OPPOSITIFOLIAE

RUBIALES

AL TERNIFOLIAE

ORCHiOALES
~
MYRTALES !;
C~
.
t
~

I RIDA L ES ROSALES

RANALES GERANIALES
 ~
~
RELATIONSHIPS of the ORDERS of DICOTYLEDONS ~
c
(a fte r H ul chi n son) "m
N
,
Cosuarinales Polygolales 1
Sopindoles
Apocynales
00
w

t
Viololes
Meliales
\ Rubioles Verbenales
Bolonops idales R utole s \ ~
Tomoricales ..,..,Signonioles Ebenales Asteroles
ug an a es~ ~ontololes
J I d I
Logonioles ../

LeitneriOI~ V Rhomno~
Urlicoles Myrslnoles Goodenioles

es\
Myrtales
I
Cappo rl doles \
. ' Olocolesl
tI
"-
.
Mol pighiales --Euph orb loles
Personoles
t
My ricales Fago les Gu t tife roles Cit I - r ~ ;> L om iol es
.......... e as ro es - - -~:
~ ~ I
.
Aro l loles . \ \ - Eri coles
sal lcale~ " . l' T ilioles .. Ma lvale s Va lerianol es Sol onoles

C----';" I
unonlQ es
~/
I ,,,,,,umelldales
-f
Plttasporales

/
V

' \
.
/ cu curb,to,ei
Ca clales

Primuloles
Pl ontog inales
"'-

"Campana les
~
Bo raginales
I
Polemon ;ol es
I
.
Legummole s

Coriariales
----..,
Rosales
\
T heales
~
1/.
/
Och nales

Bixoles - - - -
V
:..,--Looso les
Po sslf loroles

~Thymelaea\eS
p
__ Protea les \
~
Cheno odiole s
.
Onogroles

Gentlanales
\.
Umbelloles
\
Geron iol es
!
Podostemonoles
.
Sarracenlole s
~ Dilleniales C ophullales
Polygonales - - - ar y , . Resedales
Saxlfraga les ,
Louro les Annonales Piperales • Berber idoles Bro s ~ ic oles

" / /' '-.. Rh oeodoles

'----. \ Ari sloloch iales / " ...........
Magnoliale s Ronoles-""""' IO MO NOCOTYLE DONES
L1GNOSAE ______ HERBACEAE
( fUndamentall~ ______ (funda menta l ly herbaceous)
DICOTYLEDONES
I ~
a
w
HYPOTHETICAL PROANGIOSPERMS
604 28

considered "fundamentally and predominantly" woody (Lignosae) and those con-

sidered "fundamentally and predominantly" herbaceous (Herbaceae). Parallel
lines of development He re considered to have occurred within these two main
groups .

l:Ierbaceae Lignosae
- . - - - - -__ Ranales Hagnolia1e, __- - -
(Ranunculales)

This is considered an unnatural division by most workers (note wide separa-

tion of Umbelliferae f rom Araliaceae, Labiatae from Verbenaceae. Bignonia-
ceae from Scrophulariaceae. etc.). Gamopetalae considered polyphyletic,
rather than diphyletic as in Bessey's system. Revision of the monocots more
widely accepted, similar to that o f Lindley , based more on nature of perianth
than on ovary posit i on . Often did not give reasons for changes in the dis-
pos i tion of orders and families (hm..rever, here He do not downgrade the value
of a good taxonomi c i ntuition) . Relied for the most part on morphological
characters, to the exclusion of much recent information from the fields of
anatomy , embryology, chemistry , cytology, palynology, etc. Excellent knowl-
edge of the \..rorld flora . Accepted a large number of orders and families--a
"splitter." Se e chart of Hutchinson's system, p. 603.

21. Benson, 1957

Basically a Besseyan-type system . Dicots divided
Lyman into f i ve informal polyphyletic grades "representing
stages in floral evolution." (See chart, p. 605).
OTHER recent Ranalian-based systems include those of 500 (1953), Novak (1954),
and Emberger (1960). Dey1 (1955) and Kimura (1956) have presented classifi-
cations for the monocotyledons .

22 . Takhtajan, 1943 A system in the Bessey- Hallier tradition ; strongly

Armen 1953 influenced by Hallier. The' division into subclasses is
1959 considered a major conceptual advance by many (for
1966 characters of Subclasses see discussion of Cronquist ' s
1969 system below) .

MAJOR GROUPS IN TAKHTAJAN'S SYSTEH (1969)

CLASS MAGNOLI ATAE (dicots)
SUBCLASS HAGNOLIIDAE SUBCLASS ROSIDAE
Superorder Hagnolianae Superorder Rosanae
SUBCLASS RANUNCULI DAE Superorder Rutanae
Superorder Ranunculanae Superorder Aralianae
SUBCLASS HAMA}illLIDIDAE Superorder Celastranae
Superorder Hamamelidanae Superorder Proteanae
SUBCLASS CARYOPHYLLIDAE SUBCLASS ASTERIDAE
Superorder Caryophyllanae Superorder Lamianae
SUBCLASS DILLENIIDAE Superorder Asteranae
Superorder Dillenianae
Superorder Er icanae
Superorder Malvanae
CLASS LI LIATAE (monocots)
SUBCLASS ALISNIDAE SUBCLASS COHHELINIDAE
Superorder Al ismanae Superorder Juncanae
SUBCLASS LILIIDAE Superorder Commelinanae
Superorder Lilianae SUBCLASS ARECIDAE
Superorder Arecanae
 ~
~
DICOTYLEDONS MONOCOTYLEDONS ~
C

•"
~m0D2-
.
~
COROLLIFLORAE THALAM I FLORAE "0

o~ ~
"'Ulo\.~ "<>~ "
e \~~
""O"laq,~a ""'~ .
0 / .,

",<1'O<hO'\'o\~~
0

E
,~~

0
01.,

, C>
alli T r ; OhO\~~
>~
,
,0
B
i)-~~
~
, kCY/
Vo'~

..tJ./l
,
"'~."QI~~
~<><,,,<:::::> - '. ' eo,{fIO"
~''''~.
~"illllirr>
s~>;i~;mm~:;~
OI>~"'a, icIC'
i'
0", C
<>"~"i~. r-::.~
'-...)~.
%,

G~"iI. ,al"

(arn tole'
~ ~ ;QIO\~' ~\ .~

0 , ," o,o~~., p <::::> ~.,

=
Ca',oa\~ po~<I .. ~ta p',ge O<1o,, ~mo\

~
OVA RIFLOR AE CALVCIFLORAE
, '0,
'0
00

"
---<1.,
~ 4f0 ,
r!!ll
Flu~ial"
wy'c'O-
Coto'\>'\
~"
B ~ QO~'
,,0 ,\'"\J
C? ..

~
\~ ~

~ ~
l~

~
,
El oet><l"<>
~' -0.'-1
Co"'90"~' s~
""
,
""'"
o~

W,o
,0'
Umb.\\tI l e~ \~~,(0)S-",'"
~;
~\~t
\<>
O\.~
COr~Q
A" , toI OO >
%",0
CHART of RELATIONSHIPS AMENT !FERAE
, 0,
"
\ ~()
of ANGIOSPERM ORDERS
GO"~
~\~~
o o~.t> (; "dO\'~ ,"
-.
00
M1" ~ ~~ ~" %0\J
Hypo~ynous and - PeriQynous Or Ep i ~1nou s FoqO ", o n ",,~~
0
l'il~eriO i 00

'"
Jui lO
O
Choro peto louS or 3I:"':'1ChO'ipetoto~5 or
Aperclcus .~ Apetalous
t:..Y~Q~ O \~~ (;1 ;;'"
o ~,~.. sol;~o
JuOIQ~ 8a IO~c'<
CO, yO "" All., 8M,on . 1957
Symp~tolous
606 28

A " splitter " at the ordinal and family levels who takes much recent palynologi-
cal , anatomical, embryological, paleobotanical. chemic al, and cytological data
into account. An English s ununary of this important sys tem i s given in Takhta-
jan (1969). More complete expositions of his system are given in the 1959 and
1966 references (i n German and Russ ian respectively) . Active system-making
(often in cooperation \"rith A. Cronquist) is ongoing .

23 . Cronquist , 1957 Concept ually similar to Takhtajan's system , differ-

Arthur 1965 i ng in detail . Subclass Ranuncul1dae merged with
1968 Ha gn oliidae. Neither much of a lumper n or much of
a splitter. Gr oups conside r ed polyphyletic are
split up and placed with taxa to \.;rhich they appea r to be phyletically related
(note treatment of Amaryllidaceae), and many smal l, somewhat anomalous or un-
usual groups are retained in the larger families to which they appear most
closel y r e lated. Huch moder n data taken into account (see Table 28-2, below) .

CHARACTERS OF THE SUBCLASSES (Cronquist , 1968. Evolution

and Class ification of Flowering Plan ts)

Dicotyledon s Honocotyledons

MAGNOLIIDAE--Apocarpous, pol ypetalous ALISHATIDAE--Hostly apocarpous ;

(or apetalous); perianth evident , aquatic , always herbs; pollen tri-
stamens numerous, centripetal; pollen nucleate; e ndosperm wanting or not
binucleate, o f ten uniaperturate; ovules starchy; stomate subsidiary ce lls
bitegmic, crassinucellate. mostly 2 .

H~~LIDAE- -Fl owers reduced, often ARECIDAE--Flowers numerous, small,

unisexual, often in catkins; perianth with spathe, sometimes on spadix;
poorly developed or wanting; never leaves broad, petiolate , usually
with numerous seeds on parietal pla- net-veined; ovary superior or
centae . sometimes sunken in axis of spadix;
subsidiary cells 4 (3-2).
CARYOPHYLLIDAE--Stamens centrifugal ,
pollen trinucleat e ; ovules bitecmic , COl'lt-IELINI DAE--Flowers with well-d efined
crassinucel l a t e , often campylotro- sepals and peta l s , or perianth chaffy
pous or amphitropous; seeds often or bristly or lacking , not on a spa-
with perisperm; plants usually Hith dix; subsidiary cells 2- 4 or more .
betalains or free-central to basal
placentation; mostly herbs . LILIIDAE-- Flowe rs few to numerous,
often large, not usually sub tended
DILLENIIDAE-- Stamens cent rifugal, by spathe and never on spadix ;
pollen binuc l eate (exc ep t Cr ucife rae); l eaves linear and parallel to
ovules seldom campylotropous or amphi- broader and net-veined; ovary
tropous; seeds seldom with uerisperm; superior or (more often) inferior ;
no betalains; many species t rees . subsidiary ce lls usually 0 , less
often 2 , rarely more.
ROSIDAE--Stamens numerous, centri-
petal, or fewer and opposite or
alternate petals; ovules bi t egmic
or crassinuce lla te ; rarely sym-
petalous .

ASTERIDAE-- Sympetalous; stamen s few,

never opposite petals , ovules unitegmic
and tenuinucellate; carp els usually 2 .
Figure 28-5 607

RELATIONSHIPS of DICOTYLEDONS
(Adapted from Cronquist , 1968)

Asterol e s
AS1(R I(ME Scraphulorioles Companulales \
Plonloqincles --- I Qlpsoeole
Lomioles- Polemonioles Ru~'oles
Genlionoles

ROSIOAE ASTERIOAE

801 0 ceoe
Plumb09inaceae

'''''''Z''''
COryOphyiloceoe
1""'""00"" A,zooce oe

"':i",,,,~, 0"""'"
Umbelloles / Geronioles 80selloceae Caclaceoe

I y Linoles

Sapindol e~ - - PalYQololes Amaronlnoceo e - ·Chenopodioceoe - Phyloloccaceao:- - NyCIOQinoceae

Rhomnoles
CARYOPHYLUOAE
CeloSlroles - EuphOrbial es

~
Sontalales - Rofflesiol e s

COrnoles

Podoslemoles PrOleoles
'-Rosoles Myrloles
/
Holo.OQOles
s':''''''

~
E ""'"
Oiapensioles ViOl ales

I~
ROSIOAE
Coppa roles
Sorroc eni ales
£b en a l e s > - -
Theales Molvales
,,,,,,,,,,,,
Pr i mulales I
Leilneriales Myricales Oilleniales

Urlicoles ':,
\"'''"'''''~ FOQol es
Eucommiales
J
Hamamelidales
Cosuorinales
OILLENIIDAE

Trochodendral e s I
HAMAMEUOAE - - _...._

/ Popoverales

Nymphaeales Ranunculales

Ar;stoloChiO leS , 1 / ""

Pi pe.oles ~ MOCjnolioles
MAGNOLIIOAE
 608 Figure 28-6

RELATIONSHIPS ~MONOCOTYLEDONS
(Adopted from Cronqu ist, 196 8)

O,en,d oles
Z i ngibera les
/
Lilioles

Juncoles Cype,oles /
LlLlIDAE
ReSliOnOleS---Y ~OleS
(rioeaul a les - -- Commelinoles

~===~
COMMELINIOAE

Nojodole.
~H~ droc~ortta les
~

,,;.m,c/
\ Triuridoles

LlLlIOAE
ALlSMA110AE

~ L ISMAT I DAE
28 609

24 . Thorne, 1968 A system in the Bessey-HalHer tradition presented

Robert with the fol lowing major taxa (from Thorne, 1968):

SUBCLASS DICOTYLEDONEAE
Superorders : AnnoniElorae Rutiflorae
Nymphaeiflorae Chenopodii f lorae
Sarraceniiflorae Hamamelidif lorae
Theiflo rae Rosiflorae
Rafflesiiflorae Proteiflorae
Cistiflorae Hyrtif lorae
Nalviiflorae Gentianiflorae
Santaliflorae Corniflorae
Geraniiflorae Lamiiflorae
Asteriflorae
SUBCLASS HONOCOTYLEDONEAE
Superorders: Alismiflorae
Triuridi florae
Liliiflorae
Ariflorae
Commeliniflorae
Takes a broader view of families than most . Aims at having a uni form size of
morphological gap separating groups of equal rank (thus places Amaryllidaceae
in Liliaceae . etc . ) .

II. EXAMPLES OF RECENT OVERALL CLASSIFICATIONS OF VASCULAR PLANTS

A. ~. O. 1942. A modern classification of the plant kingdom .

Chronica Botaniea 7: 203- 206 .

Phylum Tracheophyta (Tracheata) Order Ophioglossales

Sub-phylum PSilopsida Order Marattiales
Class Psilophytinae Order Filicales
Order Psilophytales Class Gymnospermae
Order Psilotales Sub - class Cycadophytae
Sub-phy l um Lycopsida Order Cycadofilicales
Class Lycopodineae (Pteridospermae)
Order Lycopodiales Order Bennettitales
Order Selaginellales Order Cycadales
Order Lepidodendrales Sub-class Coniferophytae
Order Pleuromeiales Order Cordaitales
Order Isoetales Order Ginkgoales
Sub-phylum Sphenopsida Order Coniferales
Cl ass Equisetineae Order Gnetales
Order Hyeniales Class Angiospermae
Order Sphenophyllales Sub-class Dicotyledoneae
Order Equisetales Order Magnoliales (Ranales)
Sub-phylun! Pteropsida Order Ericales . etc .
Class Filicinae Sub-class Monocotyledoneae
Order Coenopteridales
610 28

B. Banks , H. P . 1968 . The early history of land plants . Symposium on

Evolution and Environment . E. T . Drake (ed . ) . Yale University
Press . New Haven .

Division - Tracheophyta
Subdivision - Rhyniophytina
Order - Rhyniales
Fami l y - Rhyniaceae
Cooksoniaceae
Subdivision - Zosterophyllophytina
Order - Zosterophyllales
Family - Zosterophyl laceae
Cossl i ngiaceae
Subdivision - Psilophytina - Psilotales
Subdivision - Lycophytina
Subdivision - Spenophytina
Subdivision - Tr imerophytina
Subd i vision - Pterophytina
Class - Cladoxylopsida
Class - Coenopteridopsida
Class - Filicopsida
Class - Progymnosperrnopsida
Class - Cycadopsida
Class - Coniferopsida
Class - Gnetopsida
Class - Angiospermopsida

c. Bie r horst , D. H. 1971 . Morphology of Vascular Plants . The Nacmillan Com-

pany . Nel" York.

Divis i on - Tracheophyta Class - Ophioglossopsida

Class - Rhyniopsida Orde r - Ophioglossales
Order - Rhyniales Class - Narattiopsida
Cl ass - Zosterophyllopsida Order - Marattiales
Order - Zosterophyllal es Class - Aneur ophytopsida
Class - Lycopod i opsida (Lycopsida) Or der - Aneurophytales
Orde r - Asteroxyla l es Order - Archeopter idales
Order - Lycopodiales Order - Protopityales
Order - Protolepidodendrales Class - Cycadopsida
Order - Selaginellales Order - Pteridosperrnales
Order - Lepidodendrales Order - Cycad ales
Order - Isoetales Order - Cycadeoidales
Class - Cladoxylopsida Order - Caytoniales
Orde r - Cladoxylales Class - Conife r opsida
Cl ass - Equisetopsida (Sphenopsida) Orde r - Cordaitales
Order - Hyeniales Order - Coniferales
Order - Pseudoborniales Order - Taxales
Or der - Spenophyllales Order - Ginkgoales
Order - Equisetales Class - Gnetopsida
Class - Coenopteridopsida Order - Ephedrales
Order - Coenopteridales Order - Gnetales
Class - Filicopsida Order - \-1elwitschiales
Order - Noeggerath iales Class - Angiospermopsida
Order - Filicales Subclass - Dicotyledonidae
Order - Narsileales Sub class - Nonocotyledonidae
Order - Salviniales
28 611

Table 2B-1. Comparison of treatment of Cronquist I S Subclasses l~ith some

other recent systems .

I. DICOTYLEDONS:

Subclass Hare or less compa rable to group traditionally

known as :
HAGNOLIIDAE Ranalcs
HAHAMELIDAE Ament iferae ~. ~.
CARYOPHYLLIDAE Centrospermae
Seen as natural groups separately derived from
DILLENIIDAE Hagnoliidae . the same sorts of evo lu tionary ad -
ROSIDAE { vances occ urring in both groups , but with dif -
ferent freque ncies - Cronquist
ASTERIDAE Sympetalae

A. Takhtajan (1969) , Hagnoliidae divided into two subclasses - Magno-

liidae, Ranunculidae (includes Sarracen iaceae)
Ilamamelidae - Takh taj an accepts mure orders , some
differences in sequence .
Caryophyllidae - Takhtajan includes Theligonales
Dilleniidae - Takhtajan places Euphorbiales and
Thymel eales here.
Rosidae - Takhtajan places Oleales here, includes
Umbellales in Co r nales, includes Nepenthaceae
and Droseraceae he r e .
Asteridae - Takhtajan gives Dipsacales a basal
position, includes Rubiaceae in Gentianales ,
shuffles some families between Po lemoniales and
Sc rophulariales .

B. Thor ne (1968) : The identities of the Nagnoliidae , Caryophyllidae,

and Hamameli dae are essentially maintained ;
Rosidae , Dilleniidae , Asteridae mixed .

C. Hutchin son (1959): Al l the dicot subclasses , except for the Hamame lidae,
are dispersed throughout the system.

D. Nelchior (1964) : Informa lly recognizes , in broad outlines, the Hag-

noliidae , Hamamelidae, Caryophyllidae , and
Asteridae as natural groups ; Rosidae and Dilleni -
idae mixed together .

II . l'lONOCOTYLEDONS: Subclasses! ALISHATIDAE. ARECIDAE, COHHELINIDAE , LILIIDAE

A. Takhta j an : Takhtajan's treatment of the mon oco ts is similar to

that of Cronquist. Some differences: In the Alis-
matidae, Takhtajan removes the Tri uridales to t he
Liliidae; Takhtajan places the Typhales i n the Are-
cidae rather than in the Commelinidae ; includes the
Zingiberales in the Liliidae , rather than in the
Commelinidae.
 612 28

Table 28-1, continued

B. Thorne: Thorne's 5 superorders of monocots compare closely with

Cronquist ' s subclasses, thus :
Alismatidae ± equivalent to Alismiflorae +
Triuridiflorae
Ar ecidae equivalent to Ariflorae
Cornmelinidae equivalent to Commeliniflorae
Liliidae equivalent to Liliiflorae
C. Hutchinson: Alismatidae kept together , but Cronquist's other three
monocot subclasses are mixed together.

D. Melchior: Hel ob i ae comparable to Alismatidae of Cronquist, but

Cronqu i st's other three monocot subclasses are mixed
together.

Tab l e 28-2. Examples of "Hodern" Criteria Considered by Cronquist (from

Cronquist , 1968) .

Criterion Level or groups where used

Anatomi cal--

Stratification of phloem into fibrous

& non-fibrous tissue Sarcolaenaceae/Halvales
I nternal phloem Caryophyllales; Halvales vs . Lecy-
thidales; in Rosidae (Hyrtales);
Gentianales
Xylem vesselless Hithin Magnoliidae; Nymphaeales
Ray Hidth Ranunculales
\~ood anatomy Columelliaceae
Wood parenchyma diffuse vs. vasi-
centric Umbellales vs . Cornales
Unusual vascular bundles Thurniaceae
Nodal anatomy subclass. ordinal levels; Umbellales
vs. Cornales
Stem with anomalous secondary
thickening Caryophyllidae
Central axis of gynoecium ~'ith or
without vascul ar bundles Umbe l lales vs. Corn ales
Schizogenous secretory system Myrsinaceae ; Umbellales VS. Cornales
Laticifers Musaceae
Secretory canals Guttiferae. Sapindales
La tex sys tern Papaverales/Capparales; Sapotaceae;
Celastrales; within Gentianales
Elongate idioblasts Trochodendrales
Ethereal oil cells Magnoliidae
Foliar glands Leitneriales , Hyricales, Juglandales
Myrosin cells Capparales ; Tropaeolaceae
Table 28-2. continued

Hucilage cel1s Malvales

Raphide sacs in Zingiberales
Raphides . crystal sand Dilleniaceae
Silica bodies in l eaf epidermis Thurniaceae
Stellate hairs l-lalvales; Styracaceae
Colleters (specialized glandular
trichomes) Rubiaceae-Gentianales
Guard cell symmetry/asymme try within Zingiberales (Lowiaceae)
Subsidiary cell number subclasses of mono cots
Nectary type Malvales; monocots vs. dicots; within
Liliidae
Embryological--

General embryological studies Agavaceae

Ovules unitegmic vs . bitegmic subclass l evel and below
Ovules tenuinucellate vs .
crassinucella te subclass level and below
Endosperm vs. perisperm Magnoliidae; Caryophyllales
Endosperm cellular vs. nuclear in Polemoniales; Rubiales-Gentianales-
Dipsacales
Endosperm helobial in Alismatidae. Arecidae
Endosperm starchy vs. nonstarchy subclasses of monocots
Endosperm haustoria Ericales/Diapensiales
Integumentary tapetum within Asteridae
Chalazosperm Taccaceae vs. Cyanastraceae
Embryo sac bisporic vs. monosporic in Alismatidae
Possible free - nuclear stage in
embryo development Paeonia

Cytological--

Chromosome number Agavaceae/Xanthorrhoeaceae

Karyotype Agavaceae

Chemical --

Cyanogenetic compounds within Vinlales

Alkaloids. glucosides Gentianales
Gentiopicrin Gentianaceae
MeUat!n Menyanthaceae
Tryptophane a l kaloids Rubiaceae-Gentianales
Isoq uinoline alkaloids.
especially berber in Ranunculales. Papave rales
Betala!ns Caryophyllidae
Unusual fatty acids within Geraniales
Sero logy Papaverales/Capparales; Cornales ; Capri-
foliaceae-Cornaceae

Palynological--

Pollen binucleate vs. trinucleate subclass level; Sapindales ; in Polemon-

iales
 614 28

Table 28- 2 , continued

Pollen trlapertur ate vs. uniaper-

turate subc l ass (Hagno l iidae)
Pollen non-apertura te Scheuchzeriaceae
Other pollen characters Eucommiales, Hamamelidales. Casuari-
nales; Halvales ; Staphyleaceae ;
in Restionales; Ruppiaceae ,
Zosteraceae
Physiologica l--

Adaptation to NO) deficiency Sarracenial es

Adaptation to water stress Caryophyllidae ; some Violales
Mycorrhizal habit Ericales; Triuridales; Orchidales

Other--

Pos it ion of vessels i n pl ant Commelinidae; Liliidae

Pe t a l s sepal vs. stamin odial-
derived Paeania
Androecial ontogeny delimitation of dieot subclasses
Gynoecial ontogeny Lauraceae/Myris ticaceae
Pollination metho d subclass (Hagnoliidae, Hamamelidae):
subclasses of monocots
Fossil pollen record Hamamelidae
Carpel sealing/ unsealin g Na gnol iidae

,
Sec tion B. COHPARISON OF CLASSIFICATIONS

Pr esent ed i n this sect ion i s a comparison, in tabular form, of the

flowering p l ant c lassifica t ion systems of Cronquist. Takhtajan, Thorne ,
Engler. Hutchinson . and Bentham & Hooker . The linear sequence of orders and
families is that presented in Cronquist (1968) . All families accepted in any
of the six above-mentioned systems are accounted for , as I"ell as all families
for l"hich a description is given in Airy 5hal" (1966) .

The system of Cronquist is that presented i n The Evolution and Classifi-

cation of Flowering Plants (968). Several changes in the system wh i ch a r e
anticipated by Cronq uist are indicated in foo tnotes .

The syst em of Takhtajan is that presented in Flowering Plants : Origin

and Dispersal (196 9). Takhtajan r S Systema et Phylogenia Hagnoliophytorum
(1966) was consulted for the disposition of certain segregate families not
treated in the 1969 book .

The system of Thorne is that presented i n Aliso 6(4) (1968) . Inasmuch

as that system is as yet available only in outline form , the treatment of
taxa not spec ifi cal ly mentioned by Thorne follows that in Helchior (1964) ,
as was indica ted by Thorne himself.
28 615

The system of Engler is that presented in Syllabus deT Pflanzenfamilien

(Helchior, ed ., 1964) . Dalla Torre and Hanns' Genera Si phonogamarum (1900
1907) was consulted fo r the disposition of certain segregate families not
treated in Melchior (1964) .

The system of Hutchinson is that presented in The Famil ies of Flowering

Plants (1959) . A number of changes in the ordinal placement of certain
dicotyledonous famil ie s, as \.,rell as the addition of several new families,
\·/hich were incorporated in Hutchinson' 5 Evolut ion and Phylogeny of Flowering
Plants (1969) . are accounted for in the table and in footnotes .

The system of Bentham and Hooker is that presented in Genera Plantarum

(1862-1883) . In the Polypetalae and Gamopetalae of these authors. cohorts
are taken to correspond to orders in the modern sense. and their "orders" to
families in the modern sense. !/O\"eve r, in the case of the Nonochlamydeae
(Incompletae) and Honocotyledonae , their series are treated as orders in the
modern sense, since the series in the latter t\W major groups Here not divided
into cohorts. The correspondence of the categories of Bentham and Hooker to
those of contemporary authors is not exact. Bentham and Hooker recognized
at the sub familial (subordinal. in their terminology) level of "tribe" many
groups which are today considered families (cf . the "tribes" of Geraniaceae ,
Najadaceae , Scitamineae, etc.) .
Airy ShaH, in Hillis I A Dictionary of the FloHering Plants and Ferns ,
ed . 7. (1966). gi ves descriptions for a number of families not accepted in
any of the six systems . These segregate families are included in the table
and are indicated by the symbol (AS+) . I n most cases , description of a
family by Airy Shalv indicates his recognition of family stat us for the taxon.
A number of families not accepted in any of the six systems nor by Airy Shaw
are also included.

Families not accepted in any of th e six systems are indicated by [brack-

ets] . The generally accepted placement of these groups is indicated.

A question mark (?) indicates uncertainty regarding the position of a

taxon, or to the level at Hhich it should be received. A dash (--) indicates
that the taxon Has not treated.
The alphabetical index to 648 family names vhich follO\"s the table in-
cludes vell-kno\offi synonyms and a number of variant spellings not included
in the body of the table.

FOOTNOTES

a - subsequent to the nomenclatural sessions at the 11th International Botani-

cal Congress in Seattle in 1969, both Cronquist and Takhtajan have adopted
- opsida endings for classes. Thus : Magnoliopsida and Liliopsida , rather
than Magnol iatae and Liliatae.
b - Cronquist would nO\" refer this family to the Euphorbiales .
c - excluding Rhabdodendron? (Takh . ).
d - perhaps belongs in Portulacaceae (Takh.).
e - Hectorella in Portulacaceae, Lyallia in Caryophyllaceae? (B. & H. ) .
616 28

f - Sauvagesieae in Vi o l aceae (E . & H. ) .

g - as for Rhaptopetalum (E . & H.) .
h - affinities of Hua obscure (Cronq.).
i-including Afrostyrax? (Takh.).
j - i n Bixineae. ~"hich also includes Bixaceae (B . & H. ) .
k - as Samydaceae (Flacourtiace ae p . p.) (B. & H.).
m - including Paropsieae? (Takh . ) .
n - near Flacourtiaceae (Bixales)? (Hutchinson, 1969).
o - in Bixineae . which incl udes part of Flacourtiaceae (B. & H.) .
p - including Oceanopapaver (Takh .).
q - Oxystylidaceae (Cappara l es) is a segrega te from Capparaceae (Hutchins o n ,
1969).
r - Pr i onotaceae is a segregate from Ericaceae and Epacridace ae (Hutchinson,
1969) .
s - abberrant in Dioscoreaceae (Cronq.).
t - as treated in Takhtajan (1966), not mentioned in Takhtajan (1969).
u - Cronquist would now refer this family to the Trochodendrales .
v - excluding Polygonanthus, i ncluding Corokia ; Ca rpodetus very isolated
(Takh.).
w - excluding Stylobasium? (Takh . )
x - Cronquist would now place this f amily near th e Solanaceae.
y - placed in Saxifragales (position uncertain) in Hutchinson (1969) .
z - pl aced in Myrtales in Hutchinson (1969).
aa - pl aced in Onagrales in Hutchinson (1969) .
bb - position of Coroki a uncertain (Cronq.).
cc - excluding Corokia; Griselinia very isolated (r akh . ) .
dd - Campylostemonoideae, Cassinoideae , and Tripterygioideae very isolated
within the family (Takh.) .
ee - as treated in Hutchinson (1969) (but places Nymania i n Meliaceae) .
ff - Dapania in Geraniaceae (Oxalideae) , .~ arcotheca in Linaceae (Ixonantheae).
gg - excluding Peltanthera (Takh.) .
hh - Salpiglossidaceae is a segregate from Solanaceae (Hutchinson, 1969) .
ii - Thorne (personal communication) would now treat as distinct family .
jj - includ i ng Pe1tanthera (Takh.).
kk - includ i ng Nyctanthes (Takh.).
11 - Of tia is very isolated within the family (Takh.) .
mm - in Theales? (Takh . ).
nn - Acorus is very iso l ated within the family (Takh.) .
00 - the proper name for this family is Surianaceae (Cronq . ) .

A - as treated i n Hutchinson (1969).

B - among Genera anomal a (E . & 1-1 . ) .
C - as t r eated in Dalla Torre & Harms .
D - affinities uncertain (Cronq . ) .
E - segregation as a separate family probably justified (Helchior, 1964) .
F - among Genera dubia (E . & H. ) .
28 617

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CRO:lQUI ST TAKHTAJi\ll mORNE ENGLER HUTCHlNSON BE)"'THAM S. HOOKER

a l!! C:y mph aeaceae 25 Cabombaceae l!!. Nyrnph a eaccae i i in 1'>ymphaeaccae 253 Rana l es l!!. Nymphaea"-"a,,
I"ympnaeales)
b J!l Nymphae""-ca,, 2 7 Barcl.~vac eac l!! Nymphae"cea e i n Nym;>hacaceae in Nymph aea"-e",, i n Nymphaeac,,-ae
(Nymphae"!,,s)
Ie Eurvala c eae (A5+) : In Npphaea c c ae ]
[d Nupha racea e ; In Nympha ea c"ae ]
25 Ne lumbo naceac 31 Nelumbonal e s in Nymphae"c" a ,,! i l!!.
Ny::,.ph"e"ccac in Nymp 'H. C<lC <' Ie i~. l'y,.ph,waccae
26 C" rat opb vl laceae 28 Nymphae"\es 32 Nymplt,,<>ale s 68 Ranuflclliaies 252 R.;nales 165 " Ano .... lou,; fa""

a in Ran unculnceac 3 6 Glaucid ia c"" " in '<anllfl(:lllacca c in Ran unculaceae i r. "

l i~ l l e bo t a, eac in ~,nuncula ccae
-(Ranuncula l e.'1)
b in Ranunclliaceae 37 Ilvrlra s ti daceae in '<anun c uL",:eac l!:!. Ranunc u lac eae I n Hcll c bcracea c in Ra n un c u lnccac
- (Ranun,,-ula l es)
c in Kanunculacea" 39 Podaphvl l "-"-""" itl Berb e r l.;!"cca" i n Berbe rid aceae 2); Ra na lc s in llerberldacea"
(r.a nun c u l ::!lcs)
d .!.D. ,<anuncntaccae in ~anunculaccac in Ra n uncul"ceae in Ranun c utac cac 24 8 Ilell.>borar.c u<; in Ranuncula cene
( Rana1,,-,-'--
28 (;1 r cacasteraccae J8 R'ln unculales in R::!nuncul"ccac i I in ;(3nu"cuL1ce'''' 258 Berberidales i" Chluru n thaceae
[a Kingdon taceae (AS+ ) : in Ci r c a easterace""(C ron,!.); i n R:lnuncu l aceae? (YOlk!, ,) ]
29 Be r b"rid"cca~ 41 Ranuncu lal cs 2 9 Be ,-bc ,- idalcs 6 3 Ra nuu c ul alcs 259 BC'rL~rlcl" les 7 Runales
." '. ~.v~ ,- idace ae 4 0 N" ndJnaccac in Be rberl.~acea~ ~ Bcrbe,- i daceae 257 11. r hnidal .. s in B er~er idaccac
- (Ranuncula les )
[ b Leontic" c~ae (,\S+ ) : \" Berberidaceael
,0 L, r di;:ubal"cc3c 32Ranunc u lales 25 Be r bcri dal es 65 R"nuncll l aics 2 55 Berh.,r~dal"s in B.. r b .. ridace"e
a I n La r di zaba lac , 33 Sarg e nto doxacc3c 26 B.. rbcridales 6 4 Run un culal~s 251, B"r bcridalcs
- (Ranuncuhles)
J1 ~~ n isD"rmacca~ 3 4 Ranu nc ul alcs 27 Be r Leridal .. s 66 R"nuncu la!es 25 6 B,:, rbnidales 6 Ra n,~les
32 Cor\3riacea.. 241 R" lalcs? 209 R o~ ,des 144 Sapinda l es 23 Co ,-i ariahs 56 "AnotnulollS far.! , "
33 Con"n oc ar? aceae 296 Celastrales 208 Ro~ales 158 Celastrales 181 Cela~trales in Anaca r diaceae
34 Sub:iuccac 250 Sapindalcs? 155 Rutales 150 Sapi n da l cs 217 Sn p lndales 54 Sapindalcs
[ a Mcli os!:laceae (AS+): I n Sabiac cae ]

Order 6, PAPAV£RAL ES
35 Pap,n/erac",," 1,2 Pap avcrales 30 Hc rbcrida lc s 95 Papaver a le s 267 Rho " ad"les 10 Pa rL ~ta l es
[a Pte ridop!'v llaceae (A5+): ill Pap"ve r aceae I
36 FUr.!'lriac"a,:, 41, Pap averalcs .!.D. P"pa"CI"nCC,'C in Papa\'er.~~e"c 268 Rh oe a d ales !.!:!.
Papavc ,-a ceae
a 1n 'umariac .. ac 4 3 !!)' pcconccae in P,lp ave r" ceac ~ Papavcraceae .!.!!. Fu:Mriaceae in P,w averaccac
(Papavcrales)

[Ta khta jan ' s orders lll!clale s , /<:elur..!lOn alcs, Ranunc ulalcs , Papavera l cs and ~ar racen ia l es
constit ut e. his subclnss Rnnuncu ildae]

~
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mrn:uwsol> BE~TII!\M (, IIOOn:R

CRO:<QU1ST TJ\KHTAJM TTf0RliE E!lGLER

Subclas~ 11 ; 1IA.'W".ELlDAE

1.6 T ro cb oden 'lra l u~ 17" 11"r:lm!1 ,'l lduies 59 MnKnoI1 3.1~s

, In Ma;;""l lac~.,,'

"
IMl lb=,..dldal~ ~ ,·0 ~~,s",oJlal ca In Trochod"ndra c " '''' 1. In ~:ar;n,,1 L1 r , ' ;'('
:·9
"" ~, ! l a",~..,lld"I~ ,_
~{) llldy".,Jal" ~
18~ IL~ ...~ "", IL d "h· !I
101 F... phorbhh· ~
1('2 ;I. .,sat,, ~
(, Lelt""rI"1 ,, s
~ ! II "",.. =1 [d3\"" 15(, Unisexudh''''
near IA'itn,'ria ?

""I. ; ] !i.lma..elld., l,, ·:

In Hama.,..\id ,,,; <>n,,l
184 lIam.' .... ]1<1.11 "~ 11) 3 ~<)~al~ " S1I 11 .' ..... ",..1 i d.~1"'5 6~ Rosal,,~

",
Ie
1" lIAm::,--;;e l lda <: ~ ael
(AS+) , in na -n., c e ll J"c. '".' I
.')l!a"'a,....clld"l~~ 21~_ Pl tt,"'po r3 lea ]01. ko""ie -' 51 I I'H,,"~,,,,11cl,,1" H l!!. lIa "';!1Il~ llJ"c. ' a~

"
12 rrt!c~ l " s
Order 9 : F.UCOmlII>US
4S Euco:mla,,-"''', ~4 r.uc o ...-lal cs
18Jlf; "'3""I H:>\,'''' i J,"C,C,C,CeC",C.'.,-------CC---- - - -
< 10: \ ~ RT I r."'I.r.S _
~~ u ... le a l "-" 9~ Unlr~\ ~~ I I t Tl :,," 1,' " -i.-~- -Urll " ales l!C Un 1","c"U
U! ...~ce~t:}
'':I lla "be y~h~ unkal~ ~ l l !n nr.., c~ u' n \Jrttcale~

51, U rrl c31~ s

~7 U nlcale ~
HlO UrU<:,11~"
!!1 )lor~('''''' <:
C~nnabloac"-,,e. (:nnna],!<! aceae '
~8 Urt i cate. 10\ UTtI ~ 3h,~
lJ
~
l'rtJc8 1 ~8
;-!or ',ce he

'I. Unlc.,h,"
11]
~~
t:Tllcal~"
Urtl ca l i's

71 Uri icalcs
}."
Unl~px"al ~"

Ord .. . 11, l.F.lTllER 1 ALE~ _ _ _ ,_ _ _ _~ _ __ _ __

~

S! l.el ln ~r l" c"'~" .. 7 U:ilnt'rl ni " F t6~ 1",ll';~r ; 3J 11 S I ."ltn,·r!.d,, ~ ~ 8 I.cicncrlah,,, IS] i!nl""xua\"5

1.!'. Eupl", r ]) I ,1 c, ~,' 115 RUlal e~ f.& Ju ); lanJ"le~ In Sl,....ar""bll~cac~

\i! S
16 7, kUlal,," 1 J u g l a1\d.,le~ ('5 .J " gL,ndnles nB Un isewaJrs
J " f; hnd" te~

Ordu 11, MYR1':ilLFS IS9 Un isexunl ....

6~ ~;d<alTs 16J llyri c a!';;,,' --~ 2 Jugbnda\" " ~9 Myricalcs
S5' lli:ricac"n.·

Or der 14, fA("..I\I.r.S 154 Unlsexu"h,s

JII9 !l.aianopal., s " llaiauupnles 60 8.,1.,nopsida.\cs
S6 lIa lanopac .. ae 63 Kalanopal .. s
S 7 fasa c""e
SII ~dulac ..., ~
.1 i n Het" l ac".1~
" !'~&ai .. ~
62 Iletulnles
1.!'. lIe t uincenc
187 Fagal<,s
11111 F.1l1ale~
(~ tI lI"tul.,c".';') til
q Faga\es
8 faga l "s
!I('\UhC~.l"
62
61
63
f agale"
fngal"s
\,:o r vl"c".'"
1J161 Untse xua l".
( a s f!!.!>.ullf .'r.1<')

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CR~UIS' · TAKHTAJ t\ll THOR."E ENGLER IlUTCIlINSON BE)I;,IIAM ~ IlOOKEtt

Orde r 17: MmU:S

71 Sa tac .. ae -- - -70 C"r;·-o phylla les 178 lIatidalcs >0, Ba tale s 286 Chc nopoclt alc~ 1 35C-u-~ve-",b-r)'ae

Order 18 : roLyc;o;MLF.S
J2 Po lynonacc.~e 84 Po fy&ona l es 177 Ch"nopodL~les 25 Polygonalu 176 Poiygon>!lc s D6 cu r v .... bryae

Orde r 19 : PLU~IRAG I NALf.~

'i3- Pl umb,,&jnM .. a~ 85 PJ.Ull'\> a glnallls 67 Plu~baginales 237 Pl umbng inaics 19G-Pr\t:'.uL!. les 100 Primul a le s

Subd3SS IV: DILLENIID.'E

Ord e r 20 : DI LLENt ALES

H D1l 1e ninc ea" 87 Dtl i~n-i~- J~ Th ..<lles -- ---- -----r6(;;;-tc iTeral es -- 19 u1l1 enT:llei---- 2 g,,';-;"-fu
75 P""oniaccac 89 Pneonialcs 35 Thcales 77 Gu tUfera les 24 7 Ranale~ In R.~nunculacene
16 CrossosocataCeae 88 01 11" nh.l .::.. 191 ~osales 78 Cuttiferales Zl IH llenlalea Tn Dillenlaceae?

Ordl' r 21 : TII EAI. f.S

f7' Ochnaee ae 90 The ate s 1, 5 Tl,., a l es 82 Gu ttifer ales 1 4S0chnalu 42 Ge r anlales 1
a !!!. Och naee.le 0;1 !,.oph; r <lce:ul !!!. Oehnaeeac !!!.Ochnaceae !!!. Oehnaceae !!!.
Dipterocarpaceae
(Theal .. ,,)
b In Oc h""ce"e 9 3 S ~ra~b ur&erl ace3e 49 Th~alcs 84 Guttifcra les 144 Ochna hs
(The a1eR)
[c '.'allac~ac~ac: \n Odln~c ...~e J
78 Rhopalocarpacea~ 11,3 :i..~lvales 94 :'1alval"5 194 Violale . 147 Ochnal cs
(or: Sphacro sepala~ea.· )
79 Sa rcola~na ecae 16 2 :i..~lvale& 48 Th""les 172 Xalvahs 146 Ochnale& 31 Gut t Herales
(or Chl acnaceac , RhodoIB(maeeae)
80 Dipl~ rocarp~ ceae 9, Thealeb SO The •• l es as Guttif erli l es 148 Ochnale s )0 Guttlfera l cs
a1 Carvocar a eea e 101 T~eale 5 41 Theales a7 Cuttiferales 142 Thea l u in Theaceae
a2 Thucea" 97 The al" " Ja Theales 86 Cuniferales 13S Thealu i9 Gu t tlferales
[3 Siadeniae .."e (A S+) : .!.I). Theace"e]
[b Ternst ro c",laccac (AS+ ) : !E!. Th~lIc el\c J

,
[e C ~t.'.e lliaceae. (,.,S+): .!!!. Thea c eae ]
.!!!. Theac""e 99 Penta!,;\wllleeae 44 Thealcs 156 Ce la stralcs 139 Thea le s .!!!. Theatea ..
(Theales)
, .!!!. Theaceae 100 TetrallOCr16 t"eeae .!.!l Theaeea .. .!!!. Thea e""" 140 Thealcs .!.!!. Oehnae""'''
(Th~al c6)
.!!!. Thcaceac 102 As teroE "laceac .!.!!. The"ceae i!l Theaeeae .!!!. Theaee.,e. A 1£ rlac ourt i acca"R
(Thea1cs )
g .!.!!. The 3eeae. 10 3 Pell1cicr.leeae In Theaee.ae !.!!. Theaeeae 138 Thea} e" .!.!! Theaeeae
(The ales)
lJl i!l .!.!!.
" .!!!. Theaeeae

83 Stlchyu ra eelc
107 Bonnet iaecae
(Theal" s)
11 ) Vio lales
The aee lle

37 Theale"
Thea ecac

186 Violale s
134 Thea1es

S3 lI a!ll3mel1dales !!!.
Thc aeenc

Th""ce",e

~
N
~
 ,

~
N
N

CRONQUIST TAKHTAJAN THOR);E ENGLER HUTCH INSON BENTHA.'l & HOOKER

84 Act in idL1ccac 14\ Ericales 36 Thea l es 81 Guttifcrales 137 Thea l"" in Theae.,ae
a in Actlni di aceae 140 Sa urauiacca e in Actinid iace ae ~ Actlnldiaceae lJ6 The ales in The ace""
(F.rical"s)
8 5 Mares,-a 98 Thenles 40 Thea les 88 Guttiferales 141 Theal es in Th ea eeae
86 Quiinac 104 Thc ales 46 Theales 89 Guttiferales 161 Guttifcrales
87 El atina ___ _ in Gut tiferae
110 Thea l cs? 53 Thea l es
88 Hedus"gvnace ae
198 Violales 271 CarYQphyllales 26 Guttife r alcs
105 Tl1 e31"s? 206 Rosa l es 80 Cuttiferales
•
89 Gutliferae
106 On cothecaccac?n:., ,< U3 Theales
in Ebenacca,,?
108 The"les 52 Theales 90 Guttiferales 159 GutlHeralcs 28 Cuctifernles
(or: Clus;aceae)
a in Guttifentc 109 Hvper!ca"e"" ~ Guttlferae in CuttHera e 158 Gu ttifera les 27 Guttiferales
(Theales)

""n Malvale s
160 Malvalcs
164 Malvales
47 Thea l es
96 !1alvales
93 !1alvales
177 Malvales
173 ~:alvales
176 ~alva le s
T iliales
116 Tillales
117 Tiliale.
01acaceae8
Ma l .... ales
Malvalcs
91, Bombacaceae

."
165 t'.alval es 97 Malva l es 175 ~!alvalcs 119 Tilla!es In l'.a l vaceae
95 I-:alvace"e 166 :1alvaJes 98 Mahales 174 Y",lvales 1 20 !'~~l v ales J2 Hah'ales
167 Hua ccac i In st yraeaceae? 1 26 Haipighiales
(? Ma l"ales)

Order 23 : UCYTH I !JAU:S

96 Le"ythlda"eae 220 ~yrtaies --9: Thca l es 211 llyrtlfloTa e 163 Myrtal es in flyrta~eae
[ a Astera"thaceac (AS+) : In l.ecythldaceae; 163a Myrtales? (Hutc h.)A]
{h Bar dnst onl"~.ceae (A5+) :ln Lec yt hidacea~ : 162a Ilyrtales ? (Hutch.)A]
k Foetidiaceae (A5+): in Le,,-ythidaceae]
( d Napole ona ccae (AS + ) : In Lecythidaceae]

Ord er 24: SARRACE:>IAI.F.S

97 Sar ra ""ni aceae 45 Sarrac enia les 33 Sa rra,,-eni,il cs 92 Sarra ceniales 308 Sarrace ni ales 9 Parietales
98 Nepenthace<,e 212 Nepentha le s 55 Kepen tha l es 93 Sarracen lales 263 Ar istolochia l es 138 ~ultiovul. terr .
99 Droserace .. e 211 Nepenthales 198 Rosales 9~ Sar ra~ e n iales 307 Sarraceniales 63 Rosales
Orde r 25 : VIOLALES

100 Flacourtiaceae I I I Vi o lales 70 Clsta!e. 183 Viol ales

·[Y7 Pad etalesJ
76 II lx.l l es 73 Passlflora! es k
[ a Ho"",- i L'ceac , b S"",ydaceae , c Neuman " laccae (AS+ ): In Fl acourtiace.,e ]
d in Flacourtiaceae 112 Laclstcmatace a e In Flaco-;;-':t1accae i ll na,,-ourtiaceae 80 Bixales 1 163 " AnOl:la l ous r.1rn . "
- (Violales) -
101 Peridiscaceae 114 Viol ales n
Cis ta l e. 18~ Violales 118 Til1al e s
n
In Bixin eae
102 Sc v phos t e gi aceae 119 Viol a les 73 Cis ta les 187 Vlolales 187 Ceh"tralcs
10 3 Violaceae 115 Vi o lal es U. Cista ie s 185 Vioh l es 99 Violales 15 Parietales
10~ Tu rner;.ceae 121 Passjflo ral es 77 Cis tale s 188 Vio la les 104 Loasale. 7S Passiflorales
N

'"
106 ~ alesher bia eeae
107 Sixaceae
122 Pa~sif1orales
116 Viol ales
78 Cistale"
7,} Cis tale"
189 Vio l ale s
193 Viobles
",74 Sixa l es
l!! Six"ce"e 1 1 7 Co chlosEermace"e in ~ixace a eii 195 Viob l es 77 Bixales !!! Bix i ne30e
" (Viol a les)
108 Ci st accae 118 Vio l a l es 76 Cistales 1 92 Vio l a l es 75 Bixales 14 Pa r ietales
109 Ta~ ri caceae 136 Taloa r lcale" 87 Tanaricalcs 196 Vio l a l es 97 Tamariea!es 25 Caryophyllinae
110 Fran ke ni30 c eae 138 Tarnaricales 88 Tamarica les 197 Viola l e.Q 96 Tama r ieales 22 C3oryop h yl l inae
111 Dloneophdl3or.e3oe 95 Thea l e s (in flacourtiaceae) 83 Guttifcralcs in l'lacourtiaceae
96 Die ~ ode n draeeac 14 530 Ochnales A
" (?Theales)
112 An c ist r oclad a ceac 94 Theales 131, Gc r a n ia l es 91 Gu tt ife r ales 149 Ochna 1e .. i n Dipterocarpaeeae
113 fouguie r ia c eae 1)7 Ta mar!eales 114 Sol a naies 2'} 4 Tubif l orae 98 Tamarica!es i " Tama riea c eae
l!op1esti gmatac . O
114
lIS Achari"ee"e
338 Polemonia1e"
123 Passifln r "lcs
263
80
Lamia!e ..
Cistales '" Ebenales
19 1 Vi olales
78
108
Bixales
Pa$sifloralcs ~ P3ossiflor3ceae
116 Caricaee;)e 1 24 Passiflora1cs 81 Cista l es 199 Vlol<ll~ s 112 Cueurbitales in P "ssiflor .~ce.~e

117 Loasaee"" 339 Polewoniales 85 Cista le s 200 Violales 105 I.oasales 74 P""sifloralcs
118 Begon taeeac 127 Begoniales 83 Cis t ales 202 Vi ola!cs 110 CucurhH"les 78 PassifloTalcs
119 Datisc"ee"e 126 Begonialcs 84 Cis tales 201 \'101a1"s III Cucu r hlt"les 79 Passiflora!cs
la Tctramel"ce.~e (AS+) : iIl Da t isca c cac)
120 Cucurbit"ee"e 12,} Cueurbitales 82 Cistales 203 Cucurbitales 109 Cucurhitales 77 Passiflorales

Or der 26 : SAL I GALES

1 21 S"licace"e 139 Saliealcs 86 Salie"les 7 Sa lie ales S7 Salicalcs 162 "I\nomalous fa m. "
I" Populaeeae: in Salicace.,e ]

Order 27: CI\PPARALES

122 Tovadaceae 131 Capparalcs ;n Capparaecae 98 Pnp"veTales 95 Capparidalcs in Capp"raceac
123 CnppaTaceae 128 C"ppar"les P 89 Cappaddales 96 PapavcTales 93 C.~pp"ridales 12 P"rieta1e s
(or : CJPpa r id.~ceac)

,
a in Gappar.,ceae
in C3PP'lraeeac
i~
!!!
CappaTaeeac
Capparaee30e
in
in
Capp~raeeae
C.~pp"racc"c
l!!
!!!
Capparace"e
Cappar30cea e
Oxvstvlidaceaeq
Cleomact' ac A
.!.!l
~
Capparaceae
Cappar"ee"e
(Cruclales)
c ~ Capparaceae 129 Koeberlini.~ce~e .!..!!. Capparaceae in C,1PpaTaeeae 172 Celastrales in Slmaroubaceac
(C"pp"r"lcs)
Id Canoti"ecae (I\$-t) : i!!. Koeberlini"cc<lc (C<lpp~Taccae) J
e l!! Capparaceae 130 Pcntadipland Tac. in Capparaee.'e .!..!l Capparace"e 179 Gelastr"l~s
(c.1pp.1r" l es)
(~ Cappar"cea~) l J5 fmbli"!\laceae in Cappa r aceae in C<~pparaeeac i!:!. Fl"courtl.,ecae A in CapP il r.1ceae
(Capraral"s)?
124 Cnlciferae 133 Capp.Hales n
1 25
(O T: II r assica c eae)
Resednee~c 134 Capparales
Cappadd"les

91 Cappar!dates
" P,'paverales

99 P.'paver"les
269 Crucial~s

270 Rcsed"les
" Parietales

13 P.1riet" l es
126 )1orincaeeae 132 Cappara l es 90 C"PP.Hidales 100 P;, p averales 94 C"pparidales 5 7 "Ano,,", lous fam . "
624 28

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 ~
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IlUTCHIN.'ON BE-NTlL\M & HOOKER
i!!
a !!l Byb lt daccae 192 Rori du laceac 214 Pittosporales J 13 Ro sales ,,,
89 Pittosporales Droserae"""

(S " xlf ,- tl saies)

350 Scrophu iar iaies in Sa xif ra ga ceac 274 Tuhifiorac 329 re r son.~les 120 Personalcs
\1.8 Columelliaccue
149 Hydrangeacc,," 190 Sa xif r agaJe s in Sa><ifragaceae in Saxifragace.~cE 33 Cunonlales i!! Saxif r agace~e

.~ in Hydrangeae""e in Hydrangeaccac !!l Saxifragaceac in Sax i fragac<'n"E 32 Ph iladelEhac cac i n Sad f raga,,-eae
(Cunontaies)
150 Grossu l a,-i"ee"" 189 Saxlf r aga l cs in Saxlf rasaceac in saxifragaccae E 34 CUllolliaies in S"xifragacea e
a 1E Grossu l ariac . 182 Escalloniace ae v 1E Saxifragaccae in Saxifra gaceae E 37 CunonL. l es i!! Saxifragaceae
, in Grossulariac. 18~
(Sa xl fr aga lcs)
T~tracar E acaceae
(Sa~ifI"aga 1 es)
in Sax if ragaceae in Saxifragace~eE in Es callonlaceae in Sa ~i fragaceae

, I teacea e in S.,~ Jf ragac"-""- .!..'! Sa~ Huga~eaeE .!..'! Esc<~llolliaccac in Sa~i[I""g"ceae

in Grossula~ia c .

in Gro.sulariac .
'" (Saxifragales)
186 Brexiaccac in Saxifragac"-ae in Sa xi.fra g a~eaeE ill Escalloniaceae in Sa~ifTagaceae

,"
(Sa"ifragales)
~ Grossula ri ac. Ig7 Phvll onoma ceae i n Saxifragaceae .it.!. Smd [ r agaceae E ~ Escal10niacea c in SaxifragDee.,e
(OT' Dul ongiac. )

, in Grossul"ria c .
(?Saxifra gales)
188 Pterostemonace"e
(?Saxif ragales)
in Sa x Hr"g aceae in Saxifragaceac[ 30 Cllnonia l cs .!!l Rosace ., e

g ill Gros$llia r inc . m )Iontin ia ceae i n Saxifragaceae in saxHraga ccacF. i n Esc a llonlaceae .it.!. onagraceae B
(Sa ~ifr ~g~ l es)
195 Saxlf ra ga les 215 Pittos pora l es 11~ Rosales 56 I\,,,,,a,,.cll da les 65 Rosales
lSI llruniaccac
152 A\scuosmiaceac ~ Caprj[ollaceac' (i n Caprifollaceae) in Cap r ifoliaceae; in Caprifoliaceae A .it.!.
Caprlfollaceae
153 Crassulaceae 197 Sa~ifragales 1 9~ Rosales 105 Rosales 298 Saxif r agales 6 2 Rosales
198 Sa ~if Tagales 195 Rosales 106 Rosale s 299 Saxlfragalc8 in Saxif ragaceae B
15~ CeEhnlotaccac
Ro~a le s 300 Sax ifragales 61 Rosales
155 Saxlfragaccac
a !-". SaxifTagDceae
199 SaxJ( r "gales
196 Penthora~eae
(Saxifragales)
196 Rosa les
in Sa~if Tag aceae in '" Saxifragaceae E In Crassu l ac(!<,e A .!!l Crassulaceae

b .it.!. Sax\fragaceae 200 Vahli~ceae in Saxlfragaceae in Saxifragaccac E 302 Saxihagal"-s in S.,xlfragaceac

(Sadfragalc s)
c in Sa x ifragaceae 201 Fr., ncoaceae in Sax\ [r.,ga ceae i n Saxifragaceac E 303 Sa~i[Tagnles In Saxlfragaccae
(Sa~ifragales)
d ~ 202 Eremos"naceae
Saxif ra gaceac in Saxifragaccae in Saxifragaccae E JOI Sa xlfrasales .it.!. Saxifragacca~

(Saxifrag.1ies)
e in S"xif~agacea" 203 Pa rnass iacene in Saxlfragaccac in Sa~ifragaceacE 305 Saxif r agales in Sa~ifragaccae

(SaxHrag ales)
f (~ Saxifragac.);U 179 ParacrvEhiaceac
Sa~ ifragales)
, "
(ill Saxl fra gaceae) 183 Trlbelaccae
(Saxlfr ag ales)
(~ Saxifragaceae) ill Saxifragaccac in Escal lon laceae" in PittospoTaceac F

Lc~uToEe t alaceae in Saxi frag ac cae in Saxihagace"c in Sax i fragaceae in Saxifragaceae

~ Saxifragaceae)
" I .!.!!. P" rn a ss iac.

~
N
~
 ~
N
~

CRO~QUIS T TAKHTAJA): TlIOR,-:E El{G1.ER HUTCH INSW BENT1C",'! -b- -H6ok-I(R-

156 Rosacea e 201, Rosa l es 190 Rosal <> s -115 Rosales 24 Rosales 60 Rosa l"s
F~ ~alaceae }
~ Po"'acea" in Rosaceae
!b Prunace a e . )
Anwpdalac<,ae: ~ Rosaceae
l Drupaccae
157 ~e ur a d~ ccac 206 Rosa l es in Rosaceae 11 6 Rosales in Rosace"e ~ Rosacea e
158 Ch rvs obal ,~n "c<,a(' 205 Rosale s'" in Rosaceae 117 Rosa l es i n Rosace"e 11\ Rosaceae
J
159 Leg"minos"e
207 Hi ::-.osac(>ae
(Faba l es)
20~ Cac~alpin;acea"
(fabalcsl
193 Roso l es J 19 Rosa l es
r"
i 7.7
)!i<:I0.,,, cc a e
(Legumin,~ l cs)
Caesalplnlaccac
( L eguminal"-~)
59 Rosales

209 Fub acc""

(Faba l es) l" P"pil ionaceae
(Lesumina ies)

Order 33 : PODOSTEft~l.f.S
160 Podost"",aceae 213 Podast e0 a le s 200 R06,1 1cs 122 Podostcwa l es 309 Pod o st,,"~l~s 137 ~:\lltiovulatac
la Tri~ti c h acea<: ('\5+1 : in P adoSl~maccae j aq u,1t lcac

OrJer 31, : llALO RAGA LES

161 )lalo r,lgaccae 227 Hippurld"lc~ 252 Carnalcs 217 :ly nHla rac 29 1 Lythr al es 66 Rasalcs
(ar : llalo rrhagirla<:cac)
l a ~"riaph"l l aeeae : i..'l Hala l-agaeeae)
I f. 2 (;m:neraccac 22& Hil'pur lda les in nalorasaccaeil in Ha l aragaceae in flaloragaceae i..'l
Halaragace.le
163 Hj pJ!uridaccac'" 229 Hl p p\l ~;da lcs 253 Corn,,- l es i19 ~IHt1flora" in Iialaragace,~c in l!ala r,l g~ceae
I" ~ i" d il\o naceae D R6 Thclig(Yna l es Chcnopodiaiesi i 2 18 ~l~-r t I f larae 285 Chc nopodial c6 i.n Unic ac e ae
(or : Cynacr"",nace.1e)

Order ) 5 : ~lYRTALfS
165 Son~e r"ti"ceac
166 b -th ra ceac
215 ~lyrta l es in Lythraceae 209 ' Iy r tlflarae 165- Myrta!cs in' Ly th r acca c
214 ~lyrtaJes 221 Myna le s 204 :Iy r tiflarae 288 Lythral es 71 ~lyrtale6
167 P~":le.~c<:ac 22 4 ~l yr taJes 227 ~Iyrta l ('s 179 Thymdaea l es 84 ThYI:".e lae:ll es 148 Da~hnales
168 Crvp teron l"c,"a(' 180 S"xHrag a l e~ 123 flyrtales 206 ~lyrt:!f la r ae 39 r.unoniales Y in Ly t hraceae
I~ <) Th ,-rr.ciacac('a<e 175 Th yrr.elaea lcs 109 Eup hcrbia l cs 181 Thyr:te l aeales 85 ThymelaeaJes 147 IJap hn ales
" in Th,c:tel acaceae in Thyme 1 " Cacc,'c .f!!c ThYt:1elae,,-ceac i..'l Thymc laeac eae 82 Agui l ariacene in Thyt:1e lae a ceae
(Thyme la ca l es)
b in Th)'T.'. elacace,,~ '.n Thyme laeace ne I n Thyr.c la cacca e .!E. Thy,"cl"caceac 8 1 Gan""tvlaccac in Thy::tc l a cnccaeB
(Thymelaea le s)
170 T r aEac('ae 226 )Iynal es 224 ~Iyr tales 205 , ly r Hflorae 29 0 Ly t hralcs aa in OnagT accac
(or : HyJ r ocn rya ccac)
171 lJia l vee talanth" c . 330 Gcntianales 208 :Ivrtifl a rne 116 Rubiales
172 ~l" r tacpae 221 Myrtales 228 ~lyrt[]l cs 207 ~t~'nif l orae 162 ~ lyrtalcs 69 f!vrt alcs
n in )Iyrtaceae in Nyrtaceae (in ~l y rt aceae) ~ ~lyrtaceae 199 Hctc roE,"x idaceac !:lL;·thraceae B
(Rhamn"lcs)
[b P$ll o xdac c"e ("S+) : in or nea r 'Iyr t"ccacj
N
~
 225 ~y~ ~ <ll"~ 230 Myrul .." 215 ").".",,, .. ~
In L~lh~acea,,&
~> .." - ' - -
223 Myrtales 226 ~lyrtale 5 216 ~yrtHlonf! 35 Cunonlal .. "
222 Myrtales 229 ~ynales 112 ~\· rt1fl or." 168 Myrt31f!" 70 ~iy rt" l e 5
, In Mf!i.stoo:.H,,,:eae1
68 '{"nale s
ii"9 Mynales 225 Myrt.:lles 211. ~I)·ntflorae 167 Hyrta,""

Or dor 36 : PROTF.,\LES -(~9Daphnales

315 f.laeagnaies 103 Rhar.llla\"" 182 ihYIT.el3e<ll"s 200 ~hnl:mal,."
17B Ehe8gnaee .• ~
316 Proteal"s 220 Proleal"" 15 Protu l "" 87 I'roteal~s 146 Daphnales
"9 PrOl",\e~<I"

Order 37 : COR~o\1. ES
217 Xyrt31e s 2~6 Cornale s 213 )Iyn lfi orlle 161, tlyrtal,.s -6-' ~yrtale s
180 Rhl ~o phnrn eeaf
~ (l.!!. Rhlzophor .• e . ) LI S Ani,oehvlleaeeac (l.!!. Rhizophor~c ea~j ..!..!l Rhlzophor"eeae ..!..!l RhllOphorace<l" l.!!. R h izophora e"a~

(Myrtale~)
[b Po l "8cn.'nlha c~3": In " nl"ophy l l e a ce.'" (Rhlzophorace .• ej (T"kl1 . ) [
181 Davldinccne 278 Ccrn"lu 1" ;;,·s 5acc.,,, 22) U".boI 11rl orac l!l Ny$~ace,'c

182 :-;'· ~~3C~3e 279 Cornales 248 C~ r nal e" :122 Umbe ll1fl orao 1. 6 MlIllalcs in Cornae"ac
280 COTndes 250 Co rnal<" 22 11"t'lhe l lifloT"" I.~ Ar(l \i ~lcs in Co rna<:ea"
183 AhnBI~(en~
184 COT1\acue b6 276 Cornal ~ .cc 2~9 Corn a les 22.1, t!r.:bel l jflor~u 4) Ar.,l1a1"s 84 ij"b"lla l e~
a l.!!. Corn .• cenf! 281 ~~~slixiaccae in CO Tnaceae i!!. Corn~e".,~ 1'1Cornacc"" in Co rn .,ceae
TCornnle~)
b (.!..!!. Co r nateao) 282 Ilchdn gtaceae in Corn3ce3e l.'!. CornaCCIIC 1'1 ATalla~e ,, ~ in Ar"llaccac
(Cornaln)
c 1n COrrlac,",lIe 283 Tnrlcell13c"8c in Cornaccac ..!..!l Cornaceac l!!. CornaC~ 3 e in Co r n a eeae
(Co rnlll es)
18S Garr"ac~n~ 277 Coroales 2S1 Comales 225 Umbcll!flora" ~5 Arallalc5 l!!. Corn"ce~e

Or der )8: So\l;TA~Lf.S

186 v.edusandracc,""D 310 Snn td ales 122 San tala ie5 24 ~edusandrnlc .. 19) Olac ... l"s
181 Dlp~nlodontaee~cO )09 Santalale5 71 Cis tal"" 17 Santn\al"s 192 01;'1ealc5
302 Santaldes 12) Santalales 16 S""t;'llalu 18!' Oheales ~6 Olacales
188 Olac <l ccae
~ Ola~"ee"e !'l Olacace,,~ in Olaeaceae
•.!..!!. 013caceae )0) ~pf lace"e
(San t nlales)
in Olacac~a~

b .!.!!. Ohellee&<'. )05 Octokner_~ceae (l!!. Olllc" e<'3") .I.!!. Olacacuf! 190 Olacales
(santalales)
c .!.!!. Olacaeeae )06 f.rnhropaiaceae (1" Olacace<l~ ) !n OlnCncelle 185 CeI3"nale~ l.!!. Olacac e ae
(S.~nt3lal"s)
.!.!!. (in 01.:tcaeeac) in Ol<leaccllc I'll flp u"d~ hI Olaeaceae
d .!.!!. Ohclleeae Oillcace<le
(Olaelllcs)
lO~ Sanca\31es 124 Santalales 18 S<lnt<l\ales 189 ()lacales in Olacaceac
189 Opllilleeae
190 C rub bI8Ce(\~ J~ 9 r:Tie.~\~~ 217 Pill O"pora l es 1<) S.'nl~la\es 195 Sn olt.1b1 c£ in Sa!llal aceae
s;;;tiiT:i;:;,;;; )08 ~ant .,h1~s 125 Santa\a I"5 20 S.'n\al·, I ~·, 196 S,,!ltalalcs lSI ;'ch l~"ydos~onal!
'" .,
192 I.or(lnthnc~n<'
ITI LOrnnll". celle
312 Sa ntnla l cs
!'l l.ornn tl ' .~ce :tc
127
128
S",'ta l ales
Viscaceae
22 S;\T\t alnl es
l!:. Lorll1\th"c~""
19~ S.' Tlt"la l es
l!!. l..or"nth"cc.~c
150 ,\chla")"da~por,,.l"
in LCr <lll thllcea e
(Santal.11 e~ )
 ~
N
co

194 BaianoEhoraceae 314 Santal"l"" 129 S,mt.~l.,l"" 23 Balanophoralcs 198 San talal es 152 Achl"mydos~oreae
195 C\,,,o",orl,,ceae 313 Santalnles 130 Santa l" l"" 220 MyTtiflorae ~ Baianophoraceae in SalanophoT3ceac

Order 39 : RAFFLES IALES

196 ~itrastemon aceae in RaffJeHlaccac in Rafflesiaccae in Rafflesiaceae In Raffles!""e""A
197 H\'dnoraceae 2~ Raf fles!al", 69 Rafflesialc" 75 Aristolochlales 261 Arlstolochlales in Rafflesiaccae
198 Rnfflcsiaceac l) Rafflesiale" 68 RafflesL11cs 74 Arlstolochiales 262 Aristolochiales tJ9 ~lult iovu l . teTT .
(or : Cytinaceae)

40 : CELAST RALE S
olamat.'Ce"" 29~ Celastrales
200 Hippocraleaceae 291 C"l"s tra le s in (c l as traccac 162 Cclastrales 184 Cel astra\cs ~ (clastracca"
201 Ce lastrac cae 290 Cel astrales dd 118 Santalales 160 Cc[ast r ales 180 Ccla stralc" 49 Cclastrales
[a Chl nRlth a mn "ccae. in Celastraceac (Cronq.)]
[b Lophopvx idaccae (AS-+-) : in Cel.,straccac ? ( Takh .) 1
, in Cclastraccac 295 Goupiaeeae in Ceias tra ceac in Celasaaccae IS3 Ce l astra l es ~ Cc b straccac
(CclastTales)
202 Siphonodontaceae 292 Celastra l es .!n Celas tr ace"e in CeLaslTaccae IS~ Celastrales in Cclast"'ccac
(or : Ca~ u ,ia c eae)
203 Staek housiaccae 293 C.. lastrales 119Santalales 163 Cclastrales 182 Celastrale" SO Cclastrales
20 4 Salvador"ce"e 289 Cc\astralc" Ol ealc" 1M Cel"strale"
205 A9uifoliaceae 286 Ce lascra les '"
)9 Thea l es 157 Cclastralcs~
171 Cclastra l cs
170 Cclastrales
*incl . Sphenos tmnon"cc"c?
107 GenU ana ! c"
47 Olacalcs

" lEe Aquifoliaceae 287 Phc l li ncaceae (itl Aquifoliaceae) ~ Aqulfoliaccae ~ Aqulfol i"o eae ~ Rutac cae?
(Cclast:ralcs)
Ieacinacea e 288 CeiaHrales 120 Santa l ales
'"
[a Irvingbaileva~cac: in I cacinaeeae ( Ta~h . ) ]
207 Card 1op t eridace ,1 e 307 S.,nt"la l es 12 1 Santalabs
166 Celastr"les

167 Ce la s~ ralcs
176 Celastrales

171. Celas tral es

in Ol ac3<:eac

in Ola eaec ac
208 Dichapet"laeeae 172 Euphorbiales 108 Eu?horbiales 180 Thymel"eales 25 Rosales 45 Geran1.)les
(or: Cha il l eUaceacl

Order 4 1 : EUPHORBIALES
209 Buxaccac 168 Euphorbi" lcs 110 [uphoTbiaics 165 Celastrales S~ Hama",clidales in Euphorbiaecae
a in Buxace ac D 169 Sl"'""ondsiaceae (in Bux"oeaej? ~ Bu xaceae i n Buxaccae in Euphor blac e"e
(Euphorbiale.)
[ b Styloeera ta oea c (AS-+-): in Buxaceae]
210 Eupho,biaceac 171 Euphcrb l"lcs 104 EUp!o oTbial cs 130 Ge ranial es 133 Eup horb ia lcs 153 Un isexua les
[a ,\ndrostachvdaceae (AS-+-) : In Eu phoTb iac eae i
[b Blse ho fiaccae (AS-+- ): in EuphoT biaceae ]
[ e H...."enocaTdiaeeac (AS -+-,)",,: in Eupho r b i a ceae ]
[ d Penceae (,\ 5-+-): in Eup horbiaceae]
[e Slilaglnaceae (AS-+-): i n Euphorbiace a e (near Ic ~d na,,-eae ? (Airy-Shaw)]
[f Uapacacc a e (AS+): i n Eup horbia"-e",, (ne ., r An,'c a Tdiaeeae?)]
[8 Sce p.1c cae (AS-+-) : in F.uphorb iaceae]

N
CO
 CI«):lQUIST TAJalTAJA~ THORll:E ESGLER lIiiTClffN-SOS 8E/I.'TIIA.'t ~ Hom::ER
211 o..phnlphviia .. c"",, 170 f.upho r b1ales -ilif Plnosporalrs III Ceran!..t"" S5 tln"",,,,,, ll dales !!!. Eupho r bIaceae
212 AeKtOxl .. a .. .,ac 297 Celast ralc8 ? 106 EUl'ho rblales 152 Sa p lndal ... 178 Ce ta sl rales In Euphorhla"<lae
211 !'''nd4''~~'' 17l Eupl\orbla.les 105 Euphorbiales 159 Celastu.tes 169 CelAst rale"

\'itaceae 170 RII<l.U13les In Vl tae" ",,-

Comales 169 Rhn .. n31,, ~ 52 Celastra~e5

Orde r /oJ: SAP1)-;OALES

217 Suph'·\Car.Uol 2L2 Saptndales 207 Rosales 161 Cel" ~l ralea 221 Snpindalcs In 5a plndac ., ,, e
216 lIeltanthac/!me 24e S"plnd"les 156 Rutales 151 Snplndnles 214 Saptnda l e9 In Saplndace., e
219 Grevl"cea~ 249 Saplnda l .,. 199 Rosales In ~ .. I1""t l".ce,,,, 36 Cun oni"l@ " In Sapindace a e
220 Conn"r"c~a" 210 CO\ln~r~lca 192 Ro .• al ,,_. ITs Ros3 1 e~ 200111"0\,, 1..- 58 Rnsa l es
221 Stdnha~lac<'A~on 235 Ruuln? (in Rosaceae) !.!!. Chry~obalnnacuef:!!l Ro~aceaeA In Ro""c",,"
3 ~ StylobasiacQne ~ Sl~a roubaceac? 149 Surian nc<"ne In Slll'l.~roubneeae in SI m.uoubnceae In S l::tarouh ~ceac
(Ruta l es)
222 "'b. n iac~3e 245 S"plnda l e~ 157 Rutales U8 Rutalu 222 Sapl ndal c8 ~ Saplndace3e
221 S.,plndaccu 2U Snplnd~les 154 Rutales 148 S ap!nd.~IC8 215 Saplnd al u 53 Saplndales
[a Dodonaeacea": ~ Saplnd ac e""j
b ~ Saplnd ~ cuc 247 8rat~chnelde ra c. 160 ~utal.,s g7 Saplndalea ~ S"p lndac e3~
(Saplndalu)
22~ Hlppoca5tnnaceae 246 Saplndalu 159 Rutales 149 Saplndaiea 220 Sap lndal u in Sapindacea ..
225 Acetacea .. 243 Saplnddu 158 Rutales U6 Saplndalu 219 Saplndnlu b S3plndaceae
226 Burscraeu., 233 !!.ulate. 152 Rut",les 136 Rulaies 2tl Ruules t;] Geranlales
227 An a ca rdlac"." 230 RUlates 153 Rut al es 145 S~p lnd"l u 218 Saplndales 55 Saplndales
a .!!!. Ana .. "rdl",ce",., 232 Podo~cea" (~ Anacardiac""c) ~ Anacardlacea,,? 216 Soplndah~ In Sapindaceac
(Rut&!.,s>
I' (AS+): In Anacardlac.,ae(?) [
1< In ",,;tardlaceae]
no iJI Rutdes In Anaeardlacca" 154 Jullanlales 223 Saplndaiu in Anaeardiace",,?
234 Rutal ... 148 RU~31"s Il~ Runic,. 2 10 Runlet; 41 Ger,,,,lales
'"• 240 Klrkiace .... .!.!l S;OI3 roubacC3., ~ SI .... roub3c"ae[ !!!. SI"",roubac.,ae'"
(?Ru t &!eal
b In SII'a r oubac"ae 260 Balanl'a"eae !.!!. Sl.... roubacea" In 2ygopl\ylhceae 131 ~.alplfl,htal"" In Sl .... rnubar.e .."
(Ceranlales)
c .!!!. SI"'aroubae.,ae in tKonanthac.,ae~ !.!!. Stn:arnubaee3e ~ SI ..... roub.'e., ~eE .i!!.
230 Cnenr" .. ""e 237 Rutaln no Rutales 1)) Runl",.
'"
173 C el,,~ual es
IrvlnshccBc
()lalp Ighlaies)
SI=rnubaceae?

!!!. SI"",roubacca..
231 Ruucea., 236 R"tales 147 Rutales 132 lIu tftlu 209 Ru tale s 40 Geranl3les
[3 Rhabdod"ndr.ceae (AS+): In Rutaeea" (?) (Takh.) ; near Centrosper"",,,? (Alr)·-Sh.w) 1
232 M;i'1" .. e.u 238 Rutal "" 151 Rutalcs lJ7 Rulale. 2\3 :-lililllleR 44 Geran1ale.
• (!.!!. l~eliaceae) in )!dia .. ~,,~C .i!!. Sa plndar.eae ee !!!.
Sa pl ndaceae

I' (AS +): Rutnc.cacl

I, (AS+): Sapindaceae; i n Simaroubac"ae (IIutch.)A j
 •

~
w
o

;T TAKHTAJAN THORNE ENGLER HUTCHINSON BENTHAM f. HOOKER

~upu,uaccac 2S9 Geraniales 136 Gerantales 127 Gerani31es 132 )1alpighlales 38 Cc ranLl.les
Zygophyllaceac 258 .~ inariaceae _~ Zygopbyllace.,e .!.!!. l ygophyllac"3" .!!!. Zygophyllaceae 1£ Zygophyllacea e
(Gcran{al~s)
b !!!. Zy gophyllace3" 261 Peganac"a" in ZygophyUaccac in ZygoplJyUacea c in Zygophyllaccac in Rut aceae
(Geraniaies)

m p1dob.O.t.r .r (.!.!!. Oxal idaceael is! Ox"l1d " ce3e no

b in Oxa lJda ceae in Oxa l idaceae (.!!l Ox a I idac""e) in Ox~lidaceae 212 Avc rrhoaceae in Cer.:lni.:lceae
(Rutalcs)
c (i.!'.. Oxa lidaceael 263 Hv p s"och,H1tacea~ (in Ox,,1 idacea,,) in Oxalidaceae in Oxalidace"e ~ Ceraniaceae
(Geranialcs) -
23 5 Gc raniacea e 264 G"raniales 138 Gcranial cs 12S Geraniales 330 C"rnniales 39 Geran iales
a.!!!. Gaaniaceae 265 Dirach~ace~e (!.!!. Ceraniacea,,) i!'. Ccranlace"e lH Til ialcs
(Geraniales)
b lD. (;(,ra niaceae 266 Vivianiace.1e (iE, f,c.ra n laceae) i!'. Ce raniaceae 91 Pitto~por"les i n Ceraniaceae
(Geranialc s )
in Cer an;',,,eae 267 Bicbersteln1ace~c (in Gcrani.:lceae) in G~ra nl aceac in Geraniaceac A J~ Ceraniaceae
(Gc nm bl~~)
d .!.!!. C<.> ran l accae .!.n Geraniacca~t (in G~rania c ~ae ) ill Ccranlaceae 127 Ledocar paceae In Gcnwiaccae
('1alpighialcs)
2)~ L;",.nant i1ac,,<.e !711 Gc rani ales 11,1 Ge ran ia l es 12 3 r. .. ranial~, 331 Gcrar.ia le s in Gcrani~ceae

217 Tro2 ,,,,olac<,,,c 2f.8 Ge ran ia l "" 140 (,pr"ninlcs 12f. Gerania l cs 333 G~ra"iales in Geraniaceac
)3 ~ Ceraniales in Geran iaeeae
2 38 Ba 1 ,.",j nacca,' 269 Geranl a l es 139 Ger3nialc, lS3 Sapindales

Ord er ~5 : LI~ALE S
239 I lu"'i r iacca~ 254 G" Tania1e s in 1.in ,,~ eae in l. inac~a,,"F." 123 '1a 1 pighia les 36 Gcraniales
2~ O[ p·throxvl accae 255 G~ ranial es 115 Gerania1cs 129 Gc raniatcs 128 ~!alplghjal es tn Llnaceae
241 Un"cca<' 252 Cc~"niales I JJ f,~r.~ntale" 128 Geraniale s 124 :'lalp i g hla le~ 35 Gentnln1e s
a i!'. LIna cea e 2;'1 Ilup"niacca.> 1~ Linac c ac ~ Lina ccac i n Linaceae in Llnaceae
" .... 'dnia l ".)
l> in Linac eae 253 lx"nanth~ ce a'· l.!:!. Linacca c ~ Linaceae 12 1 '!alpighialcs in Unaceae
(f,erania l "s)
c 1..n_ Un3cea~

Order 46 : POI.Yf,.\L,\ LE S
in :lalp ighiac~ae in Lina cc. ac in Linaccac
'"
242 ' l alpig h iac~M 257 (:" r~n iDl~s 1',2 ~cr"ntales 139 Rutales l22 Hal.p lghi alcs 37 Ge ra niRlcs
!4J T risoniaceae 27 1 PoJygdLll~~ 145 Ger,mial cs 140 Rutales 102 PO!\'ga bles in '!o<:ilvsiaceae
2M. ~·si"c~a.· 272 Polygala 1e, 146 GCHmL,les l ~l k\lta1 ,,~ 103 l'olygal~I"s 21 Po lyga1in~ c
245 T r~mandra c~ a c 175 l'olygala l ", 213 Pit ~ os porales 1 ~2 ~ Ll ta 1 "s 92 Pittos?orales 19 Polygallnae
2~6 ~anthopilv1l:;!-e a c in Po lv galac e ar (.!.ro Pd yr,alace a c ) in Polyga l ~" e" ~ in Po1ysabcea e'\ in Polvgalacea c
2~ 7 PO lvgd 1.::!.~ 273 Polygala l e~ 143 (;c~"nial"" 143 Ruta l ~s 100 Polyr,a la ]cs 201' o1 v salinac
[a Di c lldant hHa cea~ : in Po 1y g<ll"ced~1
248 ;:raT1><'ri3CC,,,' 27 4 Polyga l a 1es 140 Ge ranl~ l cs 120 Rosa l es 10 1 Pol\'~31alc~ l!! l' olyga l ac"a e
N
~
 CRONQ-U-'(ST T-IlKIlTIIJIlN THORNE ENCLER HUTCHlNSO~ BE-"'THIIM & HOOKER

'der 47 : UMBELLALf:~hVRALIIILES2 83 Umbel131 ,

Cornaics 254 Comalcs 226 Umbell1florae 47 Arallales
Comalo. in ATal13ceac 227 Umbclliflorae 311 Umbellalcs 82 Umbellales

~ Umbcllifcrael

Subclass VI : ASTERIDAE

Or der 48 : CENTIANALES 110 Gcn tian;fes

322 Centianales 231 Gentianales 246 Gentiana l es 226 Logan l a l es
251 Loganiaec<,e ~ Loganiaccae
a in Loganiaccae 323 Anloniaecac gg (;n Loganiaecae) ~ Logan laccac 228 Logania!e.
(Gentianales)
(in Loganlaceae) ~ Loganiaceac 225 Logania l cs in Loganiaccae
b ~ Loganiaecae 324 ~,llaccac
(Gonthnales)
~ l. ogan i.,ceae 247 Gentiaoales ~ Potal iacea e ~ Loganjaceae
c ;n Logan;a cc ae 321 n csfo"tai n iaceae
(~ntianales)
d in Loganiaccae .i.!'c Lo>;.,o;aceae (~ Loganiaceae) .i.!!. Loganiaccac 229 SE;geliaceae ~ l.oganiace ae
(Logan;alo.)
(.i.!'c Loganiaceae) in Loganiaccae 2)1) St nc hnaceac ~ Loganiaceae
c in Loganiaceae in Loganiaeeae
(Logan ial"s)
234 Centia<lales 248 Gentianales 293 Gentianalcs III Ge n tianales
252 Geotianaceae 328 Gentian ales
326 Gentian ales 233 Gentianales 250 Gentianales 233 Apocynal cs 108 Gentianales
253 Apocynaccae i!'. Loganiaeeae
a in Apocynaeeae 325 Ploc?~per"alaCeac (i!l LOBania ceae) in 1.0ganiaceae C 232 Apoeynal~"
(Gentlanalcs)
327 Genti an des In Apocynaceae 251 Gentianales 235 ,\pocynales 109 Gentian •• le.
25 4 Asc lepiadaceae
~ Ascle~iadaceae
a 10 Asclepiadaccae l.!! '\sclepiadac~ae .i.!!. Apocynac.ea e ~ Asc l epiadac c,e 234 PerlE l ocaceae
(Apo~ynales)

,lanales ~ Con voh'ula

111 So l ana l cs 263 Tubif lo ra" 321 SolanaJes 116 Polenonia l es

256 Solanaceae 340 Sc roph ula r i.d es
~ Solanaceae i!! Solanac eae ~ So lanac eae Salpiglossidaceae ~ Solanaceae
a.i.!'c Solanaceae
(Solanalcs)hl
i!! ;';olanaceae?t .i.!!. Solanaceae 264 Duckeodendraceae i!'. Ehretiaceae
0
(Tublflorae)
[e Goet?caceae (AS+) : In Solan aceae (Takh . ) I I1 S Polemoniales
333 Polemoniales 112 Solan ales 255 Tubiflorae 322 Solanales
257 Convolvulaceae
[a Dlchond.aceae, b Hu:nbeniaeeae (AS+): i!l Convo l vulaceae )
[c CardiopteTygaceae (,'.5-+): near Con volvulaceae (A i ry-Shaw) I in Convolvu13cea e
3)4 Polcrnonlales in Convolvu l aceae in Convolvulaceac 337 PolemonL,les
258 Cuscutaceae In Genl1anaceae
329 Gentia n ales 235 Gentl .. nales 249 Gentianales 791,Gen tianalcs
259 Menvan thaccae 112 Polcmon iales
332 Po leJ:lOoia l es 11 3 Solanales 253 Tubiflorae 335 Palemontales
260 Pole~ont3ceae
in Polemon iaccae
3 in Pole ~ o n iace.,e in Polemonlaceae (in Polcmoniaceae) in Polemoniaceae 238 Cohoeaceae
(JHgnoniales)
335 Polemon ialcs 260 I.a", i ales 256 Tubiflo~a" J36 Po l Hlonlalcs 113 Pol~mo n ialcs
26 1 H~droEhvl1aceae
[337 Polemonialcs 262 Lal:liales 258 Tubiflorac 156 Eric •• les 99 Edcal es
262 Lennoaceae
632 28

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 CI/OSgUIST TAKliTAJ AS THORNE ,;N(;i:-ER HUTCI II NSON I\ENTH A.'I ~ HOO KER

o rd c T S3: C,\)IP ASU LALES C:;:"':;;;;-;;;C;C;;--_"CCC;;C;C;;;;C-;;-;;;;-~"'"C;:;::;;;OCC;:,-;:;---"C7"';CO;:";C;"-;C---"'7~.~;;C:C"':;;;:;;;0--

281 pc~hragmat"c. I J i';campan n lnceae 115 Carc.panu i a l es 286 C"rc.pan u lal es 1,1, C~"p<,,,ulaceac .!.'2 C, mpanu l aceae
282 Sphenoc l e 3ceae 3M C~mpanulalc' in Canpa n nlaceae lBS CaT:lpnn ulai cs i n Cal:1pa n UL1ceae i n Campnn n i aceae
281 Car> pnnu l acc"e 362 C" "'panu b les 116 CampHlHdalcs 284 C~ ",p ,,,'ublcs 315 CIl:1 pan a l es 93 C"mp"n a lcs
a J. n C~"? 3n u lac~ae 163 Lobe l l a ceac i a C~mpanu l ac~ae i .n Cal:lpan ul aceae 316 Ca,,~an ., l es in Camp"nulaceae
(Camp" nu la i es) - - -
197 Rosales 289 C""'pan u l,des 319 Coodenin l "~ 9 i Ca!:lpn n a Les
Ca C1 p" n nia l es
284
a
~ l i d i 3 cene
In Styll d iaceae '"
M Ikmal L ace~c
(Ca mp a nul a l es)
in ~tv li ~i"r."a" in Sty i idiace " e ) 0 1. S a" if r ~g"les i n Sa xi f T" g.1ceae

i n Goodeniac~a.· 288 Ca,.,. panulnle~ 3 18 Goodenia l cs irl Gcod en i~c"""

Ca m p"n u l al('~
285
286
B r unoni"L~ae
Coode n iaceae ""
367 C" m p a nu l ~le~ 117 C"I:lpanul!J l es 287 Campan ul .-. I es ~1 7 Gocd<'nia l "s 92 Cat:l p a n a l es

'.1rdn S4 , Rl'Hj,\L CS
287 Rub;ac~~e 3n r;"ull.-.nal,,~ 2)2 G~nt i ann l e. 252 C;e" t iana l ~s 237 Ru bial~s 86 Rubia l es
A
., ill RlIhi.>ceae In Kuhiaceae ' i'l Ruhiaceae 269 ~e n riq"cda~e<l~ in Rnbiac<'ac i n Rubiaceae
(Tubiflom,,)
[ h ~;;",cl"aceae (,\S + ) : in i(Hbi"ceae]

Ordp r SJ : l)lPS ACA LES

288 Ca p rif o li~~"ae 3\7 Dipsaca~l e s 255 IHp~ac:"les i80· lJ ip-;;;~cales 48 Aril l iale~ 85 R\Jb i alcs
1.1 Sambllc"cea~ (,\S+ ) : in C,,?ri f oliace3e , (?) Taklo.]
Ib (a r lem3nni"ce"" (,; 5+) : in Cap r ifol i aceac: (?) Ta k h . ]
281 Dipsacale~ 306 S 'J.x i r r ag~les i n Capdfo l iaceae
289 ,\do~.~ceac 31~ Di~s"calcs 2S6 Dips"c" l es
282 llipsac" l es 312 Va l"da n"l"s 87 Aste r a l c5
290 Valeri<lnacca" 319 !lipsaca l cs 2S7 D L rsac~ l es
283 Dipsac" l es Jl3 V" l e r Lana l ~,; 88 Aste r a l es
2~1 Dipsn c aceac 320 Di p ,;aca l cs 2)8 Di?sa c a l es
1 .~ :o1o ri naceae (AS'>-) : in f)jpsacaceac)
Ib Tr i plc'tq;iaceae (AS+) : ill lJipsact,,:ea,,]
290 Carr.pan lll a i es 314 V.~ l er i 3na l "s 89 Astera l es
292 Cal v ceraceae 309 Ca l yceralc' ~S9 Dipsaca l es

Or d<- r 5 6: ,\STE R,\LES

2~3 ~os i tae 370 Aster . lies 267 ,bteralcs 29 1 Ca".panula \e s no A~ t eralcs 90 Asteralcs
(o r: As tC r ~ce.,e ,
Car<\uac cac)
[ a A::thro~iac"De (AS +) : ~ Coc:pos i t a<' 1
Ib Ci cbo ri ace"e : ~ Co~pcsi t ae ]

CLASS B: l.lLlATAE o (~o nocoty l eJonae)

Subc lass \' 1 1 : AL I S~jATI DA~

Ord e r 57: ALlS~WrA1.ES

111 Alis mn l es 268 ,\ 1 isn:a l es 293 lIelobiae 343 llutomales ll'. :'1 i smaCeaC
29' flu t o rr,<)ceae in ll u tom~ceae .!.'2 Butom..' ~ e"e l'.!. A l is""~,,-eae
295 Li mnochari t acea" 372 Alisrr,alcs i n A lis m .~tacc a c
269 Al i smai cs 292 Hdobiae ]I.S ,\Ii,;""ual c s 196 Apocarpoc
2~6 Ali~;;"1tacc"c 373 Alisna l e~
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p"doaeeae
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~
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~
 ,

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o

INDEX TO FA)HL Y NAMES, continued

Trapaceae 170 Tropaeolaceae 237 Velloziaceae 344 Winteranaceae 9

Trapellaceae 2788 Turneraceae 104 Verbenaceae 265
Tremandraceae 245 Typhaceae 323 Violaceae 103 Xanthophyllaceae 246
Tribelaccae 155g Viscaceae 192a Xanthorrhoeaceae 343
Trichopodaceae 350h Uapacaceae 2l0f Vitaceae 216 Xyridaceae 309
Trigoniaceae 243 Ulmaceae 46 Vitidaceae 216
Trilliaceae 340k Umbelliferae 250 Vivianiaceae 235b Zanichelliaceae 304
Trimeniaceae 14 Urticaceae 50 Vochysiaceae 244 Zingiberaceae J29
Triplostegiaceae 29lh Zosteraccae 305
Tristichaceae 160a Vacciniaccae 1298 \"allaceaceae 77, Zygophyllaceae 2JJ
Triuridaceae 307 Vahliaceae lSSb Wellstediaceae 263c
Trochodendraceae 38 Valerianaceae 290 Winteraceae 4

AUTHOR INDEX TO CLASSIFICATION SYSTEMS

Adanson 585 , 592 Caesalpino 585, 588 Gerard 585, 588

Albertus ~Iagnus 585 Novak 586
Camerarius 585 , 589 Gundersen 586, 601 Pliny 585, 587
Baillon 586, 596 Cord us 585, 587 Hallier 586 , 600
Bankes 585, 588 Pulle 586 , 601
Cronquist 586, 606-608 Hayata 586, 601 Ray 585, 589
Banks 586, 610 Dan~in 585 Hofmeister 596
Bauhin , G. 585, 588 Rendle 586 , 600
de Candolle , A . 585 , 593-594 Hutchinson 586 , 601 Rivinus 585 , 588
Bauhin, J . 585, 5HS de Jussieu , A . 585 , 592-593 Jung 585, 588
Benson 586, 604 Skottsberg 586, 601
de Jussieu, B. 585 , 592 Kimura 586 506 586
Bentham & Hooker 585, Deyl 586 Lamarck 585
594-596 Takhtajan 586 , 604
Dioscorides 585, 587 l'Ecluse 585 , 587 Theophrastus 585, 586
Bessey 586 , 598-600 Dodoens 585 , 587 Lindley 585, 594
Bierhorst 586, 610 Thorne 586. 609
Eichler 586 , 596 Linnaeus 585, 589-592 Tippo 586, 609
Bock 585, 587 Emberger 586 1 ' Obel 585 , 587
Braun 585, 594 Tourne f ort 585, 589
Endlicher 585, 594 Magnol 585, 589 Turner 585 , 588
Brongniart 585 , 594 Engler & Diels 596 Hattioli 585, 587
Bro\olll 585, 594 van Tieghem 586, 598
Engler & Gilg 596 Helchior 586 , 596 1,.,Tarming 586, 597
Brunfels 585 , 587 Engler & Prantl 586 , 596 Mez 586, 601
Burtt-Davy 586, 601 Wernham 586 , 598
Fuchs 585 , 587 />Iorison 585 , 588 Wettstein 586 , 597

~
00
 CLASS I FICATION LITERATURE

1. General References

Airy Shaw, H. K. 1966. \.Jillis' A Dictionary o f the Flmlering Plants and

Ferns . 7th ed ., revised . Cambridge University Press . Cambridge.
Arber, A. 1938. Herbals, Their Origin and Evolution . Cambridge Univer-
sity Press. Cambridge .
Benson, L . 1957 . Plant Classification . D. C. Heath and Company. Boston.
Core, E . L . 1955 . Plant Taxonomy . Prentice-Hall, Inc. Englewood Cliffs ,
New Jersey.
Cronquist, A. J. 1968. The Evolution and Cl assif i cation of Flm.;ering
Plants . Houghton Mifflin. Boston .
Dalla Torre, C. G. de, and H. Ilarms. 1 900-1907. Genera Siphonogamarum
ad Systema Englerianum Conscripta.
Davis, P. H., and V. He)'\.JOod . 1963 . Principles of Angiosperm Taxonomy.
D. Van Nostrand Co ., I nc . Princeton , New Jersey .
Gundersen, A. 1950 . Families of Dicotyledons . Chronica Botanica.
l.Ja1tham, Hass .
Hutchinson, J . 1959. The Families o f Floweri ng Plants . 2 vols. Oxford .
Lawrence, G. H. N. 1951. Taxonomy of Vascular Plants. Nacmi1 l an . New
York .
1955 . An Introduction to Plant Taxonomy. Macmillan. New York .
Porter, C. 1. 1959. Taxonomy of Flol>lering Plants. W. H. Freeman and
Company. San Francisco.
Rendle, A. B. 1967 (reprinting of 1930 book) . The Classification of
Flm.;ering Plants . 2nd ed . Cambridge.
S,.;ingle , D. B. 1934. Textbook of Systematic Botany . NcGraw- Hill. Net~
York .
Takhtajan, A. 1969. Flowering Plants--Origin and Dispersal . (English
translation by C. Jeffrey) . Oliver and Boyd . Ed i nburgh.

II. Ancients

Albertus Nagnus . ca . 1250. De Ve getab i lis.

Dioscorides . 64 AD. Materia Nedica.
"Gart der Gesundheit ." 1485. Hainz .} many editions and printings
Hortus Sanitatis . 1491 . Mainz .
Pliny. 77 AD. Natural History .
Theophrastus . ca . 300 BC. Enquiry into Plants and the Causes o f Plants
(Historia Plantarum) .

III. Herbalists

Bankes, R. 1525. Herba11.

Bock, J . 1539. Neu Kreuter Euch .
Brunfels, O. 1530-1 536 . Herbarum vivae eicones.
Cordus, V. 1561. Historia Stirpium .
Dodoens , R. 1583. Stirpium Historiae Pemptades; Cruydeboeck.
Fuchs. L . 1542 . De Historia Stirpium; Neu Kreuter Buch.
Gerard, J. 1597 . Herba1l or General Historie of Plants .
l'Ecluse, C. de. 1601 . Rariorum p1antarum historia.
IIObel, N. de. 1581. Stirpium adversaria nova.
t-lattioli , P. 1544. Commentarii in sex libros Pedacii Dioscoridis (plus
many editions) .
Turner, \.J. 1551, 1562, 1568 . A New Herball.
 642 28

IV . Hechanical 5ys terns

Bauhin , G. 1623 . Pinax th eat ri botanici .

Bauhin , J . 1650 . Histor i a plantarurn un i versali s (posthUlllOUS) .
Caesalpino , A. 1583 . De plantis libri .
Camerarius , R. 1694 . De Sexu P1antarum Episto l a .
Linnaeus , C. 1735 . Systema Naturae .
1737 . Genera P1antarum (and subsequent edit ions) .
1751 . Ph i losophia Botanica .
1753 . Spec ies Plantarum.
Hagnol , P . 1689 . Pr odromus historiae genera l is .
Morison , R. 1680 . Plantarum Histori a Universalis .
Ray , J. 1686-1704 . lIistoria p l antarurn .
1703 . l'lethodus plantarum .
Rivinus, A. Q. 1690 . Introductio generalis in rem herbariam .
Tournefort , J . P . de . 1694 . Elements de Botanique .
1716. Institutiones Rei Herbariae .

V. Natural Systems

Adanson , M. 1763 . FamilIes des Plantes.

Bentham , G., and J . D. Hooker . 186 2-1883 . Genera P1 anta r urn . 3 vols .
London .
Braun, A. 1851. Betrachtungen liber die Erscheinung der Verjiingung in
der Natur . Leip zig .
1853 . Das Individuum der Pflanze in seinem Verhaltniss zur
Species . Berlin .
Brongniart , A. 1843 . Enumeration des Genres des Plantes, &c. Paris .
• Brown, R. 1866-1867. (The misce l laneous botanical works of Robert
Brotm . Edited by J . J . Bennett. Published for the Ray Society by
R . llard..Ticke . London . ).
Candolle , A. P . de . 1813 . Theorie Elementaire de la Botanique . Paris .
Candol1e, A. P . , A. , and C. de . l82ll- 18 73 . Prodromus systematis
natura1is regni vegetabilis. 17 vo1s. Paris .
Endlicher , S . L . 1836-1840 . Genera Plantarum . Vi enna .
Hofmeister , H. 1851 . Vergleichende Untersuchungen ... F . Hofmeister .
Leipzig .
Jussieu, A. L . de . 1789 . Genera plantarum secundum ordines natural e s
deposita.
Lamarck, J. 1778 . Flore Francoise ; and other Harks .
Lindley, J. 1830 . Introduction • to the Natural System of Botany . London.
1846 . Vegetable Kingdom .

VI . Phylogenetic Period

Arber, E. A. N., and J . Parkin . 1907 . On the origin of Angiosperms .

Journal of the Linnean Society (Botany) 38: 28-80 .
Bai11on , H. 1867-1895 . Histoire des Pl antes. 13 vo1s . Paris .
1871-1888 . Natural History of Plants . 8 vo1s . London .
Benson , L . 1957 . Plant Clas si fic ation . D. C. Heath and Company .
Boston .
Bessey , C. E . 1897 . Phylogeny and taxonomy o f the Angiosperms . Botan-
ical Gazette 24 ; 145-178 .
1915 . The phylogenetic taxonomy of flm~ering plants . Annals
of the Hi ssour i Botanical Garden 2 ; 109-16 4.
28 643

Boivin , 8. 1956 . Les Famil I es de Tracbeophytes . Bulletin de la Societe

Botaniq ue de Fr ance 103: 490-505 .
Burtt-Davy . J . 1937 . On the primary g r o ups of dicoty l edons . Annals of
Botany , New Se r ies 1 : 429-437 .
Clements. F. E .• and II . N. Hall. 1923 . The Phylogenetic Hethod in Taxonomy.
Carnegie Institute of \~ashington PuhI. No. 326 .
Copeland, IJ . F . 1957 . Forecast of a system of the dicotyledons . Hadrono 14:
1-9 .
Cronquist , A. 1957 . Outline of a new sys tem of fa milies and orders of di-
cotyledons . Bulletin du Jardin Botanique de 1 ' eta t, Brussels 27 : 13-40.
1965. The status of the general system of c lassification of flm.;er-
ing plants. Annal!:; of the Hissouri Botanical Garden 52 : 281-303 .
H68 . The Evolution and Classification of Flowering Plants .
Boughton lIifflin . Boston .
Darwin , C. 1859 . Origin of Species .
Deyl, 11 . 1955 . The evolution of p l a n ts and the t a xonomy of the Honocotyledons .
Acta Nuse i Nat. Pragae, V. 11 , 3 , no . 6 : 1-143 .
Eichler , A. B. 1876 . Syllabus der Vo rles ungen Uber Phane rogamenkunde . Kiel .;
1 890 . Sy llabus, 5 Aufl. Berlin.
187S~1878 . B1Uthendiagramme. 2 vo 1 s . Leipzig .
F.mbe·rger, L . 1960 . Les Vegetaux Vascu1aires ( Vol. 2 of N . Chadefa ud and L.
Emberge r, Trai te de Bo tan ique Systematique) . Haisson et Cie . Paris .
Engler , A., and K . Prantl. 1887-19 i5 . Die na turl ichen P flanzenfamilien . 23
va1s . Leipzig .
Syllabus der Pflanzen f amilien e d . 2 : 1898; ed . 3 : 1903; ed . 4: 1904 ;
ed . 5 : 1 907 ; ed. 6 : 1909; ed . 7 : 1912 ; ed . 8 : 1919) .
a nd E. Gilg . 1924. Sy llabu s der Pf1anz enfamili e n. (eds . 9, 10) .
Berlin .
_-;;-""",".' and L. Diels . 1936. Syllabus der Pflanzcn(amilien . (ed . 11) .
Bcrlin .
Gundersen , A. 1950. Families of Dicot yl edons . Chro n i ca Botanica . Waltham,
Mass .
Hallier , H. 1905 . Provisional scbeme of the natura l (phylogenetic) sys tem of
flowering plants . The Nel. Phytologist 4: 151-16 2 .
1912. t ' orgine et Ie systeme phyletique des Angiosperms . Archives
neer1andai ses des sciences exact es et natul'elles , ser . 2 , I : 146~234 .
Hayata , B. 1921. The natural c lassif ication of plants according to the
dynamic system . leones Plantarum Forrnosanarum 10: 97~234 .
Hutchinson , J. 19 26 , 1934. The Families of Flowerin g Plants . 2 vo 1s . Hac~
millan . London.
1959. ibid . cd . 2 . 2 vols . Cla r en don Press . Oxford .
1969 . Evolu t ion and Phylogeny of Flowering Plants . Academic Press .
London and Ne w York .
Kimura, Y. 1956 . Sys teme et Phylogenie des Honacotyledones . No tula e Syste-
maticae 15: l37~159 . Paris .
Kun tze , O. 1891 . Rev isio Generum P1 antarum. Ii. Stu rtz . \~ iirzburg .
}lel chior , H. (ed .). 1964. A. Engler ' s Sy llabus der Pflanzenfamilien . (ed .
12) . Vol . II . (Vol . I, 1954. Bacteria to Gymnosperms, E . \~erde rmann, ed . )
Gehriider Borntraeger. Berlin .
}!ez , C. 1926 . Die Bedeutung de r Serodiagnostik nil' die Stammesgeschictliche
Forschung . Botanisches Archiv 16 : 1~23 .
Novak, F . A. 1954 . System Angiosperm . Preslia 26 : JJ 7~J64 .
Pool , R. J . 194 1 . Flm.ers 9.nd Flm~ering Plants . NcCra l.~lIill . New York .
Pulle , A. A. 1938. Compendium van de Terminologie, Nomenc1atuur en Systematiek
der Zaadplanten .
 644 28

Pulle, A. A. 1950. ibid ., ed . 2. Utrecht.

Rend l e , A. B. 1930, 1967 . The Classification of Flowering Plants.
ed . 2 . Cambridge .
Schaffner, J. H. 1932. Revised catalog of Ohio Vascular Plants. Ohio
State University. Columbus.
Skottsberg, C. 1940. Vi:ixternas Liv, vol. 5 . Stockholm .
S06 . R . 1953. Die modernen Grundsatze der Phylogen ie i m neunen System
der Blutenpflanzen . Acta biologica academiae scientiarum Hungaricae
4: 257-306 .
Takhtajan, A. 1943. Sootnosheniia ontogeneza i philogeneza u vysshikh
ras tenii. (English summary) V. H. Molotov Universi ty Press . Erevan.
1953. Phylogenet ic principles of the system of higher plants.
Botanical Review 19: 1-45.
1959. Die Evo lution der Angiospermen . Jena .
1966 . Systema et Phylogenia Nagno1iophytorum . Soviet Sciences
Press. Hoscm.,r and l.eningrad.
1969 . FloHering Plants--Origin and Di spersal. (English trans -
lation by C. Jeffrey) Oliver and Boyd. Edinburgh.
Thorne, R. 1963. Sane problems and guiding princip l es of angiosperm
phylogeny. American Natu ralist 97(896): 287-305 .
1968. Synopsis of a putatively phylogenetic classification of
the flm.,rering plants. Aliso 6: 57-66.
van Tieghem, Ph. 1898 . Elements de Botanique. ed. 3.
Harming, E. 1895. A Handbook of Systematic Botany (English edition by
H. C . Potte r ). Hacmillan . New York.
h'ernham, T . J. 1911. Floral evolution. New Phytologist 10 : 109- 120.
Hettstein, R. von . 1911 . Handbuch der Systematischen Botanik. GebrUder
Borntraeger . Berlin .
1935. ibid. ed . 4. Leipzig and Vienna .

VII . Recent Systems for Vascular Plants

Banks, H. P . 1968 . The early history of land plants. Symposium on

Evolution and Environment. E. T. Drake (ed.). Yale University
Press . NeH Haven, Conn.
Bie rhorst, D. \~. 1971. Horphology of Vascular Plants . Hacmillan.
New Yo rk.
Lam , H. J . 1948 . A new system of the Cormophyta. Blumea 6 : 282-289.
Tippo, O. 1942. A modern classification of the plant kingdom. Chronica
Botanica 7: 203-206 . Haltham, Mass .
29 645

Chapte r 29 . FAMILIES OF VASCULAR PLANTS

Traditionally t he family has been the basic study unit in systematics

teaching a nd tra ining . Expedience in identifica tion comes ~"i th the recogni-
tion of family characteristics . Comprehension of present classification
systems is dependent upon a f undamental understanding of the fami l y diag-
nostic features. Three approaches a r e used in this chapter for understanding
the family conceptuall y and identifiably : (1) diagnostic characters are g iven
in summar y form as aids in field identi fication , (2) synoptic keys to sub-
classes . orders and families are presented as conceptual aids for the funda-
mental understanding of families, and (2) plates illustrating features of the
structurally more difficult famil i es have been designed for visual help. For
convenient reference the fam ilies, orders , and s ubclasses have been indexe d.
Detailed family descriptions will have to be procured from other sources (see
referen ces at the end of the chapter) . See Fami l y Plate Illustrations, pages
852 - 877 .
Section A. AIDS FOR FIELD IDENTIFICATION OF
eOi'U-ION FAMILIES OF FLOWERING PLANTS
IN NORTH AMERICA NORTH OF MEXICO
(Arranged according to the Engler System)

Honocotyledoneae

1. Typhaceae . Flowers densely crO\"ded in a terminal spike ; perianth of

brist l es .

2. Zosteraceae . Aquatics; flowers in spikes; no distinct perianth.

3. Najadaceae. Aquatics ; flower solitary in axi l s of leaves; no perianth .

4. Alismataceae . Perianth present ; parts in 3 ' s; carpels form disk or ring

on r eceptacle.

5. Poaceae . Culm hollO\", terete ; leaves flat , 2-ranked ; sheath usual ly

open ; f ruit a grain .

6. Cyper aceae . Culm sol id, triangular; leaves channeled , 3-ranked ; fruit
a nutlet .

7. Araceae . Inflorescence a spadix; perianth wanting or inconspicuous .

8. Xyridaceae . Plant scapose; f l owers in dense globose to cylindrica l

heads .

9. Commel i naceae . Herbs ; leaves with a tubul ar sheath; f l oral parts in 3 ' s;
calyx green ; corolla colored; ovary superior; capsules several- seeded .
(3; 3; 2-6; 2- 3) *

*Floral formula for flower ; with the first number referr i ng to the U of sepals ;
second, to the /I of petal s ; third, to the If of stamens; and f ourth, to the If of
carpe ls. Formula is given only when it is considered of some va lue in famil y
identification .
."
646 29

10. Juncaceae . Grass-like; floral parts i n 3'g; fru it a capsule.

(3; 3; 3-6 ; 3)

11. Li l iaceae. Perianth conspicuous; floral parts in 3'$ - stamens 6; ovary

superior . (3; 3; 6; 3)

12. Amaryllidaceae. Periant h conspicuous; plants scapose; Eloral parts in

3'g - stamens 6; ovary inferior . (3; 3; 6; 3)

13. Ir idaceae . Perianth conspicuous; flora l parts in 3 ' 5 ; ovary inferior .

(3; 3; 3; 3)

14. Orchidaceae. Perianth conspicuous; floral parts in 3' s - stamens 1 or 2;

gynandrium and pol1inia presen t; corolla zygomorphic; ovary inferior.
(3; 3; 1- 2; 3)

Dicotyledoneae

Apetal ae (without petals)

15 . Sal icaceae . Tree s or shrubs; dioecious; EloHers in catkins; bracts of

female catk in deciduous.

16. Juglandaceae. Trees; monoecious; fruit with a thick husk; leaves pinnately
compound .

17. Betulaceae . Trees or shrubs ; monoecious; female flowers usually in ca tkins;

2-carpellate.

18. Fagaceae . Tree s or shrubs; monoecious ; female flm~ers usually not in

catkins; 3-carpellate.

19. Urticaceae. Herbs; ovary superior ; stinging hairs usually present.

20. Santalaceae. Semiparasitic herbs, shrubs, or trees; leaves alternate or

opposite; ovary in ferior; fruit a drupe .

21. Aristolochiaceae . Herbs or vines; calyx petaloid; fruit a 6-locular cap-

sule.

22. Polygonaceae. Herbs o r vines; nutlet trigonous; stipules sheathing.

23. Chenopodiaceae. Herbs; perianth and bracts not scarious; filaments sepa-
rate; fruit a utricle.

24. Arnaranthaceae. Herbs; perianth and bracts scarious; f ilaments usually

connate b e low; fruit a utricle .

~popetalae (with separate petal s)

25. Caryophyllaceae . Herbs; leaves opposite; inflorescence cymose; placenta-

tion f r ee-central . (5; 5 ; 5- 10; 1-5)

26. Nymphaeaceae. Aquatic; leaves long-petioled, peltate; floral parts many

except sepals, 3; vessels absent; placentation laminar .
29 647

27. Ranunculaceae . }lastly herbs; floral parts spirally arranged ; stamens

numerous; pistil unica rp e l late ; endospe rm present . (5; 5; "" ; co)

28. Magnoliaceae. Trees; floral parts numerous - spirally arranged; gynoe-

ciuffi apocarpous.

29 . Papaveraceae. Herbs; sepals 2, caducous or fugacious ; juice milky or

colored; perianth conspicuous; gynoecium syncarpolls .

30. Brassicaceae. Herbs; floral parts in 4'5; fruit a silique or silicle;

stamens tetradynamous. (4; 4; 6; 2-4)

31. Sarraceniaceae . Insectivorous perennials; l eaves hollow, basal; flot-rers

5-merous; style umbraculate, ovary superior; fruit a capsule. (5; 5; "'; 5)

32. Saxifragaceae. Herbs, shrubs, or vines; f loral parts in 5's except car-
pels mostly 2; endosperm present. (5; 5; 5-10; 2)

33. Hamamelidaceae. Monoecious trees or shrubs; leaves alternate, petiolate;

perianth present or absent; stamens numerous; carpe l s 2, ovary partly
inferior; fruit a capsule . (0-7; 0-4; 8- 2) 00
;

34. Rosaceae. Herbs, shrubs, or trees; floral parts in 5 's; ovary superior
to inferior; leaves stipulate; endosperm absent. (5; 5; 5- 00 ; 1- 00 )

35. Fabaceae. Herbs. shrubs, trees, vines; floral parts in 5 's - carpels 1;
corollas mostly papilionaceous; stamens monadelphous or diadelphous; fruit
a legume . (5; 5; 5-10; 1)

36. Geraniaceae. Herbs; floral parts in S ' s; carpels coi l ed at maturity;

leaves lobed or dissected. (5; 5 ; 10; 5)

37. Polygalaceae. Herbs; flowers showy . irregular; petals 3 ; stamens 6-8.

(3; 3; 6-8; 2)

38. Euphorbiaceae . Herbs, shrubs, small trees; apetalous; monoecious 3-car-

pellate; inflorescence a cyathium in Euphorbia .

39. Anacardiaceae . Shrubs or small trees; leaves pinnately compound; stigmas

3; fruit a drupe.

40. Aquifoliaceae. Shrubs or trees; floHers dioecious or polygamo-dioecious.

small; inflorescence axillary.

41. Aceraceae . Trees; flm.,rers dioecious or polygamous. small; leaves pal-

mately lobed; fruit a samara .

42. Rhamnaceae. Shrubs or small trees; stamens alternate with sepals ; leaves
pinnately veined; fruit a drupe.

43. Vitaceae . Woody vines; leaves palmately veined; fruit a berry .

44. }~lvaceae. Perianth conspicuous; stamens monadelphous; sepals persistent;

anthers I-locular; carpels 5 or more; pubescence frequently stellate .
(5; 5; 00 ; 2- 00)
 648 29

45. Hypericaceae. Herbs or shrubs; leaves opposite; stamens usually in fasci-

cles; carpels 3 or 5; petals mostly yellmol; conspicuous secretory canals
or vescicles in all organs.

46. Violaceae. Herbs; corolla spurred; placentation parietal; fruit a capsule.

(5; 5: 5; 3)

47 . Cactaceae. Fleshy, prickly plants; f loral pa rts many; ovary inferior.

48 . Elaeagnaceae. Shrubs; l e aves a l ternate, simple, usually lepidote below;

flowers perfect, 4-merous; c arpel 1, ovary superior; f ruit drupaceous
with a fleshy hypanthium. (4; 4; 4; 1)

49 . Lythraceae. Herbs or shrubs; l eaves opposite or whorled; ovary enclosed

in a persistent calyx.

50. Nelastomataceae. Herbs; petals conspicuous; floral parts in 4'5; anthers

poricidal: ovary infe rior. (4; 4: 8; 4)

51. Onagraceae. Herbs; f loral parts in 4's; ovary inferior; anthers longi-
tudinal ly dehiscent . (4; 4; 8; 4)

52. Araliaceae. Herbs or shrubs; flowers in umbe l s; leaves compound; ovary

inferior; f ruit a several seeded berry-like drupe; carpophore usually
absent.

53 . Apiaceae . Herbs; f lowers in umbe l s; stylopodium present; f ruit a schizo-

carp; ovary infer i or; carpopho r e usually present. (5; 5; 5; 2)

54. Cornaceae . Shrubs or small trees; leaves usually opposite; flowers 4-

merous; ovary i nferior; f ruit a drupe. (4; 4: 4; 2)

Sympetalae (with fused petals)

55 . Ericaceae. Mostly shrubs or trees; floral parts in 5's; petals usually

conspicuous. (5: 5; 5- 10; 3-5)

56. Primulaceae . Herbs; p l acentation free-central.

57. Styracaceae . Leaves simple; hairs stella te.

58. Oleaceae. Trees or shrubs; leaves opposite, frequently pinnately com-

pound; fruit a samara or drup e ; stamens 2. (4: 4; 2; 2)

59. Gentianaceae . Herbs; leaves opposite, exstipulate; ovary I- locular;

flOi.Jers showy; pl acentation parie tal.

60. Apocynaceae. Herbs or shrubs; l e ave s opposite: mi lky l atex p resent;

fruit a follic l e; po llen not in poll inia. (5; 5; 5; 2)

61. Asclepiadaceae. He r bs or v ines; crown. gynosteg i um, pollinia present:

milky latex present; f ruit a f ollic l e. (5; 5 ; 5; 2)

62 . Convolvulaceae. Twining vine s; corolla plicate: fl owers axi llar y.

(5; 5; 5; 3)
29 649

63. Polemon i aceae . Herbs ; co r o lla r egular ; ovary 3- 1ocul ar ; carpels 3 ;

corol la lobes convolute in bud . (5 ; 5 ; 5 ; 3)

64 . Hydrophyllaceae . Herbs; corolla regular; ovary one-locular ; carpels 2 ;

co r olla lobes usually imb r icate in bud. (5 ; 5; 5 ; 2)

65 . Verbenaceae. Herbs , shrubs, or small trees ; leaves opposite ; ovary 2- or

4-1ocular separating into 2 or 4 nutlets; style terminal. (5: 5; 4-5; 2)

66 . Boraginaceae . Herbs; ova r y 2-1ocular developing into a 4-nutlet fruit;

leaves a lt ernate . (5 ; 5 ; 5 ; 2)

67 . Lamiaceae . Herbs or shr ubs; corolla zygomorphic ; style gynobasic ; leaves

opposi t e , aromatic . (5 ; 5; 2- 4 ; 2)

68 . Solanaceae . Herbs or shrubs; anthers frequently po r icidal; ovary 2-

locu l ar or falsely 3-5 locular . (5; 5; 5; 2-5)

69 . Scrophul ariaceae ." Herbs ; cOTolla zygomorph i c; stamens frequently 2;

fruit a capsul e . (4 - 5 ; 4- 5; 2-5 ; 2)

70 . Plantaginaceae . Herbs; flowers inconspicuous ; scapose; perianth scarious;

f r uit a cir cumscissile capsule . (4 ; 4 ; 4 ; 2)

71. Rubiaceae. Herbs o r shrubs ; floral parts mostly in 4 ' s , carpe l s 2 ; ovary
infe r ior; leaves opposite or whorl ed and stipulate. (4 ; 4; 4 ; 2)

72 . Caprifoliaceae . Nostly shrubs; floral parts in 5 ' s , carpels 2 or 3;

ovary inferior; l e aves exst i pulate. (5 ; 5 ; 5; 2- 3)

73. Cucurbitaceae . Tendril-bearing succulent vines; monoecious ; ovary 3-

locular; fruit a berry .

74, Campanul aceae . Her bs ; corolla usually zygomorphic; anthers occasionally

united into a tube ; ovary infe r io r. (5 ; 5 ; 5 ; 2-3)

75 . Aste r aceae . Herbs or shrubs; inflorescence a capitulum; pappus , bracts,

syngenesium present ; carpels 2; ovary inferior ; ovule 1; fruit a nutlet
or cypsela. (pappus; 5 ; 5; 2)

Section B. '~SYNOP T ICAL KEY TO FAMILIES, ORDERS, AND SUBCLASSES OF

PINOPHYTA (GYMNOSPERMS) NATIVE AND CULT I VATED
IN NORTH AMERICA NORTH OF NEXICO

1. Leaves pinnately compound , terminating a simple or seldom-branched stem ;

seeds borne on sporophylls . CCycadopsida , Cycada l es)
2 . Leaflets ,,,,.i th single midrib and no lateral veins; megasporophylls
crowded on an indete r minate axis; cultivated only .... 1. Cycadaceae
2 . Leaflets with numerous par a llel longit udinal veins dichotomousl y
bran ched near the base ; megasporophy ll s form i ng a dense deter-
mi nate cone; cultivated and native .. . , .... . .. . . . ...... 2 . Zamiaceae

*Contributed by James W. Hardin, North Car olina State University , Raleigh .

"
650 29

1. Leaves simple, throughout branched stems ; seeds terminal or borne on

ovuliferous scales in cones .
3. Leaves fa n- shaped, bilohed at apex , dichotomously veined , deciduous ,
borne on ends of long shoots or on indeterminate short shoots ; seeds
1 - 2 t e rmin ating an elongated stalk and \·lith fles hy outer coat; tree ,
cultiva t ed only (Gnetopsida , Ginkgoales) ............. 3. Ginkgoaceae
3. Leaves acicular, linear, al"l-shaped , or scale-like , evergreen Or
deciduous; seeds terminal on short lateral branches or in cones .
4. Stems jointed and photosynthetic; secondary wood \~ith vessels ;
l eaves reduced to scales. opposite or in \"horls of 3; pollen
cones compound ; seeds dry l-lith l.,ringed bracts or fleshy \~ith
\~ing l ess bracts (Gnetops ida , Ephedrales) . ........ 4. Ephedraceae
4. Stems not jointed and \~itho llt vessels ; leaves photosynthetic and
variabl.e; pollen cones simple; seeds terminal without bracts or
borne on scales in cones . (Pinopsida)
5. Seeds one or rarely 2, drupe-like or berry- like; leaves
linear and p ers ist ent .
6 . Pollen-sacs 2 per sporophyll; leaves mostly 1-4 in. l ong;
seed on a lateral branch , drune-like t~i.th f leshY coat ,
also bearing an e nlarged fleshy aril at base of seed ;
cultivated only (Pinales) .. .. . . ........ . 5 . Pod ocarpaceae
6 . Pollen-sacs 3- 9 per sporophyll ; l eaves 1/2 to J in . long .
7 . Leaves 1- 3 in . long \~ith 2 broad g la ucous lines beneath ;
seed drupe-like with f leshy outer coat ; basal a ril
lacking or rudimentary ; cult i vated only
(Pinales) . .... ... . ......... ...... 6 . Cephalotaxaceae
7. Leaves 1/2-1 1/4 in . long , either lVith pale green to
tawny bands beneath or with 2 very narrot~ glaucous
lines; s eeds part ially or complete l y enclosed by a
fleshy aril (Taxales) . . . . . . . . . . . .. . . ... 7 . Taxaceae
5. Seeds one to many in a ~~oody or berry-like cone ; leaves
variable , p ersistent or dec i duous (Pinales) .
8 . Leaves ovate-lanceolate , 1/3 in . or more wid e at base ; one
ovule per ovulate scale ; scale and bract fused ; c u lti-
vated only .. ...... ..... . ... ...... .... . . 8 . Araucariaceae
8 . Leaves usually narrower at base ; ovuliferous scale lVith 2
o r more ovules ; scales and bracts separate o r partially
to completely fused .
9 . Ovul i ferous scales spirally arranged, imbr icate and
flat to someHhat peltate ; leaves usually al te rnate
(opposite or apparently whorled in cultivated
Hetaseguoia and Sciadopit ys) .
10 . Female cones lVith flat scales and 2 rounded
seeds; brac t sub tending ovuliferous scale
distinct ; pollen Hith 2 ai r- bladders; l eaves
acicular or linear . .. . . ... .... ..... 9 . Pinaceae
10 . Female cones Idth some\~hat peltate sca l es and 2 9
triangular seeds ; bract fused to scale ; pollen
Hithout air-bladders; leaves and tHigs deciduous
or persistent, leaves linear , alVl- or scale -
like ....... .... ....... . . ....... 10 . Taxodiaceae
9 . Ovuliferous scales opposite or \~horled and pc-Itate or
flat ; bract fus ed with scale ; po llen without air -
bladders; leaves opposite or \~horled , aHl- or scale-
like, persistent. .. . .............. . 11. Cupres s aceae
29 651

Section C.

SYNOPTICAL KEYS TO FANILIES, ORDERS, ANn SUBCLASSES OF ANGIOSPERNS

(Adapted from Cronquist I s "Evolution nnd Cl assification of

Flowering Plants [1968]; with permission . )

Class : ~~gno l iopsida (Dicots) Class : Liliopsida (Hanocots)

Cotyledons 2 (seldom 1, 3, or 4) Cotyledon 1 (o r the embryo sometimes

undifferentiated)
Leaves most l y net-veined Leaves most l y parallel-veined
Intrafascicular cambium usually Intrafascicular cambium us ual l y absent ;
present usually no cambium of any sor t
Vascular bundles usually borne in Vascular bund l es generally sca t tered.
a ring I.;h i ch encloses a pith or in 2 or more rings
Floral parts, I~hen of definite num- Floral parts , ,.,.hen of defin ite number
ber, typically borne in sets of 5, typically horne in sets of 3 , seldom
less often 4 , seldom 3 (carpels 4, never 5 (carpels often fewer)
often Eel.;er)
Pollen typical ly triaperturate, or Pollen of uniaperturate or uniaper-
of triaperturate-derived type, turate-der ived type
except in a fel'; of the more pr imi-
tive fam il ies
Hature roo t system eithe r primary or l-Iature root system I.;holly adventitious
adventitious, or. both

SYNOPTICAL KEY TO THE SUBC LASSES OF' HAGNOLIOPSIIM (DICOTS)

1. Plants r e latively primitive, the flOl. . ers typi ca lly apoca rpOlls, always poly-
petalous or apetalous and nearly always \<lith an evid pn t per i anth , usually
I.; ith numerous, centripetal stamens, the pollen ah.;ay s binucleate and
often uniaper turate ; ovules ah.;ays bite gmic and crassinucell a te ... ... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . I . Hagnoliidae
1. Plants more advanced in one or more respects than the Nagnoli i dae, and
never with uniaperturate pollen .
2 . Flowers more or less st rongly reduced and often uni se xual , t he peri-
anth , poorly developed or I.;anting, often borne in catkin s , bu t never
forming bisexual pseudanthia, and never wit h numerous seeds on
parietal placentae . . .. . .. . . .. . .. . . .. .. .. . . ... . . . .... II. Hamamelidae
2 . FIO\.;ers usually more or less I"ell developed and '-'.'ith an evid ent
ped anth, but if no t so , then usually either grouped into bisexual
pseudanthia or with numerous seeds on parietal pL'Icentae , only
rarely with a ll of the charac ters of the Hamame l idae as g i ven above .
3 . Flowers polypetalous or less often apeta l ou s or sympeta l ou s , if
sympetalous then u s ually either with more stamens t han corolla-
lobes, or \~ith the stamens opposite the corolla lobes , or with
bit egmic or crassinucellate ovules; ca r pels 1 to many , f r ee or
more often united.
4 . Stamens, ,-,.'hen numerous, developing in centrifugal sequence;
placentation various , often parietal or free - central or
basal, but often also axile ; species with few stamens and
axile pl acentation usually either bearing several or many
ovules per locule, or ,.;ith a symp eta l ous corolla , or both .
652 29

S. Pollen trinucleate; ovules bitegmic, crassinucellate, most

often campylotropous or amphitropous; seeds very often
with perisperm; plants usually either with betalains
instead of anthocyanins , or with free-central to basal
placentation , or both; only a few species are ordinar y
trees ....... ... .... .. .......... .. . .. III. Caryophyllidae
S. Pollen usually binucleate (notable exception: Cruciferae);
ovules various, but seldom campylotropous or amphitro-
pOllS; seeds seldom with perisperm; no betalains; placen-
tation rarely free-central or basa l, except in the
Primulales; many species are ordinary trees ... . . " .....
. . . .. ..... .... .. ..... .. . ......... .. . IV. Dilleniidae
4. Stamens, when numerous. developing in centripetal sequence;
flowers seldom with parietal placentation (notable ex cep-
tion: many Saxifragaceae) and also seldom (except in para-
s i tic species) with free-central or basal placentation in
a unilocular. compound ovary, but very often (especially
in species with few stamens) with 2- several locules that
have only 1 or 2 ovul es each; flowers polypetalous or less
often apetalous. only rarely sympetalous .. ....... V. Rosidae
3. Flowers sympetalous; stamens generally isomerous with the corolla
lobes or fewer , never opposite the lobes; ovules unitegmic and
tenuinucellate; carpels most commonly 2. occasionally 3-5 or
more ................. ... ..... . •. ............ . ..... VI. Asteridae

1. }1AGNOL IIDAE

1. Plants \·lith ethereal oil cells.

2. Plants woody; flowers normally developed. with an evident perianth of
separate tepals which mayor may not be differentiated into sepals
and petals .......................... ....... .......... 1. Nagnoliales
2. Plants differing in one or more respects from the above, most species
herbaceous .
3 . Flm..rers with much reduced or no perianth. tending to be crowded
into a spadix; ovules orthotropous; seeds often with perispe r m.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . Piperales
3 . Flowers with a well developed perianth typically consisting of a
gamosepalous , more or less corolloid calyx. not crowded into a
spadix ; ovules anatropous; seeds never with perisperm ......... .
............. .... .. . . .......... ...... ...... .. 3 . Aristolochiales
1. Plants without ethereal oil cells.
4 . Plants aquatic. lacking vessels; p l acentation laminar ; most species
\..rith uniaperturate or uniaperturate-derived pollen ... 4 . Nymphaeales
4. Pl ants terrestrial or occasionally aquatic, \-lith vessels; placenta-
tion marginal; pollen triaperturate or triaperturate-derived .
5 . Gynoecium mostly apocarpous or seemingly or actually unicarpel l ate,
seldom evidently syncarpous; sepals usually more than 2 ....... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. Ranunculales
5. Gynoecium syncarpous; placentation parietal; sepals 2 o r occasion-
ally 3 ... .... .................................... 6. Papaver ales

1. HAGNOLIALES

1. Flm~ers hypogynous; endosperm ,veIl developed; most families with several

or numerous ovules per carpel; pollen mostly uniaperturate or triaper-
turate .
29 e3

2. Pollen uniaperturate; stamens often more or l ess laminar .

3. Nodes unilacunar, two-trace; stamens 6-9.
4. Stamens laminar; tepa 15 numerous; carpels 6-14; lianas Hith
opposite. exstipulate leaves .. ... .... .. . 1 . Austrobaileyaceae
4. Stamens \.,tith normal filament and anther: tepals 3; carpels 3;
erect shrubs with alternate , stipulate leaves . ... . ......... .
.. . . .... . .. .. . . . .... ..... . . ..... . .. ... . . . . .. 2 . Lactoriclaceae
3. Nodes trilacunar or multilacunar; stamens few to more often
numerous.
5. Leaves stipulate .. . ... ... ... . .. ... . . .. . ..... ' , . . 3 . Magnoliaceae
5. Leaves exstipulate .
6. Vessels absent: stamens usually laminar: endosperm not
ruminate .. . . . .. .... ....... .. . . . .. . . .. . . ... 4. Winteraceae
6. Vessels present; stamens and endosperm various .
7. Stamens laminar; perianth not trimerous .
8. Carpel so l itary, pluriovulate, unsealed; endosperm
ruminate ... .. . ..... . ..... . . . . ... 5 . Degeneriaceae
8. Carpels several, uniovulate. largely or wholly
closed ; endosperm not ruminate. 6.Himantandraceae
7 . Stamens not laminar ; perianth most commonly tri
merous .
9 . Stamens distinct, lvith short. thick filament and
l.,rith the connective surpassing the anther and
enlarged above it; endosperm ruminate; ovules
I-many .. ... . . .. ... ..... ... .. ... . ... . 7. Annonaceae
9 . Stamens with the filaments united; endosperm not
ruminate.
10. Carpels separate; ovul e solitary ; flowers uni-
sexual; filaments united into a solid solumn
... . . . . . ..... ...... . ..... . .. 8. Myristicaceae
10. Carpels united i nto a compound pistil with
parietal placentation; ovules 2-many ;
f lowers perfect; filaments united into a
tube ..... . .. . .... .. . .... .. .... 9. Canellaceae
2. Pollen triaperturate; stamens Idth normal filament and anther; nodes
uni1acunar; leaves alternate, exstipulate .
11. FIOl.,rers perfect; trees; carpels in one l.,rhorl; fruit follicular
. ....... .. .. . . . .. .. .. .... . ..... . . . .... ... . ......... 10. Illiciaceae
11. FIOl.,rers unisexual; lianas ; carpels spirally arranged. becoming
fleshy in fr uit ..... .. ... .... .... . . . . .... . .. . . ll . Schisandraceae
1 . Flowers perigynous to epigynous (except in Amborellaceae and Trimeniaceae);
ovules only 1 or 2 per carpel (except in Eupomatiaceae) ; pollen various
but seldom either uniaperturate or triaperturate; stipules none.
12 . Nodes multilacunar; ovules numerous; endosperm ruminate; leaves
alternate; pollen biaperturate ... . . .... . .. . .. . .. . .. 12. Eupomatiaceae
12. Nodes unilacunar: ovule 1 (or 2 but only 1 maturing) ; endosperm . when
present , not ruminate .
13. Vessels absent; stamens numerous; leaves alternate .. . . . . . . . . . . .. . .
. . . . . .. ... . . . . ... .. . . . ... . . ... . . . . . . ... . . . . .. . . . . 13 . Amborellaceae
13 . Vessels present; stamens numerous or often fel'; .
14 . Endosperm present; leaves opposite.
15. Ovary superior; flowe rs mostly unisexual .
16 . FlOl.,rers hypogynous; carpel 1 (2) . ... . . 14 . Trimeniaceae
16 . FIOl"ers perigynous; carpels (1) several to many .. . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15. Monimiaceae
"
654 29

15 . Ovary inferior; flowers perfect ; carpels 2- 3 .. . .. .... . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 . Gomortegaceae
14. Endosperm wanting.
17 . Carpels nume ro us ; pollen biaperturate o r rarely triaper-
turate; leaves opposite .. ... ..... .... 17. Calycanthaceae
17. Ca rpel 1 (or seemingly so) ; pollen nonapertura te or
rarely uniaperturate; leaves mostly alterna te .
18 . Ovary supe r ior . .... .. .. .... ........ . .... 18 . Lauraceae
18 . Ovary i nfe r ior . .. .. ..... .. .......... 19 . Hernandiaceae

2. PIPERALES

1. Nodes un ilacunar ; leaves opposite; ovary more or l ess inferior , unicar-

pellat e , with a single ov ule ; seed with copious endospe rm and no peri-
sperm ...... . .............................. . ........... 1. Chlo ranthaceae
1. Nodes tri l acunar or multila cunar ; l eaves mostly alternate ; ovary
sup erior , excep t in some Saururaceae ; seeds with copious pe r isp erm
and scanty endosperm .
2 . Ov ules ( I ) 2-10 per carpel ; carpels distinct or united into a com-
pound ovary wi t h parietal or modif i ed laminar pl acenta tion . . ... ... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . Saururac eae
2. Ovule sol itary i n t he single l ocule of the compound or pseudomono
merous pistil .................. . ...... .. ....... .. ... .. 3 . Piperaceae

3. ARISTOLOCHIALES

Represented by .. ...... .•. .. .... ... .. . . .. ..... .. ... .. . ... .. L Aristo lochiaceae

4. NYMPIIAEALES

1. Pla nts with roo t s ; flowers per fec t: carpe l s I - many ; leaves a l te r nate,
entire to dissec ted ; integuments 2 .
2. Polle n of uniaperturate or un i ape r turate- de rived type ; seeds with
smal l embryo , some endosperm , and ab undant perisperm ........ .. ....•
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . Nymphaeaceae
2. Po llen tr1aperturate ; seeds with large embryo and no endosperm or
peri sper m.... ...................................... 2 . Nelumbonaceae
1. Plants rootless . free - floating; f l owers unisexual; carpe l 1; l e aves
who rled , c l eft with slender segments ; integument 1; pollen uniaper-
turate . ...... .. . .. .................................. 3 . Ce r atophyllaceae

5. RANUNCULALES

1. Gynoecium mostly of separate carpels , o r seemingly or actua lly of a sin-

gle carpel; endospe rm copious to scan ty or none ; herbs and woody vines,
seldom shrubs .
2. Leave s mostly alterna t e ; pe t als not adhering to th e carpels ; flm.ers
variously organ iz ed, often trimerous in part or wit h numerous sta-
mens and carp els. never pentamerous throughout .
3. Flowers pe rfect; herb s (includ ing herbaceous vines) o r shrubs;
endosper m wel l devel oped .
4. Leaves wi th dicho tomous , free venation; nodes unilacunar;
ovul es orthotropous; flowers r educed , wit hout petal s or
petaloid nec taries ...................... 2 . Circaeas t e ra ceae
29 655

4. Leaves with net vena tion ; nod es trila cunar to multilacunar;

ovules anatropous ; fl owers usually we ll developed , often
Idth necta r iferous petals .
5 . Carp els usually 2 or more and distinct , seldom so l itary
or weakly united ; stamens usually numerous , the
anthers opening by longit udina l sl i t s; he rb s , rar ely
shrubs ... ... .. . .. ... .. ... .. . . . . . .. . ..... 1. Ranuncul aceae
5 . Carpel seemingly solitary ( the pi stil probably actually
2- 3 carpell ate with a l l hut one carpe l obsolete) ; sta-
mens typic ally I-several sets o f 3 , usua ll y opening
by valves; herbs and sh rub s . . . .......... 3 . Be rberidaceae
3. Flowers unisexual ; nearly all woody plan t s , most l y vines .
6. Leaves mostly compound; seeds with s mall embryo and abundant
endos perm; petals when present nectariferous; mos t l y
monoecious ... .. . . ............... . .... .. . . . 4 . Lardizabalaceae
6 . Leaves mostly simple ; seeds mo st l y with l ar ge embryo a nd
scanty endosp e rm; petals not nectarif e rous; dioecious . .. .. . .
..... ..... .. .. ... .. . ........ . . . .. . ... . . ... . 5 . Henispermaceae
2 . Leaves opposite or l.;rhorled; peta ls persisten t , enlarging i n fruit and
adhering to the i ndivid ua l carp el s t o produc e a set of pseudodrupes;
f lowers wholly pentamerous ; nodes unilac unar ... ...... 6. Coria r iaceae
1. Gynoecium obviously of un ited carp e l s ; endosperm scan t y or none; t rees
ar,d shrubs .
7 . Flm.;rers di plo s temonous, the inne r set of stamens modifi ed into stami-
nodes; carp e ls 2 but only one fertile , this ,,,Hh one ovule . . .... ... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 . Cor ynocarpac eae
7 . Flowers haplostemonous; carpels 2-3, each with 1 or 2 ovules ..... ... . .
. ..... . . . ...... . . . .... . . .... . ..... . ..... .... .. .. . ..... . . 8. Sabiaceae

6. PAPAVERALES

1. Stamens numerous; f lowers regular ; petal s neither spurred nor saccate;

la t ex s ystem more or less we ll developed . . . .... . . . ... . . .. 1 . Papaveraceae
1. Stamens 4 or 6; f l owe rs usually irregular; usually some of the petal s
s pur red or saccate ; l atex sys t em wan ting or occasiona lly rep r esented by
scatter ed elongate secretory cel l s .... . ... . . . . . .. .. . ... . . . 2 . Fumariaceae

II . lIANAHELIDAE

1. Flowe rs eithe r with distinct carpels (a single ca rpel in Didymeles , of th e

Hamamelida l es ), or \.;rith dehiscent fruits, or both, var io usly perfect or
unisexual ; ovules I - many per car pe l; seeds endospermous, l.;rith small to
l a r ge emb r yo .
2. Vessels wanting ; some or all of the flowers perfect ; carpels 4 or
more , in a single ,.,thorl , laterally coherent but scarcely forming
a compound pis til , the group of carp els ripen i ng into a f o l licetum;
ovules several to many in each carpel; s tipules wanting or repre-
sented only by a pai r o f flanges on the pe tiole ; leaves wit h unique .
elongate (often branched) icliob l asts ....... ' . ..... 1 . Tr ochodendra1es
2 . Vessels present, f l ot-lers perfect or more often unisexual ; ca rpel s
either I-several and separa te , or 2-4 and united (at l east below)
to form a comp ound pist il; ovules I -many ; s tipules nearly a l ways
present excep t i n Euptelea and Didymeles ; no consp icuous idio-
blasts . ..... . . .. . .. ..... .. ... .... .. .... ... .. . . .... . . 2 . Hamamelida l es
656 29

1. Flowers either with united carpels, or with a pseudomonomerous , seemingly

unicarpellate pistil; fruits indehiscent; ovules not more than 2 per
carpel ; f10Hers in most families unis exual ; seeds with or without endo-
sperm, but the embryo always fairly large.
3. Leaves pinnately compound or trifolio1ate; ovary 1-2 locular. Hith
1-2 ovules per 1ocu1e, or one locu1e empty . . . .. ...... 6 . Juglandales
3. Leaves simple (though sometimes deeply lobed or cleft); gynoecium
various .
4. F1moJers , or at least the male fl owers , not in catkins; ovary uni-
locular or rarely biloc ular; calyx often present; plants woody
or herbaceous.
5 . Ovules 2, unitegmic , intermediate betHeen crassinucellate and
tenuinucel1ate; stipules none ; nodes unilacunar; flowers
so l itary, ,oJithout a pe rianth .. .. ......... .. . . 3 . Eucommiales
5. Ovule solitary , bitegmic , crassinucellate; stipu1es usually
present; nodes tri1acunar to multi1acunar; flowers clus-
tered, usually with a vestigial perianth . . .... . 4. Urticales
4 . F1m"ers, or at least the male flowe rs, usually in catkins (except
some Fagaceae); ovary ,,,ith I-several locules; calyx mostly
\"anting or very much reduced; plants \"oody.
6. Pistil pseudomonomerous, seemingly composed o f a single car-
pel, with a single style and locule; ovary superior . .. .. . . .
. . ... . .. . . ... ... .. . . .. . . . . . ... . .. .. ..... ... . 5 . Leitneriales
6. Pistil composed of more than one carpel, as evidenced by the
presence of more than one sty l e and often also more than
one locule; ovary in ferior or nude .
7. Leaves opposite or alternate, seldom whor l ed, always more
or less Hell developed; ovules each with a single
embryo sac .
8. Ovule solitary , orthotropous (except in Canacomyrica),
unit egmic ; plants aromatic , the leaves resinous-
dotted ... .. . . . . . .... .... .. . .. . ... . . . .. . 7 . Myrica1es
8 . Ovules more than one, anatropous, bitegmic or uni
tegmic; plants not notably aromatic . . .... 8 . Fagales
7. Leaves whorled, reduced to scales; ovu l es with multiple
embryo-sacs . . .. ... ... .. ................. 9 . Casuarina1es

1. TROCHODENDRALES

1. Perianth present; stipular flanges present; stamens 4 , opposite the sep-

als; leaves palmately veined; idioblasts simple or branched, secre-
tory .. . . . .. . . . . . .. . .... . ... . . . . . . . .. ..... . . . . . ..... . . 1 . Tetracentraceae
1. Perianth none; stipu1es none; stamens numerous; leaves pinnately veined;
idioblasts branched, not secretory . .. . . . .. . . ... . . . .. 2. Trochodendraceae

2. HAMAHELIDALES

1. Carpels separate (or solitary) .

2 . Ovules numerous; fruit follicular; plants dioecious; leaves stipu-
late, palmately veined, entire . ........... . .... 1. Cercidiphyl1aceae
2. Ovules 1- 3; f ruit indeh iscent; f lowers var iously perfect or uni
sexual; leaves various .
3. Carpels several; stamens 3-many; plants monoecious or with per-
fect flow ers .
 Stamens numerous; ovules more or less anatropous; flm~ers per-
4.
fec t; connective of ordinary type , not apically enlarged
and peltate; leaves exstipulate , pinnately veined .......... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Eupteleaceae
4. Stamens 3-4 (7); ovules more or less orthotropous; plants
monoecious; connective apically enlarged and peltate;
leaves stipulate, palmately veined ............ 3. Platanaceae
3. Carpel solitary; stamens 2, united by their fi laments; plants
dioecious ........................................ 4. Didymelaceae
1. Carpels united (at least below) into a compound pistil.
S. Leaves alternate (rarely opposite); carpels 2 (rarely 3); flowers
perfect or unisexual, with at least a vestigial calyx and often
with a corolla as ,.,rell; f ilaments free; ovul es I-many ; trees or
shrubs ............ ... .............................. 5 . Hamamelidaceae
5. Leaves opposite; carpels 3-4; plants dioecious; perianth none; fi l a-
ments united; ovules many; undershrubs ............. 6 . }!yrothamnaceae

3. EUCOMHIALES

Represen ted by ....................... .... ...................... 1. Eucommiaceae

4. URTICALES

1. Trees, Hithout milky juice; seeds l.,rith straight embryo and no endosp erm ;
ovule pendulous. anatropous; anthers e rect in bud.
2. Leaves alternate, stipulate; styles 2 ...................... 1 . Ulmaceae
2. Leaves opposite, exstipulate; style 1 ................... 2. Barbeyaceae
1. Plants either not woody, or with milky juice, or with endospermous seeds,
often with 2 or even all 3 of these features; anthers in flexed in bud ,
or less often erect.
3. Styles or style branches 2, sometimes one of them reduced ; ovules
mostly anatropous and pendulous; embryo straight or more often
curved.
4 . Woody plants, seldom herbs, mostly \o.'ith milky juice; anthers
usually in f lexe d in bud .............................. 3 . Horaceae
4. Herbs. without milky juice ; anthers erect in bud .... 4. Cannahaceae
3. Style I , unbranched; ovule orthotropous, basal ; embryo straight; herbs,
seldom shrubs or sof t-wooded trees, without milky juice ............ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. Urticaceae

5. LEITNERIALES

Represented by . ............................................... 1. Leitneriaceae

6. JUGLANDALES

1. FloHers in triplets, the middle one perfect and fertile, wi th 4 sepals, 6

stamens, and a superior , bilocular ovary, one locule empty, the other
with a single ovule attached to the partition; lateral flowers female
but sterile; leaves pinnately compound , stipulate; f ruit samaroid ...... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Rhoipteleaceae
1. Flm.,rers unisexua l , the staminate ones l.,rith 3-many stamens, borne in cat
kins, the calyx wanting or adnate to the bract; female flm.,rers solitary
or spicate, with 1- 4 sepals or calyx teeth; fr uit drupaceous or nut-
like.
,
658 29

2. Ovules 4 (2 in each of 2 locules) , apical ; ovary superior : leaves

trifoliate, ninutely stipulate ................. .. 2 . picrodendraceae
2. Ovule soli t ary , basal, in a unilocular , inferior ovary ; leaves pin
nately compound , exstipulate ....... . .. ... ........... J . Juglandaceae

7. HYRICALES

Represented by ..... ... ........ ............ ... ..... •. . . . . . •• .•... 1 . Hyricaeeae

8. FAGALES

1. Seeds with endosperm; leaves exstipulate ; ovules erect , basal or nearly

so, unitegmic; plants dioecious ; pistillate flmver solitary i n a multi-
bracteate involucre \"h i ch ripens into a hull subtending the fruit ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Balanopaceae
1. Seeds Hithout e ndosperm; leaves stipulate ; ovules pendulous , apical or
nearly so; plants monoecious or seldom dioecious, or the floHers rarely
perfect.
2 . Styles and locules 3 (seldom 6) ; int e guments 2 ; female floHers mostly
not in catkins, sub tended individually or in small groups by an
involucre or many bracts Hhich are concrescent at maturity into an
indura ted hull ...... ... ............. .. .................. 2. Fagaceae
2 . Styles and locules 2 (se ldom J); integument 1; female flO\.;rers mostly
borne in catkins; r ruits Idthout a hull of bracts , or sometimes
Hith a f oliaceous hull derived from 2 or a fel" bracts . 3 . Betulaceae

9. CASUARINALES

Represen ted by ... ........ ... .................... .. ... . • ... • .. 1 . Casuarinaceae

III. CARYOPJlYLLIDAE

1. Seeds \"ith very scanty or no true endosperm, but very often Hith an
evident perisperm; betalains often present; plants often \"ith
anomalous secondary thickening .
2 . Ovules campylotropous or amphitropous, seldom anatropous; see ds \"ith
a curved embryo that is typically peripheral to a \.;ell defined
perisperm; pollen trinucleate; flOl-Jers seldom very much reduced
or borne in catkins, but if so (some Chenopodiaceae) then Hith
unilocular , uniovulate ovary .. .. . . .... . ........... 1. Caryophyllales
2. Ovules anatropous; seeds Hith nearly or quite straight embryo and
neither endosperm nor per isperm; pollen binucleate; flO\.;ers very
much reduced, borne in unisexual catkins ; ovary 4-1ocular , Hith one
ovule per locule; plants lacking both anthocyanins and betalains ...
......... . .... .... ........ ....... ..... ... .. . ............ . 2 . Batales
1. Seeds with more or less copious endosperm, but without perisperm; embryo
straight or curved , peripheral or often embedded in the endosperm ;
ovary unilocular, Hith a single basal ovule; no betalains ; no anomalous
secondary thickening.
3 . Perianth undifferentiated, or with 2 sets of more or less similar
tepals, not dichlamydeous and sympetalous ; carpels (2) 3 (4); sta-
mens mostly 6-9, in 2 or 3 cycles; plants often \.;ith sheathing
s tipules ........ ..... ...... .... .. .. ...... ........ .... 3. Polygonales
3 . Perianth differentiated into calyx and corolla , the corolla sym
petalous; carpels 5; stamens 5 , opposite the petals ; leaves mostly
exstipulate .. . .. .. ................... .. ............ 4. Plumbaginales
29 ~9

1. CARYOPHYLLALES

1. Gynoecium apocarpous , monocarpous , or sometimes syncarpous , if syncarpOll$

then usually with as many locules as car pels; ovules 1 per locule , or
some l ocules empty .
2. Locules a ll ovulifero us (though not a l ways fertile) ; perisperm wel l
developed; flO\·,ers perfect or less often unisexual ; plants not
cactuslike .
3. Sepals mostly free or nearly so , seldom petaloid; carpels mostly
2- several , occasionally only one ; inflorescence usually race-
mose or spicate ; leaves alternate ; stem seldom with anomalous
secondary thickening . ....... .. .......... ....... l . Phytolaccaceae
3 . Sepals joined into a tube \-Ihich resembles a sympetalous corolla .
and often sub tended by sepaloid bracts; carpel 1; inflorescence
most l y cymose ; leaves opposite or sometimes a lterna te; stem
very often l.,rith anomalous secondary thickening .. 2 . Nyc taginaceae
2 . Two locules empty , the third with a singl e ovule; perispe r m apparently
\~anting; flowers unisexual; inflorescence cymose ; spiny, cactuslike
trees . . .. . .... . ..... . ... . ... . ... ...... ..... . . . . .. ... . 3. Didiereaceae
1. Gynoecium mostly syncarpous. either unilocular or with the ovules more
numerous than the l ocules. or both .
4 . Tcpals and stamens usually more or less numerous; plants succulent;
ovary very often inferior.
5. Ovary unilocula r, nearly always inferior, \~ith parietal (rarely
basal) placentation ; mostly spiny stem-succulents l.,rith much
reduced leaves; Nc\~ t.lorld . . . ...... ...... ..... . ..... . 4 . Cactaceae
5. Ovary plur i l ocular , or occasionally unilocular with a f r ee-central
placenta ; ovary superior to inferior; leaf-succulents, not
spiny; chiefly Old Horld .. . . . .............. ....... .. 5 . Aizoaceae
4 . Tepals and stamens mostly few and cyclic ; plants sometimes with succu-
lent leaves . but not spiny or cactuslike; ovary superior .
6. Perianth usually dichlamydeous or seemingly so (except notably
the Holluginaceae, with axile placentation) .
7 . Sepa l s 4 or 5 ; plants without betalains , but usual ly with
anthocyan ins ; stamens not at once of the same number as
and opposite the petals; plants sometimes Idth anomalous
secondar y thickening .
S . Ovary 2-S-locular , with axile placentation; petals much
reduced or commonly wan ting; leaves opposite , alter-
nate or ,... horled ..... .. .. ... . ... ... . . .... 6 . Nollugjnaceae
8 . Ovary unilocular (sometimes partitioned toward the base.
~1ith free-central or basa l placentation ; petals usually
more or less well develop ed ; leaves opposite ....... .. . . .
. . . . . .. . ... ..... . . . . ..... .. . . . . . . . .. . . 7. Caryophvllaceae
7. Sepals (or sepaloid bracts) mostly 2, seldom more ; plants I~ith
betalains , lacking anthocyanins; stamens usually as many as
and opposite the petals (o r pe t a l oid sepal s) , o r sometimes
more numerous; no anomalo us secondary thickenin g .
9 . Ovules (1) 2- many; plants not climbing; internal ph l oem
usually \...anting ; fruit usually capsular . B. Portulacaceae
9 . Ovule solitary; climbers ; internal phloem present ; fruit
indehiscen t. . . . . . .. . . . . . ... . . . . . . . . . . . . . . . 9 . Basellaceae
600 29

6. Perianth monoch1amydeous. seldom peta1oid , often small or some-

times even obsolete; ovules mostly solitary (rarely several)
on a basal placenta; stem nearly alHays with anomalous secon-
dary thickening.
10 . Perianth not scarious , o fte n green; filaments mostly free;
flowers perfect or not infrequently unisexual . . . ..... . .... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. Chenopodiaceae
10 . Perianth more or less scarious; filaments mostly connate
belm. (the androecial tube sometimes even simulating a sym-
petalous corolla); flowe rs most ly perfect, seldom unisexual
... ... ... ..... .... . ... ... . ... . . . . . . ... .... 11. Amaranthaceae

2. BATALES

Represented by .. ...•.• • . . . • •• • •. ... . . ....... ...... . . .. . . ... • . ... . . 1. Bataceae

3. POLYGONALES

Represented by . .. .. • • • • • • .. .. • .. .. . .. . . .. . ... ....... .... . . ... . 1. Polygonaceae

4. PLUHBAGINALES

Rep resented by ........ .. • . . ... . .. .. ...... . ...... ... ... . . .• .. 1. Plumbaginaceae

IV. DILLENIIDAE

1. Carpels separate; stamens numerous; ovules bitegmic, crassinuce11ate;

seeds arillate and with a \.,rell developed endosperm . ..... . 1. Dilleniales
1. Carpels unit ed to form a compound pistil ; other characters various, the
stamens numerous to feH, the ovules bitegmic or unitegmic, crassinucel-
late to tenuinucellate, the seeds Idth or Idthout an aril and with or
without endosperm .
2 . Flowers mostly polypetalous or apetalous, seldom (mainly some Vio-
lales) sympetalous; stamens numerous to feH; ovules chiefly biteg-
mic and crassinucel1ate, but sometimes unitegmic, or tenuinucel-
late, or both.
3. Plants insect ivorou s herbs or shrubs .. .. .... ..... 5 . Sarraceniales
3. Plants not insectivorous; habit various .
4 . Placentation mostly axile, seldom parietal.
5. Sepals imbricate; filaments free or connate in groups;
mucilage cells and sacs mostly wanting .. . . ... 2. Theales
5. Sepals va1vate; filaments often connate into a tube or
column; mucilage cells or sacs very often present .
6. Ovary superior; pubescence very often stellate or
lepidote . . .. ... .. . ... ........ . . ..... . . . . 3 . Halvales
6 . Ovary inferior; no stellate or 1epidote hairs . .... .. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 . Lecythida1es
4 . Placentation mostly parietal , seldom axile .
7 . FloHers normally developed, l.,rith an evident perianth.
perfect or less often unisexual, not borne in catkins.
8 . Plants usually without myrosin cells , very often
\"roody; f101.,rers hypogynous to ofte n perigynous; or
epigynous; l eaves mostly simple: carpels mostly 3
or more, seldom only 2; perianth seldom tetra-
merous . . .. ....... . . .. .. ..... .. . . .. . . ... . 6. Vio1a1es
29 661

8. Plants commonly with myrosin cells, herbaceous or less

often \wocly; flowers hypogynous; leaves simple to
more often compound or various l y dissected; carpels
most commonly 2: perianth often tetramerous . ..... . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 . Capparal es
7 . Flowers much reduced, un i sexual , \.,rithout an evident per
iaoth, borne in catkins; woody plants .. . ... . 7 . Salicale s
2. Flowers mostly sympetalous (except mainly some Ericales, these with
unitegmic. tenuinucellate ovules) ; stamens seldom more than 2 or 3
times as many as the corolla lobes.
9 . Placen tation axile or less often parieta l; stamens various, often
more numerous than or alternate with the corol l a lobes .
10. Plants with the pollen in tetrads (sometimes only one grain of
the tetrad maturing), or with poricidal anthers , or lacking
chlorophyll , often with more than one of these characters;
ovules unitegmic, tenuinuce l late ..... . .. . ........ 9 . Ericales
10. Plants otherwise , the pollen in monads, the anthers opening by
longi tudinal slits, the shoot always chlorophyllous ; ovules
bitegmic or unitegmic, crassinucellate or tenuinucellate.
11 . Plants dwarf (commonly prostrate) shrubs; functional sta-
mens as many as and alternate with the corolla lobes;
ovules unitegmic ... .. . . .. .. .. . . .. ... .... 10. Diapensiales
11. Plants shrubs or trees; stamens usually either more num
erous than or opposite the corolla lobes; ovules biteg-
mic or unitegmic . ... . . . ..... ....... ... .... . . l l. Ebenales
9 . P l acentation free- central or basal; functional stamens as many as
and opposite to the coro l la lobes; ovules chiefly bitegmic .... . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12 . Primulales

1. DILLENIALES

1. Flowers hypogynous , with neither a hypanthium nor an intrastaminal disk;

raphides and crystal sand present , often in long sacs or tubes; leaves
mostly entire; trees , \wody vines, and shrubs, seldom herbs , of tropi-
cal and subtropical regions , especially Australia . . . . . . . . l. Dil1eniaceae
1. Flowers perigynous, or with an intrastaminal disk; raphides and crystal
sand absent ; North Temperate regions.
2 . Seed \o1ith copious endosperm and smal l embryo; flowers with an intra-
staminal disk, but without an evident hypanthium; mesophytic shrubs
or herbs with large , usually cleft leaves .. . . ... . . . . .. 2. Paeoniaceae
2 . Seed with thin endosperm and large embryo ; flowl2rs with an evident
hypanthium , but without an intrastami na1 disk; xe rophytic shrubs
with small , entire leaves .. . ..... . ..... . . . ...... . 3. Crossosomataceae

2. THEALES

1. Leaves mostly alternate; endosperm present or absent.

2. Leaves mostly stipulate .
3 . Leaves simple , or seldom pinnately compound ; embryo of normal
proportions.
4 . Connective not exserted; calyx not winged; endosperm present
in most species.
5 . Style mostly gynobasic; anthers mostly opening by termi-
nal pores ; plants lacking mucilage cells; mostly not
of Madagascar ... . .. ..... . ... .. ....... . ... ... 1 . Ochnaceae
662 29

5.Style not gynobasic (except in one sp . of Rhopalocarpa-

ceae) ; anthers open i ng by l ongitudinal s l its ; plants
Hith mucilage cells; confined to Madagascar .
6 . Flm..rers Hith a prominent intrastaminal disk; fila-
ments irregul arly connate toward the base; po l len
in monaus .. ......... .... .. ...... 2 . Rhopalocarpaceae
6 . FloHers without an intrastaminal disk, the stamens
inserted within a ring of staminodes ; filaments
free; pollen in tetrads .... .. ... . . 3. Sarcolaenaceae
4 . Connect i ve prominently exserted; calyx mostly winged in
fruit ; endosperm mostly "'anting . .. .. .... 4 . Dipterocarpaceae
3. Leaves palmately compound; embryo with enlarged , thickened hypo-
cotyl and reduced cotyledons . .... ... ....... .. .. 5. Caryocaraceae
2 . Leaves exst i pu l ate , except in some Theaceae .
7 . Erect trees or shrubs .
8 . Petals usually 5: stamens mostly numerous , seldom only one
or 2 cycles; stigmas (often also the styles) distinct .... . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. Theaceae
8. Petals 4 ; stamens 8; style \..rith a capitate , merely lobed
stigma ..... .. . . ............................ 7 . Stachyuraceae
7 . Trailing or climbing shrubs or vines .
9 . Endosperm copious; integument 1 ; bracts not highly modi fied ..
.... . ........ . ........... .. ... ... . . ..... .. . 8. Actinidiaceae
9 . Endosperm scanty or none; integuments 2; bracts highly modi
fied, becoming pi tcherlike , saccate , or spurred ... .. ...... .
. ... ... . . .. . ... . . .. . .. . .... . . . . .. . .. . .... . 9 . Marcgraviaceae
1 . Leaves mostly opposite or whorled; endosperm Hanting .
10 . Leaves stipulate .
11 . Trees ; stamens 15 or more ; fruit a berry ..... .. ... . 10 . Qui i naceae
11. Herbs or low shrubs; stamens 10 or f ewer; fruit
capsular ......... .. ... .. ..................... ... 11. Elatinaceae
10. Leaves exstipulate .
12 . Ovary l7 - 25- celled; ovules 2 per cell; no secretory canals ..... . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. Hedusagynaceae
12. Ovary (1) 3- 5 (15) - celled; ovules I-many per cell; conspicuous
secretory canals or cavities in a l l organs ... .... IJ . Guttife r ae

3. HALVALES

1. Peta l s valvate; plants mostly t,ithout mucilage cells and ah..rays Hithout
mucilage sacs or canals; pubescence not stellate or lep i dote .
2. Leaves stipulate; sepals separate to the base or near l y so ; petals
mostl y separate; integuments 2 ...... . . . ... . . . .. ... 1. Elaeocarpaceae
2 . Leaves exst i pulate; sepals connate into a cup or tube; petals often
connate belm..r ; integument 1. ................. . .. . 2. Scytopetalaceae
1. Petals imb ri cate or contorted ; plants mostly with mucilage cells or often
\,ith mucilage sacs or canals; pubescence mostly stellate or l epidote.
3 . Anthers bilocular ; filaments free or connate; mostly trees and
shrubs, a few herbs .
4 . Stamens numerous , most l y free , arranged in a single whorl ...... . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . Ti liaceae
4. Stamens (including stami nodes) IO - numerous, in two or more whorl s,
mostly connate by their fi l aments, seldom only 5 and then
always connate . . . . ... .. .... ... ... ... . . . ........ 4 . Sterculiaceae
3. Anthers most l y unilocular; filaments connate.
29 663

5. Pollen smooth, tripo rate; fr uit loculicidal1y dehiscent . or inde-

hiscent; mostly trees and shrubs , a fe\-l herbs ... .. . 5 . Bombacaceae
5. Pollen mi n u tely spiny, mostly multiporate; f ruit typically septi
cidally dehiscent or $chizocarpic, seldom l oculicidally dehis-
cent; plants herbaceous, or occasionally softly woody ... . .. ..... .
........ . .......... ... ......... .... ................. . 6. Malvaceae

4. LECYTHIDALES

Represented by .•••• ••• •••••••..••••..•••...••••..••• •• •• •••••. 1. Lecythidaceae

5. SARRACENIALES

1. Leaves, or some of them , modified to form pitchers; placentation axile ;

style solitary , or sometimes very short or none .
2. Flowers perfect ; filaments distinct; pollen grains i n monads; ovules
unitegmic , tenuinucellate ; herbs , not c l imbing; New World .......... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Sarraceniaceae
2. Flm..rers unisexual; filaments united into a column; pollen grains in
tetrads; ovules bitegmic, crassinucellate; herbs or shrubs, often
climbin g; Old "\.]orld .. . ...... ... .. . ................... 2 . Nepenthaceae
1. Leaves not forming pitchers ; placentation parietal; styles severaL ...... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Droseraceae

6. Vl OLALES

1. Ovary usually superior; plants of various habit , most families with a well
developed endosperm.
2. Flowers polypeta1ous or sometimes apetalous; endosperm mostly well
developed.
3 . Endosperm oily ; many species cyanogenetic.
4. Stamens mostly numerous, seldom only 5 or fe~..rer; trees or
shrub s, seldom climbing .
5. Ovules mostly on parietal placentae; petals usually pres-
ent (sometimes scarce l y distinguishable from the
sepals); carpels 2-10, with free or united styles . ..... .
.. . . ...... . ..... . ...... . .... . . . ...... . . 1. Flacourtiaceae
5. Ovules hanging from an apical placenta; petals none; car-
pels 3-4; styles free .... ...... . ........ 2 . Peridiscaceae
4. Stamens mostly numerous , seldom only 5 or fewer; trees or
sometimes climbing.
6 . Carpels 9-12; stamens 3; flowers un isexual; ovules numer-
ous on a broad , basal placenta; woody plants ........... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . Scyphostegiaceae
6. Carpels 2-5 ; stamens 5; flowers mostly perfect; ovules on
parietal placentae .
7 . Flowers without a corona ; trees, shrubs, or herbs, not
climbing.
8 . Flowers hypogynous, with a single style , often
irregu1ar ........................... 4 . Violaceae
8 . Flowers perigynous, with separate styles, regular .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. Turneraceae
7. Flowers t..rith a corona; herbs and shrubs, often climb
ing.
664 29

9. Seeds arillate; plants often cl i mbing by tendrils;

flowers hypogynous to somewhat perigynous;
sepals imbricate ; petals imbricate or seldom
wanting ....... . ... . .. . ... .. ... 6. Passifloraceae
9. Seeds exarillate; undershrubs or herbs , not
climbing ; flm"ers strongly perigynous; sepals
and petals valvate ........... 7 . l'lalesherbiaceae
3. Endosperm starchy ; few species cyanogenetic .
10 . Plants variously woody or herbaceous. but not climbing;
ovules several or many, included in the fruit .
11 . Stamens mostly numerous; style 1; leaves small or more
often of ordinary size.
12. Leaves alternate , stipulate, pal mately lobed or
veined; ovules anatropous; upright shrubs or small
trees ................................... 8. Bixaceae
12 . Leaves opposite, exstipu1ate , entire, not palmate;
ovules orthotropous or seldom anatropous; herbs
or low shrubs . ... . . .. .. .. .... .. ...... .. 9. Cistaceae
11. Stamens mostly 4-10 , seldom numerous; style s I-severa l ;
leaves small , often ericoid or reduced to scales; herbs
or more often shrubs or trees.
13 . Leaves alternate: sepals connate belm,,; styles 3- 4 ...
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 . Tamaricaceae
13 . Leaves opposite; sepals free ; style 1 . . .... . ... ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. Frankeniaceae
10 . Plants woody climbers with stout , hooked branch tips or leaf
tips.
14. Ovules numerous, on 2-5 parietal placentae, conspicuously
exserted from the capsule at maturity; styles several;
plants climbing by hooked leaf -tips ... •• • •••.• .. • ....•.
.. .. .... . . . . . .. . . . . ... . . . .. .... . . . . 12 . Dioncophy11aceae
14. Ovule solitary on a basal placenta , included in the
mature nut; style solitary; plants climbing by hooked
branch-tips ....................... 13. Ancis troc1adaceae
2 . Flowers sympeta10us; endosperm scanty or none except in the Acharia
ceae .
15 . Flowers perfect; leaves simple, entire or nearly so; stamens 10
or more .
16 . Thorny, xerophytic shrubs with small leaves; carpels 3;
fruit capsular; stamens 10-17 ............ 14 . Fouquieriaceae
16. Unarmed, large- leaved trees; carpel s 2 ; fruit drupaceous;
stamens numerous ...................... 15. Hoplestigmataceae
15 . Flowers unisexual; leaves simple or often lobed or compound;
stamens 10 or less.
17 . Endosperm well developed; plants herbaceous or nearly so and
\~ithout a latex system; stamens 3-5; style 1. . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 . Achariaceae
17 . Endosperm scanty or none; plants woody and with a well
developed latex system; stamens 10 (5); styles free ....... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17 . Caricaceae
1. Ovary mostly inferior; plant s mostly herbs or herbaceous vines, a fet-I
trees and shrubs; endosperm scanty or none, except in some Loasaceae .
18 . Flowers perfect; stamens mostly numerous; style 1; integument 1 . .... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18. Loasaceae
18 . Flowers mostly unisexual; stamens few or less often numerous; styles
I-several ; integuments 2.
29 665

19 . S tamens 4-many , a ll with bilocula r an t he r s ; plant s without ten-

dr ils; s t yles fr ee except in some Begoniaceae ; pe tals s epa -
rate or none .
20 . Sep als 2 (- 5) ; leaves st ipula te ; carpels 2-5 . . . 19 . Begoniac eae
20. Sepals 3- 9; leaves exstipul ate ; car pels 3- 8 .... 20 . Datisca ceae
19 . S tamens 1-5, typically 3 with one unilocular a nd two bilocular
anthers; mostly tenclriliferous vines; s tyle most ly s ingle ;
corolla mostly sympetalous . ... .... .... . . .. .. ... 21 . Cucurbitaceae

7. SALICALES

Repr esented by . ... .. • ..... ... .. ................ ... .... .... ...... . 1 . Salicaceae

8. CAPP ARALES

1. Placentation axile or seemingly SO ; sepal s, petals , and stamens 8 each ;

ovar y closed, with 5- 8 carpels ; endosper m present; coarse herbs o r
sof t s hrubs with trifoliate l eaves ... . ..... ... ..... .... ... l. Tovariaceae
1. Placen tation nearly a lwa ys par ieta l (some t imes with intruded placentae) or
seemin gl y so; sepa l s, pet al s, and carpe l s usually fewer; seeds wit h
l i ttle or no endosperm.
2 . Flowers hypogynous .
3 . Flowe r s mostly r egula r or on l y s li ght l y irregular; s epa l s most l y
4 ; carpels mos tl y 2 (or seemingly so); style s hort or e longa te ,
with cap i tate or sligh tly l obed s tigma .
4. Ovary most l y I- ce lle d; stamens 4-many , never tetradynamous
as in the next family ; flowers almost always \"ith an evi-
dent gynophore or androgynopbore ; pollen binucleate ; leaves
simple to trifolio la te or palmately compound, not much di s-
sected; herbs , sh rub s , or trees, chief l y tropical and sub-
tropical .. ... ... ... ..... .. ..... ... ....... . . ... 2. Capparacea e
4. Ovary mos tly 2-cell ed , with the ovules attached to th e margi ns
of the partition; stamens typically 6 and tetradynamous (the
outer ones shorter than the 4 inner ones) , seldom fe\"e r;
flm"e rs rarely with a gynophore o r androgynophore ; pollen
trinucleate; l eaves often more or less dissected i n pinnate
fas hion, but without distinct , articulated leaflets; herbs ,
se l dom shrubs, chiefly of temperate regions ... . 3. Cruciferae
3 . Flowers us ual l y evi dentl y i rregular ; sepals (4) 5-6 (-8 ) ; carpels
2-6; st i gmas sessile , sepa r ate , as many as the carp els ; ovary
of ten open at the top ; he r bs ..... .. ..... .. .. . . . .. . . 4 . Resedaceae
2. Flowers peri gyno us; sepa l s , petals a nd f un ctiona l stamens each 5; car
pels 3 ; trees with pinnat ely decompound leaves ... .. ... 5 . Horingacea e

9. ERI CALES

1. Embryo normally deve loped , \" it h 2 co t yledons ; plants mo re or less ~lOody,

always chlo r ophyllous.
2 . Pollen gr ains borne singly; s t amens mostly twice as many as the
peta l s , commonly opening by terminal pores .
3. Ovul es I (-3) per locu le ; petals commonly joined at the base ; seed
without a seed coat ; car pels 2-5 . ... ... .. .. .... . . . 1 . Cyrillaceae
3 . Ovules numero us; petals free ; seed with a see d coat; carpels 3 ....
. . . . . ..... . . ... . .. . . . ... . . . ... . . . . . ...... . . .. ..... 2 . Cl ethraceae
666 29

2. Po l len grains nearly alvays borne in tetrads (in some Epacridaceae

only one grain of the tet rad matures).
4 . Sepals and petals each 4·-7; corolla mostly sympe talous; habit
various .
5 . Stamens mostly twice as many as the corolla lobes; leaves
rarely palmately veined ; antho::!rs mostly opening by termi-
nal pores , of ten appendicula te; \-lidesp read .. ... 3. Ericaceae
5. Stamens mostly as many as the corolla lobes; leaves mostly
palmately veined; antho::!rs opening by longitud inal slits,
not appendaged; chiefly Australian .. ........ 4. Epa cridaceae
4 . Stamens and petals each 1-3 , the petals separate; stamens opening
hy long itud inal sli ts, not appendaged; dwarf, often prostate ,
evergreen shrubs ... .. ............ .... ..... ... .... 5 . Empetraceae
1. Embryo very small and scarcely d ifferentiat e d, \dthout cotyledons ; herbs
or half-shrubs, often Hithout chlorophyll.
6 . Plants usually with green leaves; anthers opening by pores; polle n
usually in tetrads; po::!tals separate ; placentation axile ......... . . .
· .. .... ......... . ... . ........ ... ...... .. ....... . ...... 6 . Pyrolaceae
6 . Plants without chlorophyll , the leaves reduced to mere scales;
anthers opening by longitudinal slits ; pollen grains borne singly ;
petals separate or united; placentation axile or parietal ....... .. .
· .... .. ........ .... .................... . ........... 7 . Nonotropaceae

10 . DIAPENS IALES

Represented by .........•• • . ... ....... . .... . ...... .... .••..... 1 . Diapensiaceae

11 . EBENALES

1. Plants \-l H h a \.,rell developed latex system; pubescence of 2-armed hairs,

one arm sometimes reduced or obsolete; nodes mostly trilacunar ........ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . Sapotaceae
1. Plants \-lithout a latex system; pubescence not o f 2-armed hairs except in
some Ebenaceae; nodes unilacunar .
2 . Flowers mos t1y unisexual; styles sep<\rate a t leas t dis tally ..... . ... .
· .......... .. ............. . ........... ... ......... . ... . 2 . Ebenaceae
2 . Flm-lers perfect; sty le solitary , Hith a capitate or slightly lobed
stigma .
3 . Pubescence of stellat e or peltate hairs; fruit most ly dry; ovary
superior to less often inferior; anthers more or less linear .. .
. . . . . . . . . . . . . . . . . . . . . . • . • . . . . . . . . . . . . . . . . . . . . . • . • 3 • .Styracaceae
3 . Pubescence of simple hairs (or none); fruit more or less fleshy ;
ovary inferio r (somet imes only half inferior) .
4 . Anther s linear; flO\,;ers \,;ith a corona; stamens 8 (tHice as
many as the corol la lobes) . ....... ... ..... 4. Lissocarpaceae
4 . Anthers ovate or rotund; flm-lers without a corona; stamens
usually more than 8 .... ... ............ . ..... 5 . Symplocaceae

12. PRINULAl.,ES

1. Plants woody, mostly arborescent, largely tropical; fruits mostly fleshy,

often I-seeded (o::!ven though there are seve ral or many ovules in the
ovary .
2 . Flowers with staminodes alternate with the corolla-lobes; plants
Hi thout an evident secretory system , the leaves not gland - dotted ...
.............. ... ...... ... ..... . .. . .. . ........ .. . 1 . Theophrastac eae
 2. Flowers Hithout s t aminod es ; plants with a schizogenous secr et ory
sys tern and gland-dott ed leaves .... ... ... ..... ......... 2 . Nyrsinaceae
1. Plants herbaceous or occasionally half-shrubby , chief l y of temperate
regions or altitudes; fruit capsular, wi th (2 - ) many seeds . . . . ......... .
.. . ......... . . . . . . .... . . . . ... . . . . .... . .. ..... . ...... . . . . . . ) . Primul aceae

V. ROSIDAE

1. Plants relatively primitive , the flowers commonly either apocar pous or

Hith the carpels united only tOlvard the base or by their styles , or
....'ith more or less numerous ovules per carpel , or with more or less
numerous stame ns, often witll more than one of these features ; vascular
bundles never Hith internal phloem; plan ts never parasitic ( t ho ugh some-
times insectivorous) . .. . .. . ............................. ...... 1. Rosales
1. Plant s more advanced in one or another respect , the floHers mos tly syncar
pous (or pseudomonomerou s), seldom apocarpous , typically with not more
than twice as many stamens as sepal s or petals (stamens numerou s in many
Nyrtales an d in some others), very often wi t h only I or 2 ovul es per
carpel; vascular bundles sometiflles with internal phloem; plant s auto-
troph ic or somet imes para sitic .
2 . Vascular bundles mo stly with internal (as well as external) phloem;
flow ers strongly perigynous to epigynous , often with numero us sta-
mens and numerous ovules .... .. . . .... .... .... .. . . ... . . . ... 4 . Hvrtales
2. Vascular bund les \dthout internal phloem; f l owers various .
) . Flmlers pseudomonomerous t strongly ~erigynous .•.. • ...• 5 . Prot eales
3 . Flowers evidently \dth more than one carpel, or not strongl y peri
gyn ous, or both .
4 . Pl an t s mostly parasitic or hemiparasitic (wholly autot ro phic
in some Santalal es) .
S . Ovules 1 - 8; plants with or \,'ithout chlorophylL ......... . .
... .. . . . . ... .. . . .. ... . . . .... .. . . ... . . . . . . . . 7 . Santal ales
S . Ovules very numerous; plan ts t~ithout chlorophyll . . . .... .. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 . Raff l esiales
4 . Plants (Hith few exceptions ). autotr ophic .
6 . FlOl~ers of ordinary type , not marked l y reduced except in
a feH small families such as the Ga rryaceae (Cornales) .
7 . Leaves mostly compound or conspicuously lobed or
cleft, less often simp le and unlobed .
8 . Ovary supe rior; pollen variously binucleate or
trinucleate.
9 . Plants mostly woody ...... . ...... 12 . E,apindales
9 . Plants mostly h erbaceous or nearly so ..... ... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 . Geraniales
8 . Ovary inferior ; pol l en trinucleate .. 16 . Umbellales
7 . Leaves simple and entire or merely toothed, only sel-
dom compound .
10. Ovary most l y superior.
11. FIOt~ers regular .
1 2 . Stamens basa l ly connate , usually more than
5 ; disk usually wanting; pollen tri-
nuclea te . ...... . . ........ . . .. 14 . Linales
12 . Stamens fre e from each other, seldom more
than 5 ; d i s k often present .
668 29

13 . Stamens alternate \·l ith the petals, or

the floHers rare l y diplostemonous;
pollen variously binucleate or tri-
nucleate . ..... , . " .. . 9 . Cel as t rales
13 . Stamens opposite the petals; poll en
binucleate ... , . , ...... 11. Rhamnales
11 . Flowers irregular (except in Tremandraceae)"
, , , .. ....... ... , ........ . .. . 15 . Polygalales
10 . Ovary inferior , ........ , .. . ... . .. , . . . . 6. Cornales
6 . Flm.,rers more or less reduced and often un i sexual , the
perianth poorly developed or wanting .
14 . Endosperm wanting ; submerged aquatics . 2 . Podostemales
14 . Endosperm present; habitat various .
15 . Herbs I·lith an inferior ovary and ..Jitho u t milky
juice; flm.,rers not form i ng pseudanthia ........ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . llaloragal es
15 . Trees, shrubs, or herbs, \.;ith superior or naked
ovary and often with milky juice: f l owers often
grouped into pseudanthia ....... 10 , Euphorb i ales

1. ROSALES

1. Seeds mostly \.,rith endosperm (except notably the Davidsoniaceae and a fe\.,r
Grossulariaceae); leaves wi th or Hithout stipul es .
2. Plants more or less distinctly woody, not succulent (herbaceous only
i n some Byblidaceae, marked by the conspicuous , g l andular , insec t -
catching ha i rs); carpels nearly ah.,rays united into a compound pis-
til (some exceptions among the Cunoniaceae) (segregate order
Cunoniales) .
3. Leaves mostly compound (simple in some Cunoni aceae and Eucryphia-
ceae) , stipulate ; flowers hypogynous, with axil e placentation,
or the carpels rarely free.
4 . Leaves opposite or whorled ; seeds with e ndosperm.
5 . Carpels 5-12 (18) , multiovulate; stamens numerous .. ..... .
. . . . . . . . . . . . . , ....... , . ........ . .. . .. . . 1 . Eucryphiaceae
5 . Carpels 2-5, l - many-ovulate ; stamens as many or twi ce as
many as the sepal s , seldom numerous . ... . . 2 . Cunon i aceae
4 . Leaves alternate : seeds without endosper m... 3 . Davidsoniaceae
3. Leaves simple , entire to deeply cleft , exstip ulate , or seldom
with vestigial stipules ; flowers hypogynous to epigynous , with
axile or parietal placentae .
6. Flowers hypogynous or slight l y perigynous ; ovules unitegmic
and , so far as known , tenuinucellate .
7. Stamens 5; co r olla polypetalous to mor e or less sym-
petalous.
8. Leaves \·,rithout insect-catching hairs ; co r olla usually
more or l ess sympetalous ; half- shrubs o r more often
shrubs or trees; anthers opening by elongate s l its
or l ess often by pores .......... .. 4 . Pittosporaceae
8 . Leaves with insect-catching hairs ; corolla strict l y
polypetalous; herbs or hal f-shrubs ; anthers opening
by apical pores or sho r t s l its ...... . 5 . Byblidaceae
7. Stamens 2: corolla sympetalous .......... 6 . Columell i aceae
6 . FloHers usual ly partly or wholly epigynous. seldom hypogy
nous ; ovules diverse.
H 669

9. Leaves opposite; stamens lO-numerous, seldom only 5;

ovules few to many, unitegmic, tenuinucellate . ••. , .. ,' .•
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. Hydrangeaceae
9. Leaves alternate; stamens 4- 6 , sometimes accompanied by as
many staminodes; ovules various.
10 . Petals separate , or sometimes wanting .
11 . Ovules mostly numerous , on axile or parietal pla-
centae, variously unitegmic or bitegmic , crass i-
nucellate or tenuinucellate; leaves normally
developed , the plant not heatherlike .... ...... . .
. .. . . ..... ...... .. ......... .. . 8 . Grossulariaceae
11. Ovules 1- 2 (8) in each cell of a 2-3-celled ovary ,
axile , unitegmic , crassinucellate ; leaves mostly
small, the plant often heatherlike.9. Bruniaceae
10 . Petals connate, the corolla sympetalous .............. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. Alseuosmiaceae
2. Plants either herbaceous, or succulent, or both, without insect-
catching hairs; leaves exstipulate, alternate or seldom opposite;
stamens as many or twice as many as the sepals; ovules mostly biteg-
mic and crassinucellate (segregate order Saxifragales).
12. Plants succulent ; carpels as many as the petals, free or united
only at the base ................................ 11. Crassulaceae
12 . Plants scarcely or not at all succulent; carpels seldom the same
number as the petals.
13. Carpels 6 , f r ee , uniovulate; sepals 6; petals none; stamens
12; some of the leaves modified into pitchers ... ......... .. .
· .......................................... 12 . S;ephalotaceae
13 . Carpels 2-5 , usually more or less united, at least below, and
usually with numerous ovules on parietal placentae; flmolers
mostly 4-5- merous, with 1 or 2 sets of stamens and usually
with petals; leaves not modified into pitchers ........ .. ... .
· .......................................... 13 . Saxifragaceae
1. Seeds mostly without endosperm, except in a few Rosaceae; leaves mostly
st ipulat e .
14. CarpQls I - many , when solitary the plants tdth simple leaves and only 1
or 2 ovules per carpel; stamens free , numerous or less often only 5
or 10 (Rosales, sens. strict . ).
15. Gynoecium of I-many free or less often united carpels, never with
the carpels united only by a common style, nor with a solitary
carpel that has a gynobasic style .
16 . Gynoecium apocarpous, or syncarpous with 2-5 styles; stamens
usually more than 10 ................. .. ......... 14. Rosaceae
16 . Gynoecium syncarpous, with 10 styles; stamens 10 ............. .
· ...... ..... .............. ..... .... . .... .... . 15. Neuradaceae
15. Gynoecium of 2-3 carpels united only by a gynobasic style , or
seemingly of a single carpel with a gynobasic style ............ .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 . Chrysobalanaceae
14. Carpel nearly always solitary, and usually with more than 2 ovules;
leaves usually compound; stamens (5) 10-many , often connate by the
filaments (segregate order Fabales) ............ .... .. 17 . Leguminosae

2. PODOSTEMALES

Represented by .. ... . .. . ...... ... ................. ............ . 1 . Podostemaceae

 670 29

J. IIALORAGALES

1. Ovule apical ; embryo straight ; stamens 1-8 .

2 . Styles 2-4; ovule bitegmic ; stamens (1) 2-8.
3 . Ovary 2-4-locular; terrestr i al or aquatic herbs \.,ith small , al-
ternate or more often oppos i te or Hhorled, exstipulate leaves
and small inflorescences; f ruit not drupaceous; petals present
or absent .. ... .. ....... . ... . .......... . .... . .... 1 . Haloragaceae
3. Ovary unilocular; te r restrial herbs with large , alternate , stipu
late leaves and large inflorescences; f ruit drupaceous ; petals
none ... .... ............. . ........... .. .. ... ...... 2. Gunneraceae
2. Styl e 1; ovule unitegmic ; stamen 1 ; emergent aquat i cs ; leaves
whorled , exstipu1ate · ... ........................... 3 . Hippuridaceae
1. Ovul e basal, unitegmic; emb r yo st r ongly curved; stamens 6-28; terrestrial
herbs with stipulate leaves that are oppos i te below a nd alternate
above ; ovary unilocular, Hith a single style ....... . ... 4 . The ligonaceae

4. HYRTALES

1. Ovary superior .
2. Plants woody or less often herbaceous , most l y terrestrial, Hhen
aquat i c not free -floating.
3. Ovules 2-many per carpel ; ovary 2-severa1-10cular ; petal s present
or absent ; fruit capsular .
4 . Stamens numerous (seldom only 12) ; carpels 4-20; ovu les
numerous ... .... ... ...... ....... ..... ...... 1 . Sonneratiaceae
4 . Stamens 4-8 (12 in some Lythraceae ).
5 . Flowers perf ect; petals present or absent .
6 . Filaments elongate ; carpels 2-3, seldom more numer-
ous , each with 2-many ovules ; mostly dip l ostemo-
no us . . ... ... ..... .. ........... . ... . . . . 2 . Lythraceae
6 . Filaments very short : carpels 4 , each Hith 2 4
ovules; hap los temonous ...... ... ....... 3 . Penaeaceae
5 . Plant s polygamo - dioecious; petals none ; carpels 2. multi
ovulate .............................. 4 . Crypteroniaceae
3. Ovule solitary in a pseudomonomerous pistil , or the ovules soli
tary in each of 2 or more l ocules ; petals scale- like or none;
fr ui t seldom capsular ; mostly diplostemonous . . . 5. Thymel aeac eae
2 . Plants free - floating annual aquat i cs ; ovary bilocul ar, with a single
pendulous ovule in each locule, only one l ocu1e maturing; haploste-
monou s .. .. . ...... ...... ..... ....... . ....... .. ...... . ... 6 . Trapaceae
1. Ovary in most spe cies infe rior.
7. Placentation axile, seldom parietal; fru it seldom I-seeded and inde-
hiscent.
8 . Stamens numerous .
9 . Carpels 2-3 , seldom more ; leaves glandula r -punctate (incon-
spicuously so in Dialypetalanthaceae) .
10. Fruit a many-seeded capsule ; anther s opening by terminal
po res; no intern al phloem ........ 7 . Dial ypetalanthaceae
10 . Fruit few- seeded (even when the ovary has many ovules), a
berry , drupe, or caps ule; anthers opening by slits or
less often by termi nal pores; mostly with internal
phl oem ..... ... .... ..... . . ... ...... ... .. . ... 8. Nyrtaceae
9 . Carpels more or less numerous, commonly about 9; leaves not
glandular-punctate ; fruit a many-seeded berry . 9 . Punicaceae
29 671

8. Stamen s up to t\.,rice as many as the pe t als (or sepals).

1 1. Anthers opening by longitu dinal slits; c onn ective no t much
modified; l eaves mostly without subparallel longitudinal
ribs .
12 .
Embryo sac 4- nucleate; endosperm dip loid ; most species
herbaceous; ovules usually many ; fr uit usually capsu-
lar , sometimes a berry or drupe . .......... l0 . Onag r aceae
12. Embryo sac 8-nuc leate; endosperm triploid ; plant s woody ;
ovules 2-3 per carpel; fr ui t drupaceous ... 11. O!iniaceae
11 . Anth ers mostly opening by terminal pores; connective variously
mod i fied; leaves mostly \.;rith prominent subpar allel longi-
t udin al ribs ................... . ...... ·· . 12 . Helastomataceae
7. Placentation apical i n a I-celled ovary ; ovules 2 (-6); fruit I-seeded,
indehiscent ......................................... 13. Combretaceae

5. PROTEALES

1. Plants with a pubescence of peltate scales; flowers not aggregated into

Compositae-l ike bracteate heads; f ru it a pseudodrupe , the dry achene
surrounded by the persistent fleshy hypanthium or calyx-tube ; nodes
unilac un ar . .. ............... . ..... .. ....... ...... . .... ... 1 . Elaeagnaceae
1. Plants \·.rithout peltate scales ; flowers often aggregated into Compositae-
like bracteate heads ; fruit diverse, but not as in the Elaeagnaceae;
nodes trilacunar .... . ....... . ..... . .. ···· ............ · · · .•. 2. Proteaceae

6. CORNALES
1. Flowers perfect or unisexua l, \"ith more or less well developed sepals or
petals or both, not borne in catkins.
2 . Ovules 2 (seldom more) pe r locule; petal s folded or convolute in bud;
leaves opposite , stipulate; stamens more numerous t han the petals;
fruit a berry ...... . ....... ...... .......... ........ 1 . Rhizophoraceae
2 . Ovules solitary in each 10cule (or some 10cules empty); petals valvate
or imbricate; l eaves opposite or alternate , exstipu1ate ; fruit
mostly drupaceous.
3. Stamen s more numerous than the petals (or sepals) excep t i n some
Alangiaceae ; leaves strictly a l ternate; embryo large.
4 . Ovary 6- 9-locular .......................... · .. · · 2. Davidiaceae
4 . Ovary l - 2- locular .
S . Petals imbricate ; flowers perfect o r often unisexua l;
ovary unilocular .... . .............. ·.· ...... 3. Ny ssaceae
5 . Petals valvate; flowers pe r fect ; ovary l-2-lo cular .. .. ... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 . Alangiaceae
3 . Stamens as many as the petals; leaves opposite , seldom alternate ;
embryo sma l l ............................. ·· .. ... ···· 5 . Cornaceae
1. Flowers unisexual (p l ants dioecious) , the pisti llate ones \"ith vestigial
or no perianth, th e staminate ones with a perianth of 4 sepals , both
types borne in catkins ....................... ··.· ..... ··· · . 6 . Gar r yaceae

7. SANTALALES

1. Fruit capsular; ovules 6- 8 ; leaves with small stipules ; pl ants a uto-

t rophic .
2. Fe rtile stamens opposite the petals, alternating with well developed
staminodes ; petals and sepals well di fferent iated; f lowers in
slender , catkin-like racemes .... . ................. . !. Nedusandraceae
 672 29

2. Fert i le stamens alternate with the petals, alternating with a set of

nectary glands; petals and sepals very much a l ike; flowers in glo -
bose umbels ..................................... 2 . Dipentodontaceae
1. Fruit indehiscent; ovules 1-5; leaves without stipules ; plants autotro
phic or more often partly or lv-holly parasitic .
3 . Plants with chlorophyll; flowers perfect or so metimes unisexual ;
fruit dry or fleshy .
4. Ovules mostly well differentiated; leaves alternate except in the
Grubbiaceae and most Santalaceae; flmv-ers mostly small and not
very showy; plants rooted in the ground .
5 . Petals present; stamens , when i n a single cycle, opposite the
petals; ovary superior to occasionally i nfe rior.
6. Ova r y 2-5-celled below, with 1 ovule per locule; integu-
ments 0-2 .................................. 3 . Olacaceae
6 . Ovary I - celled, with I apical or basal ovule ; integument
none ...................................... 4. Opiliaceae
5 . Petals absent; stamens , when in a single cycle , opposite the
sepals; ovary inferior .
7. Plants fully autotrophic; stamens 8 , in 2 cycles ; ovules
unitegmic .....................•.......... 5. Grubbiaceae
7. Plants partly or wholly parasitic; stamens 3-6, in a sin
gle cycle ; ovules without integument ..... 6. Santal aceae
4. Ovules scarcely different i ated f rom t he large , centra l placenta
of the I - celled ovary, without integument; leaves opposite or
whorled; perianth often large and shm.:y; plants rarely rooted
in the ground ................................... 7. Loranthaceae
3 . Plants without chlorophyll; flowe rs mostly unisexual ; f ruit dry, nut
l i ke .
8. Plants aerial, attached to the s tem of the host; fruit 3- winged;
ovules without integument .. . .................. 8 . Misodendr aceae
8. Plants terrest rial, attached to the roots of the host ; fruit not
winged.
9 . Ovary superior; ovules 1-5 , without integument ; stamens
usually more than one ................... 9 . Balanophoraceae
9 . Ovary inferior; ovul e solitary, unitegmic ; stamen 1 ......... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 . Cynomoriaceae

8. RAFFLESIALES

1. Ovary superior; scale-leaves opposite ; flowers perfect ............... . .. .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . Mitrastemonaceae
1. Ovary inferior or half -i nfe rior; scale-leaves alternate or none .
2. Flowers mos tly perfect; leaves none .................... 2. Hydnoraceae
2 . Flowers unisexual; scale-leaves alternat e ......... . .. 3. Rafflesiaceae

9. CELASTRALES

1. FlO\v-ers diplostemonous ; petals none; disk none; hab it ericoid ........... .

. ....... .. .. ....... . ...... . . ...... .... .. ........... . 1. Geissolomataceae
1. Flowers haplostemonous (except a few Celastraceae) ; petals generally well
developed; disk often present; habit not ericoid .
2 . Ovules erect and basal, or several and superposed in 2 r ows ; disk
present exc ept in Salvadoraceae; integuments 2 except in many Sal-
vadoraceae; leaves opposite or less often alternate .
29 673

3. Flowers \.,tith a disk ; carpels more than 2 except in some Celastra-

ceae; flowers pentamerous except in some Celastraceae .
4 . I~oody plants; flm"e rs hypogynous ; leaves opposi te or a1 ter-
nate.
5 . Disk extrastaminal; stamens mostly 3 (2-5); leaves mostly
opposite; seeds without endosperm ; l atex system \.,tell
developed .... .. ........ . .............. 2 . Hippocrateaceae
5. Disk intrastaminal , or the stamens seated on the disk;
stamens 4-5 (seldom 6-10); seeds mostly with endosperm;
latex system seldom \.,tell developed.
6. Disk of ordinary type, merely surrounding the ovary
at the base; leaves opposite or sometimes alter-
nate; locules 1-5 .............. ····· . 3 . Celastraceae
6 . Disk much enlarged, enclosing the ovary; leaves alter-
nate; loc ules 5 ............... ·· .4. Siphonodontaceae
4 . Herbs; flm.,ters distinctly perigynous, the disk lining the
hypanthium; l eaves alternate; seeds \.,t ith endosperm ......... .
. . ....... . ....... ...... .. ....... . . . ....... 5. Stackhousiaceae
3 . Flowers without a disk , but the stamens s ometimes alternating with
nectariferous glands; carpels 2; flm.,ters tetramerous ; seeds
Idthout endosperm ..................... ···· ...... 6. Salvadoraceae
2. Ovules pendulous, apical or nearly so, 1 - 2 per locu l e, unitegmic
(except in Dichapetalaceae); leaves alternate .
7. Stipules \.,tanti ng or vestigial; seeds with endosperm except in some
Icacinaceae; disk wanting.
8 . Corolla mostly polypetalous; fruit drupaceous ; woody plants
without milky juice.
9 . Locules 4-6 or more; petals imbr ic ate; pollen binucleate ..
. . ....... . ...... .. .. ... . . . ......... ..... 7. Aquifoliaceae
9. Locules 3, or more often 1 by abortion; petals valvate;
pollen trinucleate ... ............... . · .... 8 . Icacinaceae
8 . Corolla sympetalous; fruit samaroid; climbing herbs with milky
juice ................................... 9 . Cardiopteridaceae
7. Stipules present; endosperm wanting; disk present and intrastami
nal, or rep r esented by nectariferous glands alternating with the
stamens ...................................... 10 . Dichapetalaceae

10 . EUPHORBIALES

1. Fruit mostly dehiscent and with more than 1 s eed , seldom drupaceous; seeds
usually with a caruncle.
2. Ovules apotropous, the raphe dorsal; fruit a 10Clllicidal capsule , or
sometimes drupaceous; sma l l trees and shrubs, \dth neither stipules
nor a milky juice; disk wanting .......................... !. Buxaceae
2 . Ovules epitropous, the raphe ventral; fr u it mostly schizocarpic, each
carpel eventually dehiscing ventrally to r eleas e its solitary seed;
plants of very diverse habit, often with a milky juice. and usually
\d th s tipules ; disk of ten present ...... . ............ 2 . Euphorbiaceae
1. Fruit indehiscent, drupaceous, often I-seeded; seeds mostly without a
caruncle.
3 . Ovules 2 in each lo cule , or 1 locule empty; leaves \.,tithout stipules.
4. Ovules epitropous , present in each of the 2 (-4) locules ; petals
none; stamens 6-12; di sk \.;ranting ; embryo minute .. 3 . Daphniphyllaceae
674 29

4. Ovules apotropous, borne in only 1 of the 2 locules; petals 5 ;

stamens 5, alternating with the disk-glands ; emhryo not espe-
cia l ly smalL . ....... ... ... . . . ........ .. ..... .. 4 . Aextoxicaceae
3. Ovules solitary in each of the 2-5 locules ; leaves wit}l stipules;
petals 5 ; stamens 5 , 10, or 15; disk none .. ... ...... .. . 5 . Pandaceae

11 . RHA}!NALES

1. Ovule solitary in each locule; carpels mostly 3-8, seldom only 2; shrubs
or small trees, sometimes climbing, but seldom with tendrils .
2 . Stamens free; leave s simple, entire or merely toothed, usually sti-
pulate; embryo large; flm.;ers hypogynous to pGrigynous or epi-
gynous; fruits diverse. ····· ... ....... . ..... .... ...... 1. Rhamnaceae
2. Stamens with the f ilame nts un ited into a tube ; leaves simple to com
pound, exstipulate; e mbryo small; flol.;ers hypogynous; fruit a berry
. ..... ..................... . . .. ....... .. . . . . ..... .. .. . .. 2. Leeaceae
1. Ovules 2 in each locu1e: carpels 2; mostly tendril- bearing climbers;
leaves stipulate, usually compound or lobed ; fru it a berry . . 3 . Vit aceae

12 . SAPINDALES

1. Ovules usually more than 2 per locule or carpel; endosperm I.;e ll developed;
ovules apotropous; flowers usually perfect .
2 . Leaves stipulate, most ly pinnately compound; f lower s haplostemonous,
I.;ith 2-4 carpels and axile placentation.
3. Leaves mostly opposite ; disk annular, intrastaminal, or the sta -
mens seated on the disk; fl owers regular; carpels 2-3, seldom
4 ; pollen binuclea te ....... . .... . .. .. ...... . .. . 1. Staphyleaceae
3 . Leaves alternate; disk unilateral , extrastaminal; flOl.;ers irregu-
lar ; carpels mostly 4: pollen trinucleate .. .... 2 . Nelianthaceae
2 . Leaves exstipulate, simple , alternate; flowers diplost emonous , I.;ith
mostly 5 carpels, the placentation parietal; disk annular, extra-
staminal ...... . ......... ... .... .. . . ............... . . .. 3 . Greyiaceae
1. Ovules seldom more than 2 per carpel or locule ; endosperm in most famil
ies wanting or scanty; ovules variously apotropous or epitropous;
f l olJers perfect or unisexual.
4. Flowers strictly apocarpous .
5 . Style terminal; fruit dehiscent; seeds arillate; leaves mostly
compound ...... . . ... .... . . . .. . ..... .. .. . .......... 4 . Connaraceae
5 . Style basal; f ruit indeh iscent; seeds exarillate: leaves simple ..
........ . ... .. . . ..... . . . ......... . .... .. ... . . . 5. Stylobasiaceae
4 . Flm.;ers nearly always more or less strongly syncarpous (some excep
tions in Simaroubaceae) .
6 . Disk mostly extrastaminal (and often unilateral) or I.;anting
(intrastaminal in some Aceraceae); ovules apotropous except in
Akaniaceae; flowers very often irregular .
7. Leaves mostly alternate; stamens typ ica lly 8 (5-many) .
8. Seeds with endosperm and without an aril; IJood rays very
wide; disk Hanting; ovules epitropous , 2 in each locule
... . . .. ..... . . . .......... . . . ...... .. ...... 6. Akaniaceae
8 . Seeds \Jithout endosperm and cormnonly Hith an aril; ~.;ood
rays narrow; disk present, often unilateral; ovules
apotropous , 1 or less often 2 in each locule .. . ....... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. Sapindaceae
7 . Leaves opposite .
 9. Leaves palmately compound; fruit a tricarpellate , usuall y
I-seeded capsule ; flowers irregular ; stamens 5-8 .... .. . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 . Hippocas tan a c eae
9 . Leaves simple and palmately lobed , or pinnately comp ound ;
fruit a bica r pellate , 2-seeded samara ; flowers reg ular ,
stamens 4-10, most often 8 ..... . ............ 9 . Aceraceae
6 . Disk intrastaminal , annular or often developed in t o a gynoph o r e
(\.mnting in the Julianaceae) ; ovules epitropous except i n t he
Anacardiaceae and Julianaceae .
10 . Bark and h'ood with prominent , specialized resin duct s .
11. Ovary 2-5 celled, \,rith 2 (seldom only 1) epitrop o us ovules
per locule .. .. .... . .. ....... . .......... · .10 . Burseraceae
11 . Ovary I-celled (by abortion), with a single apo tropo us
ovule .
12 . FlOl'Jers normally developed , perfect or sometimes un i -
sexual , with an evident perianth ; intrastaminal disk
rresent . .. ... . .. . .... . .. . .. . . . .. . .. 1 1 . Anacardiac e ae
12 . FIOl>,ers reduced, unisexual , the pistillat e on es wi th-
out a perianth; disk none .. . ....... .. 1 2 . Julianaceae
10 . Bark and wood \,rithout prominent resin ducts .
13 . Leaves mostly exstipulate, the stipules, if any , small a nd
soon deciduous ; leaves mostly alternate .
14 . Stamens mostly free .
15 . Leaves not glandular-punctate .
16 . Seeds Hithout endosperm; leaves mostly com-
pound, seldom simple; bark b i tter ; sty l e s
often separate; floHers often S-merous ;
pollen binucleate .. . . . ... .. 1 3 . Simaroub ac eae
16 . Seeds \.;ith endosperm; l eaves simpl e; bark no t
bitter. (?) ; styles united ; flOl,rers 3-mero us
or seldom 4-merous; pollen t r inucleate ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 . Cn eoraceae
15 . Leaves mostly p;landular-punctate .... .. 15 . Ru taceae
14. Stamens mostly connate by their filaments ........ .... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16 . Meliac ea e
13 . Leaves \~i th well developed, persis ten t s tipules , mo stly
opposite ; seeds of ten \>lith endosperm ... 17 . zy g ophyllaceae

13 . GERAlnALES

L FlOl·.'ers regular or slightly irregular , not spurred, or the spur inconsp i-

cuou s and adn ate to the pedicel; stamens 2 or 3 times as many as the
sepals or petals, sometimes sO'r:Je of them staminodial; leaves mos tly
deeply cleft .
2 . Style not gynobasic ; integuments 2; ovules, Hhen f e\,r , pendu lous ;
filaments basally connate ; annual or more often pe r ennial herbs ,
o r seldom \,rQody plants .
3 . Fru i t a beakless , loculicidal capsule or seldom a berry, Hith (1)
2-many seeds per carpel ; styles free; endosperm wel l deve l oped ;
pollen binuc l eate ; leaves \,rith or more often Hithout st i p u les .. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Oxalidaceae
J . Fruit a beaked, mostly septicidally dehiscent capsule or a schizo
carp , the mericarps I-seeded and mostly again d e hiscent ; styles
usually united, at least belm>' , into a prominent column ; end o-
sperm mostly scanty or none; pollen trinucleate ; leaves Hith
stipules ... ....... . ... . ... . . . . . . . .. . ..... ... .. .... 2 . Ger aniaceae
 676 29

2. Style gynobas i c : i ntegumen t I: ovules 1 per carpel, erect: filaments

f ree; annual herbs; seeds without endosp erm... . . . .. 3. Limnan thaceae
1. F lO\~e r s st rongly irregular, one of the sepals with a conspicuous f ree
spur; stamens 5 or 8 , les s than twice a s many as the basic (unmodified)
numb er of sepal s or petals: leaves s imple , not deeply lobed .
4 . Leaves pe ltate , palmately nerved; stamens 8 , carpel s 3 , each uniovu-
late , the fruit a schizocarp; plants with my rosin and without
raphides . .... .... .... . . . . . . . .... ... .... .. ... ... .... 4. Trop aeolaceae
4. Leaves not peltate, the venation pi nnate ; stamens 5; carpels 5 , each
with several or many ovu l es ; fru it an e l astically dehi scent cap-
s ule, or se l dom a berry ; plant s with raphides a nd without myrosin . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 . Balsaminaceae

14 . LINALES

1. Plants woo dy, often arborescen t; fruit drupaceous .

2 . Peta ls exappendi culate ; carpels 4-7 , commonly 5; flowe rs with an
intrastaminal disk ; leaves withou t stipules ........ . . 1. Humiriaceae
2. Petals internally appendicu l ate ; carpels 3. 1 fertile: flowers with-
out a disk; leaves wi th s tipules . ... ... .. .... . ... 2 . Erythroxy l aceae
1. Plant s he rba ceous or occasionally shrubby , sel dom arborescent; fruit
mostly ca psular , seldom drupaceous or nutlike ; petal s exappendiculate;
leaves mostly stipulate: flo ral disk usually wanting .. . . . . .. 3. Linaceae

15 . POLYGALALES

1. Carpels mostly 3, seldom 2-5; st ame ns mostly dehiscent by longitudinal

slits. seldom by terminal po res; l eaves us ually st ipulate.
'.
2 . Fertile stamens several; filaments usually connate toward the ba se .
3. Petals 5 , commonly fringed or toothed; f r uit ind ehiscent; ovules
1 per ca rpel ; stamens 10 , some ste rile .... .... . 1. Malpighiaceae
3 . Petals 3 , entire; frui t capsular: ovules I-many per carpel; sta-
mens 3-12 . . ... . . . . ... .. .... . . . .. . .. ... ... ..... . . 2. Trigoniaceae
2 . Fertile stamen 1. but staminodes also usua l ly presen t; fi l aments
free ; fruit us ually caps ular; ovule s I -many per carpel . .. . . . ..... . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . Vochysiaceae
1. Carpels 2 , or seemingly only 1; st~mens dehiscent by termina l pores;
leaves usual ly exstipulate.
4 . Pistil more or l ess evidently bicarpellate . but not always bilocular;
seeds with endosperm; stamens 8-10 .
5. Stamens f ree .
6. Flowers regular ; placentation axile in a bilo cular ovary ;
heatherlike shrubs or s ubshrubs with small leaves ........ . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Tremandraceae
6 . Flowers irregular; placentation ap i cal or parietal in a uni-
locular ovary which may show traces of a partition; small
trees with normal leaves ... . .. .... .. . .. . 5 . Xanthophyllaceae
S . St amens mostly monadelphous ; flowers i r r egular; ovary evid en tly
bilocular ; herbs . vines, or shrubs ....... . ..... . 6. Polygal aceae
4. Pistil pseudomonomerous and with a pair of api cal ovules ; seeds with -
out endosperm; s tamens 4 or seldom 3; low shrubs or seldom herbs ...
......... . . .. .... . . . . . . .. . . . ... . . .. .... . .. . ...... . . . 7. Krameriaceae
29 677

16 . UMBELLALES
1. Carpel s I -many, typicall y 5; f ruit f l eshy or dry, usually a berry, the
carpels somet i mes separating, but without a carpophore; trees and
shrubs . sel dom herbs o r lianas . ... .......... .. ... . . .. ...... 1. Araliaceae
1. Carpels consis t ently 2; fru it a dry schizocarp, the me r icarps s eparated by
a usua lly persisten t carpophore: herbs , rarely shr ubs or trees ... .. . . .. .
................ . ............................ . . .. ... .. .. . 2. Umbelliferae

VI. ASTERIDAE
1. Ovary chiefly s uperior.
2 . Plant s usually \"ith well developed internal phloem: leaves opposite ;
endosperm nearly always nuc l ear; no i ntegumentary tapetum; corolla
regular. its lobes usually con tor ted in bud . ... ... , . .. 1. Gentianal es
2. Pl ants usually either wi tho u t i nt e r nal phloem, or with al ternate
l eaves , or both; endosperm nuclea r or more often cellular; integ u-
mentary tapetum often presen t; c orolla regula r or irregular, \"it h
various so rts of aestivation.
3 . Fruit typically of 4 or more or less distinct (or separating) nut-
lets, derived from a bicarpellate ovary that often has a gyno-
bas i c style . . . . ...................................... 3. Lamiale s
3 . Fruit othen..r!se ; style rarely gynobasic.
4 . Corolla scarious , persistent; fl ower s mostly anemophilous,
te tramerous as to calyx, co rolla, and androeci um; leaves
usua lly all basal ........................... 4. Plantaginales
4 . Corolla otherwi se ; flow ers most l y entomophil o us or ornithophi-
lous, variously pentamerous o r t et r aroerous, wi t h i somero us
o r anisome r ous stamens ; l eaves seldom all basal.
5 . Flowers mos tly regular and with as many f unctional sta-
mens as corolla lobes, typically 5 of each .... .. ....... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . Polemonial es
5 . Flowers mostly irregula r and with fe\o,'er func tional stamens
than corol la l obes , o r sometimes with regular, tetra-
me r o us cor oll a and 2 o r 4 stamen s, or the co rolla rarely
wan t ing ........ ... .................. . . 5 . Scrophulariales
1. Ova ry chiefly i nferio r.
6. Fl owers borne in various sor t s of inflorescences, bu t if bo rne in
involucr ate heads then the ovules either pendulous o r mo re than one;
e ndospe rm present or absent .
7. Leaves nearly always alternate; stamens free from the corolla, or
at t ached at the base of the tube ; anthers i n most families con-
nivent or connate around th e style , which pus hes out the poll en;
plants nearly alway s herbaceous .. .. .......... . ... 6 . Campanu lal es
7 . Le aves usually opposite or whorled (notable except ion: Calycera-
cea e ); s tamens attached to the co r o l la tube, gene rally well
above t he base; flowers mos tly witho ut a specialized pollen
presentation mechan ism (notable exceptions: Ca l yce raceae, some
Rubiaceae); plan ts woody or herbaceous .
8 . St ipules usually presen t and i n te rpet iolar (sometimes reduc ed
to a mer e interpetio lar line , or enl arged into l eave s);
co rolla typically regular and with i somerous stamens; endo-
sp erm nuclear; plants chie fly tropical and woody, but some
members herbaceo us , or of temperate regions , or both .. ... .. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7. Rubia l es
678 29

8. Stipules typically none, ...,hen present usually smal l and ad-

nate to the petiole; corolla regular or often irregular,
the stamens often fewer than the corolla lobes ; endosperm
cel l ul ar; plants woody or often herbaceous , of tempe r ate
or less of ten tropical regions ................ 8 . Dipsacales
6. Flowers borne in invo lucrate, centripetally f lOlolering heads; ovary
I-celled, \.,rith a solitary , erect ovule; anthers connate or conni-
vent into a tube around the style, Hhich pushes out the po l len ;
endosperm ,,,ranting from mature seeds ..... . ... .......... . 9 . Asterales

1. GENTIANALES

1. Plants without a latex system ; s ty le not espe cially thickened and modi-
fie d dis t ally; carpels fu lly united, only the stigmas sometimes sepa-
rate .
2 . Plan ts mostly Hoody and with more or less vlel l developed stipules,
often containing alkaloids , but l acking gen tiopicrin ; placentat i on
a xile ............ .... ..... .. .. ...... ......... ...... .. 1 . Loganiaceae
2. Plants mostly herbaceous, without stipules, containing the bitter
glucoside gentiopicrin but lacking alkaloids; pl acentation mostly
parietal ....... . ..... .... ........ .... ........ ....... 2. Gentianaceae
1. Plants {."ith a well developed l atex system ; style usually more or less
thickened and modi f ied at the tip; carpels typically unit ed only by the
style and/or stigma, but sometimes wholly united ; stipules none ; a lka -
loids often present, but gentiopicrin lacking .
3 . Androecium free f rom the stigma and without translators ; pollen gran-
ular, not forming pollinia; carpels often united by part or all o f
the style as well as by the stigma , or even wholly united; corona
mostly poorly developed or want ing; trees , shrubs , vines , or less
often herbs ... . ............ .... . .. ...... ............. 3. Apocynaceae
3. Androecium concre scent to the stigma and provided {dth translators ,
the pol len grains bo r ne in dense masses (pollinia) ; carpels united
only by the stigma ; corona more or l ess \."ell developed ; herbs ,
vines , o r shrubs , rarely trees ..... ....... ... ..... 4 . Asclepiadaceae

2. POLEHONIALES

1. Plants with in ternal phloem , and always autotrophic .

2. Carpels 5 ; ovar y with a terminal style and ripen i ng into a schizo-
carp , or with a gynobasic style and ripening into 5 or more 1 - to
several-seeded nutlets; en dosperm cellula r; more or less succulent
herbs and shrubs , confined to the Pacific slope of S . Am •••••••••••
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . Nolanaceae
2 . Carpels 2 (-5) ; fruit not schizocarpic and style never gynobasic ;
plants {"idespread, seldom succulent .
3 . Ovules and seeds mostly numerous, on axile placentae; cotyledons
not pl icate; style barely or scarcely lobed; endosperm mostly
cel lular ............................. . ............ 2. Solanaceae
3 . Ovules 2 (rarely many) per carpel , basa l, erect , the partition
present or absent; cotyledons plicate ; style entire or often
cleft , or the styl es sometimes \"holly distinct; endospe r m
n uclear ....................... . ............... 3 . Convolvulaceae
1. Pl ants without internal phl oem, and e ither autotrophic o r pa r asitic .
4. Plants parasitic , \."ithout chlorophyll; embryo scarcely differentiated .
H 6n

5. Carp els 2; fruit capsular ; twining stem-parasit es , not rooted in

the ground at maturity; endosperm nuclear; pollen trinucleate ...
. ..... . . . ....... . . ... . ..... ... .. . ....... . . .... . ... 4. Cuscutaceae
5 . Carpels 6-14; fruit of l-seeded nut lets ; erect root- paras i tes;
endosperm probably ce llular ; pollen binucl eate ..... 8 . Lennoaceae
4. Pl ants autotrophic; embryo normally deve loped.
6. Corolla lobes valvate or induplicate-valvate in bud; plants mor e
or less aquatic; placenta tion parietal; endosperm cellular . ... . .
... . . . ..... . . . ..... . . ............ ....... ... .. .. . 5 . Henyanthaceae
6. Corolla lobes convolute or imbricate in bud; plants terrestrial ,
though sometimes of wet places.
7. Carpe l s 3 (2-4); corolla l obes convolute in bud ; placentation
axile; endo sperm nuclear .......... .... .... .. 6. Polemoniaceae
7 . Carpels 2; corolla lobes imbrica t e or rarely convolu te i n bud;
placentation parietal (often with i ntruded placentae) or
occas ionally axile ; endosperm nuclear or cellular . ...... ... .
. ....... . .. ..... . ..... . .. .... . . .... . ...... 7. Hydrophyllaceae

3. LANIALES

L Leaves mostly alternate, usually entire; flowers mostly r egular and witt
5 stamens ; sty le typically but not always gynobasic, the fru it typi-
cally but not ah;rays of 4 distinct nutlets; stems not sq uare ....... . ... .
.. . ....... .. .. ..... . . ... ... ..... .. .. ..... .... . .......... . 1 . Boraginaceae
1. Leaves mostly opposite (or whorled) , and often toothed or clef t; flowers
mostly more or less irregular (or apetalous) and \;rith 1-4 stamens , the
exceptions (regular corolla a nd/or 5 stamens) being fo und chiefly in
the Verbenaceae; stems commonly square.
2 . Flowe r s apetalous , uni sexua l; styles 2, separate; stamen solitary;
endospe rm wel l developed; plants mostly aquatic, often free-float -
ing ............ . ..... .. ... . . ..... . . ..... ... .. ..... 2 . Callitrichaceae
2 . Flowers petaliferous, usually pe r fect; style usually soli tary; sta-
mens 2 or mo re; endosperm usual ly scanty or wanting ; mos tly terres-
trial plants, never free-floating aquatics .
3 . Style terminal, t he ovar y on l y slightl y o r not at all lobed;
plants seldom aromatic .
4 . Ovary with 2 (seldom more) usually biovulate carpels, with a
false par tition bet\;reen the ovules of each carpe l, ordi-
nari l y ripening into 4 separating n utlets or into a 4-
stoned drupe . .. . .... . . . . .. ... . .... ..... . . .. ... 3 . Verbenaceae
4. Ovary pseudomonomerous , wi th a single ovule and locule ... . ... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 . Phrymaceae
3. Style mos tly gynobasic, th e ovary more or less deeply 4 cleft and
ripening into 4 nutlets; plants mostly aromati c ...... 5 . Labiatae

4. PLANTAGINALES

Represented by ......•• ... ••. .. ............................... 1. Plantaginaceae

S. SCROPHULARIALES

1. Flowers perfect or less often un isexual, almost a lways with a calyx and
usually also with a corolla; stamens mo stly 2-4 (5); habit and habitat
various .
680 29

2. Placentation axile to parietal or apical; plants not insec tivorous .

3 . Corolla with mostly 4 (5) lobes and only slightly (or not at all)
irregul ar, or sometimes ",anting ; woody pl ants (rarely herbs)
with mostly opposite or whorled leaves,
4 . Stamens 4 , rarely 2 or 5 ; ovules mostly numerous in each
1 0cule ........ ... .. .... ....... ...... ........ 1. Buddlejaceae
4 . Stamens 2, rarely 5; ovules mostly 2 (seldom 4-many) in each
locule .. ... . ...... ....... , ...... . ... .. . , ....... . 2 . Oleaceae
3. Corolla with mostly 5 (seldom 4) lobes , or 2-lipped I"ith scarcely
lobed lips ; habit and leaves various .
5 . Mature seeds usually Idth a well developed endosperm .
6 . Placentation mostly axile or apical .
7. Ovu les 2 or more in each locule; anthers (1) 2-
locular .
8. Fruit dry and most l y dehiscent , rarely f leshy ;
herbs , rare l y shrubs; ovules mostly more or
less numerous i n each l ocule, seldom only 2 ;
leaves opposite or al te rnate . . . .. ..... ... . .. , . .
.. ... . . . ... . , ............... 3 . Scrophulariaceae
8. Fruit drupaceous, indehiscent; woody plants ,
sometime s arborescent; ovules 2 (4 - 8) i n e ach
locule ; leaves mos tly al ternate .. 4 . .t!..voporaceae
7 . Ovules mostly solitary in each locule , somet ime s on ly
one locule developed ; anthers uni l ocu l ar at least
at maturity; herbs or shrubs , mo sr: ly with alternate
leaves; fruit small, indch i scent, the carpel s some-
times separating ......... .. . , ..... 5 . Globulariacea e
6 . Placentation mostly parietal .
9. Plants chlorophyllous , not parasitic , with \"ell
developed, mostly opposite or Hhor l ed leaves ; ovary
often more or less inferior; embryo well developed .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6 . Gesneriaceae
9 . Plan ts parasitic, \"ithout chlorophyll, the leaves
reduced and alternate ; ovary superior; embryo
mi nute, scarcely different i ated .... 7 . Orobanchaceae
5 . Hature seeds mostly with little or no endosperm ; leaves
chie f ly opposite or whorled.
10. Trees or large shrubs, seldom herbs ; leaves mostly com-
po und, seeds usually \~inged ........... .. 8 . Bignoniaceae
10 . Herbs, rarely shrubs or trees ; leaves mostly simple ;
seeds not \.inged .
11 . Seeds with an enlarged and sr>ecialized funiculus ;
plants without specialh:ed mucilage hairs; fruits
fleshy to more often capsular , with axile placen-
tation ......... . ... .... .... .... , . . ... 9 . Acanthaceae
11. Seeds with normal, unspecializ ed f un i culus; plants
\"ith a unique t ype of mucilage hair ; fruits capsu-
lar or drupaceous to more often nutlike , often with
consp i cuous app en dages .... . ......... 10 . Pedaliace a e
2 . Placentati on f ree-cen tr a l; herbs, aquatic or of wet places, mostly
insec tivorous . ..... . .. .... . ... ... .. . .. ... .... ... . . . 11. Lentib ulariaceae
1. Fl owers unisexual, I. it hout periantb; stamen solitar.y ; aquatic herb s . ... . .
. . . . . . , ......... ....... ........... ... .. , . ... ...... . 12 . .livdros tach yacea e
 6. CAMP ANULALES

1. Style without a n indusi um, b u t often with collecting hairs.

2 . Stamens as many as the corolla lobes. typica l ly 5 , free or connate
on ly by thei r a nt he r s ; anthers i n trorse .
3 . Style glabrous , with a solitary s t igma ; plan ts apparen tly without
a latex system .
4. Petals valvate ; fruit a berry ; leaves asymmetric : polle n bi-
n ucleate .. ...... . ......... . ... ... . .. . .. l. Pentaphr agmataceae
4 . Petals imb ri ca t e ; fruit a circumscissle capsu l e ; leaves not
notably asymmetric; pollen trinuc l eate .... . 2. Sphenocleaceae
3 . Style with wel l developed collecting hairs below the usually sepa-
rate stigmas; plants with a 1.o.'ell developed latex system .. . .. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . : ... . . ... .. .... 3. Campanulaceae
2. Stamens 2 or 3 , the filaments adnate t o the style or connate in t o a
tube around i t : a nt hers extrorse: no latex system ... . 4 . Stylidiaceae
1. Style with a more or less cup ulate indusium just beneath the stigma(s),
but without collecting hairs: no latex system .
5 . Flm"ers regular or nearly so , borne in involucrate heads: endosperm
none; ovary superior , unilocular , with a single basal ovul e . ... .. .. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 . Brunon iaceae
5 . Flowers mostly ir r egular , not borne in invo l ucrate heads ; endosperm
,,,ell developed ; ovary generally inferior, 1 - 2 locular, ,.;rith 2 or
more ovu l es ..... .... ... ... .. .... ... . .. . ... ... . .. . . . . . 6 . Goodeniaceae

7. RUBIALES

Represented by . ..... . .. . .. . ..... ... ... ... .... . .. . .. . . ... .• •••. .... 1 . Rubiaceae

8. DIPSACALES

1. Fil aments and anthers distinct; leaves nearly always oP?osite : pollen tri-
nucleate ; inflorescence various .
2 . Plants mostly woody , seldom herbaceous; stamens mostly as many as the
corol la lobes , typica l ly 5, seldom only 2; ovules often more than 1
per locule; endosperm well devel oped ; fruits diverse, often "lith
several o r many ovules per locule ............ . ... . . l . Caprifoliaceae
2 . Plants herbaceous or nearly so; stamens sel dom (except some Dipsaca
ceae) of the same number as the corolla lobes; ovules pendulous ,
not more than 1 per locule .
3 . Stamens 8-10 , twi ce as many as t he c o r o l la lobes , paired in the
sinuses of the c orolla, each ,dth o n ly a single pollen s a c;
fruit a dry drupe 'dth several stones; endosperm well devel oped;
flot.,lers in compact heads . ... .. ..... .. .... .... ..... .. 2. Adoxaceae
3 . Stamens 1-4 (rarely 5), as many as or usually fewer than the
coro l la lobes, each with 2 pollen sacs; fruit dry , I-seeded;
endosperm scanty or none .
4 . Ovaries not individually enclosed or subtended by a gamo-
phyllous i nvolucel; flOl.,lers not i n compact , invo l ucr ate
heads ; ovary mostly trilocula r, with 1 fertile and 2
sterile locules ; corolla mostly 5-10bed . . . . . 3 . Val e rianaceae
4 . Ovaries i ndividually enclosed or sub tended by a gamophyll ous
invol ucel ; flowers mostly born e in compact, invol ucrate
heads; ovary strictly unilocular; corolla mostly 4-5 lobed . .
.... .. ... .. . . .... . .. . ............. . . . .. ... ... . 4 . Q.i psacaceae
682 29

1. Filamen t s and anthers connate i nto a tube around the style; le3.ves alter-
nate ; polle n binucleate; f lO\.Jers In involucrate . centripetally fl o~..'er-
ing heads, with re gular co rolla ........... .. ........... . 5 . Calyceraceae

9. ASTERALES

Represen t ed hy ....... ... ..... ... ................... . ........... . 1 . Compositae

LILIOPSIDA

1. FlO\.Jers mostly apocarpous, the syncarpous forms Idth a mod ified type of
lamina r placentation ; plants mostly aqUatic or subaquatic, al\.Jays her-
baceous; pollen consis tently trinucleate ; endosperm mos tly ~·.'8n t ing , not
st archy I·,hen present; subsidiary cells mostly 2 .•. ....... 1 . Alismatidae
1. FIO\.Jers mostly syncarpous , the apocarpous forms borne on terrestrial ,
woody plants; plants te rrestrial or. less often aquatic ; pollen binu-
c leate or tr i nucleate; endosperm present or absen t, often starchy , sub -
sidiary cells none to several.
2 . Seeds mostly Hith a starchy endosperm; floHers either ,dth \VeIl di f -
ferent i ated sepals and petals (the sepals mos tly not pe taloi d), or
more or less r educed and Idth chaffy or brist ly (or no) perianth,
not aggrega t ed into a spadix ........... .... ..... ... II. Commelinidae
2. Seeds mostly Hith a nonstarchy endos perm (notable excep t ions : Ara
ceae, Pontederiaceae , Philyd raceae) or without endosperm; fl o\.Jers
usually either Hith the pe tals and sepals much alike (commonly both
petaloid ), or reduced , collectively subtended by a spathe , and
aggregat ed into a spadi x.
3 . Flowers usually numerous, small , sub tended by a prominent spathe,
and often aggregated into a spadix ; plan ts often arborescent;
leave s in most species broad , petiola te, and not t.Jith typical
parallel venation ; supporting cells typically 4, less often 3
or 2 ; ovary superior (sometimes sunken in the axis o f the
spad i x) ................ . .......................... III . Arecidae
3 . FlO\.Jers fe\~ to numerous and small to large , but generally not
sub tended by a spathe a n d never aggregated i nto a spadix ;
plants seldom arbo r escent; leaves narrm.J and parallel- veined,
or le ss often b r oader and net-veined ; suppor ting cells typi-
cally none , less often 2 , rare ly more ; ovary superior or more
ofte n inferior .. ... ............ .. .................. IV . Liliidae

I. ALISNATIDAE

1. Seeds l.Jithout endosperm at matu r ity ; plants mostly aqua ti c or semi-

aquatic , autotrophic .
2. Perianth dif feren t ia ted into evident sepals and petals ; f loHers of t en
bra ct ea te.
3 . Flowers hypogynous ; carpels free or nearly so ; embryo sac , except
in Bu tomaceae, b i sporic . ....... . ....... . ......... l . Alismatales
3 . Flm.Jers epigynous , with compound ovary and modified laminar pIa
centation; embryo sac monosporic ............. 2. lIyd rocharit ales
2. Perianth , when pre s e nt, not di f ferentiated into sepals and petals;
bracts usually wanting, exc ept in Scheuchzer iaceae , but the peri-
anth s ometimes consisting of only a Single , bractlike tepal ; embryo
sac monosporic ....... .. .......... ... ...... ... . . ........ 3 . Naj adales
1. Seeds l.Jith \.Jell developed endosperm ; plants terrestria l, mycot rophic ,
t.,rithout chlorophylL .... . .......... ... ............. .... .. 4 . Triuridal es
29 683

1. ALISHATALES

1. Plants Hi thout secretory canals; pollen uniapertura te ; embryo sac mono-

sporic; embryo straight ; placentation laminar; leaves linear , not
differentiated into blade and petiole .. ........ . . . ......... 1. Butomaceae
1. Plants Hith schizogenous secretory canals; pollen multiaperturate; embryo
sac bisporic; embryo horseshoe-shaped; placentation and leaves various.
2 . Ovules several or many, scattered over the inner surface of the car-
pel (Le., the placentation laminar) ......... . ... 2. Linmocharitaceae
2 . Ovules I or 2, seldom mor e, basal or on a marginal placenta ... ....... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3. Alismataceae

2. llYDROCHARITALES

Represented by ..... .• • .......... ... ........................ 1. Hydrocharitaceae

3. NAJADALES

1. Ovules 2 or more, basal , anatropous; stamens 6 or more, f ree , often more

numerous than the tepals; f ruit s dehiscent, follicular; endosperm
helobial.
2 . Tepals 0-3, sometimes petaloid ; inflorescence a simple or basal l y
forking spike, bractless, although the perianth sometimes consists
o f a single bractlike tepal; pollen uniaperturate. the grains borne
singly; aquatic plants with submerged or floating leaves ........... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . Aponogetonaceae
2 . Tepals 6 , never petaloid ; inflorescence a raceme , each pedicel sub-
tended by a bract; pollen nonaperturate, in dyads; emergent marsh-
plants ...... ... ..................... . ....... . .. .. 2 . Scheuchzeriaceae
1. Ovule solitary; stamens either fe':ler than 6 , or as many as and adnate to
the tepals , or both; f ruits indehiscent or seldom schizocarp i c or irre-
gularly dehiscent; tepals 0-6 , never petaloid; pollen mostly nonaper-
turate.
3 . Ovule basal , erect , anatropous; endosperm nuclear.
4 . Hare or l ess emergent marsh-plants, scapose or subscapose , Hith
chiefly or ,,,hol1y basal leaves and a terminal raceme or spike;
carpels and stamens usually more than 1 .. .. ..... 3 . Juncag i nacea e
4. Submerged aquat i cs '\lith branching stems and opposite or whorled
leaves, the flOlvers solitary in the axils ; carpel 1 ; stamen 1. ..
. .... ...... . .. .. .... ... ........................... . 4 . Najadaceae
3 . Ovule ap ical or lateral, pendulous, more or l ess orthot ropous.
5 . Carpels usually 2 or more, separate ; pollen not threadlike; endo-
sperm chiefly helobial; plants not truly marine .
6 . FlOlvers in spikes or racemes, perfect or less often unisexual;
habit various.
7 . Tepals 4 ; stamens 4; spikes axillary; fruit in g carpels
sessile; pollen ellipsoid or spheroid; plants of fresh
water ............. . .................. 5 . Potamogetonaceae
7 . Tepals 0 ; stamens 2 ; spikes terminal; f ruiting carpels
st ipit ate; pollen of a unique , bilateral type; plants of
brackish Hater .............. ...... ..... .. .. 6. Rupp i aceae
6 . FloHers in axillary cymes, or solitary in the axils , unisex-
ual; submerged plants of fresh or brackish '''ater .. ......... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7 . Zannichelliaceae
5 . Carp e l 1; pol len threadlike; endosperm often or usually nuclear;
submersed marine plants ........ . .. .. ... .. . ........ 8 . Zos teraceae
684 29

4. TRIURIDALES

1. Flowers perfect; carpels 3; seeds numerous .............. 1 . Petrosaviaceae

1. Flowers unisexual; carpels severa l ; seeds 1 per carpel .... 2 . Triuridaceae

II . COMMELINIDAE

1. Flowers \"ithout nectaries or nectar ; ovary superior; vessels well devel-

oped in all vegetative organs .
2. Flm"ers Hith more or l ess shm>'y petals that are well di fferentiated
from the sepals. ne arly ahmys perfect; pollination most l y by
insec ts ............................................. 1 . Commelina1es
2 . Flm"ers perfect or often unisexual, Hithout showy petals, the peri
anth sometimes in 2 ser i es , but the 2 series much alike and not
petaloid.
3 . Ovules various , but never at once solitary , pendulous from the
apex of the ovary, and anatropous ; pollen mostly trinucleate;
embryo embedde d or more o f ten peripheral to the endosperm ;
p l ants terrestri al or less often aquatic; flowers perfect or
unisexual. borne in various sorts of inflorescences .
4 . Ovary wi th 1-3 f ertile locules and as many stigmas , each
fertile locule \"ith a single, apical , pendul ous , ortho-
tropous ovule; embryo p e ripheral to the endosper m; flowers
in most species and families unisexual .
5. Flm"ers aggregated into dense , pseudanthial, involucrate
heads , o f ten pollinated by i nsects ; ovules tenuinucel-
late; anthers usual l y bilocular ; leaves all basal ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2 . Eriocaulales
5 . Fl owers seldom aggregated into dense heads , normally wind-
pollinated or s elf-pollinated; ovules crassinucellate ;
anthers often unilocular ; leaves cauline or (in some
sma l l fam i lies) all basaL ............... 3 . Restionales
4. Ovary otherwise, either with more stigmas than locules, or
\·1i t h more than one ovule per locule , the ovules of various
form and position; embryo peripheral (Gr amineae) or em-
bedded (Juncales , Cype raceae) in the endosp e r m; flowers
per f ect or unis e xual .
6. Ovary \.;ith 1-3 locules and 3-many ovules; fruit capsular ;
pollen in tetrads; flowers in open or compact inflo r -
escences unlike those of the Cyperales ...... 4. Juncales
6 . Ovary wi th a single locule and ovule; f ruit indehiscent;
pol len in monads; f l owers in characterist ic spikes or
spikelets .......... . .. .................... . 5 . Cyperales
3 . Ovules solitary (or solitary in each locule), pendulous from the
apex of the ovary , anatropous; pollen binucleate ; embryo em-
bedded in the endosperm; floHers unisexual; borne in dense
spikes or heads; plants aquatic ..................... 6 . Typhales
1. Fl m>'ers with septal or septal- derived nectaries ; ovary inferior except in
some Bromeliaceae; vessels chi e f l y confined to the roots , except mainly
in some Bromeliaceae .
7 . Stamens 6 ; flo\"ers regular or sometimes somewhat irregul ar ; xero-
phytes and epiphytes \dth firm , narrm" , often spiny-margined leaves
. .. .................. . ....... ... .. ................... 7 . Bromeliales
7 . Stamens feHer than 6 (6 only in Ravenala, of the Strelitziaceae) ;
flowers distinctly irregular; mesophytes \"ith unarmed , pinnately
veined, oft e n broad leaves .. . ........ ... ...... . ..... 8 . Zingiberales
29 685

1. COHHELINALES

1. Leaf sheath open, often not well differentiated from the blade.
2 . Plants mostly terrestrial, often of \.,ret or mars hy !)laces , but seldom
truly aquatic; leaves generally all clustere d at the base, the
f l owers in a compact inflorescence terminating a l ong scape or
pedunc l e.
3. Stamens 6 , open ing by termina l pores; inflorescence a head of
spikelets , each spikelet with several brac ts sub tend ing its
single flowe r . .. ... .... . .... ... ......... ... . . . .. . . 1 . Rapateac:eae
3 . Stamen s 3 , often accompanied by 3 staminodes ; anthers open i ng by
longitudinal slits; inflorescence a simple racemose head . ..... . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. Xyridaceae
2 . Plants aquatic; leaves n umerous , lin ear or th r eadlike, distributed
a l ong t h e stem; stamen s 3 , the anthers openi ng by terminal pores or
short slits .................................. ... ..... .. 3 . Hayacaceae
1. Leaves differentiated into a closed sheath and a \.,rell defined , commonly
somcHha t succulent blade . ... .. ........... . . . . ... . ... ... . 4 . Commelinaceae

2. ERIOCAULALES

Represented by ... . . ....•• . . ... .. ... ... ..... .. . ... .. ••• .•. . . . .. 1. Eriocaulaceae

3. RESTIONALES

1. Flowers perfect; stamens 6; leaves chiefly or \.,rho1ly cauline, wi th closed

sheath ........ .. ... .. ... ... .. . ....... .. ... ... ......... 1. Flagellariaceae
1. FlOI.,ers unisexual; stamens 1-3; leaves ca ul ine or basal , the shea th mostly
open or poorly developed .
2. Stamens (1) 2- 3; tepa Is usually present; most l y coarse perenn ials.
3 . Leaves chiefly or wholly basal ; anthers bilo cular, free .
4. Inflorescence open, cymose ; flot... ers bracteate; ovary trilocu-
lar; plants mostly dioecious ..... .. ... .. .... 2. Anarthriaceae
4. Inflorescence a dense spike suggestin g t hat of Xyris; flowers
ebracteate; ovary bilocu l ar; plants monoecious ... . .... .... . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . Ecdeiocol eaceae
3 . Leaves "Iholly cauline; anthers unilocular, or rarely bilocular but
then laterally connate; inflorescence more or less open .... .. .. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Restionaceae
2 . Stamen 1; tepals none; diminntive, annual herbs with chiefly basal
leaves; anthers unilocular ....................... 5 . Cen t rolepidaceae

4. JUNCALES

1. Inflorescence of diverse sorts, bnt not as in the Thurniaceae; vascular

bundles of ordinary nature , \.,rith abaxial phloem, not as in the Thu rnia-
ceae ; cells I·dthout silica bodies ... ..... . .................. 1. Juncaceae
1. Inflorescence of aIle o r more dense heads sub tended by spreading , leafy
bracts; vascular bund l es of the l e af in ver t ical pa irs , the l ower (and
smalle r ) bundle of a pair with the phloem on top (adaxial) , facing the
phloem of the upper bundle; some cel l s of the leaf epidermis containing
silica bodies ... ... . .. .... .... .. . . .... . .... ... . ...... . .... 2 . Thurniaceae
 6H 29

5. CYPERALES

1. Flm..rers spiral l y or less often distichously arranged on the axis of the

spike or spikele t. usually each f lOHe r seemingly or actua l ly subtended
by only a single bract. without an evident bract be tween the flower
and the axis; seed coat generally free from the pe ri carp ; leaf sheath
usually closed, s tem usually so l id, often triangular; flowers often
with a perianth of evident bristles ; carpels 3 or less of t en 2; embryo
embedded in the endosperm ........................... . .... . 1. Cyperaceae
1. FI Ol"ers distich o usly arranged on the axis of the spikel et (or only one
per spikelet), each floHer ordinar i l y sub t e nded by a pair of bracts
(lemma and palea), the palea inserted between the flower and the axis ;
seed coat generally adnate to the pericarp; leaf sheath usually open;
stem usually ho l lm~, never triangul ar ; f l owers without a perianth,
unless the lodicules are so interpreted; carpels 2 , rarely 3; embryo
peripheral to the endosperm ................................ .. 2 . Poaceae

6. TYPHALES

1. Inflorescence of globose heads; vestigial perianth usually present;

achenes sessile or nearly so, not Hind-distributed . .... 1. Sparganiaceae
1. I nflorescence of dense, cy l indrical spikes; perianth apparent l y none;
achenes s l enderly long-stipitate, \~ i t h long hairs on the stipe, \~ind -
distributed .. .. . . ...... ... .. . .............................. 2 . Typhaceae

7. BRQl.JELIALES

, Represented by ... ....... .. .. ..............................•... 1. Bromeliaceae

8. ZINGIBERALES

1. Functional stamens 5 or rarely 6 , each I~ ith 2 pol l en sacs; plants with

raphide sacs ; guard c e lls symmetr i cal except in Low i aceae .
2. Flowers perfect ; leaves and bracts distichously arranged; plants
I~ithout laticifers; fruit capsular or schizocarpic .
3 . Ovules numerous in each locule; fruit capsular; s e eds arillate ;
stigma trifid .
4. Flowers Io/ithout an evident hypanthium; leaves obviously pen-
niveined; guard cells symmetrical : most species caulescent
or even arborescent ....... .... .. ......... 1. Strelitziaceae
4 . Flm~er s wi t h a long hypanthium much surpassing the ovary :
leaves seemingly parallel-ve i ned (actually penniveined with
narrm.!ly ascending veins); guard cells asynunetrical; acau-
lescent herbs ............ . ..................... 2. Lowiaceae
3. Ovules solitary in each locule: fruit schizocarpic; seeds not
arillate ; stigma capitate ...................... 3 . He liconiaceae
2. Flowers unisexual; leaves and bracts spirally arranged ; plan ts with
laticifers; fruit f l eshy, indehiscent, the seeds not ar illate .. ... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. Nusaceae
1. Functional stamen 1; plants without raphide sacs; guard cells asynunetrical
except in Cannaceae.
5. Stamen with 2 pollen sacs, not petal oid ; endosperm helobial.
6 . Leaves and bracts distichously arranged : sheaths open ; plants
a roma tic, wi th abundant oil cells .............. 5 . Zingiberaceae
29 687

6. Leaves and bracts spiral l y ar ranged; sheaths closed; plan ts with-

out o i l cell s ...... .. ... . . .................. . .. ... .. 6. Costaceae
S. Stamen lJith a sin g le functional pollen sac , the other sac modif ied
and petaloid ; endosperm nuclear .
7 . Ovules numerou s in each of th e 1-3 locules; st em 1"1th mucilage
canal s; emb r y o st r a ight; s eeds Ilot ari l l a t e . .. .... .. 7 . Cannaceae
7 . Ovul es sol itary in each of t he (1-2-) 3 l ocules; stem witho ut
muc ila ge canals; embryo fol ded; seeds most l y arill ate ... .. ... . . .
...................... . .... .... ....... . ... . . . .. .. . 8 . Marantaceae

Ill. ARECIDAE

1. Flowers we ll developed (though s mall). \~ith an evident, biseriate perianth

of 6 members , not c rowded in to a spadix; carpel s f r ee or united ; pl ants
very often arborescent and l.Ji th a t e rminal crown of large leaves, the
leaves i n any case with sheath, pet iole , and exp anded , pli cate blade;
endosperm n uclear . ...... .... . . . ... ..... .. .............. .. .... 1 . Ar ecales
1. Flower s more or less reduced and very often crowded into a s pad ix; carpels
united t o form a compound ovar y ; plants o f various habit , but never at
once arborescent a nd with a term ina l crm.Jn of broad IC!aves; l eaves not
plicate except in some Cyclan tha ceae .
2. Stamens most l y lO-numerou s ; vesse l s generally p r esent in l eaves and
stems as we l l as in roots; e ndosper m nuclear or hel obia l; flowers
unisexual ; leaves usually eithe r with b ifid b l ade or no t divided
into blade and pe tiol e .
J . Leaves I. ith bl ade , petiole , and basal s heath , the blade us ually
bifid ; pl a nts monoecious ; endosperm helobial; neotropical herbs,
se l dom woody .. . . . ..... .. ..... .. ... . . . .. .......... 2 . Cyc lanthal es
J. Leaves e l onga te and narrOH, somel.ha t sheathing at the base , but
not divid ed into blade and petiole , not bi fid; plan ts dioe-
c ious ; endosperm nuclear; paleo t ropical woody p l ants .... .. ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3 . Pandanales
2. Stamens most l y 1-8 (flowe r s rarely produced i n Lemnacea e ); vessels
confined t o t he r oots , or l.Jho l ly Han t ing ; e ndosper m cellula r;
floHers perfect or uni sexual; leaves (when p r esent) often divided
into blade and petiole; herbs , or se ldom \lIOody c limbers .. .. 4. Arale s

1. ARECALES

Re presen t ed by .....• ••• .. ..........................••••• . •••• ... .. 1. Arecace a e

2. CYCLANTHALES

Repre s ented by . .... . •. •••... ..... ... ... .... ... . . ... . •.•• • . • • .. 1. Cyclanthaceae

3. PANDANALES

Represented by .. . . . • • ••••• . ... . . ... .. . ..... .. .. . .... • • • •• .. . . • .. 1 . Pandanaceae

4. ARALES

1. Plants with roots , stems , and leaves , terrestria l or sometimes more or

less aquat ic, but only r arely f ree-floating ; inflorescence a spadix ;
plant s usually with ves sels i n the roots a nd tracheids i n the roots,
stems. and l eaves ... .. . ... .. .... . ...... ... .... .. .... . . . . . .... . 1. Arac eae
688 29

1. Plants thalloid, free- fl oating , with or without I-several short, slender

roots; f l owers (rarely produced) not forming a definite spadix ; plants
lacki ng both vessels and tracheids ............... .. ........ 2. Lemnaceae

I V. LILIIDAE

1. Plants not obviously mycotrophic; seeds of ordinary number and structure ,

usually with a we ll d i fferentiated embryo and 11ell developed endosperm;
most fa~ilies and genera 11ith septal nectaries . sometimes with other
kinds of nectar i es in addition or instead; ovary superior or inferior ..
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. Liliales
1. Plants strongly mycotrophic , somet i mes 11ithout chlorophyll; seeds very
numerous and tiny , 11ith undi fferentiated embryo and very litt l e or no
endosperm; nectar i es diverse , but not (or at least not usually) septal;
ovary inferior ..... .. .............. . ... . . . ................ 2 . Orchidales

1. LILIALES

1. Endosperm starchy (some times also 11ith some fat); stomates \odth 2 subsi-
d i ary cel ls (4 o r mo re in sane Philydraceae); perianth more or less
distinctly irregul ar; plants either aquatic , or with only a single
stamen.
2 . Tepals 4 (one member of the outer set derived by connation of 2, one
memb e r of t he i nner s e t missing) . not united into a tube ; plants
terrestrial; sepal nectaries ",anting; stamen solitary ............. .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 . Philydraceae
2 . Tepal s (4) 6, usually connate be l m1 into a perianth tube; plants
aquatic; septal nec tarie s present; stamens (including staminodia)
3 or 6 .............. .... .......................... 2 . Pontederiaceae
1. Endosperm (,,,hen pr e sent) t.,rith reserve ce l lulose and/or fat or protein ,
sometimes also with some starch; stomates in most families usually
I"ithout subs i diary cells; flowers I" i th more than one stamen ; plants
seldom aquatic .
3 . Leaves typically narrow and parallel- veined, sometimes broader and
more net-ve ined , but on l y seldom with both a b r oad , net - veined
blade and a dist i nct petiole .
4 . Vessels chiefl y or I"holly con f ined to the roots; stomates typi -
cally I·,it hout subsidiary c e lls ; perianth and ovary seldom
conspicuous ly hairy or glandular.
5 . Habit lilia ceous (most l y so f t-leaved herbs f r om a bulb or
r i1i zome); stamens 6 or J .
6 . St amens (including staminodes) mostly 6, seld om more
than 6 or only 4; ovary superior o r less often in-
f erior ............ . ............ . .. . ........ 3 . Liliaceae
6. Stamens 3 ; ovary inferior .... ..... ........... 4. Iridaceae
S. Habit agavaceous (mostly coarse , often shrubby or arbores-
cent plants with firm , perennial leaves) ; stamens 6.
7 . Perianth usually more or less corolloid , not chaffy ; X
16 or more , usually Hith a few l arge chromosomes and
more numerous small ones .......... ... ...... 5 . Agavaceae
7 . Perianth dry and chaffy ; X = 11 ....... 6. Xanthorrhoeaceae
4 . Vessels well distributed in all vegetative organs; stomates
mostly with 2 subsidiary cells; perian t h , or ovary, or both ,
commonly densely hai ry or glandular .
29 689

8. Sh rubs or s ub shrubs with the leaves crowd ed at che top of a

usually branching stem , the old leaf-bases persistent and
cloching the stem; plants without branched hairs ..... .. .... .
· .... .. ..... .. ...... . ................ . . .. . . .. 7. Velloziaceae
8 . Scapose o r subscapose herbs ; inflorescence often with branched
hai rs . ...... .... ... . .... ... .... . ............ 8 . Haemodoraceae
3. Leaves with a definite peciole and an expanded (often large) . mo r e o r
l ess net-veined blade .
9 . Plancs acaulescent; vessels confined to the roots .
10 . Ovules n umerous on parietal placentae ; endosp e rm \~ell devel-
oped. rich in protein; chalazosperm wanting ; leaves often
much cleft . .. . .. ........ .... ............... ... . . 9 . Taccaceae
10 . Ovu l es 2 in each of the 3 l ocules (but usual ly on l y one loc ule
fertile) . on axile placentae; endosperm non e; chalazosperm
abundant . rich in starch and fat ; l eaves undivided .. . .... .. .
· .. ... . . .... . ...... ... .................. .. . 10. Cyanas traceae
9 . Plant s leafy-stemmed . usually climbing.
11 . Flowers dime rous; vessels conf ined to the roots ..... ...... ... .
· .. .......... .... ... . ..... .. ...... . ..... .. .. . 11. Stemonaceae
11. Flowers t r imer ous; vessels well distributed in all vege tat ive
organs .
12 . Ovary superio r; nectaries staminal or staminodia l; plants
not t wining . mostly climbing by tendrils and without
prominent tubers ....................... .. 12 . Smilac ac eae
12 . Ovary i nferior; nectaries septal; plants mostly t\~in ing ,
without tendrils. us ually with large tubers ... .... ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 . Dioscoreaceae

2. ORCHIDALES

1. Stamens 3 or mo re . free from the style; pollen grains not cohering in

pol1in ia; flowers regular or less often irregular; terrestrial plant s.
most species with reduced leaves and without chlor ophyll; nectar ies .
at least i n the Burmann i aceae. ovarian .
2 . Flowers regular or near ly so; stamens 3 or 6 .
3 . Stamens 3. opposi te the sepal s; sepals valvate .. .. 1 . Geosi ridaceae
3 . Stamens 6. or more often 3 and opposite the petal s; sepals imbri
cate ...... ... ..... .. . . .. ... .... .. . .. ... .. .. .. . .. 2 . Burmanniaceae
2 . Flowers highly ir regular; stamens 6 . . . . . .... .. ... ... .. .. . 3 . Co rsiaceae
1. Stamens I or 2 , rarely 3 . mostly adnate to the style ; pollen grains i n
most gene r a cohering in pollinia; flO\~ers more or less stro ngly irregu-
lar; terrest r ia l or often epiphyt i c plants. \~ith or l ess often wit hout
chlorophyll; necta r ies diverse . but sel dom ovarian ... ..... 4 . Orchidaceae
Section D.
DIAGNOSTIC KEY TO THE FANILIES OF LOUER VASCULAR
PLANTS IN NORTH AHERICA (NORTli OF MEXICO) '"

1. Plants cons i sting of a dichotomously branching system of sterns; leave s

and roots la cking . but possessing mi nute enations scattered on aeria l
stems ; sporangia la r ge (1-2 rom acr oss) . tri l ocular . ....... 1. Psilotaceae
1. Plant consisting of stem . roots (except Sal viniaceae). and leaves; spor
angia large or small. but never tril ocular .

*Contributed by John T. Micke l. The New York Botanica l Garden .

690 29

2. Leaves with a single unbranching vein , small (less than 1 . 5 em long) or

l ong grass-like aquatic ; stem protostelic o. eustelic , lacking leaf
and b ranch gaps (except Equiset aceae); sporangia bo rne in cones , on
stem in axils of leaves , or embedded in the bases of grass - like leaves .
3. Leaves ",harled , reduced to toothed , generally non - green sheaths at
nodes ; stems h ollow , jointed , flu ted; sporangia borne under pel-
tate scale-like structures that compri se the cone . .. 2 . Equisetaceae
3. Leaves green, spirally arranged or rarely whorled with four leaves
per ,,,horl , never for ming sheaths ; stems n ever holl ow, jointed , or
f lut ed ; sporangia ahlays borne in association Idth leaves .
4. Plants terrestrial ; leaves less t han 1 cm long; sporangia in the
a xi ls of eithe r vegetative leaves or spec i alized l eaves in
cones; homosporous or heterosporous ; spores all te trahedral .
5 . Plants homosporous; sporangia in axi1s of vegetat ive leaves
or in cones that are round in cross section ..... ... .. . . .. . ,
... ... ..... . . ...... . ...... .. . , ... . ......... 3 . ],ycopodiaceae
5 . Plants heterosporous , megaspores fo ur p er megasporan g ium;
sporangia borne in cones that a re four-sided in cross sec-
tion , rarely round ........... .. .......... 4 . Se1agine11aceae
4 . Plants aqua t ic or semi-aquatic; leaves grass-like (3 40 cm long,
0 . 5 - 2.0 mm \.,ride) , tufted, arising f rom a short , erect , corm-
li ke stem ; s porangia embedded (adaxia11y) in the leaf near its
base; heterosporous; megaspores tetrahedral , hundreds per mega-
sporangi um; microspores bilat eral .. .... ... .. .... .. 5. Isoetaceae
2. Leaves gen e rally I·lith branching veins; leaves small to large ; stems usu
a lly solenostelic I·lith l eaf and branch gaps ; sporangi a never borne in
cones or in axi1s of leaves but on the leaf, rarely in nut-like sporo-
carps .
6 . Plants aqua tic or semiaqua t ic; sporangia enclosed in nut-like sporo-
carp s ; heterosporous.
7 . Plants rootin g in mud; leaves alternate , petiolate, more than
2 cm long , \-1ith fou r leaflets at tip of petiole or l.,r i th no
l eaflets; sporocarps 2-6 nun across, arising from base o f
pe tio1e ... . ... .. .............. .. .... ... . . ....... 6 . Harsileaceae
7 . Plant s free floating , rarely on mud; leaves simple , sessile ,
round or bilobed, opposite or 1'lhor1ed, less than 1 . 5 cm long ;
sporocarps 1 mm or less across , borne on leaf or root-like
skeletonized leaf .
8 . Leaves 1 mm or less across , opposite and bilobed; sporocarps
borne on lm'ler leaf lobes ; hairs lacking from leaf surface ;
plan ts inhabited by blue-green alga Anabaena . . 7 . Azo1laceae
8 . Leaves 6-15 mrn across , Hhorled with one member of whorl
trailing beneath , appearing as a root and bearing the
sporocarps ; upper leaf surface covered lolith h ai r s t hat are
fused in clusters ; plants not inhab ited by Anabaena .. . . .. . .
. . . . . . . . . . . . . . . . . . . . . . .... . . . ............ .. . 8 . Salviniaceae
6. Plants terrestrial, rarely aquatic ; sporangia borne on under s i de of
leaf or on upr ight spike or panicle arising from t he petiole , never
en c l osed in sporocarps ; homosporous .
9 . Sporangia borne on uprigh t spikes or pan icles that arise from the
petio le .
10 . Fertile panicles tl>'0 per leaf; sporangia ca . 0.3 nun acros s ;
rhizo me creeping at or just belO\ol ground level ; strengthen-
ing tissue present thro u ghout plants, making plant parts
tough .... .. ..... .... ........ . . . ........ .. . .. .. 9 . Anemiaceae
29 691

10 . Fertile spike or panicles one per lea f ; sporangia ca . I mm across;

rhizome upright, fleshy , \1ell beneath the surface of the ground ;
strengthening tissue generally lacking , plant parts soft ........... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10 . Ophioglossaceae
9 . Sporangia borne on the leaf blade, either on no r ma l vegetative leaves or
strongly modified leaves ; sori on t he blade marg i n or the under surface .
11. Leaves extremely delicate and filmy , only one cell thick , 1-15 cm
long ; sporangia borne on elongate receptacle i n cups or tubes on
l eaf margin ; rhizome often fine and hair-like , 1 mm o r less in
thickness ....................................... 1 1. Ilymenophyllaceae
11. Leaves more substantial, more than 1 cell thick ; sari dorsal or mar
ginal , but if in marginal cups , leaves more t han 30 cm long ; rhi-
zomes generally more than 1 mm thick.
12 . Leaves regularly dividing in half \-lith dormant bud at the po i nt of
forking ; pinnae pectinate ; sari with few sporangia, (4-7) all
developing at one t i me ....... .... . ............ 12 . Gleichenia c eae
12 . Leaves not regul arly dividing dichotomously ; sari \'lith many spor
angia or on ly 1, usually not develop i ng simul taneously .
13 . Leaves climbing; sporangia borne singly under ep i derma l f laps
on constricted segment lobes . ........... ..... 13 . Lygod i aceae
13. Leaves not climb i ng ; spo r angia not bo r ne singly under epid er
mal flaps .
14 . Leaves filiform ; spo r angia borne on p i nna t e ( 2 - 4 mm long)
or digitate (7 - 8 mm long) divisions at end of leaf ..... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14 . Schizaeace a e
14 . Leaves not filiform ; sporangia not borne on sma l l projec
tions from end of petio l e .
15 . Fertil e leaves or fertile portions o f l eaves lacking
any leaf tissue ; brown fer t ile portion at tip or
middle of vegetative fronds o r on totally distinct
fronds; sporangia sessile with annul us a n irregular
lateral patch of thick-\,'alled cells .. 15 . Osmundaceae
15 . Fertile leaves or fertile portions of leaves possess
ing at lGast a small amou nt of leafy tissue whi ch
may be green or brmm ; sporangia stalked, \1ith a
vertical annulus .. .. . . . .. . (Polypod i aceae ~ lata)
go to 16 couplet
16 . Rhizome clothed \dth hairs ; rhizome l ong- creep-
ing ; branches arise regularly either by dichoto-
mous forking of rhizome and/ or from buds on the
base of the petiole ; sari marginal , pro t ected
either by an outer <lOd inner indusium o r only by
a recurved marginal flap .... 16. Dennstaed tiaceae
16. Rhizome clothed \-lith scales; rhizome erect or
creeping ; branches arise only from the stem ,
never from the pet i ole ; sari marginal or on the
l ower side of the leaf, protected by a true i n-
dusium, a recurved marginal flap , or n ot a t a l l .
17 . Sari marginal, or if dorsal , elongate along
veins or completely cove r ing I m,er surface
and 11ithout true indusium; spores tetrahe-
dral ; petio l e bundles I, 2 , or rarely many
(20 or more) .
 692 29

18 . Plants aquatic, either floating or rooted in mud; leaves less

than 0.5 mm lon g ; often proliferating by buds on the leaves
.... . . ..... ... ... .... .. ................. . .. 17. Parkeriaceae
18 . Plants terrestrial, or if rooting in mud then leaves more
than 1 m long; not proliferating by laminar buds .......... .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 . Ad i antaceae
17 . Sari dorsal (marginal in Paltonium); spores bi latera l (tetrahe
dral in Grammit i s); petiole bundles I (x shape), 2 , or 3-7.
19 . Ind usium l acking ; sari round (elongate along margin in Pal-
tonium); blades mostly simp l e or pectinate, less commonly
pinna te; spores lacking pe rispore . . ................. .... .. .
..... ........ ..... .. . .. ..... . ... Polypodiaceae ~ strictu
19 . Indusi um generally present; sori round or elongate along
veins or midrih, rarely cover i ng blade surface ; blades
mostly once pinnate or more dissected, rarely pectinate or
simp l e; spores wi th perispo re .. ... ......... 19 . Aspleniaceae*

*The name "Aspidiaceae" has more often been app lied to this group , but that
name is illegitimate.

FM-IILY REV I Etol EXERCISES

Family Key Exercise : An excellent revieH for diagnostic family study is to

take the family key in a manual, guide , or synopsis , appropriate to your area ,
copy or mark the key leads to the families you have studied and then summarize
'. the critical characters used in delimiting each family studied .

Family Comparison Chart Exercise: Hake a list of families studied accor ding
to a system or alphabet i ca l ly in a left hand column a n d across t h e top list
characters used in your family analysis and then fi ll in columns with appro-
priate character states from your fami l y analyses . See e xample below .

Familv Inflores cence Sex Floral Formula Corolla Type Etc.

Alismataceae I
Poaceae

Cvoeraceae

Etc .
29 693

FAMILY LITERATURE
(See references at end of Chapter 28)

Bessey , C. E. 1915 . The phylogenetic taxonomy of flowering plants . Annals

of the Hissouri Botanical Garden 2 : 109-164 .
Cronquist. A. 1968. Evolu tion and Classificat ion of Flowering Plants.
Hought on-Nifntn Company . Boston .
Lawrence. G. H. H. 1951. Taxonomy of Vascular Plants. The Hacmillan Com-
pany . New York.
Melchior , H. 1964 . A. Engler ' s Syllabus Oer Pflanzen-fami11en II . 12th ed .
Gebriider Borntraeger. Berlin .

FAMILY LABORATORY EXERCISES

Dur ing the course of a student ' s training in systematics he should become
acquainted \dth the (0110\,,10g families due to the i r phylogenetic or economic
significance: Hagnoliidae- (1) Nagnoliaceae. (2) Annonaceae, (3) La uraceae,
(4) Saururaceae, (5) Aristolochiaceae, (6) Nymphaeaceae , (7) Ranuncul aceae,
(8) Berberidaceae , (9) Papaveraceae, (10) Fumariaceae; Hamamelidae-{ll) Hama-
melidaceae , (12) Ulmaceae , (13) Urticaceae, (14) Juglandaceae , (15) Hyricaceae,
(16) Fagaceae , (17) Betu l aceae , (18) Casuarinaceae ; Caryophyllidae-( 19) Cary -
ophyllaceae , (20) Portulacaceae, (21) Amaranthaceae , (22) Bataceae, (23) Poly-
gonaceae , (24) Plumbaginaceae; Dilleniidae-(25) Dilleniaceae, (26) Theaceae,
(27) Clusiaceae , (28) ~Ialvaceae. (29) Droseraceae, (30) Violaceae, (1) Cista-
ceae, (32) Cucurbitaceae , (33) Salicaceae , (34) Brassicaceae , (35) Ericac eae ,
(36) Diapensiaceae, (3 7 ) Symplocaceae, (38) Primul aceae; Rosidae-(39) Saxifra-
gaceae , (40) Rosaceae, (41) Fabaceae, (42) Podostemaceae , (43) Lythraceae , (44)
Onagraceae , (45) Elaeagnaceae, (46) Cornaceae , (47) San t alaceae , (48) Celastra-
ceae, (49) Aquifoliaceae , (50) Buxaceae, (51) Rhamnaceae, (52) Vitaceae , (53)
Aceraceae, (54) Rutaceae , (55) Geraniaceae, (56) Polygalaceae, (57) Araliaceae,
(58) Ap i aceae ; Asteridae-(59) Gentianacea e , {60} Asclepiadac eae , (61) Convolvu-
laceae , (62) Polemoniaceae, (63) Boraginaceae, (64) Verbenaceae , (65) Lamiaceae,
(66) Plantaginaceae, (67) Oleaceae, (68) Scrophulariaceae , (69) Lentibularia-
ceae, ( 70) Campanulaceae , (71) Rubiaceae , (72) Caprifoliaceae, (73) Asteraceae;
Alismatidae-(74) Alismataceae, (7S) lIydrocharitaceae , (76) Najadaceae, (77)
Po tamogetonaceae; Commelinidae-(7 8) Xy r idaceae , (79) Commelinaceae, (80) Erio-
caulaceae, (81) Juncaceae , (82) Cyperaceae , (83) Poa ceae , (84) Typhaceae, (85)
Bromeliaceae, ( 86) Naran taceae; Arecidae-(B7) Arecaceae, ( 88) Araceae, (89)
Lemnaceae; Liliidae-(90) Pontederiaceae , (91) Liliaceae, (92) I ridaceae , (93)
Dioscoreaceae, (94) Burmanniaceae , (95) Orchida ceae . At least 35-40 of these
families should be studied i n the basic course in systematics .
Each of the families listed ahove s hould be analyzed for diagnostic and
phylogenetically significant characters . A sample family work sheet is pro-
vided which can be reproduced as it is or appropriately modified. Each family
analysis should pe rtain to the species studied in that family with supplementary
remarks to be added from the l iterature or other observations \~hen the species
does not adequately r epresent the diagnostic or phylogenetic c harac teristics
of the family. A median longitudinal section of the fIOHer and a cross section
of t he ovary s hould be drawn for each family. Unique features fO T the family ;
e . g. , syngenesia , pollinia , arillate seeds , should be illustrated . Fl oral dia-
grams should be optional .
A desirable aid for family study is to have a class herbarium a vailabl e
with t he famili es arranged alphabetically Idth mounted specimens of at least
one species of each genus within each family from the area. To sho,,", i ntra-
familial relationships , demonstration material could be arranged according to
s ub-family and/or tribe.
694 29

SAMPLE FAHIL Y HORK SHEET

Species Family

l. P lant Type 13 . Stigma Type

2. Plant Duration 14 . Style Type

3. Plant Sex 15. Ovule Type

4. Root Type 16 . Fruit Type

5. Stem Type 17. Seed Type

6. Leaf Type lB. Embryo Type

7. Inflorescence Type 19. Locat ion/P lant Organ

B. Flower Type 20 . Position/P lant Organ

9. Androecial Type 2l. Arrangement/Plant Organ

10 . Stamen Type 22 . Floral Herosity

ll . Gynoecial Type 23. Venat i on

12. Carpel Type 24 . Sex

Flower 1.S . Special Features

General Family Comments:

 FANILY AND HIGHER TAXA IND EX

H llr un onla ceae 61H C"pressaceac 650 Flagellariaceac 685

Acanthaceae 680 Buddlej<lceae 680 Cuscutaceae 679 Fouquier!aceae 664
Aceracea.:! 647, fi75 Bu~mannla"eae 689 Cyanastraceae 689 Frankenlaccae 664
Acha ria eeae 664 Bursera"eae 675 Cycadaccae 6t,9 Furnariaceae 655
Actinidiaceae 662 Butollk~ceae 683 Cycadales 649 G-K
Miantaceac 692 Buxaceae 673 Cycadopsida 649 Garryaceae 671
Adoxaceae 681 Bybl1daceae 668 Cyclanthaceae 6117 Gelsso1omataccae 672
Aexto xicaceae 674 C Cyclanthflles 687 Gentianaceae 648 , 678
Agavaceae 688 Ca " taceae 648 , 659 Cynomoriaceae 672 Ge ntianales 677, 678
Aizoaceae 659 Callitrichac eae 679 Cyperaceae 645 , 686 Geosiridaceae 689
Akaniaccac 6ll; Calycanthaceae 654 Cyperales 681" 686 Geraniaceae 647, 675
Alangiaceae 671 Calyceraceac 682 Cyrillaccae 665 Gc ranlales 667, 675
Alismatac eac 61,S, 683
Alismatales 682 , 683
Campanul"ceae 6t.9 , 681
Campanulales 677 , 681
0-' Ge~neriaceae 680
Daphnip hyUaceae 673 Ginkgoflceae 650
Alismatidae 682 Canellacea" 653 Datiscaceae 665 Ginkgoa les 650
Alseuosmiaceae 669 Cannahaceae 657 Davidiaeeae 671 Gleicheniaceae 691
,\maranthaceae 646 , 660 C,mnaeeae 687 Davidsoniaccae 668 Globulariaccae 680
Amaryllidaceae 646 Capparaee"e 665 Degeneriaceae 653 Gnctopsida 650
Amborellaceae 653 Capparales flol, 665 Dennstaedtlaceae 691 Gomortegaeeae 654
.~nacardiaceae61.7, 675 Caprifo l iaeea e 649 , 681 Dialypetalanthateae 670 Goodeniaceae 681
Anarthriaceae 685 Cardiopteridaceae 673 Diapensiaceae 666 Greylaceae 674
Ancistrocladaceae 664 Ca ri caceae 664 DiapensLales 661, 666 Gro~s ular iaceae 669
Aoemiaceae 690 Car yoca rae eae 662 Diehapetalaceae 673 Grub blaceae 672
Annonaceae 653 Caryophyllaee ae M6, 659 Didiereaceae 659 Gunneraceae 670
Apiaceae 648 Caryophyllales 658 , 659 Dldymelaceae 657 Cuttiferac 662
Apacynaceae 648 , 678 CHryophyllidae 652 , 658 Dilleniaceae 661 Haemodoraceae 689
Aponogetonaccae 683 Casuarinaceae 6.58 Dilleniales 660 , 661 Haloragaceae 670
Aquifotiaceae 64 7 , 673 CasuarLnales 656 , 658 Dilleniidae 652, 660 Ilaloragales 668, 670
Araceae 645 , 687 Celastraccae 673 llioncophyl1aceae 664 lIamar'le lidaceae 647 , 657
ATales 687 C '>Jastr,~les 668, 672 Dioscoreaceae 689 Ilam"melidales 655 , 656
Araliaccae 61.8 , 677 Centrolepidaceae 685 lJipentodontaceae 672 llamamel1dlle 651, 655
Araucariaceae 650 Cephalotaceae 669 Di psacaceile 681 Ilelieoniaceae 686
Arecaeeae 687 Cel'halotaxaceae 650 Dipsacales 678, 681 Hcrnandiaceae 654
Arecales 687 Cer atophyl laceae 654 Dipterocarl'aceae 662 Himantandrace"e 653
Arecirlae 682 , 687 Cerddiphyllaceae 656 Droseraceae 663 Hip pocast"naceae 675
Aristolochiaceae 646 , 654 Chenopodlaceae M6, 660 Ebenaceae 666 Hippocrateaceae 673
,',ristolochiales 652, 654 Chloranthaceae 654 Ehe nales 661, 666 Ill ppuridaceae 670
Asclepiadaceae 648, 678 Chrysobalanaceae 669 Ecdeiocoleaceae 685 lIop lestigmataceae 664
Aspleniaceae 692 Circaea~teraceae 654 Elaeagnaceae 648 , 671 lIumiriaceae 676
Asteraceac 649 Cistaceae 664 Elacocarpaceac 662 lIyd nora" eae 672
As terales 678, 682 Clct hraccae 665 Elatinaceae 662 Ilydr"ngeaceac 669
Asteridae 652, 677 Cn"orflceae 675 Empetraceae 666 Hydrocharitaceae 683
Austroballeya"eae 653 Columelliaceae 668 Epacridaceae 666 Hydrocharitalcs 682, 683
Azolla"eae 690 Comb r etaceae 671 Ephedraceae 650 Hyd rophyllaceae 649, 679
Balanopa"eae 658 Commelinaceae 645, 685 Ephedral es 650 Hydrostachyace"oo 680
Balanophoraceae 672 Commelinales 68t, , 685 Equisootacooae 690 lIymcnol'hyllaccae 691
Balsaminaceae 67fl Commelinidae 682, 684 Erieaceae 61,8 , 666 Ilyperic"ceae 648
Barbeyaceae 657 Compositae 682 Ericales 661, 665 Icacinaceac 673
Basellaceae 659 COtlllaraceae ('74 Eriocaulaeeae 685 Llliciaceae 653
Hata"eae 660 Convolvulacea.e 648, 6711 Eriocaulales 681., 685 [rida"eae 646 , 688
Batales 658 , 660 Coriariaceae 655 Erythroxylaccae 676 [soootaceae 690
Begoniaceae 665 Cornaceae 648 , 671. Eucorruniaccae 657 Juglandaceae 646, 658
Berberldaceac 655 Cornales 668, 671 EucommialcH 656 , 657 Juglandaies 656 , 657
Betulaceae 646, 658 Corsiaceae 689 Eucryp hi aceae 668 Ju l ianaceae 675
Rignoniaceae 680 Corynocarpaceae 655 Euphorhi.~c e"e 647 , 673 Juncaceae 646, 685
Bixaceae 664 Costaceae 687 Eup ho rb i aLes 668 , 673 J uncaginaceae 683
Bombaca"eHe 663 Cras~ulaceac 669 Eupo matiaceae 653 Juncales 684, 685
Bo raglnHccae 649 , 679 Crossosomataceae 661 Eupteleace"e 657 Krameriace"e 676
Brassicaceae 647 Cruciferae 665 Fahaceae 647 L- M
Bromeliaceae 686 Crypteroniaceae 670 Fagaceae 646, 658 Labiatae 679
Bromeliales 684, 686 Cucurbitaceae 649, 665 Fagales 656, 658 Lactoridaceae 653
Brunia ceae 669 Cu noniaceae 668 Flacourtlaceac 663 Lamiaceae 649
 696 29

Lamiales 677, 679 N-q Portulacaceae 659 Stcmonaceae 689

Lardizabalaccae 655 Najad«ceae 645 , 683 Potamogctonaceae 683 Sterculiaccae 662
Lauracea" 654 Najadales 682, 683 Primulaceae 641l , 667 Strelitziaceae 686
Lecythidaceae 663 Nelu:nbonaccac 651, I'rimula1es 661, 666 Stylidiaceae 681
Lecythidales 660 , 663 Nepenthaceac 663 I' rote"ceac 671 St ylo bas iaceae 674
Le"<leeae 6 74 Neuradaceac 669 Proteales 667 , 671 Sty racace" e 6~8, 666
Lcguminosa" 669 Nolanaceae 678 Psilotaceae 689 Sy mplocacea c 666
Leitneriaceae 657 :lyetagillaceae 659 Punlcaceac 670 T-Z
Leit n criales 656, 657 Nymphaeaceae 646, 654 Pyrolaceae 666 Taeeaceae 689
Lemnaceae 688 Ilymp h acales 652, 654 Quilnac cae 662 Tamaricaccae 664
Lennoaceae 679 )Tyssaccac 67.1 R-S Taxaceae 1)50
Lcntibula"daceae 680 Ochnaceae 661 Raff Jesiaccae 672 Taxa1es 650
Liliaeeae 61,6, 688 Olacaceae 672 Rafflesiales 667, 672 Taxodiaceae 650
Lili.alcs 688 Olcaccae 648, 680 Ranunculacc"e 647, 655 Tetracentraceae 656
Liliidac 682, 688 Oliniaceae 671 Ran unculalcs 652 , 654 Theaceae 662
Liliopsida 651, 682 On a graceac 648 , 671 Rapateaccac 685 TheaJe,; 660, 661
Limnanthaceae 676 Opi liacc ilc 672 Rescda ccae 665 Theligonaccae 670
Limnocharitac('ae 683 Ophioglossaceae 69 1 Restionac ea" 685 Theophra s taccae 666
LLn.weae 676 Orch.tdaccac 61,6, 689 RC!ltionales 684, 685 Thurniaceae 685
Linales 667, 676 0rc hidal cs 688 , 689 P.hamnaceae 647, 671, Thymelaeaceae 670
Lissocarpacea" 666 Orob,1nc hac eae 680 Rhmnna1e" 671, Tiliaceae 662
Loasaceac 664 Osmu nclaceae 691 Rhizop h oraccae 671 Tovariaceac 665
Loganiaceae 678 Oxalidac~ae 675 Rhoipteleaceac 657 Tra p accac 670
Loranthaceae 672 l'aeoniaceae 661 Rho paloearpaceae 662 Trcmandraceae 676
Lowiaceae 686 Pandaceae 687 Rosaceae 647 , 669 Trigoniaceae 676
Lycopodiaceae 690 Pa nd,mu ccac 67/" 687 Rosales 667 , 668 Trimeniaceae 653
Lygorliaceae 691 I'a ndana les 6117 Rosidae 652 , 667 Triuridaceae 684
Ly thraceae 648, 670 I'al'averaceae 6f,7 , 655 Rubiace"e 649 , 681 Triurldales 682 , 684
a,.gnoliaceae 647, 653 Pa paverale,; 652, 655 Ruhiales 677, 681 Trochodendraccae 656
~!agn oliales 652 I'arke riac eac 692 Ruppiaceac 683 Troc h odcndrales 655 ,
Hagnoliidae 651 , 652 l'assi floraceae 664 Rutaceile 675 656
~!"gnoliop 5ida 651 Pedali"ceae 680 Sabiaceae 655 Tropaeolaceae 676
H"lesherb iaceae 66t, Pcnacaccac 670 Salicace:le 646, 665 Tur neraceae 663
:!alpighbceae 676 Pentaphragmatacc;Je 6111 Sa l1 ca lcs 661 , 665 Typhaceae 645, 686
:lalvac"ac 647, 663 l'eridis c aceae 663 Sa 1vado ra ce:le 673 Typha1e,; 684 , 686
~lalva l cs 660 , 662 l'etrosavLaceac 684 Salviniaceae 690 Ulmaceae 657
,1a ran t;Jceae 687 Phi lydraccac 688 San ta lac cae 646, 672 Umbellalcs 667, 677
!-!arcgraviaceac 662 Ph r ymace,.e 679 Sant:ll:l1es 667, 671 Umhelliferae 677
Nan; Lleileeae f.9n Phytolacca~cae 659 Sapind,1ceae 67~ Urticaccac 61,6 , 657
~·l<lY<1caceae 6S5 I'icrodend r accac 658 Sapinda Lc s 667 , 674 Urtiea1es 656, 657
!'ledusagynaceae 662 Pinaceae 650 Sapotaceac 66A Valcrianaceae 68 1
r!cd us:mdraceae 671 l'inales 650 Sa rracc niaccae 64 7 , 663 Vel10ziaccae 689
Helastomataceae 648 , f,71 P inophyta 649 Sarraceniales 660 , 663 Vcrbenaceae 649, 679
.'te liaceae 675 P inops ida 650 Sa r colilenaeeae 662 Violaccac 6f,8 , 663
.' lc,l'anthaceae 671, Pipcrace"e 654 Saururacca e 65f, VLobles 660
tlenispe r m:lc e :le 655 P'ipcraLcs 652 , 65/, Sax i fragaceae (,1,7 , 669 Vitaceae 6t,] , 671,
:lenyanthaceae 679 Pittosporaccac 668 Scheuchzer i. accac 683 Vochysiaceae 676
~!is o dend rac eae 672 Plantagi n a c cac 649, 679 Sc hisan draccac 653 lHntcraceae 653
itilr:lsternon a ceae 072 Plantaginalcs 677 , 679 Schizaeaceae 691 Xant hoph yllaceae 676
Holl uginaceae 659 I'latanaceae 657 Scrophular i aceac 649, 680 Xanthorrhoeaceae 688
:tonirniaceae 653 Plumbaginace"e 660 Scrop h ular ia 1es 677 , 679 Xy ridacea e 6t,5 , 685
.'tollo t ropaceac 666 I'lumha!;ina\es 658 , 660 Sc yphostegi<lccae 663 Za~i.aceae 649
~Ioracea" 657 Poaceae 645 , 686 Scyto p etalaeeac 662 Zannichell i. aceae 683
:loringaceile 665 i' odocarpaceac 650 Sclaginellaceac 690 Zin~iheraceae 686
:·lusaccae 686 I'odostemaceae 669 Si~arou b aceae 675 Zingihe r ales 684 ,
Nyoporace<~e 680 l'odostemale,; 66S, (,69 Sip honodontaceae 673 686
NyrL:aceae 658 l'o l e!'1on iacc,1e 649 , 679 Smi I acace<le 689 Zostcraceac 61,5, 683
.'lyricales 656, 658 I'olemoniales 677 , 678 Solana ceae 64') , 678 Zygophyllaceae 675
:lyristicilceae 653 Po1ygalaceae 647 , 676 Sonn cratiaceae 670
: tyrotha,ml aceae 657 l'olygala1es 661l , 676 Spargil ni ac eae 686
;1vrsLnaceae 667 l'olygonaceae 61,6 , 660 Sp h enocleac eae 681
~lyrtace a e 670 l'olygonales 658 , 660 Stachyuraccae 662
~ty~tales 667 , 670 l' olypodi a ceae (,91 , 692 Stackhousi:lceae 6 73
l'ontederiace ae 688 Sta p hvleaceae 674
W 697

Cha pter )0 . TAXONONIC LITERATURE AND BIBLIOGRAPHIC AIDS

The descrip tive nature o f taxonomy , t he type method of documentation, and

the prin cip l e of priority have contributed g r e atly to the n ecessity that tax-
onomis ts be bib l iographers as Hell as plant sc ienti sts . The degree to I'/hich
one becomes acqua inted with and is able to handle bibliography wi ll affect , in
large measure, one ' s success as a taxonomist . Most probl ems to be solved by
the taxonomist will involve some ref e r e nce to the voluminous a nd poorly indexed
literature .

A classification scheme of taxonomic l iterature is presented in Section A,

and the high l y selected references in Sec t ion B are arranged accor ding t o this
sys tem. Th e aim of t he presentation of refe r ences in Section B, to bo rrow a
phrase from B. D. Jackson ' s i ntroduction to A Gui de to the Litera tu re of
Botany , is "sugges tive, no t exhau s tive ." The empha sis is on basic r eferences
and l iterature guides and type s of r eferences to be cons ulted . St udy of the
classifica tion system i n Section A,II and analys i s of a question to be answe r ed
should enable one to choose from the selec ted bibliography (Section B) eith er
specific references or types of references to be consulted in th e l ibrary .
}~ny referenc es are annotated as a guide to their cont ent s , purpos e and use.
Only taxonomi c literatur e and basic literature guide s are included here .
References specifically treating morpho l ogy , anatomy, cytology , eco logy, etc.
may be found in the appropriate chapter s in this book. In Section C some of
the ref e r ences useful in organizing a nd pre paring taxonomic papers and sug-
ges tions for t he co lle c t ion and preparation of a reference file are given .

The £0110\"ing outline ~"it h page numbers of appropriate sec t ions of this
chap ter is provided to facilitate t he location o f ref e r ences .

Section A. CLASSIFICAT ION . 699

I. Cla ss i f i cation Used in Libraries 699

A. Call Number for a Specif i c Reference 700
n. General Groups and Examples . . . . 700
II . Classification of Taxonomic Lite ra t ure 701
A. Nomenclature . ... ... .. . 701
B. Terminology and De scription of Taxa 701
C. Identifica tion 701
D. Distribution 701
E. Illustrations .
F. Cl ass i fic ation Systems 701
G. Botanical I nstitutions, Depositories and Collectors 701
H. Current Li terature 702
1. Refe r e nce Lists, Bi bliog r aphies , Guides, Cata l ogs
and Indices . . . . 702
J. General Texts . . . . . . 702
K. Ili s tory and Exp l oration . 702
L. Honographs and Revisions 702

Sec tion B. REFERENCES 702

I. Nomenclat ure 702

A. Rul es (Code) 702
698 30

B. Selected References , Guides and Ind ice s to Plant Names 703

C. Journals . . . . . . . . . 708
II . Terminology and Description of Taxa 708
III . Identification . . . . . 709
A. Floras . . . . . . . . . . 709
1. Guides to Floras and Floristic (.Jorks 709
2 . Selected Regional , State and Local Floras of No rth
America . . . . . . . . . . 709
a . Arct i c and Subarctic North Ame rica 710
b . Eastern North America 710
c . Central North America 710
d . Rocky l>!ountains, Great Basin , SouthHestern States . 711
e. Pacific States . . 711
f . Hal<laiian Islands . 712
3. Other Selected Floras 712
a. Africa 712
b. Arctic and Antarc tic Regions 712
c. Asia . 712
d. Europe ........ . 713
e. Nalaysia , Australasia and Oceania 713
f. North and Central America 713
g. Islands . . . . . , 713
h. South America 714
B. General References for Identification of Families 714
C. General References for Identification of Genera 714
D. Honog raphs, Revisions, Synopses 715
E. Selected Handbooks and Speciali zed \.Jo rks 715
1 . Cultivated Plant s 715
2 . Aquatics . . . . 715
3 . Poisonous Plants 716
4 . Trees and Shrubs 717
5. I-.Teeds . . . . . 719
6 . Selected Re fe rences to Specific Taxonomic Groups
Commonly Trea ted in Separate Handbooks and Nanuals . 719
a . Cactaceae 719
h . Fabaceae (Leguminosae) 720
c . Ferns . . . . . . . • 720
d . Orchidaceae 721
e . Poaceae (Gr;mjnt>a.,). 721
7. Popular treatments 722

IV . Distribut i on . . . . 723
A. General (Generic distribution) 723
B . Specific . . . • ...... 723
1 . 1'1onographs and Revisions (s e e section B,X) 723
2 . Floras and Checklists (see section B, IlI ) 723
3 . Atlases and I ndices of Plant Distribution 723
a . Index (Uorld) .... 723
b . Specific areas and plant groups 723
4 . Computer-Happed Floras nt,
5 . Regional Bibliographies 724

V. Illustrations 725
A. Indices 725
30 699

B. Journals 725
C. Nan uals . 725
726
VI . Botanical Institutions, Depositories anJ Collectors 726
A. Indices and Gui des to Herbaria . . . . 727
B. Guides to Collectors and Type Specimens . . . 727
C. Guides to Botanical Gardens . . . . . . . . . 728
D. Collection Centers (otber than for specimens) 728

VII. Current Literature . . . . . . . . . . . . . . . 728

A. Selected Journals of Inte rest to Taxonomists 728
B. Guides to the Titles of Periodicals . . . . . 729
C. Abstracts, Guid e s and I ndices to Current Literatu re 730

VIII . Reference Lists , Bibliographic Guides , Catalogs and Indices 737

A. Catalogs . . • . . . . . • . . . • . . •• .... 737
B. Bibliograpbies (Selected) . . . . . , . . . .... 738
1. Gene ral Bibliographies. Guides and Subject Ind ices . 738
2 . Regional Bibliographies . . . • .• . . . . 739
3. IHbliograp hics (S pecific Groups and Approaches) 742
C. Cumulative I ndic es to Journals 743
D. Index to Dissertations 744
E. Research Ind i ces 744

IX . General Texts . . . . 744

X. Nonographs and Revisions . . . • . . . . . . . . . • . 745

A. Serial }Ionog raphs and Revisions (including examples of
regional " monographs" or revisions in "floras") 745
B. Guides . . . . • . . . . . . . . . • . . . 746

Sec tion C. Su ggestions for the Preparation of Taxonomic Trea tments and
Reference Files . . . 747
Index to Authors, I ndexes and Guides . . . . . 749

Section A. CLASSIF I CATION

1. CLASSIF ICATION USED IN LI BRARI ES

Tiw systems of class i f ication are currently used in libraries--the Dewey

Decimal and the Library of Congress. In f act, many libraries are in the
process of changing from the older DeHey Sys tem to the LC System, and famili -
arity \"ith both systems is advantageous . The folloHing lists include e x am-
ples of some of the major categories of each system .

DEWEY DECIHAL LIBRARY OF CONGRESS

Natural Science 500 Natural History QH

Biology 570 Botany QK
Botany 580 Zoology QL
Zoology 590 Agricu lture 5
Agriculture 630 Forestry SO
Horticulture SB
flibliography Z
 700 30

For a comparison of the t\-;o systems and other pertinent info rma tion on library
classification systems and theory and botan ical bib liography see SHift. L . H • •
1970, Botanical Bibliographies, a guide to the bibliographic mate rials appli-
cable to Botany_, Burgess Publishing Company , Hinneapolis. Hinnesota .

Since the LC System is so widely used, the £01101-.11ng examples are given
to acquaint the student with the disposition of taxonomic literature and the
meaning of call numbers in this system.

A. Call Number for A Specific Reference

Conard , Henry S. 1905. The \~aterli1ies . A monograph of the genus

Nymp haea . Carnegie Institution of Hashington .

11K 495.N97 C7
Spermatophyta- Systematic
Science Botany Systematics Division Group Author
Q K 495 . N97 C7
B. General Groups and Examples
QK 1 - PERI ODICALS, SOCIETIES QK 97 - COHPREIIENSIVE SYSTEMATIC
QK 1.C2 - California University WORKS
Publications in Botany QK 97 . C2 - de Candolle's Mono-
QK 1.CSS - Chronica Botanica graphiae Phanerogamarum .
QK 1. F4 - Field Huseum of QK 97.E62 - Engle r's Das
Natural Hi story - Botanical Pflan zenrei ch.
Series (Chicago Natural History
Huseum - Fieldiana: Bo tany) QK 101- 474 - GEOGRAPHIC DIVISIONS
(FLORAS AND DISTRI-
QK 11 - INDICES. NOMENCLATORS, BUT:j:ON)
CHECKLISTS QK 110 - North America
QK ll .L 45 - Lemee 's Dictionnaire QK IIO .Al - North American
des Genres des Plantes Phanerogames Flora .
QK 125 - Southeastern U. S .
QK 31 - BIOGRAPHY (Individual) QK 125.S64 - Small, J . K.
QK 3l . G8 08 - Dupree , A. H. ~~nual of the So utheastern
1959 . Asa Gray. Flora.
QK 146 - Alaska
QK 91 - BOTANICAL 1·IORKS OF LINNAEUS QK l46.H84 - Hulten's Flo ra
QK 91 . 569 - Stafleu , F. A. of Alaska and Yukon .
Linnaeus and the Linnaeans. QK 211 - Mexico
QK 2l1.H3 - Hartinez , ~1.
QK 95 - CLASSIFICATION (DISCUSSION Catalogo de Plantas Nexi-
AND COMPARISON OF SYSTEMS) cana.
QK 95.L3 - LmHence, G. H. ~l. QK 281 - Europe
1951. An introduction to Plant QK 281.H4 - Hegi , Flora Von
Taxo nomy . Hittel-Europa .
QK 28l . T8 - Flora Europa ea.
QK 96 - NOHENCLATURE QK 321 - Russia
QK 96. 158 - I nternational Code of QK 321. A36 - Flora URSS .
Botanical Nomenclature .
30 701

QK 495 - SYSTEMATIC DIVISIONS QK 520- 532 - PTERIDOPHYTA

A - Z QK 524 - Systematic Divisions
QK 49S . A17 - Acer QK 525 - U. s.
QK 49S.Q4 - Quercus QK 527 - Gr eat Br i ta in
QK 49S . N97 - Nymphaeaceae QK 529 - Asia
QK 49S . N97 C7 - Conard, H. S.
The \~aterlilies . Z - BIBLIOGRAPHY
Z5356 . T8R42 - Rehder , A. - Bibliog-
raphy of Cultivated Trees and
Shrubs

II . CLASSIFICATION OF TAXONOMIC LITERATURE

A. Nomenclature . Nomenclature i s used here to refer to references deal-

ing not only l..dth rules of nomenclature (i.e . , t he Code) and works of value in
solving nomenclatural problems but also those indices which include the names
of speCies, gene ra and families . References cited in this class should assist
primarily in locating original descriptions, authors. place of publication and
synonomy and . seconda rily, for finding information on monographs, revisions,
distribution, etc .

B. Terminology and Description of Taxa. References helpful in the trans-

lation and/or writing of descriptions (e.g . , glossaries and dictionaries) are
placed here. It is often important to use the older treatments for changes in
usage of terms. For an extens ive classif i cation and definitions of phyto-
graphic terminology and the meanings of specific epithets see Chapters 4 and
6.

C. Identifi cation . Identification is one of the basic aspects o f taxon-

omy . The references cited include a variety of kinds of taxonomic l iterature
useful in identification as well as gui.des to re ferences for identification .
Se lected referenc es also inc lud e those treating native and cultivated plants;
local, state , regional and international floras; and both technical and popu-
lar ,wrks. Such references are also good sources of other information such
as distribution, variation, phenology, ecology , etc .

D. Distribution . The range or geographic distribution of taxa has long

been of concern to the taxonomists (see Chapter 1 6) , but such in formation has
become of increaSing importance in a variety of disciplines . The references
given not only include major >-lorks dealing specifically >-lith distribution but
include examples of types of works which should be examined for distribution
data.

E. Illustrations . Illustration s are particularly useful in the absen ce

of a specimen of a parti cular plant in question , i n in terpret ing type or
other descriptions and may also be selected as lectotypes (see Chapter 3) .
Several sources of information on published illustrations a r e ci ted .

F . Cla ssifica tion Systems (Phylogeny) . Classification systems are

treated in Chapter 28 and an extensive bibliography is included there.

G. Botanical I nstitutions, Depositories and Collectors . It is often

desirable and necessary to locate specific collections (e.g ., types ), borrow
representative material in preparation of monographs and revisions , locate
publications and field notebooks of collectors , compare samples of handl"ritin g ,
702 30

and secure living specimens and o ther data hous ed in bo tan ica l in stitu tions
(herbaria , botanic ga rd ens and lib r a r ies) . Selected in dices to suc h institu-
tions and type s of information are included here .

H. Curren t Lite ra ture (Period i cals and Guides) . Curr en t publications

s uc h a s books are r ather easily located i n s ub ject ca talogs in libraries,
bookse llers ' a nd pub l ishers ' catalogs , and book reviews , but periodical lit-
era ture r e mains t h e most elusive . No si ngle c urrent , comp rehensive and
r e adily usab l e index or abst r ac ting servi ce spec ifi ca l ly for taxonomic lit-
e r atu re exi s ts . The references t r eated here inc l ude the common abst ract ing
and indexing services , selected guides . and jo urna ls of in teres t to taxono-
mi sts .

I . Refe r ence Li st s , Bibliographic Guid es . Ca t alogs and Ind i ces . Included

here are references to the older literatu re, special app ro aches , library hold-
ings, etc . wi t h annotations on possible uses of specific references o r types
of references .

J. Gene ral Texts . The numb e r of general text s in Taxonomy is r ather

l imit e d , but these have a va r iety of uses . They r epresent one of t he best
ways of getting an ov erviet~ of a specific subj ec t and inc lud e valuable bib-
liographies to orig inal sources of in fo rmation. In ad dit ion to general taxo-
nomic text s several texts treating specif i c areas (ch emo taxonomy , taximetrics ,
e t c . ) are a l so i nc lud ed .

K. Histo ry and Exploration . Biograph ie s , itin eraries, co llections (her-

ba r ia) , personal libraries , diaries , f ieldbooks , corresponde nce , handwriting
sample s, etc . of botanists and exp lorers are often impo r tant to present-day
taxonomists not on ly i n loc at ing specimens , decipher i ng lab els , and deter-
mi ning type localities, but also i n unders t anding the philosophy and concepts
of t axa of ear lier worke r s (see discussion and bibliography in Chapte r 2) .

L. Monographs , Revisions , an d Synopses . These comp reh ensive works of

specif i c groups , often representing yea r s of wo r k by the most competen t
a u t h orities , al so are the most d ifficu lt t o l oca te. No sin gle index to such
works exists but some of the bas ic sources , many o f which are dis cusse d else-
whe re, are i nclu ded here.

Section B. REFERENCES

[. NO}lENCLATURE

A. Rules (Code)

Stafleu , F . A. ( chair man of edito rial committee ). 1972 . I n terna t ional Code
of Botanical Nomenclatu re , adopted by the Eleventh International Botanical
Congr ess . Seattle , August 1969 . Regnum Vege tabil e 82 . Ut recht . [The
Code is publi she d in Engli sh, Fr ench , and Gennan; in case o f inconsistency
t he English wo r ding has been a rbit rari l y decided as c orrec t . See Chapter
3 for discussion of Cod e and its use . See pp . 395-397 in t he Code for
list of e dit ions 1-10 an d the a pp endice s of the Code f o r cons erved names .]
HcVaugh, R., R. Ross , & F. A. Sta fleu . 1968. An annota t ed g lossary of botan-
ical nomenclature . Regn um Vege tabil e 56: 1-31. [An uno fficia l glossary of
the more difficult terms used in the Code ( I CBN ) --a valuable aid in inter-
preting the Code . ]
30 703

Inte r national Code of Nomencla ture for Cultivated Plants . 19 58 . Regnum

Vegetabi l e 10. Utrecht. [The rules and recommendations governing the
naming of cultivated plants are presented . The aim of this work, accord-
ing to Ar ticle 3 . p . 11. is "to promote uni fo r mity, accuracy and fix ity in
the naming of agr icultural , silvicul tural and horticultural cultivars (var i-
eties) ... wh ich are normal l y given fancy names . .. , " The IeBN (cited
above), however , sti l l governs the us e of scientific or Latin names of
plants wheth er wild o r cult i vated . ]
Synopsis of proposals on bot anical nomenc l ature . [Pr ior to an International
Botanical Congress a synopsis of proposals to amend or change the Cod e is
published in Regnum Vegetabile . A list of Nomina Conservanda Proposita is
also included (e . g ., see Stafleu, F. A., & E. G. Voss , 1969. Synopsis of
pr oposal s on botanical nomenclature- -Seattle . 1969 , Regnum Vege tab ile 60 ,
Ut re cht . ) . ]

B. Se l ec ted References , Gu ide s and Indices to Plant Names

Bentham, G., & J. D. Hooker . 1862-1883 . Gene r a P1anta rum ad exemplaria

imprimis in herba rii s kewensibus servata difinita . 3 vols . London . [A
valuable reference to names of t r ibes and genera and generic descriptions .
Hany new genera proposed . Addenda ~ Corrigenda in each vo lume . Keys and
descriptions in Latin; for dates of publication of parts an d r eferences to
commentaries see Stafleu , F . A., 1967 , Taxonomic Lit erature . ]
Burbidge , N. T . 1963 . Dictionary of Australian Plant Genera, Gymnosperms and
Angiosperms . Angus and Robertson. Syd ney. Australi a . [A d i c tionary of
the genera of Aus t ralia region including Tasmania. The i nformation treated
includes generiC names (arranged alphabetical l y) ; citation (author and place
of publication); synonomy; family or families (as treated in systems of '.
Bentham & Hooker. Engler & Prantl , Hutchinson); phytogeographi c data
(general distribution--c1assed as endemic . native or natura lized); and
bibliographies . An alphabetical list of families with their inclusive
genera is a l so i nclud ed . ]
Cando l I e , A. P., A.• and C. de . 1824-1873 . Prodromus systematis natura1is
regni vegetabili~ . 17 vo1s . Paris . [See Vol . 1 7 (1873) : 303- 314 for hi sto r y
and analysis and pp . 323-493 for general index . Many new species descr i bed
and valuable monographs i ncluded . See Stearn, I~ . T. (1939) Candollea 8 :
1- 4 ; and a l so Bueck , H. H. (1842-1874) Genera , species et synonyma
Cando Ileana a1phabetico ordine deposita. seu index generalis et specia1is
ad A. P. de Cando lle Prodromum systema tis naturalis regni vegetabilis
(ind ex to the Prod romus) ; and Stafleu, F. (1967) Taxonomic Literature ,
Regnum Vegetabi1e 52 . Utrecht.]
Christensen , C. F. 1905-1965 . Index Fi1 icum sive e numera tio omnium generum
specierumque f 1licum et hydropteridum ab anno 1753 ad finem ann i 1905
de scriptor ium, adjectis synonymis principalibus, a rea geographica . . . .
Hagerup, Hafniae . [I ssued in 12 fasc i c les 1905-1906 with 4 supplements .
This r efe r ence is to pte r i dology what Ind ex Kewensis is to the taxonomy of
flow ering plants . A taxonomic conspectus of all genera and s pe cies of
ferns with bibliographic referen ce , synonomy , and geographic distribut i on .
Sup plement I , iss ued 1913 ( treating names pub lished 1906-1912 ); Supplement
2, issued 1917 (treating the years 1913-1916); Supplement 3 , issued 1934
(incl uding t he years 1917-1933).]
The 4 t h suppl ement was prepared by R. E. G. Pichi-Sermol l i and col-
laborators (1965) an d differs from previous ,"orks in that i t treats only
generiC , infrageneric and specific names validly published between 1934 and
1960 with geograph i c distribution of new taxa ,..rith cross-refe ren ces to
704 30

basionyms . A catalog of literature and indices to abbreviations of names

of au thors and periodicals are also present.]
Copeland , E. B. 1947. Genera. Filicum . The genera of fern s . Annales Cr yp to-
gamici et Phytopathologici. Vol. 5. Chroniea Botaniea. h'altham . 1-1assa-
ch usett s . lA treatment of the f amilies and genera of the Filic ineae
arranged in a phylogenetic sequence . In addition to descriptions , keys to
genera, many references to numbers of species , synonyms, types and r e fer-
ences to the literature are i ncluded.]
Dalla Torre, C. G. , [, II. Ha'(ms (eds . ) . 1900-1907 . Genera Siphonogamarum ad
Systema Engierianum Conscripts. Engelmann. Leipzig. [An enumeration of
taxa above the rank of species arranged according to the Engler System .
Families and genera are numbered consecutively throughout and these
numbers are sometimes used in herbaria for f iling and locating materials
arranged accord i ng to thi s system . All n a me s are fol l owed by author , lit-
era t ure citation , and synonyms. The inc l usion of subfamilies, tribes, e tc.,
as well as the numb e r of s p ecies in each genus, make thi s a valuable refer-
ence . A supplement, appendix and generic index are included . Users should
a lways check the supp l ement and appendix . The index nominum has been up -
dated and entries in th e s upplement and appendix added and is sued as
.Foegister zu De Dalla Torre et Barms Genera Siphonogamarum (1958) . This
inc ludes genus, author , Dalla Torre Harms numbers , family , number of gen-
e ra Hithin family . Synonyms are also included with an indicat ion of the
genus in \~ h ich they a re placed . Thi s is one of the most useful references,
particularly when consulting an herbarium \~ith specimens filed according
to the Englerian system . ]
Dand y , J . E. 1967 . I ndex of Generic Names of Vascular Plants , 1753- 1774 .
Regnum Vegetabile 51. Utrecht . [A list of all generic names first pub -
lished f rom 1753 to 1774 whether in binomial Dr non-binomial works . The
2,998 ent ries of the index were checked with the original publication or
a photocopy of the original work . Arrangement of en tr ies (gen er ic names)
is alphabetical followed by author, date and place of publ i cation , annota-
tion, a nd name of family to \~hic h the genus be l ongs . In addi tion to a
list of ~wrks admissible as sources of generic names , a list of sources
which are not valid sourc es of names is also given. A sys tematic arrange-
ment of genera according to Dalla Torre & Harms, Genera Siphonogamarum
(\~ith modifications), is presented . Both nomenclatur al and taxo nomic syno-
nyms are indicated.]
Engler , A. 19 54 , 1 964 . Syllabus der Pflanzen familie n. Ed . 12 . Vol . 1 .
Bakterien bis Cymnospermen (H. Helchior & E . ~.Jerdcrman n, cds . ) . Vol . 2 .
Angiospermen (H . Helchior, ed . ) . Gebriider Borntraeger. Berlin . [This
is a valuable so ur ce of many kinds of i n formation . In addition to names ,
the sys tematic arrangemen t , des cr ip tion of orders . families , subfamilies ,
etc . and note s on the number of genera and the number of s pecies they in-
clude, ther e is an extensive bibl i ography in cl uding i nformation on anatomy
and morphol ogy, phylog eny, floras , monograph s and revi sions as ~~el 1 as
many valuable illustrations . This is a r eference \~hich should be widely
used by s tud e nts of sy st ema t i cs. Text in German. ]
Engler , H. G. A. , & K. A. E . Prant!. 1887-1 914 . Die naturlichen Pflanzen-
familien . Leipzig. Ed. 1. 23 vols; 2nd ed . 1924- (incomp1ete). [A
survey of all plant g rou ps (except bacteria) , with keys to gen era , descri p-
tions , selected bibliography and many notes on taxonomy and nomenclature .
A useful sununary of information on morphology , anatomy, paleobotany, embry-
ology of groups included . ]
Gray !!e rbarium Card Index (Gray Card Index) . [A card index (also available in
volumes) issued quarterly to subscribers , to all new names and neH
30 705

combinations of vascular plants, except fossils. native to the nel.. ",orid

(see area describ ed below) . The starting date for inclusion is 1 January
1886 (see exception belm") and the first issue was in 1894. The complete
set includes approximately 280,000 cards . Informat ion includes (1) new
taxa--the name, author, reference to place of publication, statement of
range; (2) new comhi nations--new name, bib liographic reference and basio-
nym; (3) base cards--basionym, bibliographic references (I·lith some excep-
tions), new name based on it and other nomenclatural synonyms based on
same type; (4) new lssues - -inclucle type, collector and herbarium.
1 . Geographic area
a . Continental Americas including the immediately adjac ent islands
( see e below) .
b . Greater and Lesser Antilles and Bermuda .
c . Canadian Arctic Archipelago , Greenland, and Iceland (coverage
not comple t e).
d . Aleutian Islands, Pribilof Islands and St. LalHence Island in
the Bering Strait (follO\"s political boundary between United
States and the Soviet Union) .
e . In southern hemisphere island possessions of American nations,
such as Easter Island, Juan Fernandez , Falkland Islands and
So uth Geo rgia. [Hawaiian Is l ands are not i ncluded.]
2. Taxa included
From its start the index has included names of genera, species
and taxa of infraspecific rank, with the inclusion at some time in
the past of subgenera . In 1945 (l.;1th iss ue 186) it I.as decided to
include names of infraspecific taxa published betNeen 1754 and 1886 .
Beginning with issue 261 the Index will include in addition to sub -
genera, new names of sec tions, seri es, and of subdivis i ons of these
(i.e .• star ting with issue 261 neH names of taxa of all ranks that
are present affected by principles of p riority Idll be included).
In addition, a taxon above the rank of species is i ndexed i f the
taxon includes genera I.ith species native to the Index area . Up
to 1970 hybrid groups were indexed i f given a name. Again , with
issue 261 , names of hybr id groups validly published according to
arti c le 40 and conforming to t he Ar tic l es of Appendix I of the
ICRN ( 1966) will be included . Nothomorphs designated by an epi-
thet preceded by a binary name I"ill also be included . (For add i -
t ional details see Shaw, in Taxon 20 : 333-336, 1971 . ) .]
Heller, A. A. 1900. Cata logue of North Anerican Plants North of Hexico .
Ex.clusive of the Lower Cryptogams .
Hooker. J. D.• & B. D. Jackson et 81. 1893-1895 . Inde~ Kel~ensis plantarum
phanerogamarum. 2 vo l s. & 14 supplements. Ox.fo rd . {An index to the
place of publication of generic names or binomials of seed plants published
since 1753. Coverage is worldwide . Volumes 1-2 include names from 1753-
1885 . Supplements are issued at 5- or 10-year intervals (more frequent
recently) . Arrangement of information : (1) genus , author. bibliographic
reference , (2) specific names arranged alphab etically followed by author,
place of publication, and native co untry of plant. Supplements include
date of pub l ication and double author entry, but i ndicat ion of synonomy
has been discontinued. Beginning \<lith Supp l ement 10 this index has taken
ove r the function of Index Londinensis, and references to illustrations are
designated by an asterisk. This is a reference to be consulted b efore pub -
lis hing a nel' name or combination and also a source to locate publication
of a name and thus the original description. The greatest disadvantage is
that names of infraspecific taxa are not included . ]
706 30

See a150 Rouleau, E. (compiler) . 1970 . Guide to Index KeHensis and

its supplements i-xiv . ~Iontrea l. [This in d ex consists of 3 sections--(l)
an alphabetical list of genera of Hagnoliophyta and references to volumes
of Index KeHensis and its supplements in Hhich the gene ra appear; (2) same
as section 1 but treats Equisetophyta, Lycopodiophyta, Polypodiophyta a nd
Psilophyta; (3) list of families and their inclusive genera and an alpha-
betical list of families. (See also the errata and the incerta sedis a t
the end of the vo lume .) This is a welcone time-saving device in that one
no longer needs to search all volumes and supplements for a name . ]
Index Nominum Genericorum . [A project proposed in 1954 to produce a f i le of
names of all genera in all plant groups , ...:ith each entry th e res ult of
painstaking bibliographic research and application of the ICBN . The first
issue (1000 cards) contributed by 14 botanists I"as i n 1955. Information
on each of the ca rds consists of the validly published generic mime, its
author, cita tion of place of publication, as preCise a date as pOSSihle,
type species and its basionYTTl (Hhen appropriate) . Numbers on each card
are a ser ia l numbe r and a numbe r designation of the person \~ho prepared
the card . Class and family in Hhi ch a genus is placed are also indicated ,
As of November, 1972 (Staf1eu, F , A. , 1972, Taxon 21 : 732) all card produc-
tion I"as stopped ! A computer printout reproduced by offset is to be avail
able in 1974 . Th i s consolidated printout l.;ill contain SO-60 , 000 generic
names .
The principles observed in the preparation of this valuabl e index
ac co rding to R. COHan (Taxon 19: 52-54 , 1970) are as £01101.s : "(1) To
avoid rep et ition of existin~ errors in ci tation s , the original literature
for every en try is revieHed I"i th great ca re , (2) Citation of type species
is undertaken only "hen evidence for doing so is inc ont estab le as, for
example. when a genus is published with only a si ngle species or. , . ,,,hen a
subsequent monographer has designated a type species , Ilhere evidence is
not so clear , the type species is not lis ted and the card is rna rked as
' provisional.' Such cards are completed and I"il l be re-issue d as taxo-
nomic studies are published that prov ide the data r eq ui red. (3) Each card
l i s ts the essential nomenclatural synonyms hut validly published taxonomic
synonyms are treated flS separate en trie s , (4) Except for the citation of
family names for each en try, the I ndex sc rupulously avo id s taxonomic deci-
sions . "]
Jackson, E , D, 1881. Guide to the lite r ature of Bo t any . London . (See des -
cription on contents in B,VIll B.]
Kelsey, H, P . , & H. A, Dayton (cds . ). 1942. Standar dized Plant Names . A
revised and enlarged listing of approved scientific and common names of
plants and plant products in American comme rce o r use . 2nd ed . J , H,
NcFarland Co . Harrishurg, Pa. (This edition inc ludes approximate ly 90,000
entries of names o f economic plants and plant products , The arrangement is
alphabetical by plant and/or product name; e . g . , drug plant names are to-
gether and include drug name , scientific ( bo tanical) name and S.P.N . common
name , Cross-referencing is good . Although us e ful in interpreting names in
agricultura l and horticultural ,,,orks, these names are not necessari l y those
in common usage,]
Linnaeus , C. 1753 . Species Plantarum . Ed . 1 , Holmi ae , [The starting point
for priority of the names of vascular plants begins with this publication .
Typification, houever , remains a problem since no types were designated and
not all specimens extant in the Linnaean Herbarium today can be taken as
ty pes . Some ref erences and gui des to the use of Species Plantarum (various
editions) are: Helle r, J. L, & I~. T . Stearn . 1959, An Appendix to the
Species Plantarum of Carl Linnaeus. app ended to the facsimile edition of
 Linnaeus ' s Species P1antarum, 2, London; Savage,S . , 1945, A Catalogue of
the Linnaean Herbar i um , London; Stearn, H. T . , 1959 , An Introduction to
the " Species Plantarum" and cognate botanical Harks of Carl Linnaeus Pre-
fixed to the Ray Society Facsimile of Linnaeus ' Species Plantarum; Stearn,
liT . T., 1961, A ne\ol photographic r':!cord of the Llnnaean Herbarium, Taxon 10:
16- 19; Stearn, \.J. T . • 1966, The use of bibliography in natural history, In:
Bibliograph y and Natural History, T . R. Buckman (ed . ) , Unive r sity of Kansas
Publicat ion Library Series 27: 1-26 . J
Nerrill , E . D. 1949 . Index Rafinesquianus--the plant names publishe d by C. S .
Rafinesque Hith reductions , and a cons ide ration of his method s , obje ctives
and attainments. Arnold Arboretum of Harvard Universi ty . Jamaica Plain,
Hassachusetts .
Pfeiffer , L . 1873-1874 . Nomenclator botanicus . Nominum ad finem anni 1 858
publici juriS factorum, classes, ordines , tribus, familias , divisions ,
genera, subgenera vel sectiones designan tium enumartio a lphabetica . Adjec-
tis auctoribus , temporibus, locis systematicis apud varios, notis literariis
atque etymologi cis et synonymis . 2 vols . in 4 parts . Kassel. [A valuable
guide to names and places of publication of taxa , particularly above the
rank of genus .
Reed, C. F . The fol10\o11ng selected references prepared by Dr . Reed are valua-
ble indices to names , synonyms, classification, bibliographies , monographs
and distrib uti on of fer ns and fern allies :
Index to Equisetopbyta . Part 1, Fossils; Part 2 , Extantes, Index Egui se-
~. Contribution of the Reed Herbarium No . 19 . Baltimore, Hary
land. 1971.
Jndex lsoetales . Boletim da Socie.dade Broteriana . Vol . 27 (2 . a ser i e) .
1953 .
!nde~~rs il eat~ et Salviniata. Bo l etim da Sociedade Broteriana. Vol .
28 (2 . a serie). 1954 . Supplement , 1965 . Vol . 39 (2. a serie) .
!..ndex 5elaginellaru.!!l~. ~Iemorias da Sociedade Broteriana . Vol. 18 5-287 .
1966 .
l.nde.,L Thelyp t eridis . Phytologia 17 : 249-328. 1968. Supplement 1 , Phyto-
logia 17 : 46 5-466 .
Stafleu, F . A. 1967. Taxonomic Literature . A se lective guide to botan ical
publications with dates. cOr.1lTlentaries and types . Regnum Vegetabile 52.
Utrecht. [The entries in this work a r e arranged alphabetically by a uthors
and include location of herbarium and t ypes, publicat ions , bibliog raphy and
references . An index to authors and to publication data i s included in
thi s volume. This publication brings together much information (particu-
larly location of types and publication dates) sca ttered through botanical
literature , much of which is not readily available to Horkers who do not
have access to a s uperior botanical library . This reference is very useful
I~hen used in conjunction \-lith other references cited in this c lass and is a
desideratum for every botan i cal library . ]
Steudel , E . G. 1821-1841. Nomenclator botanicus synonymia plantarum univer-
salis e numeran s ordine alphabetico nomina atgue synonyma tum gene rica tum
specifica et a Linnaeo et recentioribus de re botanica scriptoribus plantis
phane rogamis impo sita . Ed. I, Vol. 1 (1821), Vol. 2 (1824) ; Ed . 2 (1840-
1841), in 13 part s . Stu ttgart. [This is one of the most impo rtant post-
Linnaean indices to flO\" ering plant names (1753-1840) and \Jas the fo re-
runner of Index Kelolens is . Entries are arranged alphabetically and include
genus , au tho;:-fami1ysynonyms, habit , and refe r ences to othe r publications
(e . g ., Roemer & Sch ulte s (RS), Sprengel (5), dcCalldolle (D) . (See Stafleu,
F . , 1967, Taxonomic Literature, p . 458 f or dates of publication, biographi -
cal data and references . )]
708 30

Hillis , J. C. 1973 . A Dictionary of the FloHer i n g Plants and Fern s . 8th

ed. r ev . by H. K. Airy Shaw . Cambri dge Universi t y Press . Camb ridge.
[The 7th edition departed somel... hat from previous editions ( see p r eface of
7th edi tion ) in tha t en tr i es dea l stric tly wi t h taxonomic matters . The
aim of t he 8th edi tion, like the 7th. is to include ever v p ub lished generic
name from 1753 to the present and eve ry family name published f r om t he appear-
ance of Jussieu's Genera Plantarum in 1789. Some s upra - and inf ra-fami lia l
names are also included . Conserved gener ic name s are indicated by an ast er-
isk; reductions to synonomy and many variant s pe llings are a l so present;
brief descriptions of all a ccepted famil ies are given. The arrangement of
names is st r ictly a lphabet ical withou t regard to rank . A genus ent r y in-
cludes au tho r (s) , generic synonym (if so regarded) , family , number of
species , gene r al distribution, and i n some cases anno tations treating spe-
cial mor phological feat ures, economic value, etc . The in c luded key to the
families of flowering plants is based on Eng ler' s Classi fication as given
in Die natUrlichen Pflanzenfamil ien and revised in the 7th edition of his
Syl l abus . This is one of the most useful references f or check ing the
family to Hhich a gen us belongs, var iant s pellings , synonyms, e t c ., and
is used dai l y i n mos t her baria. In addition to the number of correc tions ,
t his e dit ion also incl udes names of inter-gener i c hvbrids of o r chids with
an indication of t heir status and a list of family equivalents be tween
this dict ionar y, t he 12th edit ion of Engler 's Syllabus and Bentham &
Hooker ' s Genera Pl antar um.
C. Journals

Taxon . This journal is published for the Internationa l Association fo r Plant

Taxonomy by t he International Bureau for Plant Taxonomy and Nomenc lature,
Tweede Tran s itorium , Uithof , Ut r echt , Netherlands , and contains many imp or -
tant papers on nomenclat ure (for indices see Ro ul eau, E. [comp i l e r], 1972 ,
I ndex to Volumes 1- 20, Part I , Scientific Names ; and Lowd en , R., 1973 ,
Index to Taxon Volumes 1-20, Part II , Authors and Subjects).

II. TERNINOLOGY AND DESCRIPTION OF TAXA (See also Chapters 4 and 6)

Candolle , A. L. P . de. 1880 . La Phytographie , ou l'a rt de decrire le s vegetaux

consideres sous differ ents points de vue . Pa r is .
Feathe rly , H. I . 19 54. Taxonomic Termino l ogy of the Higher Plants . Iowa
State Co llege Press . Ames, Iowa .
Font Quer , P . 1963. Di c c ion a r io de Botanica. Editorial Labor , S . A. Barce-
lona.
Gray , A. 1879 . Structura l Botany. Hacmil1an & Company . London .
Jackson, B. D. 1928 . A Gloss ary of Botanic Terms , 4th ed. J . B. Lippinco tt
Co . Philade l phia. [A widely used bas ic gl ossary , reissued a number o f
times.]
Lali ren ce , G. H. M. 1951 . Taxonomy of Vascular Plants . The Macmil lan Com-
pany. New York. [See the illustrated glossary of taxonomic terms, pp.
737- 775 . ]
Lindley , J. 1848 . Illus trat e d Dict i onary of Botani cal Terms . London .
[Appears as Book I II Gloss o logy in An Introduction to Botany and later
issued s eparately . Excerpt of 40 pages I>,Hh Introduction by Alice Eastwood
lia s iss ued in 1938 under the auspices of the California Academy of Sciences
and the Gard en Club of San Francisco; this excerpt was repr i nte d in 1964
with a fo rewo rd by J . J . Graham, Schoo l of Earth Sc iences , S t a nford Univer-
sity . ]
30 709

Stearn , W. T. 1966 . Botanical Latin . Thomas Nelson & Sons . London . [A

valuable aid to translation or writing of descriptions . ]
Steinmetz, E. F. 1953. Vocabu lar ium Bot anicum . 2nd ed. Amsterdam . [Sc ien-
rifie terms are in Latin , Greek, French, Dutch , German and English . A very
useful source of equivalents in these six languages.]
Systematics Associ ation Commi ttee for Descriptive Terminology . 1960 . I . Pre-
liminary l ist of works relevant to Descriptive Biological Terminology .
Taxon 9: 245 - 257 .

II I . IDENTIFICATION

A. Floras

1. Guides to Floras and Fl oristic \.Jorks

Blake, S . F ., & A. C. AttoJOod . 1942-1961. Geographical guide to floras of

the world . Parts 1- 2 . U. S . Department o f Agriculture Hiscel 1aneous Pub-
lications 40 1 &. 797. Washington, D. C. (Par t 1 reprinted by Hafner
Publishing Co ., 1963) . ["An annotated s e lected list of floras and floristic
works re lating to vascular plants , including bibliographies and publications
dealing with useful plants and vernacular names ." I ntroduction includes
many helpful notes on bibliographies, guides , dictionaries , etc . used in
compiling this work . Pa r t I --Africa, Australia, I nsular floras, North
Amer ica (including Canada , Nexico and Central America, Greenland, 1-1. I ndie s)
and South America. Part II--Western Europe, European USSR <lnd .all of Asia
including Turkey (not completed) . An index to abbreviations and to authors
is included in each part.]
Frodin , D. G. 1964. Guide to the standard floras of the world . Department
of Botany . Uni ve r si ty of Tennessee . Knoxville . [Mimeographed . This guide,
to be published later (personal communication), is for the most part limited
to publications dealing with complete floras of seed plants for a given area .
I t i ncludes much informati on from Blake and At~·wod (1942-1961), Jackson
(188 1 ) and Goodale (1879). References are grouped int o nine geograph ic
r egions--Ar ct i c and Antarctic; North and Central America; the 1·lest Indies ;
South America; Africa ; Europe ; Asia; Malaysia ; Australasia and Oceania;
Oceanic I slands (including eastern Pacific, Atlantic and I ndian Oceans) .
An index to authors is also included .]
Goodale , G. L. 1879. The floras of different countries . Bibliographical
Cont rib utions , Library, Harvard Univers i ty 9 : 1-12 .
Shetler, E. R. 1966 . Selected Guides to the Hildflo~"ers of North America.
Department of Botany , Smit hsonian Instit ut i on . llashington, D. C. (Himeo-
graphed ). [A guide to selected popular, illustrated books in print as of
1966 . 1
Shetl er , S . G. (ed.). 1973. Appendix D. Provisional List of Basic Floras . In:
A Guide fo r Contributors to Flora North America (FNA). Flora North America-
Report 65 : 01.
Tutin, T. G. , V. H. HeYlwod, N. A. Burgess , O. H. Valentine, S . H. \.Jalters , &.
O. A. Hebb ( eds . ) . 1964+ . Flora Europaea . Cambridge University Press.
Cambridge . [Volumes include list of basic and standard floras fo r Europe . 1

2. Selected Regional , State and Local Floras

of No rth America, North of Nexico
 710 30

a. Arctic and Subarctic No rth Amer i ca

Hulten, E. 1968 . Flo ra of Alaska and Neighbor ing Territories ; A l'!anua 1 of

t he Vascular Plants . Stan ford University Press . Stan ford , California .
Polunin, N . 1 9 59 . Circumpolar Ar ct ic Flora . Clarendon Press . Oxford .
Ihggi ns , I . L, & J . H. Thomas. 1962 . A Flora of th e Alaskan Arctic Slope .
Arctic I nstitute o f North America Special Publication No . 4 . Universit y
of Toronto Press . Toronto .

b. Ea stern North America

Batson, hT • T . 1972 . A Guide to th e Genera of Nat ive and Conunon1y Int roduced
Ferns and Seed Plants of the Southeastern United States e xc lud in g Peninsula
Florida . Published by the a u thor . University of South Carolina . Columbia,
South Carolina .
Fernald , N . L. 1950. Gray ' s Nanua1 of Botany . 8th ed . American Rook Com-
p any . NeH Yo rk . [Corrected pr inting 1970 , Van Nos trand Reinhold Company . ]
Gleas on , H. A. 196 3 . The NeH Britton and BrOHn Illustrated Flora of the
Northeastern Unit ed States and Adjacent Canada . 3 vols . , 3rd prin ting,
slightly revised . Published for the !'leI,] York Botanical Garden by Ha f ner
Pub lish ing Company. Nel" York .
----,c-"CC-' & A. Cronqu ist . 1903 . Nanual of Vascular Plants of Northe astern
United States and Ad jacent Canada . D. Van Nostrand Company, Inc . Prince-
ton , NeH Jersey .
Harvill , A. N •• Jr . 1 970 . Spring Flora of Virgin i a . P r ivat el y pub l ished by
the author . Farmvil le, Virginia.
Harie-Victorin , Frere . 1935 . Flore Lau r enti enne (Quebec) . I mprimede de
La Salle . 1'1ontnial. [In French.]
Radford . A. E . , H. E. Ahles. & C. R. Bell . 1968. Nanua1 of the Vascular
Flora of the Carolinas . Univers ity of North Carolina Press . Chapel Hill ,
North Carolina .
Small , J . K. 1933 . Hanual o f the Southeastern Flora . Published by the
author. NeH York . [Reissued in 1953 by th e Univers i ty of North Carolina
Press, Chapel Hill , North Carolina ; facs imile reprint issued in 2 vols . i n
1972 by Hafner Publishing Company . ]
Strausbaugh, P . D.• & E. L. Cor e . 1952-1964 . Flora of I~e st Vi rgi nia . I ntro-
duct ion and Pts . I-IV . \~es t Virginia Un iversity . ~lorg ant own, Hest Vir-
g1n1a . [Parts I , II revised 1970, 1971 , r e spectively.]
Wood , C. E . , Jr . , & Collaborators . 1958+ . Generic Flora of the Southellstern
United States . Journal of the Arno ld Arborc!tuM. [Keys to genera, illus-
trations , and superb bibl i o graphies . ]

c. Central North America

Corre l l, D. S., & N. C. Johnston . 1 970 . !>lanual of the Vascular Pl ants of

Texas . Texas Research Foundation . Renne r , Texas .
Deam. C. C. 1940 . Flora of Indiana . Department of Conservation . Indian-
apolis , In diana.
Goodman , G. J . 1 958 . Spring Flora o f Cent ral Oklahoma. Un ivers ity o f Okla-
homa Duplicating Service . Norman , Oklahoma .
Jones, G. N. 1963 . Flora of Illinois . 3rd ed. Universi ty of Notre Dame
Press. South Bend, Indiana.
Mohle nbrock , R. H. ( gen . ed . ) . 1967+ . The Illustrated Flora of Il linois .
Southern I llino is Univers ity Press . Carbondale , Illinois . [A mul t ivolume,
illustrated flo ra of I llinois to cov e r all plant gro ups--a lgae and fungi
thr ough floh1ering plan ts -- in progress; volumes appear as complet ed . J
30 711

Rydberg, P . A. 1932 . Flora of the Prair ies and Plains of Central North
America . The Ne\.J York Botanical Garden. Nev York . [Reprinted in 1965
by Hafner Publishing Company, Net.; York . ]
Scoggan , H. J . 1957. Flora of }lanitoba. National Huseum of Canada Bulletin
140 , Biological Se r ies 47 .
Shinners , L. H. 1958 . Spring Flora of the Dallas-Fort Horth Area, Texas .
Published by the author. Southern Methodist Universi ty, Box 473. Dallas,
Texas. [2nd edit ion edited by H. E. Hahler published in 1972 by Prestige
Press, For t IJo rth, Texas . ]
Steyermark. J . A. 1963. Flora of l>liss ouri. IOHa State University Press.
Ames , 1m.]3 .
Voss , E . G. 1972 . Nichigan Flora . Part 1. Gymnosperms and Hanocats. Cran-
brook Institute of Science . Bloomfield Hills , Nichigan.
lla terfall, U. T . 1969. Keys to the Flora of Oklahoma . 4th ed . Published by
the author and distributed by Oklahoma State University Bookstore . Still-
!-later , Oklahoma .

d. Rocky Hounta ins , Great Basin, Southlvestern States

Cronquist, A., A. H. Holmgren, N. H. Holmgren, & J. L. Reveal. 1972. Inter-

mountain Flora . Vol. 1. Published for the New York Botanical Garden by
Hafner Publishing Company . New York. [Other volumes in progress . ]
Davis, R. J . 1952 . Flora of Idaho. hl . C. 13 rmm Company . Dubuque, 10\.[8 .
Har rington, H. D. 1954. Nanual of the Plants of Co lorado . Sage Books .
Denver , Colorado .
Kearney, T. H. , & R. II. Peebles. 1960. Arizona Flora. 2nd ed ., with supp l e-
ment by John Thomas HOI-Jell and Elizabeth I'lcC lintock. Un i versity of Cali -
fo rnia Press . Berkeley, California.
~ioss, E . H. 1959 . Flora o f Alberta. University of Toronto Press. Toronto.
Rydberg, P . A. 1922. Flora of the Rocky Mountains and Adja cent Plains . 2nd
ed. Published by the author , New York . (Reprinted in 1954, Hafner Pub-
lishing Company, Ne1:~ York . 1
I,leber, H. A. 1972. Rocky Nountain Flora. A field guide for the iden ti fica-
tion of the Ferns , Conifers, and Flowering Plants of the Southern Rocky
Hountains from Pikes Peak to Rocky Nountain National Park and from the
plains to the Continental Divide. Colorado Associated University Press .
Boulder , Colorado.
Hooten , E . 0 ., & P . C. Standley . 1915 . Flora of NeH Hexico . Contr ibut ions
from the United States National Herbarium 19.

e. Pacific States

Abrams, L . 1923-1960. An I llustrated Flora of the Pacific States, \~ashington.

Oregon, and Cali f ornia . 4 vols . Stanford University Press . Stanford ,
Califo rnia .
Hitchcock, C. L . , A. Cronquist , N. O1:mbey , & J . \.J . Thompson . 1955-1 969 .
Vascular P lants of the Pacific Nortill"rest. Parts 1-5. University of Hash-
ington Publications in Biology Vol . 17 . Seattle, Hashington .
Hason , H. L. 1957 . A Flora of the />lar shes of Cal if ornia . University of
California Press . Berkeley , California .
Nunz , P . A. , in collaboration \<lith D. D. Keck. 1959 . A California Flora .
Supplement , 1968. Published for the Rancho Santa Ana Botan ic Garden by
the University of California Press . Berkeley, California . [A combined
edition of manual and supplement issued in 1973 . ]
712 30

Peck , H. E. 1961. A Hanual of the Higher Plants of Oregon. 2nd ed . Bin-

fords and l'- lort , Publishers. Po rtland, Oregon .
Shreve, F .• & I . L. Wiggins. 1964. Vegetation and Flora of the Sonoran
Desert . Stanford Universi ty Press . Stanford, Cal iforn i a .

f. Hawaiian Islands.

Degener, O. 1932+. Flora Hm'laiiensis or the NeH I llustrated Flora of the

Hawaiian Islands. Honolulu. [Private l y published by the author, in
progress . ]
Hi llebrand, H. 1888 . Flora o f the HaHa iian I slands ; a description of their
phanerogams and vascular cryptogams. B. Hestermann and Company . Nel']
York.

3. Other Selected Floras (not cited or indexed in Frodin's

Guide to the Florasof the \o]orld)

a. Africa

Brenan , J . P . H. , & A. Exell (eds . ) . 1961. Flora Zambesiaca . Crmm Agents

for Overseas Governmen t s and Administrat ion. London. [Issued i n parts. ]
Fernandes . A. (ed . ) . 1969+. Flora de Hocambique . J unta Investigacoes do
Ultramar. Centro de botanica . Li sboa . · [ In progress, reviev Tax~ n 19 :
430-43 1. ]
Hepper , F . N . (ed.). 1954-1972 . Flora of I']est Tropical Africa . 3 vols .
CrmVll Agents for Overseas Governments and Administration. London . [A
revised edition of Hutchinson & Dalzier (1927 -1936) Flora of Hest Tropical
Africa . Treats plants of all territories of I-Iest Africa south of latitude
l8°N to the Hest of Lake Chad and Fernando Po.]
MerxmUller , 11. 1970 . Prodromus e ine r Flora von Sud\~estafrika . J. Crame r.
3301 Lehre. Germany . (Treats 166 families of flovering plants and ferns of
Southwest Africa . This is issued in parts and is almost complete . ]

h. Arctic and Antarctic Regions

,
Love , A. 1970 . Islenzk Ferdafl6ra ... Almenna B6kafelagid, Reykj avik. [In
Icelandic; revielv Taxon 20 : 181-182 , 1971. ]

c. Asia

Davis , p . II . (ed . ) . 1972 . Flora o f Turkey and the East Aegean Islands . Vo l.
4 . Unive rsity of Edinburgh . Edinburgh at University Press. Edinburgh .
[ I ssued in parts , 4 vo l s. issued to date.]
Iconographia Cormophytorum Sinicorum. Vol. 1. 1972 . Botanical Instit ute of
the Academy of Sciences. Peking . [1157 pages--Bryophytes , Pteridophytes.
Gymnosperms and some Angiosperms ; illu strated; see Ga1ston , A. H .• 1973,
Flora North America, Science 180(4081) : 9 . ]
Komarov , V. L .• & B. K. Schischkin . 1934-1964. Flora U.R.S . S . 30 vols . Acad-
e my of Sc iences of the U.S.S . R., Hoseov and Leningrad . [Flora o f the USSR--
English translation in progress; see review in Taxon 18 : 685 - 708 , 1 969.]
Nasir , E ., & S . 1. Ali Ceds.). 1970? }'lora of \'}est Pakistan . SteHard Her-
barium . Gordon Colleg e, Rawalpindi and University of Karachi , Karachi.
[An illustrated flora issued in parts each treating a separate family . To
date 26 parts have been issued . ]
30 713

Ohwi, J . 1965. Flora of Japan . Smithsonian Institution. Hashington, D. C.

[Translat i on and revision of 1953 Fl ora o f Japan and 1957 Flora o f J apan--
Pteridophyta e dited by F. G. Meyer & E . H. Walker (Engli sh translation).
In addition to keys, description includes phytographic r e sume and histori-
cal sketch of floristic work in Japan, maps, referenc e list o f authors'
names, index to Japanes e plant names, index to s c ient ifi c names and index
to English names (mostly names of families). ]
Smitinand , T., & K. Larsen (eds . ). 1970. Flora of Thailand . Applied Scien-
tific Research Corporation of Thailand . Bangkok. [Issued in parts, incom-
plete; review in Taxon 20 : 171 , 1971 . ]

d. Europe

Clapham , A. R., T . G. Tutin, & E. F . ~.;rarburg. 1968. Excursion Flora of the

British Isles . 2nd ed. Cambridge University Press . Cambridge .
Kuzmanov, B. A., & S. 1. Kozuharov . 1963+ . Flora Reipuhlicae Popularis
Bulgaricae . Bulgarian Academy o f Sci ences . So f ia . [S e e review in Taxon
17: 205- 206, 1968 . ]
Malagarriga, T . 1965. Flora analitica de Barcelona. I . Fa nerogamas. La
Salle Bonanova. Barcelona .
Tutin, T . G., V. H. Hey\olOod, N. A. Burgess, D. H. Valentine , S . M. Halters .
& D. A. ,\"Tebb (eds . ) . 1964+. Flora Europaea . Cambridge University Press.
Cambridge. Vol . 1 (1964) Lycopodiaceae to Platanacea e ; Vol . 2 (1968) Rosa-
ceae to Umbelliferae; Vol . 3 (1972) Diapensiaceae to Hyoporaceae; Vols . 4- 5
in preparation. [Volumes also include notes on Flora Europaea organization ,
list of contributors , list of basic and standard f loras, appendices o f keys
to author abbreviations, book and periodical title abbreviations, gl ossary
and list of English- Latin equivalents, and maps.]

e. Halaysia. Australasia and Oceania

Burbidge, N. T., & M. Gray. 1970 . Flora of the Australian Capital Territory.
Australian National University Press . Canberra. [Eastern Australia; see
revie\" in Taxon 20: 182, 1971.]
Hoore, L . B., & E . Edgar. 1970. Flora of New Zealand . Vol. II . Government
Printing, Hellington, Ne\" Zealand . [Vol. I by Allan, H. H. et al.; review
in Taxon 20: 630, 1971.]

f. North and Central America

Flora Neotropica. 1968+. Published for the Organization for Flora Neotropica
by Hafner Press . Riverside , N e\~ Jersey . [A seri es o f "monographs" o f al l
tropical plants growing spontaneously in the Hestern He mi sphere . Hono-
graphs issued separately without regard to sequence; e.g ., Monograph I--
Cowan, R. S. , 1968, Swartzia (Leguminosae, Caesalpinoideae, Swartzieae);
monograph 2--Cuatrecasas, J ., 1970, Brune11iaceae. Thirteen parts issued
to date.
Sanchez, O. S. 1969. La f lora del Valle de Mexico. Editor i al Herrero
S . A' I Mexico.

g. I s l ands

Adams , C. D. , & contributors. 1972. Flm,Iaring Plants of Jamaica. University

of the Hes t Indies . Hona, Jamaica.
714 30

I-liggins, 1. 1. , & D. H. Porter & contributors . 1971. Flora of the Galapagos

Islands . Stanford Un iversity Press. St anford, California . [lntroduct ion
to physiography , so ils, climate, vegetation o f the Islands . Systematic
treatment has keys, descriptions , dist r ibution maps, illustrations , a
glossary of botanical terms , literature c i t e d and inde x to names . ]

h. South America

Co rrea, H. N. (ed . ) . 1972 . Flora Pata gonica . Instituto Nacional de Tecno-

10gia Agropecuar 1.a . Buenos Aires , Argentina. [Only Compositae . monocoty-
ledons except Graminea e publ ished to date . ]
Lass er, 1'. 1964 . Fl ora de Venezuela. Instituto Botanico . Caracas . [ Inc om-
plete . J

B. General References for Identification of Families

Bailey, L . H. 1917 . The Standa rd Cyclopedia of Jlorticulture . Vol . 1 . The

Macm i llan Company . New York .
Benson , L . D. 19 57. Plant Classificat i on . D. C. Heath and Company . Boston .
[Includes ke ys to orders and fam ilies of Nor th Amer ic an plants. north of
}lexico , but with d esc riptions and discussion of other plant families . ]
Bentham , G., & J . D. Hooker . 1862-1883 . Genera Plantarum . 3 vo1s . London .
[In Latin . ]
Davis , P . H . • & J . Cullen . 1965 . The Identi fication o f FIO\.;erin g Plant
Families , I ncluding a Key to Thos e Native and Cu lt iva ted i n No rth Temper-
ate Regions . Oliver and Boyd . London .
Engler , A. (ll. Helch ior, ed . ) . 1961, . Syllabus der Pflanzenfamilien . Ed . 12 .
GebrUder Born traeger . Ber lin. [Text in German . ]
---;-:-:~c ' & K. Prantl. 1924 . Die natUrlichen Pflanzen familie n. ~:d . 2 .
Leip zig .
Hutchinson , J . 19 59 . The Families of FloHering Plants . Ed . 2 . Clarendon
Press . Oxford .
1967 . Key to t he Fami lies of Flm·,ering Plants of t h e Hodel .
Clarendon Press. Oxford .
I\lil1is , J. C. 19 73 . A Dictionary of the Flm.Jering Plants and Ferns . 8th
ed . r evised by H. f.. Air y Sh<:"(\.;. Cambridge University Press . Cambridge.
[Synoptic key at end of the dictionary is based on Engler's Classification
as given in the Pf lanzenfamilien and revised in the 7th e d. of his Sylla-
bus .]

C. General References fo r Iden ti f ication of Gene ra

Bentham. G. , & J . D. Hooker . 1 862 -1 883 . Genera Plantarum. 3 vols . Lo ndon.

[In La t in . ]
Engler, A. , & K. Prantl. 1924 . Die natUrlichen Pf lanz enfamilien . 2nd ed .
Leipzig . [In German . ]
Hutchinson, J . 1964 , 196 7 . The Genera of Flm.rerin g Plants . Vols. I , II .
Clarendon P r e ss. Ox fo rd. [See also Hutchinson ' s Families of Flmlering
Plants cite d above . ]
Lemee , A. 1 929- 1959 . Dictionna ir e Des cr iptif et Synonimique d es Genres des
Plantes Phanero games. 10 vols. (9 and 10 are s up plements) . [See Vol. SA
for keys to genera; in French; also see appendix--Vocabulaire FranGais-
Anglais--for a French-En glish dictionary to aid in translation . ]
30 715

D. Honographs , Revisions, Synopses

(See Section B XB for guides to monographs and revisions. See also
Bibliographies (B VIII) .

E. SQlec ted Handbooks and Spec ialized Harks

1. Cultivated Plants

Bailey, L. 1I . 1900-1902 . Cyclopedia of American Horticu l ture . 4 vals . The

Hacmillan Company . NeH Yo r k . [Incl ud es keys , description s , illustrations ,
bibliographic r eferences . ]
1914-1917. The Standard Cyclopedia of Horticulture . 6 vcls . (ed .
2 in 3 vals . ); neH edition also issued i n 1933 . The Hacmi llan Company .
NelJ York . [A revision o f the reference above . Synopsis of plant kingdom
and rev is ed keys to faMil i es and genera o f cultivated plants.]
1949 . Hanual of Cultivated Plants. 2nd rev . ed . The Hacmillan
Company . NeH York . [A conveni ent manual for ident ifica tion of cultivated
plants of the United States . ]
--,c--c--' & E. Z . Bailey . 1941. Hortus second . The Itacmillan Company . Nelv
York . [An annotated dictionary of North American gardening and horticul-
tur e ; reprinted in 19t.7 . ]
Gr af , A. B. 1963 . Exotica 3 . Pictor ia l Cyclopedia o f Exotic Plants . Roehrs
Company . Rutherford , NeH Jersey . [This work has 1 2 , 025 i llustrations
(some in co lo r) ar ran ged by family . Brief descriptions , c u ltural notes,
glossary of terms, a section on plant geography , an i ndex to common names,
and an index to scientific nRmes make this a useful reference; no keys . ]
_ _ __ _ 1970 . Exot ic Pl ant }lanua1. Roehrs Company . Ruthe r f ord, NeH Je r-
sey . [Pictorial guide (3600 photo g raphs) o f exotic indoo r plants commonly
found in commercia l cultivat i on . Cult ural notes and p l ant geography , b rie f
descr ipt ions , index to names and a glossary are included.]
HcDonald, E . 1963 . The Horld Book o f House P l ants . The Hor ld Publishing
Company . NeH York . [Cult ura l notes and illustrated ency cloped ia of in -
door plants ; no k eys . ]
Nea l , H. C. 19 48 . I n Gardens of Ha\vaii. Bernice P . Bishop Huseum Special
Publication 40 . Honolulu. [Keys to genera (1.,150) or species and color
key to flOl·lers ; common names and notes on uses, f olkl or e , descriptions of
plants in lIaHaiian gardens . J
Rehder, A. 1940. Hanual of Cultivated Trees and Shrubs Hardy in North Amer-
i ca . 2nd ed . The 1'1acmi llan Company. He\>' York. [This l.oJOrk includes a
synopsis of orders and families ; keys to fa milies, gener a , species ; des-
criptions; a glossary, index, and re fer ences to illustrations . Only Lat in
named taxa are treated . Plants of subtropical and I,'armer temperate regions
are excluded . (See also Rehder , A. , 1949 , Bibliography of Cultivated Trees
and Shrubs , Jamaica Pla in, Hassachusetts). ]
\~yma n, D. 1971. I·lyman 1 s Gardening Encyclopedia . The Hacmi l1a n Company. Net.;
York. [A general reference treating cultivated p l ant s . ]

2. Aquatics

Arher , A. 1 920 . Ha ter Plants ; A Study of Aquatic Angiosperms . Unive rsity

Press . Cambri d ge, l'lassach us etts. (Reprinted as Historiae Natura lis
Classica, 1963, Ivith int roduct ion b y hI . T. Stea rn). (Not a taxonomic
treat ment, but excellent fo r interpretation of morphology and anatomy . ]
716 30

Biswas. K•• & C. C. Calde r. 1955. l!andbook of Common Hater and Harsh Pl an ts
of India and Burma . 2nd ed . Health Bu llet in No . 24 . Calcutta .
Correll , D. S , . & H. B. Correll. 1972 . Aquatic and Het l and Plants of 50uth-
I"estern United States . Envi ro nmen.t Protection Agency . U. S . Government
Print in g Office. \.]ashington , D. C. [S tock No. 5501-01 77. ]
Eyles, D. E., Ex J . L. Robertson, Jr . 1963 . A Guide and Key to the Aquatic
Plants of the Southeastern United States . U. S, Departmen t of the Inte r ior ,
Fish and l.Jildlife Service, Bureau of Sport Fisheries and Hildlife Circular
1 58 . h'ashington, D. C. [Reprint of Public Health Bulletin 286 (1944).]
Fassett, N. C. 1957 . A Nan ual of Aquat ic Plants. University of loJi sconsin
Press . Hadison , Hisconsin . [Revision of 19[10 edition ~"ith appendix by
E . C. Ogden, 1957 . )
Hotchkiss, N. 1964 . Pondl'leeds and Pond~"eed1ike Plants of Eastern North
America . Circular 187 . U. S. Department of the Int erior . Fish and ~.Jild­
life Service . Hashington , D. C.
1965 . Bulrushes and Bu1 rush1ike Plants of Eastern North America .
Circular 221 . U. S . Department of the Interior. Fish and Wildlife Service .
Hashington, D. C.
1967 . Underwater and Floating-leaved Plants of the United States
and Canada . Resource Publication 44. U. S . Department of the Interio r.
Bureau of Sport Fisher i es and Hild1ife. \1ashington , D. C.
Mason, H. L. 1957 . 1\ Flora of the Harshes of California . University of Cali-
f ornia Press . Berke l ey, California .
Natsumara , Y. , & H. D. Harrington . 1955 . The True Aquatic Vascular Plants of
Co lorado . Te chnical Bulletin of Colorado Agricultural Experiment Station
57 .
Monson , P . H. 1954 . A Preliminary Repo rt on Harsh and Aquatic Dicoty l edonous
Flora of IOHa . Proceedings of the IOHa Academy of Sc ience 60 .
Muens~her , H. C. 1944 . Aquatic Plants of th e Un i ted States . Comstock Pub-
lishing Company , Inc. Itha ca, Nel" York . [Keys , illustrations , distribu-
tion maps . ]
Sculthorpe , C. D. 1967 . The Biology of Aquatic Vascular Pl ants . St . Har-
tin ' s Press . NeH York. [Not a taxonomic treatment but a valuable source
o f information for the taxonomist . ]
SteHart , A. N. , L. J . Dennis, & H. H. Gilkey . 1963. Aquatic Plants o f the
Pacific lJorthl-Jest I-Jith Vegetative Key . 2nd ed . Oregon State University
Press . Corvallis , Oregon .
Subramanyam, K. 1962 . Aquatic Angiosperms , A Systematic Account of Common
Indian Aquatic Angiosperms . Botanical Honograph 103, Council of Scien-
tific and I ndustrial Research. New Delhi .
Hinterr i nger , G. S ., & A. C. Lopinot. 1 966 . Aquatic Plants of Illinois .
I llinois State Huseum Popular Scie nce Se ries Vo l. VI . Department of
Registration and Education. I l linois State Huseum Division and Depart-
ment of Conservation, Division of Fisheries . Springfield .

3. Poisonous Plants

Arnold , II . L. 1968 . Poisonous Plants of HaHa ii. Charles E. Tuttle , Rut-

land , Vermont.
Beath , O. A. et a1. 1953 . Poisonous Plants and Livestock Poisoning. \>,1yoming
Agricultural Experiment Station Bulletin 324 .
BroHIl, R . G. 1955. Plants Poisonous to Livestock. Maryland Agricultural
Ex t ension Service Fact Sheet 91.
Choguill, H. S . 1958 . Some Po i sonous Plants of Kansas . Kansas Academy of
Science Transactions 61(1) .
30 717

Co re , E. J.. , et al. 1961. The Poisonous Plants of \.Jest Virginia. l.Jest Vir-
ginia Department o f Agriculture . Charleston , hTest Virginia.
Duncan, W. H. 1958. Poisonous Plants in the Southeastern United States .
Published by the author. Athens. Georgia .
, & T. J. Jones. 1949. Poisonous Plants of Georgia . University of
--G~e~o~r~g~ia Schoo l of Veterinary Nedicine Bulletin XLIX , No . 13 .
Featherly, H. 1. 1945 . Some Plants Poisonous to Livestock i n Oklahoma .
Oklahoma Agr i cultural Experiment Station Circular C-118 .
Fyles , F. 1920 . Principal Poisonous Plants of Canada. Canada Department of
Agriculture, Dominion Experimental Farms Bulletin 39 (2nd ser.) .
Gilkey , H. N. 1958. Livestock-Poisoning Weeds of Oregon . Oregon Agricultural
Experiment Station Bulletin 564 .
Hardin, J . lif . 1973. Poisonous Plants of North Carolina . Agricu l tural Exp eri-
ment Station Bulletin 4 14 (rev . ed . ). Raleigh , North Carolina.
__~~~' & J . M. Arena . 1 974 . Human Poisoning f rom Nat i ve and Cultivated
Plants (ed. 2 rev . ) . Duke University Press . Durham , North Caro l ina .
Heller , C . A . 1953 . Uild Edible and Po i sonous Plants of Alaska. University
of Al aska and USDA Cooperative Extension Service Bulletin F-40 .
Hyatt , N . T ., R. G. Brmffi, & J . H. Herron . 1953 . Some Plants of Kentucky
Poisonous to Livestock . Kentucky Agr icultural Extens i on Serv ic e Circular
502 .
Kingsbury , J . N . 1964 . Poisonous Plants of the United States and Canada .
Prentice- llali. Engle\wod Cl i ffs, New Jersey .
1965 . Deadly Harvest : A Guide to ConmlOn Poisonous Pl ants . Holt ,
Rinehart and Ihnston. New York .
~~ssey, A. B. 1954. Po isonous P l ants . Virginia Polytechnic Institute Exten-
sion Service Bulletin 222 .
Huenscher, \oJ . C. 1939 . Poisonous Plants of the United States . The Nacmil la n
Company . Nel., York.
--0:---" & H. T . Hinne . 1955 . Conunon Poisonous Plants . Cornell Agricultural
Extension Service Bulletin 538 .
Panunel, L . H. 1911. A Manual of Po isonous Plants . The Torch Press . Cedar
Rapids, Iowa .
Schaffner, J . H. 1903. Poisonous and other i njurious plants of Ohio . Ohio
Naturalist IV: 32 .
Sp e rry , O. E. et a 1 . 1955 . Texas Range Plants Poisonous to L i vestock. Texas
Agricult ural Experiment Station Bulletin 796 .
Stevens, O. A. 1933 . Poisonous Plants and Plant Products . North Dakota
Agricultural Experiment Station Bullet in 265 .
l.Jest , E. 1957 . Poisonous P lants around the Home . F l orida Agr i cultural Ex-
periment Station Circular S- lOO.

4. Trees and Shrubs

Bean , \-,!. J . 1970 . Trees and Shrubs Hardy in British Is les . 8th ed. John
Nurray, Ltd. London . [Descript ions hut no keys . ]
Benson , L ., & R. A. DarraH. 1954 . The Trees and Shrubs of the South~lestern
Deserts . 2nd ed . University of Arizona Press, Tucson, and University of
New Mexico Press , Albuquerque .
Braun , E. 1. 1969 . The I~ood y Plants of Ohio . Hafner Publ ishing Company .
NeH York. [A f acsimile of 1961 edition .]
Brown , H. P . 1937 . Trees of the Northeastern United States . Boston .
Brown, R. G.• & M. 1.. . Brown. 1972. Woody Plants of Naryland . [Distributed
by Student Supply Store , University of }illryland , College Park . ]
718 30

Clark , R. C. 1971. The \-foady plants of Alabama . Annals of the Missouri

Botanical Garden 58 : 99-242 .
Coker , H. C. • & H. R. Totten . 1934 . Trees of the Southeastern States . Uni-
versity of North Carolina Press . Chapel Hill, North Carolina .
Core , E . L . • & N. P . Ammons . 1958 . \'!oody Plants in \.Jinter : A Nanual of
Common Trees and Shrubs in I>,Tinter in the Northeastern United States and
Southeastern Canada . The BOXl'JOod Press . Pittsburgh, Pennsylvania .
Dayton , H. A. 1 952 . United States Tree Books : A bibliography of tree iden-
tification . U. S. Department of Agriculture Bibliography Bulletin 20 .
[Guide to references . ]
Grimm , t.J. C. 1957 . The Book of Shrubs . Stackp ole Company . I-!arrisburg,
Pennsylvania .
Harlm~ , H. M. 1957 . Trees of Eastern and Central States and Canada . Dover
Publications . Neh' York . [Corrected edition of 1942 work . ]
1959 . Fruit Key and Twig Key to Trees and Shrubs . [Reprinted by
Dover Publications, Nelv York (trees of eastern North America).]
Harrar, E. S., & J . G. Harrar. 1962 . Gu ide to Southern Trees . 2nd ed .
Dover Publications . New York.
Hosie , R. C . 1969 . Native Trees of Canada . Canadian Forest Service, Depar t-
ment of Fisheries and Fo restry. Available from Queen ' s Printer. Ottal.;a .
( Keys , descriptions, excellent photographs and illustrations and discussion
of forest regions of Canada.]
Kur z, H. , & R. K. Godfrey. 1962. Trees of Northern Florida . Unive rsity of
Florida Pre ss . Gainesville , Florida .
Muenscher , W. C. 1946 . Keys to ~.loody Plants . 5th ed . Comstock Publishing
Company. Ithaca , Nel'; York .
Petrides, G. A. 1958. A Field Guide to Trees and Shrubs . Houghton Mifflin
Company. Boston .
Preston , R. J ., Jr. 1961. North American Tr ees exclusive of Nex ico and
Tropical United State s. 2nd ed . Iowa State University Press . Ames , Im,'a .
Rehder , A. 1940 . Hanual of Cultivated Trees and Shrubs Hardy in North Amer-
ica. 2nd ed . The Macmillan Company . NeH York. [This work excludes
those plants of subtropical and I.;armer temperate regions. Only Latin
named taxa are treated . (See also Rehder , A. , 1949 , Bibliography of Culti-
vated Trees and Shrubs, Jamaica Plain, Nassachusetts).]
Rosendahl, C . O. 1955. Trees and Shrubs of the Upper Midlvest. University
of Minnesota Press. Ninneapolis, Hinnesota .
Sargent, C. S . 1891-1902 . The Silva of North America . A descript i on of the
t rees uh i ch grow naturally in North America exclusive of Nexico. 14 vols.
Houghton Nifflin Company . NeH York .
1905-1949. Manual of the Trees of North America . 2 vols . Dover
Publications . New York . [Reprinted 1961 and 1965, the 1965 reprint I"ith
a tabl e of changes in nomenclature p r epared by E. S . Harrar . J
Standley, P. C . 1920- 1926. Trees and Shruhs of Mexico . Contributions from
the United States National Herbarium 23 .
Stephens , H. A. 1969 . Trees , Shrubs and Hoody Vines i n Kansas . University
of Kansas Press . LaI.. rence, Kansas .
Trelease . H. 1931. Hinter Botany . 3rd ed . reprinted i n 1967 by Dove r Pub -
l i cations . New York.
TreshOlv , M. T ., S . L. \.jelsh, & G. Hoore . 1970 . Guide to the Hoody Plants of
the Hountain States . Brigham Young University Press . Provo , Utah . [In-
cludes summer and winter keys to genera and key to species . ]
Vines, R. A. 1960 . Trees, Shrubs and It/oody Vines of the SouthHest . Univer-
s i ty of Texas Press . Austin , Texas .
 In the United States cooperation betHeen the U. S . Forest Serv i ce and
the s t ate forestry departments has resulted in a number of il lust r ated h an d -
books of forest trees (e.g . , Forest Trees of Flor ida , 9th ed . , Fl o rida Fo r est
Service) . !-lost contain the same illustrations and have a similar format. No
keys are included . These are available from the fo r est services of indivi dual
stat es .

5. \-l'eeds

Chancellor , R. J . 1966 . The Id12ntification of I,Teed Seedlings of Farm and

Garden . Bl achlell ' s Scientific Publications . Oxford . [A guide t o the
identi fic a tion of i mpo r tant l-1eeds of the British Isles and includes keys ,
brief descriptions and illustrations . ]
Delorit , R . J . 1970 . An I l lustrated Taxonomy Hanual of Heed Se eds . Agr onomy
Publications . Ri ver Falls, Wisconsin . [Photographs of seeds . ]
Frankton , C. 1963 . Heeds of Canada . Botany and Plant Path o logy Di v i s i on ,
Science Service , Canada Depar tment of Agriculture . Ottal-la . Publication
948 . [Descriptions , distribution , illustrations ; no k e ys . ]
Gilkey , H . H . 1957 . I>leeds of the Pacific Nor t hl-1es t . Oregon St a te Col l ege .
Corvallis . [A treatment of ,,' eedy membe r s of 46 pla nt families foun d in
Or egon . 'i·,'ashington , I daho and the weste r n por tion of Non tana p l us the
agricultural areas of British Columbia and Alaska . Keys (to families and
to sp e cies) . desc ript i o ns and i llustrations are presented . ]
Hender son, N., & J . C. Anderson . 1966 . Bo t anical Survey Hemo i r 37 . Depa rt-
ment of Agricultural Technical Servi ces . Republ i c of South Afr i ca . (Des -
criptions , d ist r ibution, illustrations ; no keys . ]
1sely , D. 1 960 . \·leed Identification and Control in the North Central St at es .
2nd ed . IOI-1a S ta te Uni versity Press . Ames , 10\.;ra . [Keys , descriptions ,
illustra t ions . notes on control . etc . ]
Kummer , A. P . 1951 . Heed Seedlings. University of Chi cago Pr ess . Chi c ago .
[Keys to 300 I-Ieed seedlings I-lith illustrat ions , des c r i ptions , and notes on
d i str i bu t ion . ]
Neadly , G. R. 10)' . 1965 . \,leeds of Viestern Australia . Department of Agricul-
ture, We stern Australia . Perth . (A genera l conside r ation of W. Aus tralian
Heeds and their control. Co lor plates and drawings and descriptions of
common I.;eedy species are included . 1
Hontgomery , F. H . 1964 . \~eeds of Canada and the No r thern United States .
Rye r son Press . Toronto . [An illustrated field guide . ]
Nuen scher , H. C. 1943 . Weeds . The Nacmillan Company . Nel-1 York . [A general
refe r ence hTith keys , descriptions , bibliography and a discussion of dissemi-
nation , importance . hab i tat and control o f ueeds.]
Reed . C. F . 1970 . Selected t.-l'eeds of the United States . Agricult u re Handbook
No 366 . Agricultural Research Servi ce . USDA . \']ashin gton, D. C. [A collec-
tion of drm-1ings. desc r iptions , and distribut i on maps of 224 species of
'·Ieeds . The twrk is index ed by both scientific and common names and a glos -
sary of phytogr aphic terms is included . No keys are included . ]

6. Selected References to Specific Taxonomic Groups Common ly

Treated in Separate Handbook s and Hanuals

a. Cactaceae

Backeberg , C. 1958-1964 . Die Ca ctaceae . 6 vols . C. Fisher. Jena .

Benson , L . 1969 . The Cacti of Arizona . 3rd ed . University of Arizona Press .
Tucson , Arizona .
720 30

Bravo, H. H. 1937. Las Cactaceas de Nexico . Imprenta Unive rsi taria . Uni-
versidad Nacional de Nexico.
Britton , N. L : & J. N. Rose. 1919-1923. The Cactaceae. Descriptions and
Illustrations of Plants of the Cactus Family. 4 vols. Publication 248 .
Carnegie Institu tion of \-lashington . (Reprinted 1963 in 2 vols. by Dover
Publications , Ne\v York) . [Keys to tribes, genera , species; well illus-
trated.]
Craig, R. T . i945 . The Manunillaria Handbook t-lith Descriptions, Illustra-
tions and Keys to the Species of the Genus Hammillaria of the Cactaceae .
Abbey Garden Press . Pasadena, Califo r nia .
Earle, H. H. 1963. Cacti of the SouthHest : Arizona, Hestern New Mexico,
Southern ilevada, and Ea st ern Cal i fornia . Arizona Cactus and Native Flora
Society Bulletin 4 . Phoen ix, Arizona .
l1arshall, 1.~. T .• & T . M. Bock . 1941 . Cactaceae with Illustrated Keys of All
Tribes , Sub tribes and Genera . Abbey Garden Press . Pasadena , California.
\!en iger, D . 1969 . Cacti of the Southwest . University of Texas Press . Aus-
tin , Texas . [Includes native cacti o f Texas , New Hex ico , Okl ahoma, Arkan-
sas and Louisiana; color plates, keys to genera and species , descriptions,
range, notes on taxonomy, l iterature, field observations , et c . ]

b. Fabaceae (Leguminosae)

Fassett , N . C. 1939 . The Leguminous Plants of 't,Tiscon sin . Unive r sity of 1-1is-
cons in Press . Hadison , IHsc onsin .
Gambill , I>r . G., Jr . 1953 . The Legumino sae of Illinois . Il linois Bio log ical
Nonographs 22(4) ; 1-117 .
Harborne , J . B. , D. Boulter, & B. L . Turner (eds.). 1971. Chemotaxonomyof
the Leguminosae. Ac ad emic Press . Ne\~ York . [A compendium o f symposium
papers I.ith considerab le information and extensive bibliographies on the
taxonomy of legumes . )
Hahler, W. F . 1970 . Hanual of the Legumes of Tennessee . Journal of Tenn-
essee Academy of Science 45 : 65-96 .
Turner, B. L . 19 59 . The Legumes of Texas . University of Texas Press . Aus -
tin , Texas .
Hilbur , R. 1. 1963 . The Leguminous Plants of North Carolina . Technical
Bulletin No . 151. North Carolina Agricultural Experiment Station . Raleigh,
North Carolina .

c. Ferns

Brown, 1'1. (compiler & editor) . 1938. I ndex to North American Fe rn s . Pub -
lished by the compiler . Orleans, Massachusetts . [A catalog of fe r ns and
fern a l lies of North Ameri c a, north of Hexico, \.lith notes on habitats and
general distribution . ]
Copeland, E. B. 1947 . Genera Filicum . Chronic a Botan i ca Company . Waltham ,
Nassachusetts .
Correll , D. S . 1956. Ferns and Fern Allies of Texas . Contributions from the
Tex as Research Foundat i on. Renner , Texas .
Ho lttum , R. E. 1968 . Ferns of Nalaya . In: A Revised Flora of Halaya. Vo l.
II . 2nd ed . Government Printing Office . Singapore .
Knobloch, 1. 1,1 . , & D. S . Correll. 1962. Ferns and Fern Allies of Chihuahua,
He x ico . Texas Research Foundation. Renner, Texas .
l'1axon, U. R. 1909 . North American Flora 16( 1 ): 31- 88 . (Schizaeaceae, Gleich-
en i aceae , Cyatheaceae (pars) . [See also Vol. 16(1) for Ophioglossaceae,
Na rattiaceae, Osmundaceae and Ceratopteridaceae by other authors.]
30 721

Ogden , E. c. 1948 . The Ferns of Haine . The Haine Bulletin 51. (University
of Haine Studies , II. 62) . University of Haine Press . Orono , }laine .
Small . J . K. 1938 . Ferns of the Southeastern States . Science Press . Lan-
caste r, Pennsylvania.
I~agne r.W. R. , Jr. 1965 . Pteridophytes . In : Bib liography. Bioscience 15: 809
810. [A good genera l b i bliography of Pt eridophytes . ]
Wherry, E . T. 1964 . The Southern Fern Gu ide . Doubleday. Garden City, new
York .

d. Orchidaceae

Case , F . H. 1964. Orchids of the Weste rn Great Lakes Region. Cranhrook

Instit ute of Scien ce 'Bulletin 48 . Bloomfield Hills , Hichigan .
Correll, D. S. 1950. Na tive Orchids of North America (North of Nexico) .
Chronic a Botaniea Company . \.j'altham , Hassachusetts .
Dodson, C. H. , & R. J . Gillespie . 1967 . The Biology of the Orchids . The Hid -
America Orchid Congress , Inc . (Includes keys to tribes and sub tribes of
cultiva ted orchids , a compar ison of orchid classification systems , a bib-
liography of "or c h id floras" and a systematic bibliography of the family . ]
Henry, L . K. , & tL E. Baker. 1955. Orchids of Wes tern Pennsylvania . Annals
of the Carnegie ~1 u seum 330: 299-346 .
Jones, O. L., & T. B. Huir (eds . ) . 1969 . Orchids of Australia . (The com-
plete edition drawn in natural color by H. II . I-licholl s \"ith descriptive
text) . Thomas Nelson (Australia) Ltd . Sydney . [Keys to genera , generic
and specific descriptions, synonomy , distribution a n d flowering periods
plus color illustrations . ]
van der Pijl , L . , (.. C. II . Dodson . 1966 . Orchid Flm"ers . their Pollina tion
and Evo l ut ion . Fairchild Tropical Garden anrl the Unive r sity of Hi ami
Press. Coral Gab l es , Florida. [Al t hough not a taxonomic treatment , this
work inc l udes a synopsis of taxonomi c r elations in orchids, a glossary ,
and a bibliography . ]
Schultes , R. E., & A. S . Pease. 1963. Generic Names of Orchids , The i r Ori-
gin and Neaning . Academic Press . New York . [ Illustrat ed and with bibl1-
ography . J
t..'allace, J . E. 1951. The Orchids of Ma i ne. Unive r sity of l1aine Bulletin
Vol. 53 . (University of Hai ne S tudies II. 65) . University of t-1aine Press .
Orono , Haine .
Winterringe r, G. S . 1967 . Ihld Orchi ds of Illinois . Illinois Sta te Huseum
Popular Science Serie s. Vol. VI . Springfield, Illinois .
Withner , C. L. 1959 . The Orchids; A Scientific Survey . The Ronald Press .
Nel" York .

e. Poaceae

Booth , H. E . 1964. Agrostology . The EndOHlnent and Research Foundation at

~lontana State College . Bozeman, Hon tana . [A text with conside rable in-
formation on morphology, anatomy, nomenclature, etc . and many useful bib-
liographies . ]
Bo r, N. L . 1960. The Gra sses of Burma , Ceylon, India and Pakistan (exclud-
ing Bamb us eae) . (A ma nual i ssued as Vol. 1 of I nternational Ser i es of
Honog raphs on Pure and Applied Biology , Division of Botany, R. C. Rollins
& G. Taylor, genera l editor s) . Pergamon Press . New York.
Burbidge , N. T . 1966 . Australian Grasses . 2 vols . Angus & Robertson, Ltd .
Sydney . [A layman ' s guide, profuse l y illustrated . ]
722 30

Chase, A. 1964. First nook o f Crasses . Jrd ed . Smithsonian Institution

Press . v:ash i ngt o n. D. C. [Good introduction to morphology of grasses and
to keys . ]
Gould, F . \ \1 , 1951. Grass e s o f the Southl·.'estern [Jnited States . University
of Arizona Bulletin . ] University of Ari zona Pr e ss . Tucson , Arizona .
1968. Grass Systemat i cs . !-!cGraH-Hill, Inc. Nel" York . [A text-
hook of agrostology, including criteria for classif i cat i on of grasses ,
tri hal arrangement, k e y to genera of U. S . grasses, extens i ve bibliography ,
specimen preparation, nomen c lature , etc . J
Hacke l, E. 1882. Honographia Festucarum Europaearum. Berlin. [Reprinted
1 964 by Hicro?-lethods Ltd ., Yorkshire , England, and Johnson Reprint Cor-
poration, NeH York . ]
Hitchcock, A. S . • & A. Chase . 1951. Nanual of Grasses of the United States .
(2nd ed . revised by A . Chase) . U. S . Department o f Agriculture Niscellan-
eous Puhlicat i on . Hashin~ton , D. C.
Kucera , C. 1. 1961. The Grasses of Hi ssouri. University of Nissouri Studies
35 . Un iv ersity of }lissouri Press. Columbia , Nissouri .
La zarides , N. 1970. The Grasses of Central Australia. Australia National
Unive rsity Press . Canb e rra.
Pohl, R . H . 1966. The gras s es of I owa . I m.,ra State Journal of Science 40 :
341-566 .
Rotar , P . P. 1968 . Crasses of Hawdi. University of HaHaii Press . Honolulu .

7. Popular Treatments

Harks of this type \.,rhich i nclud e books, pamphlets a:ld service leaflets
are so numerous and rep,ional that readers are encouraged to consult local
book stores, literatur e guides, state extension services, state museums,
university presses, and the Government Printing Offic e . The references be-
10\~ are examples of the various tYPQS of popular t.;arks availahle .

Gr eene , \.J . F . , & H. L. Blomquist. 1953. Flo~/ers of the South . University

of North Carolina Press . Chapel Hi ll, North Carolina .
Hard in, J . H. , [. D. 1>1 . Dumond . 1971 . Guide to the Literature on Plants of
North Carolina . ~lisc e llaneous Extension Publication 66 , North Carolina
Agricultural Extension Servi ce . Raleigh, North Carolina .
Justice, \J . S ., [. C . R. Bell. 1968. Hild Flm.,rers of North Carolina . The
University of North Carolina Press . Chapel Hill, North Carolina .
Lemmon , R. S ., [. C . C . Johnson . 1961. HildfloHers o f North America in Full
Color . Hanover House . Garden City, Net", York .
Noldenke, !l . N. 1949 . American Hild FIoHers . D. Van Nostrand Company, Inc.
Ne~.J York.

Rickett , H. 'I!. 1966- . \·!ild Flo\~ers o f the United States . 5 vols . }lcGrat,T-
Hill. Vol. I, The Northeastern Stat e s (2 parts); Vol . 2, The Southeastern
States (2 parts); Vol. 3, Texas (2 parts); Vol. 4, The Southl.Jestern States
(3 parts); Vol. 5, The Northwestern States; Vol. 6, The Rocky Nountain
Region (to be published) . [A series of volumes of illustrations and des-
criptions of selected ",i1J flo\·Jers . ]
Shetler , S . G., [. F . r.lontgomery . 1965. Insectivorous Plants . Smithsonian
Institution , Huseum of Natural History Information Leaflet 447. \</ashing-
ton, D. C .
30 723

IV. DISTRIBUTION

A. General (Generic Distribution)

Dalla Torre, C. G. de, & H. Harms . 1900-1907 . Genera Siphonogamarum ad

Sys t ema Englerianum Conscripta . IJ. Enge l mann . Leipzig . [Info rmation simi-
lar to that of t.a ll is given belm".1
Hutchinson, J . 1959 . The Families of FloHering Plants . 2nd ed . Clarendon
Pr ess . Oxford . [e . g . , Strelitziaceae . ... Tropical America , S . Africa and
t-!adagascar . Strelitzia (S . Africa) . ]
1964 , 1967 . Genera of Flmlering Plants . Vol s . I , II . Clarendon
Press. Oxford. [e . g . • Honopetalanthus ... 8 spp . trop . Africa . ]
\-lillis , J . C. 1973 . A Dictionary of the Flo\"ering Plants and Ferns . 8th ed .
Revised hy H. K. Airy Sha\" . Cambridge Unive r sity Press. Cambridge . [ Li sts
the number of species in each genus , name of f a mily , a nd general dis tribu-
tion ; e . g ., Harfordia Greene and Par r y . Po l ygonac eae. 2 Califo r n ia . ]

B. Specific

1. Nonographs and Revisions (See Section B X)

Host modern treatment s include either a distribution map of mos t taxa

treate d or a statement of range . In some cases a map may be prepared from
the cited specimens examined i n a systematic treatment .

2. Floras and Checkli sts (See Section BIll)

A common feature in checklists and floras is a statement of species dis -

t ribut i on or the i nc l usion of distribution maps . In some cases the genera l
gener ic and/or species distribution is indicated but the introduction or fo r-
mat of each flora sho uld be r ead carefully for criteria for inclusion an d
exclusion of distribution data.

3. Atlases and Indices of Plant Distribution

a. Index (Horld)

rralau. H. (ed . ) . 1969, 1972 . Index Iio i mensis, a world phytogeographic index .
Vol . l--Equisetales-Gymnospermae; Vo l. 2--Honoco tyledonae A-I . Scienti fic
Publishers, Ltd . Zurich . [An index to distribution maps arranged alpha-
betical l y by species but also includes family , tr ibe , genus and subgenus ,
Hhe r e appli cable . Information includes author , title , bibliographic r efer-
ence , date, page and/or map numbers , and geographic area ( area cove r ed by
map) . This is the first at tempt at a comprehensive bibliography \"ith \"orld-
\Vide coverage. The editor , in the preface to Vol . 1, cautions users that
"the aim of the index is mere literature guidance . " ]

b. Spec i fic Areas and Plant Gr oups

Critchfield , H. B. , & E. L. Little, Jr . 1966 . Geog ra phic distribu t ion of

Pines of the Hor ld . U. S . flepartment of Ag ri cul ture Hiscellaneous Publi-
cat i on 991. Hashington , D. C.
Hannig , E., & I! . \']inkler (eds . ). 1926-1940 . Die Pflanz enareale . 5 vo l s .
Fischer . Jena .
724 30

Hara . H. 1958- 1959 . Distribution maps of flm"rer in g plants in Japan. Inoue

Book Company . Tokyo .
Hulten , E. 1958 . The Arnphi-Atlantic Plants and t heir Phytogeograph ic Connec-
t ions. Almquist a nd Hiksell. Stockholm. [Reprint ed by O. Koeltz Anti-
quariat, \~ . Germany.]
1964, 1971. The Circumpolar Plants . Vo l. I--Vascular Cryptogarns ,
Con i fe r s , l·lonocotyledons (2nd ed . ); Vol . II--Dicotyledons . Almquist and
Hiksell . Stockholm . [These volumes include maps and descriptions of, or
commen ts on, variability , taxonomy , nomenc l a ture a nd distribu tion . Each
volume has an extensive b i bliography . i.e . basis for the distribu tion
shm-TTl . ]
1971. Atlas of the Distribution of Vascular Plants in No rthHestern
Europe . 2nd revised and enlarged edition . Almquist and Hiksell. Stock-
ho lm.
Jalas . J ., & J . Suominen . 19 72 . Atlas Florae Europaeae . Vol . l--Pteridophyta
(1972); Vols . 2-5 (1973-1976). Academic Bookstore. Helsinki. [Thi s is to
be an up-to-date work on t he distribution in Europe of all specie s and sub-
spe cies of vascular plants with comments on taxonomy, nomenclature and total
r a n ge . Vo lume 1 consis ts of 1 50 detailed maps showing the distr i b ution of
all species a nd subspecies of Euro pean Pte ridophyt es . ]
Litt l e , E. L .• Jr . 1971 . Atlas of the United States Trees . Vol. l --Conifers
and Important Hardwoods . Hiscellaneous Pub l ications No . 1146 . u . S . Forest
Service, U. S . Printing Office . \\'ashington , D. C. [This volume includes
distribution maps of 200 native trees of contin ental U. S . (including
Alaska) . For Ivide rang ing species , their distribution in Nexico and Canada
is also given.]
Hi rov, N. T . 1967 . The Genus Pinu s . The Rona ld Press Company . New York.
Per ring, F. I! . , [, S . M. Halters (eds . ) . 1962. At las of the British Flo r a .
Thomas Nelson and Sons , Ltd. London. [Critical supplement to the Atlas
of the British Flora, F . H. Perring (ed.) , 1968.]
Radford , A. E . • U. E. Ahles , & C. R. Bel l. 1965 . Atlas of the Vascular Flora
of the Carolinas . North Carolina Agricult ural Exper iment Sta tion Te chnica l
Bul letin 165 . [This atlas " ShONS by means of dot maps, on a county basis .
documented distribution of each species of vascular plant knmvn to occur
without cultivation in the Carolinas . "]

4. Computer-Happed Floras

Cadbury , D. A., J . G. Hal.Jkes , & R. C. Readett. 1971. A Computer-Napped Flora .

A Study of the County o f \~anvickshire . Academic Press . ~ew York .

5. Regional Bibliographies .

Regiona l bibliog r aphies often have references treating species d i stribu-

tion and maps (see Section B VI lIB for addit ional bibliographies) .

Adams , J . 1928 . A IHbliograp hy of Canadian Plant Geography I -VI . Transac-

tion s of Royal Canadian Institute 16: 293-35 5 (19 28); 17 : 103 -145 (192 9) ;
17 , 227-265 (1930) ; 17 , 267-295 (1930); 18 , 343- 373 (1932); 21 , 19-135
(1936) .
Hamann , U., & G. Hagen itz . 1970 . Bibliographie zu r Flora von I>litte1europa.
Carl Hanser Verlag . mjnchen . [References arranged geog r aphical l y .]
Senn. II . A. 1951. A Bi b liography of Canadian Plant Geography . Tr ansactions
of Royal Canadian Ins titute 1946- 1947. Vol. 26 . Publ. 863 . Department of
Ag r icu l ture . Ottawa .
30 725

v. ILLUSTRATIONS

A. Indices

Pritzel , G. A. 1854- 1855 . Iconum Botanicarum Index Locupletissimus . Berlin .

(Supplement issued in 1866). [Host of the re fe rences are included and
emended in Index Londinensis.]
Stapf , O. (ed.~929-19Jl. Index Londinensis to Illustrations of Flowering
Plants, Ferns and Fern Allies . 6 vols. with supplements by W. C. \.Jorsdell,
1941. [This work ~"ith supplements is an index to illustrations of flm..rer-
ing plants and ferns from 1753 to 1935, and includes most of the references
in Pritzel's Iconurn Bo tanicarum Locupletissimus (1855, 1866) . Arrangement
of this index is alphabetical by genus and species with some cross-referen-
cing of generic synonyms . Care should be exercised in accepting illustra-
tions as being authentic as to name. Symbols and abbreviations include:
(1) no symbols = black and \.,rhite illustration of entire plant; (2) asteri sk
colored or partially colored plate; (3) Veg. = vegetative parts (stems,
leaves, roots); (4) Fl. = floHer or its parts; (5) Fr . = fruit or part of
a fruit; (6) Hab. = habit sketch; (7) x = hybrid; (8) ! = teratological
form. Since 1935 references to illustrations have been incorporated in
the references in Index KeHens i s (beginning Hith Supplement 10) and are so
indica ted by an as terisk at the end of an en try . ]

B. Journals

Addisonia . 1916-1964 . [A publication of the Ne\.,r York Botan ica l Garden made
possible through a bequest fro m Addison Brown to establish a journal includ-
in g colored plates of plants of the United States , its terr ito rial posses-
sions and other plants in the New York Botanical Garden and its conserva-
tories Hith descriptions in popular language, notes and synonymy and a
brief statement of knoun properties and uses of th e plants il lustrated .
Publication suspended in 1964 . ]
CurtiS's Botanical Nagazine, containing coloured fig ures \·Jith descriptions and
observations on the botany, history and culture of choice plants. 1787+ .
Royal Horticulture Society . London (as of January 1971 copyright was
assigned to Bentham-Noxon Trust, Royal Botanic Gardens , Ke,.,r) . [In addition
to the excellent illustrations , descriptions, synonomy , distribution , ori-
gin of material for the illustrations, place of cul tivation, etc . are also
included. (See Chittenden, F. J . , 1956, Curtis 's Botanical Hagazine Index,
1787-1947) .J
Huntia . 1964+ . Publication o f Hunt Botanical Library (noH Hunt Ins tit ute for
Botanical Documentation , Carnegie-Hellon University , Pittsb urgh) . [Treats
Horks on illustrations and botanical portraiture . ]
Taxon . 1951+. Journal of the International Association for Plant Taxonomy .
Utrecht, Netherlands . [Often includes papers treatine illustrations and
beginning with Vol. 21( 1 } 1972 nOH i ncludes a series of portraits of botan-
ists \.,rith biographical data and selected reference s pertinent to biographi-
cal literature .

C. Hanuals

Many manuals and floras have excellent illust1."ations, and both A. Rehder ' s
Hanual of Cultivated Plants and L. H. Bailey ' s Cyclopedia of American Horti-
culture incl ude references to illustrations of cult iva ted plants .
726 30

VI . BOTANICAL INSTITUTIONS . DEPOS ITORIES AND COLLECTORS

(See also Chapte r s 31-3 4)

A. Indices and Guides to Herba ria

Clok1e . 11 . N. 1964. An Account of the Herbaria of the Department of Botany

in the University of Oxfo r d . Oxfo rd University Press . London . [An aid
in locating collec tions and h e rbaria ( e . g . , Dilline us. Sherard) ; includes
an excel lent discussion on some of th e outstanding "makers of herbar ia ,"
an index to co l lectors Hith biographica l notes , and references to bib l iog-
raphies . Areas ,. . h e re collectors collected are ind icated, as are samples
of their handl.Jriting f rom labels an d manuscripts . ]
Dandy , J . E. (ed . ) . 1958 . The Sloane Herbarium. Briti sh Huseum (Nat ural
His tory). London . [ This guide to the contents and hi storical informat ion
about cont ribut ors to the herbarium of Sir Hans Sloane preserved at the
Department of Botany a t the British Huseum (Natural History) is divided
into t t"IQ parts . Pa rt I con sists of an a nnota t ed l ist of the Horti Sicci ,
a nd Par t 2 th e biograph ica l sketches of t he con tributor s . The herbarium ,
con tained in 265 vo lumes , r a nges from impo rtant types to specimens of minor
interes t. The indices t o geographi cal loca l ities and botanical names and
the co llect ion of facsimiles of handt.,rriting of many contributors to the
herbarium make this an i mpor tant and us efu l gu ide fo r botanis ts throughout
the world . ]
Kent, D. H. ( compi l e r). 19 57 . British Herba ria: An I ndex to the Locat i on
of Herbaria of British Vascular Plant s with Biographi c al Refer e nces to
Their Collectors . Bo tanical Socie t y of the British Isles . Lond on . [Th e
division of t hi s index into the fo llm.,'ing five sections makes it a read ily
usable guide to locating collections : (1) Abbreviat ion s of names of insti-
tutions; (2) Universit i es , mu seums a nd oth er ins t i tutions ( ar ranged geo-
graphically Hith full address of i nstitut ion and notes on their herb a r ia
a n d literatur e r ela ting to collections) ; (3) Collectors ( dates , ref e r ence
to biographical details , location of i n sti tut ions where collectio ns are
preserved , details of areas wher e plants col lected); (4) Classif i ed index
to the l ocation of col l ections Idth str ong representation of local flora;
(5) Classified index to l ocation of collection s of criti cal gro ups and
genera . A bibliography is also i ncluded . ]
Lanjouw , J ., & F . A. Stafleu (compile r s ). 1964. Index Herb a riorum . Par.t 1 .
The Herbaria of the Horld . 5th ed . Regnum Vegetab ile 31. Utrecht.
[This valuable guide is divided into four section s : (1) List of herba r ia
arranged a lphabetica ll y by city in which they are l ocated and en trie s include
such i nformati on as name of institu tion, date of found ing , number of speci-
mens , type of co llection, important collection s , act i vities (e . g ., teaching ,
re sea rch). staff spec ialti es , and availability of l oans and exchange; (2)
Herhari um abbreviations (acronyms ); (3) Geo grap hical arran gement of her-
ba r ia--arran ged by country and a l so c ity; (4) Gene ral index to personal
names (cur a to r s and staff) .
The a c r o nyms fo r herbaria have become standard and are used in mos t
taxonomic paper s when specimens are ci ted. Th is is a valuab l e reference
for loca ting institut ions. specific herbar ia and spe cial ists . Although
the index is some\~ha t out o f date with respect to the siz e of collec t ions
and s taff , it is an important r eference . The s ixth edit ion is e xpected
soon ! ]
Nic r ofic he ed i tions of herba ria . The Inter Documen t a tion Company , Zug ,
Sldtzerland, has p r epa red microfiches of a numbe r of her baria (e.g., The
{.J'illdenow Herba rium a nd a Systef'latic Index to the Wi 11denow Herbarium) as
30 727

Hell as books, catalogs and lists. NeH edit i on s are announced in Taxon and
a catalog may be requested from IDe , Postrasse 14 , 6300 Zug, SHitzerland.
Th i s method of dissemination of information is gaining favor and makes it
possible f or the contents of many herbaria and books to be consulted by
those Harking outside the large botanical centers.
Savage, S. 1945 . A Catalogue of the Linnaean Herbarium. Taylor and Francis .
London . [List of specimens and inscriptions to 1828. Sequence of catalog
is the same as that of the herbarium \.. hich is essentially the Hay it \.;ras
l eft by the younger Linnaeus in 1783. The ge neric numbers are those added
by Keppst in 1836 and sheet numbers indicate the sequence of specimens in
the herbarium . This cata l og also has 70+ samples o f handwriting and a dis-
cussion of symbols and enumerations . ]
Stafleu, F . A. 1967. Taxonomic Literature . Regnum Vegetabile 52. Utrecht .
[Although basically a guide to dates o f publica ton of a number o f impor-
tant older botanical Iwrks and other information , the l ocation of herbaria
and types are also i nd i cated . The Hark is arranged by author, making it a
readily usable gu i de . ]

B. Guides to Collectors and Type Specimens

(See also refere nces cited above )

Candolle, A. de . 1880 . La Phytographie. Chapte r XXX, pp. 391-462 . Paris .

[An alphab e tical list of authors and collectors and location of their col-
lections as of 1880 . Important s ource of informat i on on l ocation of types.
Huch of this information is incorporate d into ~ Herbariorum.]
Chaudhri, H. N., 1. H. Vegter , & C. del l'la1. 1972 . I ndex Herbariorum. Part
2(3) Collectors I-L . Regnum Vegetabile 86 . Utrecht .
Hedge, 1. C., & J . M. Lamond (eds) . 1970 . Index of Collectors in the Edin-
burgh Herbarium . Edinburgh . [A valuable source for locating specimens pre-
pared by a specific collector or from a specific area . Also includes many
valuable refere nces . ]
Hepper , F ., & F . Neate . 1971. Plant Collectors i n I·lest Afri ca , A Biographi-
cal Index . Regnum Vegetabile 74. Utrecht . LInformat i on included for
collectors is name , nationality, date and place of birth, date and place of
death , references to obituaries and to itineraries, countries in Hhich col-
lections \-Jere made, and herbaria in I.;hich materi als are deposited . Selected
handHriting facsimiles are also included . ]
Hitchcock, A. S . 1934 , 1935 . Location of Type Specime.ns. List I , 19 pp .
(1934); List 2 , 30 pp. (1935). Hashington , D. C. (Hime ographed . )
[Alphabetical list o f 500-600 authors and location of their type specimens
of phanerogams and cryptogams. Although this is a list of authors, not
collectors , in many cases they are the same . The concern of the comp iler
,,,as location of holotypes, not isotypes , etc. Many large herbaria and
botanical libraries have copies of these Harks.]
Jackson , B. D. 1901. A list of the co l le c tors Hhose plants are in the her-
barium of the Royal Botani c Gardens, Ke\L KeH Eullet i n 1 90 : 1-80.
Lanjoml , J ., & F. A. Staf l eu . 1954 . Index Herbariorum . Part II(l) . Index
to Collectors A-D . Regnum Vegetabile 2. Utrecht. [Based on questionna ir e
sent to herbaria allover the t.;orld and on catalogs and other published
Horks , as I .. ell as ac tua l s e arches th r ough some herbar i a . Arrangemen t of
information consists of a list of herbarium abbreviations, general abbre-
viations , and an alphabetical list of collectors I·,ith notes on dates,
country Hhere collections \-Jer e made and herbarium in I-lhich specimens are
deposited . ]
728 30

Lanjouw , J., & F. A. Stafleu. 1957 . ~ . cit . (2) : E-H, I . e . 9 . [For collec-
tors 1 -L see Chaudhri et al., 1972 . ]
LeeuHenberg , A. J. />L 1965 . I sotypes of \-'hieh holotypes were destroyed in
Berlin . h1ebbia 19 : 861-863 .
Shetler, S . G. 1973. An Introduction to the Botanical Type Specimen Register .
Smithsonian Contributions to Botany Number 12 . I<lash i ngton , D. C. [Includes
concept , procedures and standards, and examples of files \.;rhich are designed
to become a union registry . ]

C. Guides to Botanical Gardens

(See Chapter 32 for discussion and references on botanical gardens)

Fletcher, II . R .• D. H. Henderson, & H. T. Prentice. 1969 . International

Directory of Botanic Gardens . 2nd ed . Regnum Vegetabile 63 . Utrecht .

D. Collection Centers (other than for specimens)

EHan (1970) i n his Opportunities in botanical history (Taxon 19 : 825-964)

points out the need for searching out and saving historically valuable mate-
rials as Hell as compiling informational retrieval aids and \.,rriting interpre-
tive accounts. He lists the follm..ring repositories for the preservation of the
papers of botanists and other botanical records:
"1 . Biohistorical Institute , Utrecht : Records of all aspects of the
history of botany and horticulture.
2 . Smithsonian I nstitute , l-1ashington, D. C .: Apart from its Division
of Arch i ves, the Department of Botany (Hhich includes the Hitch-
cock- Chase Agrostological Collection) separately maintains records
of a Hide spectrum of botanical topics .
3. Hunt Botanical Library, Pittsburgh: Records of all aspects of the
history of botany and horticulture .
4. University of \']yoming Archives, Laramie : Records of botanists ac-
tive in the Rocky Hountain Region.
s. Univers i ty of Texas Archives , Austin: Records of the Botanical
Society of America and of botany texts early and modern."
In addition to these sources most herbaria , many libraries and historical
societies have many valuable botanical documents such as field notes , letters ,
manuscripts, etc . which may he of consequence in taxonomic studies.

VII . CURRENT LITERATURE

A. Selected Journals of Interest to Taxonomists

( See also Lm..rrence , 1951, Taxonomy of Vascular Plants, The Hacmillan Company,
Ne\..r York, and guides mentioned belm..r)

Aliso . 1948+ . Journal of the Rancho Santa Ana Botanic Garden. Claremont,
California .
American Fern Journal. 1910+ . Published by the American Fern Society . Port
Richmond, NeH York .
American Journal of Botany . 1914+ . Published by the Botanical Society of
America . Lancaster , Pennsy l vania.
American Midland Naturalist . 1909+ . Notre Dame , Indiana .
Annals of the Hissouri Botanical Garden . 1914+ . St . Lou i s, Hissouri.
Baileya . 1953+. Journal of horticul tural taxonomy . Ithaca , NeH York.
Bartonia . 1908+ . A botanical annual. Philadelphia , Pennsylvania .
30 729

Bauhinia . 1955+. Zeitschrift der Basler Botanischen Gesellschaf t. Basel.

Biota . 1954+. Hagdalena del Mar , Peru .
Blumea. 1934+. A journal of plant taxonomy and plant geography . Le iden.
Boletim da Sociedade Broteriana II . 1922+. Coimbra, Portugal .
Botanical Revie~J (Interpreting botanical progress). 1935+ . Lancaster, Penn-
sylvania .
Botanische Jah rblicher fUr Systematik , Pflanzengeschichte und Pflanzengeographie.
1881+ . Leipzig.
Botanisk Ticlsskrift. 1880+ . Copenhagen . Bruken series .
Botha1ia . 1921+. A record of contributions from the national herbarium, Union
of South Africa. Pretoria.
Brittonia . 1931+ . Published by the Ne\.,r York Botanical Garden for the American
Society of Plant Taxonomists. Net-! York.
Bulletin of the Botanical Survey of India . 1959+ . Calcutta .
Bulle tin of the British Museum, llotany . 1951+ . London .
Bulletin of the Torrey Botanical Club. 1870+ . Lancaster, Pennsylvan ia .
Canadian Journal of Botany . 1951+. Ottawa.
Candol1ea ; Organe du conservatoire et du jardin botaniques de la ville de
Geneve. 1922+ . Geneva.
Castanea . 1936+. Published by the Southern Appalachian Botanical Club.
MorgantOlvn , Hest Virginia .
Contributions of the Gray Herbarium, Harvard University. 1891+ . Cambridge,
Massachusetts .
Folia Geobotanica and Phytotaxonomica . 1966+. Czechoslovak Academy of Science.
Praha .
Journal of the Arnold Arboretum . 1920+. Harvard University . Cambridge,
Hassachusetts.
Ke\" Bulletin . 1946+. Royal Botanic Gardens . Kew, England .
Madrono . 1916+. Published by the California Botanical Society . Berkeley ,
California .
Nemoirs of the Gray Herbarium, Harvard University. 1917+ . Cambridge, Hassa-
chusetts .
Hemoirs of the Net-! York Botanical Garden. 1900+. Ne"'l York .
Hemoirs of the Torrey Botanical Club. 1889+ . New York.
Phytologia. 1933+ . (Designed to expedite botanical publication) . Ne\~ York .
Reim.;rardtia. 1950+ . Journal on taxonomic bo tany , plant sociology and eco logy.
Published by Herbarium Bogoriense . Borgor , Indonesia.
Rhodora . 1899+ . Published by the NeH England Botanical Club . Lancaster,
Pennsylvania .
Si da. 1962+. (A taxonomic journal started by Lloyd H. Shinners and published
privately at Southern Hethodist University, H. F . Hahler and John Thieret,
eds . ). Dallas, Texas .
Svensk Botanisk Tidskrift utgifven Asvenska Botaniska Fureningen . 1907+ .
Stockholm.
Taxon . 195 1+ . Journal of the International Association for Plant Taxonomy .
Utrecht , Nether l ands.
h1atsonia . 1949+. Journal of the Botanical Society of the British I sles .
Oxford .

(See also proceedings of state academies of science.)

B. Guides to Titles of Pe r iodicals

Lawrence , G. H. N. , A. F . G. Buchheim, G. S . Dan ie ls , & H. Dolezal (eds).

1968. Botanico-Periodicum-Huntianum. Hunt Botanical Library . Pittsburgh.
 730 30

Pe nnsy lvan i a . r " Botanico-Periodicum-Huntianum (B-P-!!) is a compen dium of

info rmation on all period ical publications that r egularly con t ain (or in
some period of th ei r his tory . included) articles dea l in g \-lith plan t
scien ces an d botanical literature and Hith the persons Hha have contributed
to botany and its literature." The Hork includes more t han 12 , 000 ti t l es
in 45 languages . Info r mat i on includes a bbreviation , fu ll ti.tle a s given
on title page of Volume I , city of pub licat ion of Volume 1 . volumat ion
(numb e r of volumes published). dutes of publication , report of c hange in
titl e and location of entry in Union List of Serials . The tvo appendices
inc lude a li s t of abb reviations of Hords u sed in t itle abbreviations
(Appendix I) and:). list of language equivalents for geog ra phic names found
in th e t itles or imp rin t of periodicals treated in B- P-U (Appendix II ) .
This is an excellent source for locating cor r ect and complete titles, num-
ber of volumes , a nd s tandard abbreviat ions altd is e xtremely helpful in
deciphering abbreviations o f titles used in other publications . The cro ss-
r efe rencing Hith the Union List of Serials is an aid in locating the name
of the library Hher e a pa r t i cu l ar serial is on deposit . ]
Schl-1arten , L . > & U. H. Ricket t. 1958 . Abb r eviations of titles of serials
ci t ed by botanists . Bulle t in of the Torrey Botanical Club 85 : 277-300 .
[ See also Sup plement I, 88 : 1-10, 1961 . ]

C. Abstracts , Guides and Indices to Current Li terature

A. E . T . F . A. T.-Index . 1953+ . (Association pour l ' Etude Taxonomique de l a Flore

d' Afr ique Tropicale) . Laboratoire de botanique sys tematique de l ' Univer-
site. 28 Avenu e P . Beger. Rruxelles. Belgique . [This is an index to
t..·o rldldde pilb l icat i ons on th e taxonomic botany (phane r ogams) of the r egions
, in Africa south of the Sahara , j,ncl uding South Africa and Nadagascar.
Arrangemen t is alphabetical by fa mily, genus , and s pec ies . Pape rs o f gen-
eral interest concern ing top ics other than new names are list ed a t th e
beginning of each genus or family .
The ind ex inc ludes : " (1) all nel~ taha wit h geographic distribu tion :
fam ilies , genera , soec ies and infraspecif i c divisions, ( 2) nel-1 combina-
tions and nel-1 names . (3 ) the basionyms of the nel-1 combinations and all
other combinations in l-1h ich th e epithet has occurred , \-lith reference t o
the combination , i n orde r to avoid overlooking the n el~ comb i nation , (4)
revis ions of families , genera and critical species , (5) any r efe rences to
papers of taxonomic inte rest." This is a thorou gh and valuable index of
con siderable use to botanist s throughou t the wo rld . ]
Ascher ' s Guide to Botanica l Per i odicals . 197 3 . A. Ashe r and Company, B. \' .
Ke izersgracht 526 Amste rdam 1002. The Netherlands . [A nel~ index to the
contents of selected botanical periodicals . This guide i s to appear in
15 three-tveekly iss ues and "'ill include : (1) a list of t itles of p eriodi-
cals with page r efe r enc es , (2) a list o f the tables of c ontents of each
of the se periodicals Idth every ent r y p r ovided with a re fe r e nce numb er,
(3) an index of au thors ' names with references to en try numbers , (4) a
c umulative authors ' names index, as \~e ll as an index of all plan t names
derived for artic les listed during a yea r. The guide is to li st the con-
tents of ahout 5000 periodicals relating t o botanical studies and treat
about 100 , 000 art ic l es . The subjec t index is to contain ove r 100 ,000
references to scientific names.
30 731

Biological Abstracts - BioResearch Index*

[Biological Abstracts (RA) is a semi-monthly pub lic ation consisting of
abstracts and indexes pub li shed by BioScience!> Informati on Ser vice (B10515) ,
Philade lphia , Pennsylvania . Ahstr.acts of morc than 140,000 papers grouped
under 623 subject catego ries , derived from mo r e than 7600 journa l s origi-
nating in 1 00 coun tries are produced annually .
BioResea r ch Index (BioI) is a month l y publication which prOvides
access to nPre than 100 , 000 research reports annually , in addit ion t o
tho~e covered in Biologi cal Abstracts . Types of lite r ature repo rted in
BioResear ch I nd ex incl ude : symposia , meetings and congresses , reviews,
letters . note s , bibliographies , etc .
Each issue of both publications contains fou r indexes : Au thor, Bio-
systematic , Cross and Subject- - color coded and keyed to the abst r acts and
bibliographic references .
Each iss ue of Biological Ab s tracts contains the following in o r der of
appearance :
(1) Subj ect Guide--alphabetical listing of s ubject categories (~}ith
synonyms) .
(2) List of nel ... books received--i ncludes autho rs' names , title of pub-
lication, date published and price .
(3) Approximately 6 , 000 abstracts arranged acco rdin g to a l phabet ical
s ub ject categories and accompanied by fu ll b i bliog r aphic citations.
Subject areas of particular interest to plant taxonomists are
Bot any--General and Systematic ; General Biology-Taxonomy , Nomen-
c latu re and Terminology; l1useuflls , Botanical Gard e n s , New Journals;
Evolution; Plant Gene tics, Plant Ecology ; Plant Physiology--
Chemical Constituent s (Chemotaxonomy) .
Each issue of BioResearch Index includes i n order of appearance :
(1) Tab l e of Contents
(2) List of the Publication s Indexed wit h CODEN (an alphabet ical code
that repre~ents th at title in the comput e r files) an d Abb reviated
Title .
(3) Bibliography--each citation consists of the jo urnal source, listed
once and set o f f by do tted lines , author(s) tit le , adde d ke~... ords ,
volume number and pagi nation .
Fo rmat and use of indexes - -Indexes to both BA and BioI HUST BE SEARCHED
fo r comp rehen sive coverage of th e syst e matic litera t ure .
(1) Biosystematic Index : This index is incl uded in each issue of Bio-
lo gical Abstrac ts and l\ioResearch Index and enables retr ieval o f
references according to gross taxonomic ca tegories (e . g ., for the
vascular plants by Division , Class , and Family) . An entry appears
in t he index for each organism mentioned in a paper and art i c l es
including descr iptions of new taxa have an asterisk afte r each
re fe r ence numbe r . Re ference numbers are t o t he abst r acts in Bio-
lo gi cal Abstracts and to the bibliographic c it ations in BioResearch
Index . Common, g<!ncric an d specific names are l is t ed in the Subject
Indexes (see discussio n below) . The format of the Biosystematic
I ndex consists of a t llree co lumn e n t r y- -the Gross Taxonomic Category ,

*Adapted from BioSciences Informat ion Service, Guide to the Indexes , 1972 ed .,
Biological Abstracts, Inc . , Philadelphia, "'ith permission . The he lp of
Ann Far r en, l'rofessional RelaLtons Officer of BioScience I n for lnation
Service and her staff in adap U ng, updating and checking this sec tion
is grateful l y acknoHledged.
732 30

a Hajor Subject Area (in Biological Abstracts the subject heading

under \"hich the abstract will appear) and an Abstract or Bibliographic
Citation Number .
Use : (a) Hhere interest is in identifying all studies relating to
a single taxonomic category, check all reference numbers
next to that category in the Biosystematic I ndex.
(b) Hhere interest is i n locating all reports describing new
taxa, check the appropriate taxonomic category and note
all reports with asterisks to the right of the number(s) .
(c) Hhere interest relates to two groups of organisms (e.g.,
fungi parasitic to a vascular plant) match all numbers
appearing next to both taxonomic categories and retain
duplicates.
(d) Hhere interest relates to a gross taxonomic category and a
major subject category (e . g . , Gramineae Grm"th Studies), a
match between each Gramineae reference in the Biosystematic
Index ~dth those appearing under the Cross Index Heading--
Plant Physiology, Growth, Differentiation, offers a faster
and more comprehensive technique--numbers duplicating under
both headings will provide the desired references.
(e) Hhere interest is in locating all taxonomic studies, search
using the taxonomic category and the major subject headings
(e.g . , l'lonocot Gramineae }tonocot Syst) including the abbre
via ted name of the taxonomic category and the word ~
(Systematic) to identify 99% of these reports.
(2) Cross Index: This index is arranged alphabetically by major cate-
gories with reference numbers being repeated under every Cross head-
ing that indicates the content of a specific article. Numbers are
arranged under each Cross Heading in ten columns according to the
last digit of the abstract and bibliographic citation number . The
Cross I ndex can serve both as a bibliography for all reports on a
particular topic area or can be used for match searches between the
Subject or Biosystematic and the Cross Indexes.
I n earl ier volumes of BA and BioI, taxonomic papers were identi-
fied primarily in the Biosys tematic Index . In 1969 the subheading- -
Floristics and Distribution, under General and Systematic Botany also
appeared as a Cross Index heading in BA and BioI. Since 1970 the
Cross Index has also included all reports under the subheadings fo r
Botany , General and Systematic . Example : A search for Systematic
studies on the Dicot family Labiatae could be obtained by checking
the Biosystematic Index under Dicot Labiatae (Synonym Lamiaceae)
Dicot SYst-- tak!" all numbers . If, on the other hand, you ~"ere inter-
ested in all Systematic Studies on Dicotyledones , it Hould be faster
to check the Cross Index heading Botany , General and Systematic-
Dicotyledones . All numbers Hill be listed together in each issue and
cumulated by volume .
(3) Subject Index : This index, appearing in both Biological Abstracts
(B.A . S.I.C.) and BioResearch Index , is a natural language index com-
prised of signi f icant terms from the author's tit l e and supplemen-
tary terms such as names of organisms, geographic l ocation, compounds
and other terms added by the editor to provide indexing i n depth .
The terms are arranged alphabetically in the approximate center of
the 59 character portion of the supplemented title terms which pre-
cede and follow each key word .
 *Genus, species and common names for organisms are found in thi s
index. For those who are interested in locating systematic studies
and/or new taxa for specific organisms, match searching of the num-
bers next to genus-species names in the Subj ect I ndex Hith the higher
taxonomic category-Systematic assignments in the Biosystematic Index
would identify taxonomy studies; (e . g., peas--taxonomy studies Subject
Index--match numbers next to peas and Pisum-Sativum against the num-
bers in the Biosystematic Index next to Dicot Leguminosae Oicot Syst:
retain duplicates) .
(4) Author Index : This index to authors, both personal and corporate, is
included in each issue of Biological Abstracts and BioResearch I ndex.
Multi- authored articles are indexed under each author!s name . For
articles covered in Biological Abstracts, an author address is also
provided--for requesting reprints or other information.
(5) Combination of Indexes: It is often desirable to use several of the
above-mentioned indexes i n combination . This can save time as Hel l
as assure more comprehensive searches.
(a) Subject Index \.,rith Cross Index--effective in locating specific
information in a broad subject field.
Example: Effects of air pollution on a specific organism.

SUBJECT INDEX CROSS INDEX

Reference numbers Reference numbers
under specific term(s) Match for under broad catego ry--
*(genus-species and duplicate Public Health--Envir-
common names) numbers onmentsl Health-Air,
Hater and Soil Pollu-
tion
(b) Biosvstematic Index with Cross I ndex
Example: Cellular studies on Labiatae

BIOSYSTENATIC INDEX CROSS I NDEX

Specific Group (above Broad Subj ect Area
genus-species or common e.g . Cytology and
name) e.g . Labiatae Cytochemistry- Plant

Hatch all numbers next to Labiatae from Biosystematic I ndex with

those appearing under the Plant Cytology Cross heading. Retain
duplicates.

For more details on other searches of one or several of the indexes,

refer to the Guide to the Ind e xes 1972 Biological Abstracts and BioResearch
Index; copies available in libraries; complimentary copies may be obtained
from BioSciences I nformation Service - 2100 Arch Street, Philadelphia, Penn-
sylvania 19103. U. S . A.
Selected references used at BIOS I S for Vascular Plant Taxonomy Nomencla-
ture are Engler's Syllabus (1964), Willis's Dictionary of FloHering Plants
and Ferns (1966), Cronquist's The Evolution and Classification of Flowering
Plants (1968) , and Index Kewensis (and Supplements) .

*A fifth index--the generic index--was issued in January, 1974 . All generic

names (up to 20) mentioned in an abstract are listed here rather than the
Subject Index. The index lists taxa alphabetically and each entry includes
an abbreviated subject heading to \~hich a paper relates and a reference num-
ber . New taxa or records are indicated with an asteris k.
734 30

Biol ogic al an d Ag ricultural Index . 196 /,+. H. W. \.Ji ls on and Company . New
York . ( formerly Agricultural I ndex , 1916-1964). [A subject index to
selected lists of agricultur al periodicals and bulletins . ]
Biosystematic Litera ture; Contribut i ons to a biosyst ema tic literature index
(1945-1964) . Compi l ed by Solbrig , O. T., & T. W. J. Gade lla. 1970 .
Regnum Ve getabile 69 . Utrecht . [A source of referenc es in biosys temat ics
for 1945-1964 but not a comprehensive ind ex; wor ld coverage although cover-
a ge for some coun tries i s uneven . A list of references to chromosome num-
ber s is presented. Refe r ences are cited und er fam ily and/ or gen us names
\-lith f amilies arranged alphabetically . Genera are li sted alphabe tically
unde r each fami l y . Some cross -ref e r e nc ing of families (e . g . , Agavaceae
see Liliac eae , Amaryllidaceae) makes t his a r eadily usable r efer e nce.
Only f l owerin g plants are included ! Hop efully these a llthor s '''ill is s ue
s upplenents to bring this wo rk up to da te and keep it current . This is an
exceed ingly useful reference a nd should be available t o a ll students of
sys temati cs . ]
Excerpta Botanica . A r eference journal to literat ure of taxonomy and c horol-
ogy and of phytosociology star t e d in 1959 as a result of cooperation bet,~een
Gustav Fi she r Verlag of Stuttgart a nd the International Association for
Plant Taxonomy , Ut recht .
[The t wo major sec tions--A . Taxonomy and Cho rology , and B. Soc iology
are issued separa tely (only Sec t ion A is treated he r e) . The four maj o r
classes and s ubcla sses are given below :
Taxonomia , Phyl ogen ia: (1) Taxonomia et Phy loge n ia generalis, (2)
Nomenclature ~ t Termin ologia, ( 3) Cr yptogamae , Sc ripta misce llanea ,
(4 ) Algae, (5 ) Fungi. (n) Lichenes , (7) Br yophy ta, (8) Pte r idophyta .
(9) Spe rma tophy ta, Script a miscellanea , (10) Gymnospermae . (11)
Dicoty ledonae , (12) l!onocotyledonae, (13) Palynologia. (14) Cyto-
t axonomia.
Chorolo&ia ; (1) Scri pta miscellanea i n c l. Itin e r ar ia, (2) Euro pa,
(3) As ia, (4) Af ri ca, (5 ) America , (6) Australia et Oceania , (7)
Regiones arcticae .
Palaeobo tanica : (1) Scrip ta misce llan ea , (2) Tha llophy ta, (3) Bryo-
phyta , (4 ) Pteridophyta, (5) Spe rmatophy ta.
Varia : (1) Herbaria, Horti , Husea , Concilia . (2) Biographiae , Bib-
110graphia. (3) Pra ehis toria e t Ethnobotanica, (II) Personalia .
The material s a r e comp i l ed by geog raphic con trib utors and reviewers
and are edited under the direction of an editorial board. The referen ces
are arranged under each categor y by author and consi s t of author. title ,
b i bliographic ref e r ence and s hort r evi e" in English , Ge rman or French .
Al t hough quite useful it is incomple t e and should he used wi t h other
sou r ces. ]
Hortic ultural Ab st r ac t s . 1931+. Compi led from Horld Liter atur e on Temperate
and Tropical Fruit s , Veget ables , Ornamentals , Pl antation Crops. Common-
wea lth Agricultural Bureaux . Farnham Royal. England . [ I ssued qua r terly
\"i t h annual a uthor and subject indices . Each issue has an author inde x
and subject classifica tion. Particularly a good source for references to
taxonomy, c ultivate d plant s and to r eferences \"hich are pertinen t to hand-
ling of pl a nts for biosystematic re sear ch .]
Index to Ameri can Botanical Lite r a ture . Bul l etin of th e Torrey Botanical
Club . 1 886+ . [This index '''as s tar ted in the ear l y vo lumes of the Bulle-
tin and appeared as notes on pub li cations or under Botanical Literature
and \.as presented as a review , but beginning '~ith Volume 13 it became more
inclusive and \.as presented in the fo rmat given belm,:
30 735

(1) Taxonomy , Phy logeny and Floristics

(a) Algae
(b) Bryophytes
(c) Fungi
(d) Pteridophytes
(e) Spermatophy te s
(2) Ecol ogy and Plant Geogr aphy
(3) Pa leobotany
(4) Norphology
(5) Gene t ics (including cytogenetics)
(6) General Botany (including biography)
References are ar r anged under the above classes a lphab e tically by author .
Botan ical l iterature \Hitten or published by American s or based on Ameri-
can material is included. In 1947 the combined efforts of the e dito r s of
Taxonomic I ndex and the Index to American Lite r a ture ,,,er e acknowledged and
the Taxonomic Index as it appeared in Brittonia , Vol . 20, was the same as
that in the IABL . The index a pp ea r s in each issue of th e Bulletin of the
Torrey Botanical Cl ub and is a l so ava ilable on cards and in bound volumes .
For abb reviations of periodicals see Scln"arten. L . , & H. H. Rickett , 1947 ,
Bullet in Torrey Botanical Club 74 : 348- 356.
Index to Aus t ralasian Taxonomic Literature for 1968 . Regnum Vege tabile 66.
1970 . Compi l ed by 1. K. Fe rguson. [This index to c urren t literature , in-
cluding names of new taxa, which is thought to be relevan t to the study of
vasc ular plant systematics in Austra l asia a nd Polyne sia , is the first in a
proposed series to be issued annual l y . Th e index is mod e led on the Index
to European Taxonomic Literature (cited below) to \.;rhich Dr. Ferguson also
con t r ibutes .
The area covered by the index includes: "Austra l ia , New Zea land , New
Guinea , t he Pacific Islands west of 135°\.)" to 105°E and northl.;rard to t he
Tropic of Cancer, i . e ., including Hicrone sia, the Hawaiian Islands and
Easter Island but exc luding the Ga lapagos , San Fernandes and the Islands
of the east coast of Asia . New Guinea \"as at firs t excluded but the clo se
affinities between the flora of this area and thos e of the Solomon Islands
and northe rn Austral ia seemed to \.;rar rant its i nc lu s i on ." The New Guinea
coverage i s t herefor e probably not as comp l ete as i s desirable.
Disposition of references a nd notes is essential l y the same as that
in Index to European Taxonomic Literature for 1966 (Regnum Vegetabile 53,
1968 (cited below) . The appendix includes a list of the periodical s cit e d
and their abbreviations .
Con tinuati on of this valuable i ndex should be encourage d by workers
thro ugho ut the world . ]
I ndex to European Taxonomic Literature fo r 1965. 1966 , 1967 , 1968 , 1969 , 1970 .
Regn um Vege tabi l e 45 , 53 , 61, 70 . 80 , 83 . Compiled by R. K. Brummi tt et al.
and published under the auspic es of the Flora Europaea Or ganization by the
Interna tional Bureau for Plant Taxonomy and Nomenclature. Ut rech t .
[The goal is a convenient annual "inde x to c urrent lit e r ature relevant
to the s tudy of systematics of vascu lar plants of Europe and adjacent areas"
including references to papers, books and all proposed new names .
This i nd ex is based on books , journals a nd r ep rint s received i n the
l ibrary at Ket" , jo urnal s received at the British Nuseum (Natural Histo r y)
and no t taken at Ke\.;r , with some references being compiled fro m a b s tract i ng
journals and other i ndices . Total number of periodicals cite d in the Index
for 1965 and 1966 is 712 and a n additiona l 196 \"ere added to t h e 1967 issue.
Each subs equent iss ue lists some addi tional ones .
736 30

The language of the text of all papers is that of the title unless
indicated to the con t rary. Translat ion of titles is indicat ed and Cyrillic
scripts are transliterated .
References in each group are arranged alphabeticall y by author and
consist of title , bib l iog r aphic reference, date and brief notes ; l ang uage
of text and summary; presence of maps or photographs, figures; names of new
taxa, etc .
The geographic areas and subject classes treated in this work are as
follows:
(1) Area: "All of Eur ope as defined by Flora Europaea (Vol. 1, p.
xvi) together with Nadel ra and the Canary Islands, Africa north
of the Sahara , the countries a t the east end of the Hecliter r anean
(Israel, Syria, Lebanon, and Jordan), Cyperus, Turkey and the Cau-
casus . "
(2) Subjects a nd Arrangement :
(a) General.
(b) Biography (Personalia). This section i s arranged s ubject by
subject and "includes references to any systematic bo t anists
and collectors , native of, or Horking in, the area cove r ed by
the index . "
(c) Bibliography (Vol. 45 only) .
(d) Herbaria , Botanical Ins titutes, Botanic Gardens . This section
is arrange d alphab etica lly by city and does not include ref er-
ences to personal herbaria which are indexed in the biography
sec tion .
(e) Phy togeography . This sect i on includes papers on vegetation and
plant geography arranged alphabetically by author .
(f) Mapping (Not in Vol . 45). This section of references on map-
pin g \~as put in the General section in the I ndex fo r 1965 but
has been used as a separate class in subsequent issues. Ref -
erences are arranged alphabetically by authors.
(g) Floras and Flori stic Studies. etc . Arranged by coun try, wi t h
Europe divided as in Flora Europaea. Includes notes, i l l us-
trations and lists relevant to area covered .
(h) Chromosome Surveys. This group includes those references
Hhich are of such scope that they cannot be conveniently
listed under systematic groups .
(i) Pteridophyta. Divided into major groups--Psi10psida . Sphen-
opsida, Lycopsida and Fil icopsida with genera arranged alpha-
betically in each g r oup. General references appear at the
beginning of the section .
(j) Gymnospermae. Arranged alphabetically by genus .
(k) Angiospermae . Major divisions are based on family with genera
arranged a lphabetically under appropriate family. This section
includes some cross-referencing of family and generic names .
(1) Appendix . A key to the abbreviation of the periodicals cited
in the index in recent volumes only .
Subjects exc l uded are Phytosociological papers {but does include auteco-
logical references] and Genetics (pure) [but does include papers treating
cytotaxonomy and cytogenetics . ]
Taxonomic Index. 1938-1967 . [This s ubject index was fi r st compiled and dis-
tributed (in mimeograph form) by W. H. Camp as an outgroHth of a card sys-
tem star ted at the New Yo rk Botanical Garden. It was issued only to mem-
bers of the American Society of Plan t Taxonomists. Subjects covered
include (1) taxonomy, (2) floristics , (3 ) phytogeography and (4) phylogeny .
 It originally included only l ichens, bryophy tes and vascula r plants and was
based on literature pertaining to North American plant t axonomy Hhich \~a s
received by !'l. Y.B . G. The inclusiv e geographic area is Panama north\~ard;
i ncluding t he islands of t he Caribbean (exp anded later to H' . Hemispher e) .
Refer e n c es in e ach s ub ject class are arranged a l phabetically by author \dth
a fe\~ notes by the edi tor . The fi rst nine vo lumes t"ere mimeogr aphed . Later
a n arrange men t between t he Torrey Botanical Club an d the Ame rican Soc ie t y
of Plan t Taxonomists resul ted i n a join t staff conp ilin g and ed iting this
bibliographic d iges t of all current botanical literature either origin ating
in the Hestern Hemisphere or pertin e nt to its plants. Both the Taxonomic
I ndex and Inde x to American Botanical Literature (publish ed in the Bulletin
of t he Torrey Botanical Club) t."e r e de rived f r om this compi l ation , and the
I ndex a ppeare d a s a regular feature in Brittonia beginning with Vol. 20
(19 51). In 1967 , the ASPT voted to discon t i nue t he Taxon omic Index but the
Index to America n Bo tan i cal Literature i s sti l l pub li shed i n each issue of
the Bulletin of the To rrey Botanical Club. ]

VIII . REFERENCE LISTS , BIBLIOGRAPHIC GUIDES , CATALOGS AND INDICES

A. Catalogs (Library Hold ings)

These sources are useful in l ocating specific books a nd oth er refer ences
(particularly o l d l ite r ature) which need to be cons ulted, b ut they may also
be used as bibli ograph ic indic es .

Bibliographic Contributions from the Lloy d Library . Vo l. I (1911-1914), Bib-

liography rela t ing to the Floras ; Vo l . 2 (1914-1917 ), Bi b l iog raphy r e l a ting
to Bo tany, exclusive of Floras . Periodicals and books and pamphle t s of
authors A- H; Vo l. 3 , continuation of Vo l. 2 , Authors N- Z. CinC i nna ti , Oh io .
(See also Bull e tin of th e Lloyd Library and Nuse um o f Botany, Pharmacy and
Hateria Hedica, Vo l. 34 (1936) , Catalogue of Pe riodical Literature of the
Lloyd Li brary by Ai ken , \.1. H., & S . Haldbott. The cata l og of pe riodi cals
treats those received betl~een 1914 and 19 35 . J
Ca t alogue of Botanical Books i n the Co llect ion of Rachel NcHaste r s Hiller Hun t.
Vol . 1 . Print ed Books 1477-1700 , comp iled by Jane Quinby (1958); Vo l. 2 ,
Pa r t I, I ntroduction to Printed Books 1701-1800, Part 2 , Pr in t ed Rooks 1701-
1800 , compiled by Al lan Stevenson (1961) .
Catalogue of the Libra r y of the Arnold Arbore tum of Harvard Unive rsity . Com-
piled by Ethely n H. Tucker . Vol. 1 (1914) Serial Publications and Author
Catalogue; Vol . 2 (1917) Supplemen t t o Volume 1 and Subject Catalo gue [very
valuab l e guide to o lder literature , pa r ticular l y of Hoody plants and in-
clud es a ll t he mate rials in Vo l. I plu s the additions durin g 1914-1917 ] ;
Vol. 3 (19 33) Ser ia l Publications and Au t hor - Title Supplement Cove r ing
Access ions of 1917-1933 .
Catalogue of the Library of the Royal Botani c Gardens , KeH . Bu lle tin />1is -
cellaneous I n format ion (KeH Bulletin ) . Addit ional Series 3 (1899 and
supp lement 1898-1915 ).
Cata logue of the Printed Books and Pamphlets i n the Li brary of the Li nnaean
Society of London , 192 5. [No manuscripts catalogued ; listed by a utho r
or journal titl e , supp lemen t at end . Horks from Linnaeus's own lib rary
a re so indicated . ]
Hall , E. C. 1970 . Printed Books 1481-1900 in the Horticultural So ciet y of
New Yo rk. The Horticultural Socie ty o f Nel.J York . New York . [Al though
the 30 00 \wrks in this collection of r eferences are in the horticult ural
 738 30

fie ld, many r are and v aluab l e books o f great importan ce to taxonomic
r esearch are present . This " short-title " ca talog i s arranged a lphabeti-
cal ly by author and should be consulted \"hen tryin g to lo cate books pub-
lished betl-Ieen 1481-1900 . The library als o has hold ings of many r efer e nce
so urc es such as bibliographies, libra r y ca talogs, etc. \~hich are listed on
pp . 243-277. This list include s many of the basic bibliographic re f erences
to be consulte d in library research, many o f vhich a r e mentioned i n this
c hapter . ]
\~ood\·Ja rd , B. n.• & A. C. Tm~send . 1903- 1 940 . Ca talogue of Books , Hanu scripts .
Ha ps and DraHings i n the Brit ish tluseum (~Ia tura l History ) . 8 vals . [VoL
1-6 facsirrd l e r ep r i nt in 1964 by Stechert I!afner Service Agency , Inc . New
Yo r k . ]

IL _B ibliographies ( Selectecl.)

1. General .Bibliog raph ies . C; ui des and Subject Indices

Bihliogr aphia Huntiana (in pr epa r a t ion) . (This is to be a multivolume (H'?)

annotate d guide to the \w r1d ' s botanical li t erature published during 1730
to 1 840 . It \.rill include comprehensive t r eatments of all pertinent books ,
p a mphl ets, r eprints , a nd effec t ively publi shed ~xsiccatae and Hi ll also cite
the period i ca l literature of the authors concerned and published during this
pe riod . An attempt Hill he made to treat !1Jl books and articles dealing
h' ith plant s from a botanical point of v ie'.-l . Base l ist of refe r e nces has
been compiled from the foiloHin g sources : PritzeI, Thesaurus li teraturae
hotanicae , Eds . 1 & 2 (1851 , 1872) ; Halle r, Bibliotheca botanica (1771
1772); Nissen , Die h otanische Buchillustra t ion (19 51); Kro k . Bibliotheca
, bo tanica s uecana (1 923) ; Soulshy, A c atalogue of \"orkS of Linnaeus , ed . 2
(1933) ; Jac kson , Guide to literature o f Bot any (1 881); a nd the Catalogue
o f bo tanical books in the li b r a r Y of Rachel /'leMas t e r s Hiller Hun t.
I n addition to tit l e , biographical ske tch of authors , references to
p ub lished biographical materials , facsimi les of title pages of ce rtain
\wrks, etc . , much information on artists , p rinter s , pub li shers , e ngravers
and t he like i s also to be included . [See HcVaugh , R., 1964 , Announce ment:
Rib lio graphia 1!untiana , Huntia 1 : 17-24.]
Bo tany Subject I nd e x. Compiled by USDA library , on index cards , re produced
in 1958 i n 15 volumes by Hic r ophotor,raphy Company , Boston , Hassac husetts ,
currently avai l able f r om G. K. Hall a nd Co mpany, Boston . [InforI'lation is
Ho rldHide in scope and re fere n ces a r e ar ranged by subjec t, names of coun-
tries, scientific names and some vernacula r names . Genera a r e listed
under the family, hut are al so c r oss-refe r enced . Each volume has a table
of conten ts .]
Jackson , B. D. 1881. Guide t o t he Lite r a ture o f Botany; being a c l assified
se l ection of botanical \-Iorks in c luding nea rly 6,000 titles not given in
Pri tzel' s ' Thesaurus '. London. ( facsi mile reprint by Hafner Publishing
Company , New York, 1964 ) . [Th is classif i e d index coverS much of the lit-
erature betHeen 1871 and 1881 plus the a ddition of 6 , 000 titles to th e
period cove r ed by Pritz el ' s Thesaurus . The detai led classifica tion of
literatur e a nd the scope of literature make the index a valuable s ource
fo r informat ion i n a number of botanical disciplines . The morpho l ogy,
descr ip tive botany , a n d flo ra sections a r e part i c ularly useful . TIl is
should be used \-lith Pritzel fo r bes t coverage . Jackson cha r acterized
t his Hark as " s ugges tive , not e xha ustive ."
La\"rence , G. H. H. 1951. Taxonomy of Vascula r Plants. Chapte r 1 4 , pp . 284-
331. The Nacmillan Company . Net-! York .
30 739

Pritzel, G. A. 1851. Thesaurus Literaturae Botanicae omnium gen tium iud e a

rerum botanicarum il1itils ad nostra usque temp~ra quindecim mil1ia operum
recensens . Ed. II (partly by C. H. J . Jessen, 1871-1877) , [An ou t standing
guid e to chief botanical literature up to 1851. Part I --authors and titles
and r efer ence numbers (note addenda); Part II--systematic (s ubject) i ndex ;
Part III--author index . Ed. 1 (181,7-1882); cd . 2 (cited above) ; see also
Taxon 22 : 119-126, 1973, for biographical notes on Prit;::el and his The-
saurus by Frans Stafleu . The ' Thesaurus' should be used I"ith Jackson
(1881) . 1
Staf leu, F . A. 1967 . Taxonomic Literature . A selective guide to botan ical
publicat ions 1,,1th dates , commentaries and types . Regnum Vegetabile 52.
Ut recht. [A guide to selected older literature of significance to present -
day taxonOll1y . It in c ludes much valuable information on dates of pub lica-
tion, conunentaries on Horks and their authors, as Hell as data on t he loca-
tion of types and herbar i a . The bibliog raphies and indice~ to tit le s and
authors make this a val uable and usable reference . ]
S\~ift , L . H. 1970 . Botanical Bibliographies; A guide to bibliographic
materia l s applicab l e to botany . Burgess Pub l ishing Company . Hinneapo lis ,
Hinnesota . [An outstanding guide to bibliographic techniques and sources
for botany. Parts I an d II are particularly useful to taxonomists . l

2. Regional Bibliographies

Nany of these sources are not of as limited usage as might be cons trued
from their titles, e.g ., many include important monographs, revisions , distri-
bution maps, etc . for taxa o f interest \.,hich do not even occur in the area
surveyed (see also guides to current literature in the preceding sec t ion) .

Boivin , B. 1967(?) . Enumeration des Plantes du Canada . Provancheri a No . 6 .

Hemoires de 1 ' Herbier Louis-Harie. Universite 1.ava1. Quebec. (ex tracted
f rom Naturaliste Canadien, Vols . 93(3 , 4,5 , 6) . 24(1,l! , 5) . [ " An enumeration
of the species , varieti es, forms and hybrids kno\lll as spontaneous in Canada .
Nearly all genera are preced ed by an essential bibliography of recent mono-
graphs and other main papers relevant to each genus in Canada. Each taxon
is follOl"ed by an abbreviated sununary of its distribution north of the U. S.A .
A distinction has been made bet\~ een verified and unverified distr i bu t ions
and, in the latter case, whether or not justifying collections have b een
located yet . "
In addition to being a valuable catalog of Canadian plants , this
reference includes an excellent bibliography and is of considerabl e aid in
locating monographs , revisions and other valuable taxonomic papers . Refer-
enc es precede each genus . J
Core, E. L , \.,T. H. Gillespie , & B. J . Gillespie . 1962 . Bi.bliography of hlest
Virgi nia Plant Life . Scho lar' s Library, Hox 60 , Gracie Station , Ne\" York .
[A bibliography of more than 1,000 references on plant life of I~est Virg inia.
References listed alphabetically by author in the fo l lowing classes : a l gae ,
fun gi , lichens; bryophytes. vascular plants , fossil plants. Although se l ec-
ted for relevance to Hest Virginia, complete references are given for mono-
grapbs and revisions \~hich were serially published, even if only one part
contains a reference to the Hest Virginia plants . A valuable reference for
much of the eastern United States , par.ticularly the Southeast .
Ewan , J. 1967. A Bibliography of Louisiana Botany. In : Flora of Louisiana
by J . lL Thieret and collabora tors . Sout1n·J estern Louisi.ana Journal 7 : 2-
81. [Annotated references are arranged chronologically . An index to
authors, botanical names and subjects is included . J
740 30

Creene , \I . C .• & J . T. Curtis . 1955 . A Bibliography of \-liscons1n Vegetation .

l-lih18ukee Puhlic Museum Publications in Botany No. 1. Hih/sukee Public
~luseum. NihlBukee. [Th i s bibliography covers the per i od 1667-1953 \d t h
emphasis on post 1900 papers . Subjects treated include reports deali ng
"wit.h the na t ive plants and plant communities of \.J"isconsin , their distri-
bution . taxonomy . ecology . sociology . economic uses and conserva tion . "
The bib liogr aphy is divided into 15 sub j e ct areas Hith r eferences l is t ed
in chronological order . An author index is also provid ed .
Langman , I . K. 1964. A Selected Guide to the Literature on the Flowe ring
Plants of ~~exico. Univer sity of Pennsylvania Press . Philadelphia, Penn-
sylvania. [The aim of this guide is to bring exist i ng bibliographic
material on flO\~e r ing plants of Hexico up to date and to organize the
materials so desired references can be located with mi nimal effort . The
references are arranged by author Hith t heir publications in chrono l ogical
order Hith many entries anno tated . Anonymous artic l e s are arranged at the
end by date . The elaborate subject index (systematic group , geography ,
etc . ) makes this a very useful guide.
Lebedev , D. V. 1956. Guide to Botanical Literature of t he USSR. 382 pp .
Academy of Sc i ence USSR . Leningrad .
Neisel, H . 1924-1929. A Bibliography of American Natural History , the Pio-
neer Century 1769- 1865 . 3 vols . Premier Publishing Company . Brooklyn ,
NeH York. [Vol. 1 (192 4) An Annotated Bi bl iography of the Publica tions
Relating to the History , Bi ography and fi i b l iography of American Natur al
History and Its Institutions during Colon ia l Times and t he Pioneer Cen tury ,
which have been published to 1924, Hith a classified s ubject and geographic
index and a bibliography of bibliographies; Vo l . 2 (1926) The Insti t utions
....'hich have contributed to the rise and progress of Ame r ican Natural Hi story
which !'-Jere fo unded or or ganized betHeen 1769 and 1844 (includes cont e nts
of their j ournals and p r o ceed ings re l a t i n g to natural h istory) ; Vo l. 3
(1929) The Institutions founded or organized be t Heen 1845 and 1865 . Bib -
liography of Books, Chronological Tab l es , Index of Auth or s and Inst i tu-
tions, Addenda to Volume 1.]
Nerr111, E. D. , & E . II. \~alker . 1 938. A Bibliography of Eastern Asiat i c
Botany . The Arn old Arboretum of Harvard University . Jamaica Plain ,
r.lassachuse tt s . [This Ivo r k includes references published to December , 1936 .
The area i n cl udes China , J apan , Formosa , Korea, r.lan c hu ria, Hongo li a , Tiber
and eastern and southern Siberia in addition to major papers of adja c ent
areas such as the Philippines , Indo-China , Siam, Burma , India and Cent r a l
and North e rn Asia . Although the point of vie'" of selection was taxonomi c ,
papers "on eco l ogy , geography , exploration , economic botany, history ,
bibliography, phylogeny, pathology, agriculture. hor ticulture , forest r y ,
materia medica , pharmacy , biography" and others are i ncluded. Al l p l ant
groups are treated . Majo r categories a r e bibliography ( authors and titles) ,
a reference list of oriental serials (Hith characters ), a list of o r iental
authors (with their oriental characters) . and a subject index divided into
3 indices (genera. regional , and systemat i c) .
Supp lement I, by E. H. Halker , 1960 , American I n st i t ute of Bi ologi cal
Sciences . Hashington, D. C. This supp l ement extends cover age to 1958 . ]
Herrill , E . D. 1947 . A Bota nical Bibliography of the Is lands of the Pacific .
Contributions from the Un it ed States Na t ion al Herbar ium 30 : 1-322 . [ A r evi-
sion of the follOldng paper s : Herrill , E. D.• 1924 , Bi bliography of Poly-
nesian Botany, Bishop Museum Bulletin 13 : 1 - 68; and Pol ynesian Botan i cal
Bibliography 1773-1 935, Bishop t-luseum Bulletin 144 : 1- 194 .
The wor k therefore consists of about 3,850 author entries cove r ing the
period from 1773-1946 . The region cove r ed in this bib liography is " the
30 741

is lands of the Pacific between latitude JOoN and 30~S (excluding the Bonin Is -
lands) extendin g from Juan Fernandez and lIat.,taii in the east to the extreme
western limits of the Harianas , Caroline , and Palau Islands . It includes
all of Polynesia and Nicrone sia and the eastern parts of r-Ielanesia--as
Fiji, New Caledonia, the New Hebrides , Lord HO\.,te Island , Norfolk Island .
the Loyalty Islands, and Santa Cruz--but not the larger a r chipe lagoe s con-
tiguou s t o New Gu inea ... I I
Indexed informat i on inc lude s (1) papers in which n ew genera or species
are described from the region cove r ed ; ( 2) those papers involv ing transfers
of names of Polynesian species ; (3) species lists , all monographic Horks
in which species of the Pacific Islands are mentioned ; (4) general h10rks
(e . g., Bentham & Hooker ' s Genera Plantarum , and Engler & Pr ant l ' s Die
natiirl ichen Pflanzenfamilien); (5) appropriate papers on e cology and phyto-
geog raphy; (6) patho l ogy , forestry and some agriculture; and (7) books on
trave l. Pl ant groups treat ed include vascular plants and cel lular crypto-
gams . References a r e arranged by author chronologically . (See Halker ,
E . H., i n this same volume for a subject index \... hich is arranged by general
subject - -e.g., systematic groups.) . )
Reed , C. F . 1969. Bibliography to Floras of Southeast Asia . Bal timore , Hary-
lan d . ["A bibliography to the flora of Southeast Asia ranging from Hurma
and Tonkin, sou th through Annam, Laos , Thai l and and Cambodia to Cochinchina ,
the Hal ay Peninsula and Sin gapore ." "Publi ca tions dealing with floristic
works and taxonomic and mono graphic treatments of both vascular and non-
vascular pl ants , as Hell as those dealing \"ith the vernacular names are
inc l uded." Other subjects include cultivated, agricultural or medicinal
plants , forestry , forest floras and properties of forest timbers , climatic
and edaphic phenomena and fossil fl ora. The entr ies are arranged a l pha-
betica l ly by author. No subjec t ind ex is presen t. Note s upplement on pp.
189-191. One valuable f ea ture of t his index is the inclu s ion of author,
volumation, date of publication of specific treatments in major flora s .
Sachet, H. H., & F. R. Fosberg (compilers) . 1955. Island Bibliographies :
Micron esian Botany , Land Environment and Ecology of Coral Atolls , Vegeta-
tion of Tropical Pacific Islands . Compil ed under the auspices of the Paci-
fic Science Boa rd . National Academy of Sciences , National Research Council
Pub lica t ion 335 . '.Jashington, D. C. [A series of annotated bib liographies
arranged alphabetica lly by author. Each bibliography i s folloHed by a
subject index including geographic and systema tic classes. The bibliog-
raphies are as follm... s :
(1) Annotated Bibliography of Nicronesian Botany . Area covered in-
cludes the Marianas, Carol ine, Narshall and Gilbe rt Archipelago es , besides
the isolated islands of Harcus, I'lake , r-lapia , Na uru and Ocean .
(2) Bibliography of the Land Ecology and Environment of Coral Atolls .
For the coral atolls the area covered is that part of the tropical seas
\"here these islands a r e found, principally the cen t ral and wes tern Pacific ,
the Indian Ocean . and the Caribbean and Gulf of Mexico. Rocas, in the
Atlantic near Brazil and Clipperton Island in the eastern Pacifi c ar e also
included .
(3) Sel ected Bibliography of Vegetation of the Tropical Pacific Islands .
The coverage of this bib liography is the area extending from Guadalupe , the
Revil l agigedos. Clipperton, Cocos , t he Ga l apagos and Desventuradas \"estward
to the Bonins , Palau , Hapia, the Admiralties , Bismark Archipelago, Louisi-
ades and islands of the Coral Sea . Excluded are Japan , the Ri ukius , For-
mosa , Ph il ipp ines , East Indies, Ne\'/ Guinea, Austral ia, Kermadecs , Norfolk,
Lord Hm"e , NeN Zealand , and Juan Fernandez .
742 30

A list of ser i al abbreviations for the titles of se r.ials u sed in the

r efe r e nces for the three bibliog raphi es , \o1ith the full titl es that the y
represent a r e incl ud ed . Library of Congress numbers and libraries h'he r e
th ese vol umes are known are given in many cases . Note also addenda to
e ach bib liography and to t he l ist of serials . Taxonomic treatments are
readily located by s e a r c hing the list of ref e r ence s under the app ropriat e
family name in the subjec t index . Other subjects inc l ude bibl iography ,
expeditions a nd voyages, geology and phytogeogr aphy to mention a fe\. . .
(See also Sache t , H. t & F . R. Fosberg , 1971 , Island Bibliographies Supple-
me nt 1971 , Nationa l Academy of Sci en ces , h1ashington , D. C. } . J
Santapau , H. 1958 . History of Botanical Resea rches in India , Burma and
Ceylon . Part I I . Systematic Bot any of Angiosperms . Bangalore Press .
Bangalore City . [This wo rk includes a brief history of botany of the
area, notes on he rbaria and botanic gardens . and a bibliography of f lo r as .
hotanical explorations , and monog raph s . It includes treatments in jour-
nal s I~hich are not widely circulated . ]
Schi schki.n . B. K.• & A. A. Fedo r ov . 1963 . On the taxonomic and floris t ic
works p ubli shed i n the USSR during the las t fifteen years . !,-lebbia 18:
501- 5 62 .
Simpson . N. D. (compiler) . 1960. A Bibliogr aphic Index of the British Flora.
Pr ivate l y printed , in l i mited edi t ion of 750 copies . Bournemo u th . England .
[ " This Iwrk is devo t ed to phanerogams, vascula r crypt ogams and charo-
phytes and its aim is to pro v ide references to sources of information rel e -
vant to the identi ficatio n , history and geographical range of taxa I~hich
oc cur or have occu r red in Britain . Informa t ion is a l so p rovi ded on p l ant-
lore, l oca l names. poisonous plan ts and I~eeds . "
The book is divided into the f ollo\d ng classes : Par t l-- Cata logs of
Plant s ( mostly by author) , Distribution, Economic Plant s , Fami lies , Genera
and Spe c i es (nomenclature follows Clapham , A. R .• 1940 , i n J. Ecol. 33 ,
unless otherwise i ndicated); Floras (handbooks . keys . etc . ), Herba l s ;
Period icals ; Plant Lo r e ; References (inclu ding bibliographies . biographies .
col lec t i ng tec hniques . dictionaries , glossaries. herb aria) , Histo ry ; Tera-
tology ; Top ogr aphy ; Trees . Part II- - Plant Records (geographically
ar ranged ) . ]
S tee nis. C. G. G. J . van. 1955. Annotated Selected Bibliography in Flora
Malesiana . ser . 1 , 5, i-cxliv .
Zeybek. N. (compi ler) . 1972. A Bibliography of the Pap ers on the Taxonomy
and Ecology of Turkish Fl ora . 1841-1971 . Availab l e from Head of Sys t ematic
Botany, Un i ve r si t y of Ege , Bornova-Izmir . (~Iimeog r aphed) fArranged alpha-
betically by a uthor, language of papers indicated ; also includes names and
addre ss es of taxonomists and eco logist s of Turkey . ]

3. Bib liog rap hies (Specific Groups and App roache s)

A practic e which should be encou raged is the publication of bibliogra-

phies on specific subjects. Selected examples of such compilat ions are given
below . It is indeed unfo r t unat e t ha t the ex tensive card fi l es on various sub-
jects of ten i nvolving many year s of compilation by spec ia lists remain unavail-
ab l e to many taxonomist s . See Chapters 22 & 23 for discussion on the use of
e l ectronic data processing as one so lution to this p roblem .

Adams , J . 1928-1 936 . A Bib liog r aphy of Canadian Plant Geography . I. (1928)
Transactions Royal Canadian Institute 16 : 293- 355 ; II . (1929) 17 : 103- 145 ;
III. (1930) 17 : 227-2 65 ; IV . (1930) 17 : 267-295; Adams, J . , & H. 11 . Norend,
V. (19 32 ) 1S , 343-3 73 ; VI. (1936) 21 , 19-135 .
30 743

Bailey , L. H. 1933 . The Standard Cycloped ia of Horticulture . 3 vals . The

Hacmillan Company . Ne\" York . [Vo lume 2 include s a bib liography of horti-
culture treating books, jou rnals and s o ci eti~ s (pp . 1520-1562) and bio-
graphical ske t ches of No rth American hortic ultur ists (pp . 1563-1603) . ]
Jones , G. N. 1966 . An Annotated Bibliography of Mexican Ferns . University
of Illino i s Press . Urbana, I ll i n ois. [This Hork t reats both f erns and
fern allies of Hexico and, in addition t o author and tit le, includes "find -
ing" ind exes . This is a valuable SO U'I" ce of basic references on fe rns in
addition co the spe cific ref e rences to Mexican fern s .
Rehder, A. 1911-1918. The Bradley Bibl iography . A guide to the literature
of the woody plants of the ~<1Orld, published before t he beginning of the
twentieth century . Arnol d Arb o r etum . Jamaica Plain, Massachuse tts . [A
comprehensive s ubject and author index to books, pamphlets and jou rnal
articles publis hed to 1900 which treat woody plants . ]
1949 . Bibliography of Cultivated Trees and Shrubs Hardy in t~ e
Coo l er Tempe ra te Regi on of t he Nor thern Hemispher e . Arn old Arboretum.
Jamaica Plain , Hassachusetts. [The "purpose of this bibliography i s to give
r eferences to t he sources of the botanical names, valid names and synonyms ,
of the woody plants cultivat ed in t h e co oler region s of the temperate zone
of the northern hemisphere ." All names found in th e 2nd e di tion of Rehder's
manual (1940) are given in this treatment "~,'ith full synonymy and with addi-
tional genera, s p ecies and varieties i n t roduced or described , up to the end
of 1947 . " Lectotypes or s tandard- species are cited. In addit ion to t.he
" systematic enumera tion and synonymy of c ultivat ed trees and s hrubs , " list s
of abbrevi ations of serials and ti tles , abb revia tions and signs , additions
and emendations, and an i ndex to scient i fic names are given.]
Senn , H. A. 1951. A Bi b l i ograp hy of Canadian Plant Geography . Transactions
. Royal Canadian Institute 1 946-1947. Vol . 26 . Publication 863 , Department
of Agriculture. Ot tawa .
Solbr ig, O. T., & T . H. J . Gadella . Biosyst ematic Lit erature ; Contr.ibutions
to a b iosystema t ic literature index 1945-1964 . Regnum Vegetabile 69 .
Utrecht . [ See d iscussion under B VIlC , Current Literature . ]
Swi ft , L . H. 19 70 . Botanical Bibliographies . Burges s Publishing Company .
Hinneapolis, Hinneso ta . [See Part III, pp . 417-4 25 for "B ib liographic Keys
fo r literature on i ndividual taxa , specia l g roup s . " }
Wood, C. E., Jr . • and contribu t or s . 1958+ . Gene ric Flora of Sou t heas tern
United States . (Has exce l len t bib l iog r aphy for each g r oup treated. See
Jou rnal of Arno l d Arbo ret um , 1958+ . ]
Yuncker, T . G. 1964 . A bibliography of the fami l y Pi peraceae . Cando Ilea
19, 97-1 44 .

4. Cumulat i ve Indices to Journals

Hhen conducting a literature sea r ch these indices can save considerable

time . In botanical lib raries \"here a ser ies of specific journals is incom-
plete and funds are short , th e purchase of c umulative index volumes mi gh t be
the most expe dient solution .

Index t o Bulletin of Torrey Botanica l Club , Vo l s . 1-74 ( 1870- 1948). Compiled

by H. IL Ricke tt, 1955, Science Pr ess . Lancas ter, Pa . [Includ es index t o
authors vith title of article and a systematic index to eve r y botanical
name mentioned in the 75 volumes . ]
Index to Taxon, Vols . 1-20 . Part 1, Sc ientif ic names, Rouleau , E. (compiler),
1972; Part 2 , Auth or s and subjects , Lovden, R., 197 3 .
 744 30

Nolan , E . J . (ed . ). 1913. An index to scientific contents of the Journal of

the Proceedings of the Academy of Nat ural Sciences of Philadelphia 1817 -
1910 . Academy of Nat ural Sciences . Philadelph ia . (Index to genera, spe-
cies , etc . described or referred to in the proceedings and journal.]
Rouleau, E . 1953 . Rhodora , Index to Vals . 1-50 . New England Botanical Club .

S. I ndex to Dissertations

Dis s ertation Abstracts International , Section B. Science . University Hicro-

films, In c. Ann Arbo r, Hi chigao. (See also Index to American disserta-
tions) . [For dissertations prior to 1956 see Doctoral dissertations
accepted by American Universities, Wilson, NeH York . For discussion and
more bibliographies see S\.,rift, L . H., 1970, Botanical Bibliographies,
Burgess Publishing Company, Minneapolis, Ninnesota.]

6. Research Indices and Directories of Plant Systematists

BioResearch Index (BioI). Biosciences I nformat ion Service of Biologica l

Abstracts . [See discussion under Biological Abstracts in Sect ion B VIlC . ]
Indices to Chromosome Numbers . [See refe rences in Chapter 10 . ]
Jackson , R. C. (compiler ) . 1 966 . ASPT , lOPE I ndex of Current Taxonomic
Research. Regnum Vegetabile 43 . Ut r ech t . [This index is nOH under the
auspices of the Botany Department of the Smithsonian I nstitution and a cur--
rent ed it ion is long overdue . The research projects , including name and
address o f r esearcher , are arranged by systematic groups ; state fl oras of
the U. S . (also includes smaller areas such as counties, vegetat i on types
Hithin confines of a single state, etc .); regional floras of No rth and
, South Amer ica ; floristic studies in Europe , Asia, Africa; paleobotany;
phytogeography; indices; anatomy (arranged by systemat ic group) ; and bib--
liography , and miscellaneous studies.]
Keuken , \>,1. (compiler) . 1971. Dictionary of Plant Systematists. 1970 . Reg -
num Vegetabile 72 . Utrecht . [IAPT membership list fo r 1970 \>lith indica-
tion of membership in affiliate societies (lABG , IOPB , lOP , IDS) arranged
by country ; members are listed alphabetical ly and addresses g i ven . Insti-
tutional members (libraries, herbaria , botanic gardens , etc . ) are also
listed. ]
Roon , A. C. de (compiler). 1958 . International Directory of Spec i a l ists in
Plant Taxonomy Hith a Census of their Current I nterests . Regnum Vegetabile
13 . Utrecht. [Arranged alphabetically by specialist and by systematic
group and geography . Rather out of date but still useful.}

IX . GENERAL TEXTS

Alston, R. E. , & B. L . Turner . 1963 . Biochemical Systematics . Prentice Hall,

Inc . Englewood Cliffs , NeH Jersey.
Benson, L . D. 1962 . Plant Taxonomy: Hethods and Principles . The Ronald
Press Company . New York .
Blach"elder, R. E. 1967. Taxonomy : A Text and Reference Book . John vliley
and Sons , In c. NeH York . [A zoology text but good d iscus sion of general
aspec t s of taxonomy . J
Briggs , D., & S . N. Halt ers. 1969 . Plant Variation and Evolution . Horld
University Library, HcGra~l-Hill Book Company. Ne;.; Yo rk .
Core. E. L. 1955 . Plant Taxonomy . Prent ic e- Hal l , I nc. Engle\"ood Clif fs,
NeH Jersey .
30 745

Cronquis t , A. 19 68 . The Evolution and Classification of Flowering Pl ants .

Houghton Hifflin Company . Boston.
Da t ta , S . C. 1970 . A Handbook of Systematic Botany . 2nd ed . Asia Publish-
ing House . New York .
Dav is , P. H. , & V. H. Hey\-'Ood. 1963 . Principles of Angiospe rm Taxonomy .
D. Van Nostrand, Inc . Princeton , New Jersey.
Heslop-Harr i son , J . 1964 . New Concepts in Flowering Plant Taxonomy . Harvard
Univer s ity Press . Cambridge .
Heywood , v . H. (ed . ) . 1968 . Modern Hethods in Plant Taxonomy . Botanical
Society of Br itish Isl es Conference Report No . 1 0. Academic Press . Net~
Yor k . [A collection of pape r s gi ven at a conference on Hadern Hetr.ods in
Plan t Taxonomy and the revie'''s incl ude a major survey of the major f i elds
of taxonomy with emphasis on new technique s . ]
Hi tchcock , A. S. 1925 . De scriptive Systema tic Botany . John Wiley a nd Sons ,
Inc . New York.
Jeffrey, C. 1968 . An Introduction to Plan t Taxonomy . J . & A. Churchill ,
Ltd . Lon don.
J ohnson , A. M . 1 931. Taxonomy of the Flowe r ing Plan t s . The Century Company .
New York .
Lawren ce , G. n. H. 1951. Taxonomy of Vascular Plan ts. The "1acmillan Company .
New Yor k .
Pool , R. J . 1941 . Flowers and Fl owering Plants. 2nd ed . NcGraw-Hill Book
Company , Inc . Ne\v York .
Porter , C. L . 1967 . Taxonomy of Flowering Plants. 2nd ed . H. II. Freeman
and Compan y. San Franci sco .
Sneath , P. H. A., & R. R . Soka l. 1973 . Numerical Taxonomy : The Prin ciples
a nd Prac tice of Classif i cation. \.J . II . Fr eeman an d Company . San Francisco .
So l brig , O. T . 1970. Principles and l·lethods of Plant Biosys tematics . The
Macmillan Company. New York.
Swingle. D. B. 1946 . A Textbook of Systemat i c Botany . 3rd ed . HcGrmv-H1l1
Book Company , I nc . New York .
Ta khtajan , A. 196 9 . Flowering Plants--Origin and Dispersal . Smithsonian
Institu tion Pr ess . hTashington. D. C. [Authorized and t ran slated from the
Russian by C. Jeffrey .]

X. HONOGRAPHS AND REVISIONS

A. Ser ial Monographs and Revis ion s

( i nc l uding e xamp l es of r egional "monographs" or r evisions in "floras" )

Boissie r a , :t-lemoires des Conservatoi r e et Jardin botaniques de la villa de

Geneve . [A ser i es of monographs Hhich a r ose as a s upplement to Candollea .
See note in Taxon 22: 309. 1973 . ]
Candolle, A. P. de . 1823-1873. Pr odromus s y stematis natura l i s regn i vegeta-
bilis. .. Paris. [See discus s ion in Sect i on BIB .]
Candolle . A. de . & C. de Candolle (eds.). 1878- 1896 . Nonogr aphi ae Phanero-
gamarum. 9 vo 1 s . Paris . r.!onogr aph ic ser ies t o succeed the Prodromus . Vol .
1 (18 78) Smilac e es , Res ti aceae . Neliaceae ; Vol. 2 (1 879) Araceae; Vol. 3
(1881) Philyd r aceae. A1ismaceae , Butomac e ae , Juncaginaceae , Commelin aceae,
Cucu r b i taceae; Vo l . 4 (1883) Bu rseraceae , Anacard i ac eae , Pontederiaceae ;
Vol . 5 (1883-1887) Cyrtan dreae . Ampelidees; Vol. 6 (1889) Andropogone~e;
Vol. 7 (1891) He1astomacees ; Vo l. 8 (1893) Guttiferae; Vol. 9 (not seen) .
[Specimens cit e d . ]
Engler. A. 1900-1 968. Das Pflanzen r eich . Nos . 1-108 . Ber l i n . [A seri es
of monographs o f the p l ant ki ngdom by various authors. Although incomplete ,
 746 30

the pa r ts issued are significant Imrks. The t~ orks \.}ere published as com-
p l eted and the arrangemen t is often confusing . Basically the plan Has to
d ivid e the plant kingdom i n to four major groups (I-IV) . Under each group
fami lies \~ere also numbered (e.g •• Butomaceae IV 16). Each issue (He ft )
t~as g iv en a number (i. e .• HeEte 1·~l08) based on order of appearance . For
a de t a i led i ndex see Dav is , l'- I. T . , 1957 , ~ Guide and Analysis of Engler ' s
"Das Pflan z en r e i ch , " Taxon 6 : 161-182 , and also Sta f leu , Y . , 1967, Taxo-
nomic Litera t u r e . The various ways these monograph s have been cited in
the literatu r e are also discussed by both Davis and Stafleu .
Engler , H. G. A ., [. K. A. E . Prantl. Die natlirlichen P f lanzenfamilien . Ed .
1 (1887-19 1 5); ed . 2 (1924+) . [See discussion i n Section BIB . ]
Flora Neo t rop i ca . 1968+ . Organization for Flora Neo tropica . lIafner Press .
Ri v er side , NeH Jersey . [A serie s of monographs of tropical plants of t he
t,'e ste r n hemisphere . See discussion in section BIl l A.]
North Amer ican Flora . 190j+ . Britton , N. 1. , and contr i butors . Net·) York
Botanical Garden .
Phanerogamarum monograph iae . 1969+ . Verlag von J . Cramer . Lehre , Germany . LA
n eH s e ries of lengthy monographs, revisions and i ndepth treaLments of
limite d numbers of taxa of phan erogams . Vol. 1, Racquet, G. (1969) Revi-
sio Physolychnidum (Silene section Physolychnis) (Horld-hide coverage);
Vol . 2 , v an Royen , P . , The genus Rubus in Net,' Guinea (regional) (see
r evi etv i n Taxo n 18 : 456 . 1
Steen is, C. G. C. J . van (general ed.) . 1948+ . Flora Halesiana . h!olters-
No ordhoH Publish i ng . Groningen, The Netherlands. Series l - -Spermatophyta :
Vol. 1 Cyclopaedia of Collectors and Collections (1950); Vol. 2 Halesian
Vegetation (in preparation); Vol . 3 i'lalesian Plant Geography (in prepara-
tion) ; Vol . 4 General Chapters and Revisions (1955-1958) ; Vol . 5 Bibliog-
, r aphy, Spe cif i c Delimitation, & Revisions (1955-1958) ; Vol . 6 Systematic
Re v is ion s, 6 pts . (1960-1972); Vol. 7 Systematic Revisions , pts . 1 . 2 (1971-
1972) . [See i nside front cover of Vol. 7, pt. 2 (1972) f or Index to Revised
Familie s in Series 1; volumation and page numbers are also given . 1

B. Gui des

Nany excellent guides h ave been mentioned in other sections of this chap-
te r (pa r ticularly the sect i ons dealing 'vith bibliographies and current litera-
ture) whi c h sho u ld be consulted . The titles of some o f these and others ,.;hich
a re c o mmonly used are cited here .

A. E . T . F . A. T . Ind e x (See Section B VIIC ) .

Biologica l Abstracts (~ee Section B VIIC) .
Boivin , B. 1967(?) . Enumeration des plantes du Canada . Provancheria No . 6 .
Memoir es de l' Herb i er Louis-l'larie . Universite Laval . Quebec .
Bo t any Sub j e c t Index . Co mpiled by USDA (See Section B VIllE) .
Davis , N. T . 19 5 7 . A guide and a n alysis of Engler's DaB Pflanzenreich . Taxon
6 : 161-182 .
Ex c erp ta Bo t ani ca ( See Section B VIIC ) .
Gra y Ca rd Index . Although this index has a rather specific use (See Section B
IE) it c a n b e useful i n locating monographs and revisions since such \-Jo r ks
us ually include netv combinations and descriptions of neH taxa .
I ndex to American Botan ical Literature (See Section B VIIC) .
Index to Aus t ral asian Taxonomic Literature (See Section E VIlC) .
Inde x to Europe a n Taxonomic Literature. 1965 - 1969. (See Section B VIIC).
Ken t. D. H. 1 967 . Index to Botanical Honographs. Published for the Botani-
c al Society o f t he British Isles . Academic Press . London and NetJ York .
30 747

[ "A guide to monographs and taxonomic papers relating to phanerogams and

vascular cryptogams grm-ling \·dId in the British Isles." Nostly includes
works published since 1800.
La\~rence. G. H . ~1. 1951. Taxonomy of Vasc ular Plants. The l-Iacmillan Company .
Nel" York. [See litera ture cir.ed following description of each family . ]
Herri ll, E . D.• Eo E. H. Wa lker . 1 938 . A Bibliography of Eastern As i a ti c
Botany . Supplement by E. H. l"'alker , 1960 . (See discu ssion in Sect ion B
VIlIB) .
Reed , C. F . 1969 . Bibliography to Floras of Southeast Asia . (See Section
B VIllE) .
Sachet , N. H. , & F . R. Fosberg . 1955 . Island Bibliographies and Supp lement.
1971. (See Section B VIl l E).

Section C . SUGGESTIONS FOR Tim PREPARATION OF TAXONOHIC

TREATHENTS AND REFERENCE FILES

The preparation of any taxonomic treatmen t involves considerab le refer-

ence to the literature . Sound taxonomic l-JOrk requires not onl y an acquaint-
ance l.,rith literature but also careful documentat ion. In the prepara tion of
a monograph , revision, flora or manual, one must account for all the names
publ ished for plants in a specif ic group or in a particular area . Orig i nal
descriptions and type specimens must be located , pertinent literature sur-
veyed, repre sentative herbarium specimens examined, taxonomic decisions made ,
and results published .

SOI!lt>. means of compila tion of references, ua la and other records i s neces-

s ary . Several "how to" references of value in the organization and prepara-
tion of such treatments are Leenhouts, P . \-! . , lY61), i\ Guide to the prac tice
of herbarium taxonomy , Regnum Vegetabile 58 , Utrecht ; Bens on , L., 1962, Plant
Ta xonomy, The Ronald Press Company , Nel'l York; and Jacobs , H., 1969 , Large
families- -not alone !, Taxon 18 : 253-262 . E"'an (Taxon 18 : 19{, if . , 1969)
points out some oE the historical problems for taxonomists , and Staf l e u (Taxon
12 : ll3 ff ., 1963) discusses the dates of publication of early ",arks an d some
of the J:leans of determining the date of effect ive publication and associated
prob lems.

Students are enco uraged to begin a reference (card) file early i n their
careers . The references to be collected and the system of cata loging ar e
matters of individual interest and preference . The syStem should be one that
best complements an individual's thinkint; (c . g ., does one tend to remember
authors , systematic divisions or special subjec ts). Hhatever the system , it
sho uld be flexible , should not require that numerous refer ence cards be made
for a single reference, and should be readily lIseable . r.!ost student s Hill
keep such a file for specif ic resear ch projects (class projects , theses , etc .).
This Eile may be expanded as other interests develop , or separate research and
general reference HIes may be kept . The usual 3 x 5 or 5 x 8 card files are
conven ient and provide space for the references and for annotations. Seve r al
time-saving steps in the compilation of a bib liograph y or a reference file
include accurate citation (author , date, title, journal or publisher) , record
of call number, notes if reference is studied (comment s on contents , concepts ,
etc .), and notation on sources of references recorded but not consulted (I.,rill
exped ite interlibr ary loans) .

Systematic perusal of current issues of literature guides and indi ces

such as Index to American Botanical Literature, Excerpta Botanica , etc . , as
748 ~

',]e11 as bihliographies in journal articles, is very helpful not only in com-

piling re fe rences of interest but in directing one ' s attention to current
papers ~"hich might othen.,rise be overlooked .

Hopefully , Hidespread availability and use of electronic data processing

for retrieval o f such information will soon reduce the time and effort spent
on making and using such a primitive f orm of information retrieval as card
files, or better yet, make them obsolete.

TAXO NOMIC LITERl\TURE EXERCISE

Sel ect any species of vascular plant described 25 or more years ago and give
the f ollowing information:

1. Citation of the original publication.

2. List of synonyms (to 10) and references.
3. Copy and translate (if necessary) the original description.
4. List of references (to 10) to illustrations .
5. Location of type specimen .
6. Collector of type and type l ocality .
7. Corre ct generic name, authority and bi bliographic re f erence.
8. Type species of the genus .
9. Number of species in genus .
10 . Infraspecifi c taxa described since 1885 .
11. Distribution of this species in US and Canada and general vorld distribu-
tion for the genus.
12. Host recent monograph of the group .
13 . List the key characters used in at least 3 manuals to separate this
species (or genus) from closely related ones .
14 . An economically or ho rt iculturally important member of the group.
15. Chromosome number.
16 . List current papers or research projects published within the last 3
years treating this species (or genus) .
17 . Current authority or worker on the group .

SAHPLE FORN FOR RECORDItlG INFORNATION

Documen ta t ion
Indexes & Guides (citation of
Consulted LC or specific
No . (e.g. Index Ke\~ensis) Dewey No. Answer references)

1.
2.
3.
4.
5.
6.
7.
8.
9.
10 .
11.
12 .
31 751

Chapter 31. THE HERBARIUM

An herbarium is both a collection of dried plants and an ed ucational and

research institut ion. As a basic resource for the study of systematic botany
and related f i elds , the herbarium serves as a reference center , a documenta-
tion facility and a data storehouse . The organization, use, operation and
maintenance of herbaria are discussed below .

Section A. HERBARIUH IN GENERAL

1. HERBARIUH : CONCEPT Ai'm DEfINITION

The term herbarium , used in the strictest sense today, is simply a collec-
tion of dried specimens . Laurence (1951) and others include in their defini-
tions the arrangement of specimens in the sequence of an accepted classifica-
tion , and that the specimens are available for r eference or other scientific
study . "Herbarium" used in its original sense, hOl.ever, referred not to a
collection of p lants , but to a book about medicinal plants . Tournefo rt, in
about 1700, used the term as an equivalent to hortus siccus (St earn , 1957).
and this use \"as taken up by Linnaeus who also adopted it as a substitute for
hortus siccus, hortus mortus , and others. It \"as largely through his in fluence
that the \"ord herbarium sup e rceded the former terms .

The procedure of pressing and drying specimens for storage has been an
amazingly successful one in terms of preservation of detail and specimen lon-
gevity, and the plants so preserved provide a concrete basis for past, present,
and future studies. I n its more than four-hundred-year history the herbarium
has become an institution. Today one assoc iates the term herbarium not only
\dth preserved plant specimens but also Hith certain botanical activities .
The herbarium is the basic re feren c e source of the taxonomist and has become
a center for research as \"ell as teaching and public information .

An herbarium, a special kind of museum, can also be re garded as a data

bank with vast quantities of raw data. Each specimen has information content
and therefore value ,.hi ch \"ill , of course , vary depending on the completeness
of the specimen and data and the source of the material. Each specimen has
information about the vegetation of an area , a population, and the taxon to
which it belongs (Rollins, 1965) . The collection , therefore, rep res ents a
source o f primary in f ormation about man's explorations and observations of the
earth's vegetation and documents the results of much o f the past inquiry into
the nature and relationships of plants . Herbarium specimens are nm" used for
studies in discip lines which were probably never even dreamed of at the time
early collections were made and herbaria organized . These studies include
such fields as cytogeography, biochemical systematics, palynology and gen-
ecology .

II . HISTORICAL DEVELOPHENT

The beginning of the herbarium as a collection of dried speCimens affixed

to paper for a lasting record is attributed to Luca Ghin i (1490? -1556).
According to Arber (1938) Ghini seems to have been the sole init iator of
the art of herbarium-making and this art was disseminated over Europ e by
 752 31

his students. Gherards Cibo, a pupil of Ghini, began collecting and preserv-
ing specimens as early as 1532 and his herbarium is extant today . John
Falconer, an Englishman . is mentioned in the \.,rritings of Lusitanu$ in 1553
and \Ililliam Turner in 1569 as possessing an herharium, and i t is believed that
he also learned of herbarium making either directly or indirectly from Ghini
(Arber, 1938). Although the herbarium technique was a well-kno\-ffi botanical
practice at the time of Linnaeus, he departed from the convention of the day
(mounting specimens and binding them into volumes) by mounting his specimens
on single sheets and storing them horizontally much as is the practice today
(Stearn , 1957; Dewolf, 1968) . Although this method became general during the
second half of the 18th century, perhaps, as Stearn ( 1957) bel i eves, due to
Linnaeus's example and teaching, it Has by no means universal. In fact as
late as 1833 Asa Gray Has offering bound volumes of grasses and sedges for
sale (Dewolf, 1968) .

In the United States several herbaria are known to have existed i n the
mid-1700 ' s. Nanyof these and later ones found their \.Jay to Europe, where
they are preserved. According to LanjoU{.J and Stafleu (1964) and Shetler (1969)
the oldest institutional herbarium in the United States is that of Salem
College started in 1772 . Other early institutions include the Academy of
Natural Science of Philad elphi a, Amherst College, Boston Society of Natural
liistory, Charleston !1useum, and others, all fo unded before 1860 (Jones and
Headows, 1948).

I n addition to changes in mounting, early workers also began depos i ting

specimens in established collections, as well as exchanging specimens . It is
indee d for tunate for botanists of today that such practices developed so early.
Nany herbaria have been destroyed by fire , insects , war and ignorance , and all
'- that remains of some collections are the duplicates sent to other institutions
on exchange. An unfortunate policy l"a5 the scanty field notes made by early
collectors and the lack of concern for original labels as early herbarium
specimens Here exchanged or sold . For an interesting account of the sale,
transfer, and handling of several early private North American herbaria , see
Stuckey (1971).

III . PRESENT SIZE AND STATE OF HERRARIA

Herbaria range f rom small personal collections (mostly of a few hundred

specimens with a feH not able e xceptions) to large collections of colleges ,
universities , private foundations and governmental agencies involving mil -
lions of specimens. In 1964 ten herbaria reported collections in excess of
three mil lion specimens (Lanjouw and Stafleu, 1964; Shetler , 1969) . Table
31-1 lists the five largest herbaria in the Horld and the eight herbaria in
the United States with one million or more specimens .

Shetler (1969) est i mates that the herbarium resources of the world may
include as many as 250 million specimens , although the total number actually
reported by the various institutions \.JaS 148 million . Of this estimate about
78 million are in European collections and 36 million are in North America .
The difficulty is , of course , the number of private and small public herbaria
Hhich often do not report their holdings or .,lhose holdings are not generally
accessible.

Today collections are more accessible than ever before due to the efforts
of many individuals and national and international organizations . Two
31 753

Table 31-1. La r gest Herbaria i n the \.[orld (After Shet l e r , 1969) .

Location Abbreviation No . of Specimens Rank by Size

Kew . England K 6,500 , 000 1

Leningrad , USS R LE 6 , 000 , 000 2
Pa ris , France P 5 , 000 , 000 3
Geneva , Switzerland G 4,000 , 000 4
Lyon, France LY 3,800 ,00 0 5
Cambridge , Hass. (Combined A, A1'ffiS , ECON,
herbaria of Har va rd Uoiv.) FH, GH, NEB 3 ,540,000
New York (Bot. Card.), New York NY 3,000 , 000 B
Ha shington, D. C. (U. S. Na t. Herb . ) US 3 ,000 , 000 B
Chicago, Illinoi s F 2 ,3 50 , 000 11
St . Louis , Mo. (No . Bot . Card.) MO 1,700,000 14
Berkeley , California UC 1 ,225,00 0 21
Ann Arbor , Hich. NICH 1,000 , 000 24
Philadelphia, Pa . PH 1,000 , 000 24

ex tremely useful i ndices for lo cating collections are I ndex He rb ariorum ( Lan-
jouw and Stafleu , 1965) which inc ludes locat ion and curator s of herba ria of
the world, size and type of co llec tions, type of i nstitution , publicat ions
(serials), and important historical col lec tions a nd Index Xyl ar iorum (Stern,
1967) , an index to woo d collections of the wo rld. The principal wood c ollec-
tion s of the Un ited St a t es include thos e of the Arnol d Arbor etum (Aw) , Ca m-
bridge , Nassachusetts; Rancho Santa Ana Botanic Garde n (RSAw) , Claremont ,
California; U. S . Forest Products Labo ratory (MADw , SJRw) , Madison , \.[isc onsin;
and Smithsonian Institution (USw) , Wa sh ington , D. C. A current project under-
way which should a l so be of tre mend ous aid to systematists is the " Type RegiS-
try" being compiled by the Smithsonian I nstit ution . Variou s herba ria, part i-
cularly those with an ab undance of type material and other historic collec tions ,
often issue accoun ts of their col l ections; e . g ., The Sl oane He rbar ium in t he
Bri tish !-Iu seum (Dandy , 1958), Herbar ia o f the Departmen t of Botany i n the
University of Ox f ord (Clokie, 1964), and Li nnean Herbaria (Stearn, 1957 ) .

IV . THE CHANGING NATURE OF THE HERBARIUN AND ITS FUTURE

The nature of the he rbarium as an institution and co llec t ion remained

es sentia lly unchanged until about fi f t y y ears ago . Deve l opmen t s i n gene t iCS ,
biochemist r y , cytology , ecology, a nd o the r fields leading to less des cr iptive
and more experimental \-lo rk have been la rg ely r esponsib le for the change s that
have occurre d , but unfortunately there has been some resistance to this change .
The herbarium , ",h ile it s e rves its fo rmer goals as a center for documentation ,
a source of inf ormation for monographs and floristic works , and as a resource
in educati on and other fields , has the oppo rt unity and the responsibi lit y to
expand to meet the needs of t he fut ure . Hos t systematists now use numerous
approaches to problem solving , a nd t he herbarium shou ld remain as the central
facility , as i ndeed it i s i n many insti tut ions .

The status , ope rational aspects , and future of herbaria have been r e -
viewed by Beaman (1965) a nd Shetler (1969). Rollins (1965) has also pointed
out some of the roles of the herbarium in re search and teaching . She tler (1969)
has cast s ome doubt on the future of the herbarium as a scientific institution
754 31

and resource. It seems desirable that herbarium goals and practices he re-
viewed , par ticul arly 1n the light of the recent developments in taxonomy and
botany in general . Each herbarium administrator . s ta Ef member. and all of
those who use the herbarium should consider the role of the herbarium not
only at thei r institution bu t also on the in ternationa l scene , and guideline s
should be set up to insure maximum availability of information in herbaria
both at present and in the future . Turrill (1964) has stated that the re are
two sets of prob lems facing herbaria--one is hot" herbaria can he maintained
within reasonable limits and , tl';O, how they can be imp roved. Some of his
suggestions include division into spec ial and general types . Spec i al her-
baria would include local olles (those devoted to a given area and se r ving
l ocal inter es t s), specia l research he rbaria (e . g ., cytological and cy togene-
tic vouchers) :lIld ecologic herbada (including not only specimens but photo-
graphs, field notes, schemes of ecological analysis) . Admittedly the general
herba r ium presents additional ilnd more difficult problems, and Turrill (1964)
proposed the limitin g of such her-baria in number l"itl1 each having some spe-
c ial ization ; e . g ., systematic or r egional. He furthe r summarized the purposes
of a general herbarium as follows: "(1) to provide facilities for determina -
tion on any material, inc l uding new taxa ; (2) to enable ne\~ monographs and
floras to be prepared ; (3) to preserve specimens of historic importance, such
as those dating discoveries , introductions , and increases or restrictions of
ran ges ; (4) to ass e mbl e data f or I,'orking out ranges and ecological di stribu-
tions; (5) to bring together in a relatively permanent form specimens for
comparative morphological or phylogcmetic studies and to provide material f or
special researches, as in plant anatomy and palynology ." For an interesting
account of t he relevance of national, regional and local herbaria see Cron-
quist (1968) , fi r enan (1968), and NcNeil (1968).

The has ic probl em:'; of general herbar ia still remain . Hot... large should
they become? \fuat kind of mat erials are to be included? How can info rmat ion
and materia ls be efficiently retrieved? Shetler (1969) has pointed ou t some
of the idiosyncrasies of herbaria as wel l as their shor tcomings . He proposes
that herbaria employ business-like methods and that compute r s be used for
much of the labo r ious retrieval nm" needed and demanded by scientists and
laymen . Such reorganization and innova tion requires broad cooperation and
considerable plann ing. Cur-a tors should not blindly bec ome slaves to the "ma-
chine" as many have to " the ir collections . " The first s tep in solving the
basic problems of herbaria is to accept the challenge of the future wi th an
o p en mind and a r eso lve to deal t"ith these problems . This may mean a com-
p lete reorganization, disso l ution , expans i on , or combina t ion of exist i ng
collections. As !!lentioned above, the estab lishment of special and reg i onal
herbaria is an attractive possibility . Regional center s with adequate sup-
port fro m all level s--Iocal , r egional , national, and perhaps internationa l
sources- - a nd proper commitment could, through the conso lidation of collections
and the cons equen t reduction in us eless and expensive competition and duplica-
tion , serve as clearing houses for exchan~ es and l oans , provide safe deposi-
tories for documentation and assist local and specia l herba r ia by routing
special materi als and duplica te s to the proper instit ution. Certainly such
c e nt ers sho uld be able to supply materia l s and assist in bu ilding supe rior
t e a ching and r eference col l ections . These in stitutions should use eff i c i ent
mean s of specimen handling (loans, location of types , and sp ec ial materials)
and informa tion r et rieval and could well provide r esea rch space and eq uip-
ment for use of visiting botanists . I n addition to the collection, a super-
ior library, ga r dens, and t echn ical staff could become available to more
peop l e than is currently possible.
31 755

In July , 1972 , the Association of Systematics Collections (ASC) I,/as

established and began to address itself t o many of the problems facing all
systematics collC!ctions . The goal of the ASC , according to the con s ti tution
and by-lm"s , is "to foste r care , ma\l<l.gement, preservation and improvement of
systematics collections and to facilitate their utiliza t i on in science and
society by :

1. Providing representation for ins t itutions housing systematics

collections ;
2. Enc ouraging direc t interaction a mong those con cerned Hit h systematics
collections and their use ;
3. Provi ding a forum for consideration of mutual problems;
4. Promoting the ro l e of systellk"ltics collections in research , education,
and public service through :
a. the coordination of info r mat i on concerning needs of use r s,
b . the planning of advisory services ,
c . t he developmen t and implemen tation of national goals and
priorities ;
5. Other means and devices which shall be determined from time to time
by the Membership" (Association of Systematics Collections ,
1972 : 1) .

At the time this association was fo r med at a Systematics Biology Symposium

held in July , 1972 , by the Smitlll:lOnian I n stitu t ion in col l aboration Hith the
National Aca demy of Sciences , sjx report s \vere presen ted by a Commi tte e of the
Conference of Directors of Systematics Collections . The six reports and re-
lated mate r ials , including considerable i npu t f r om the symposium parti cipants ,
\Je r e consider ed by a writing committee of tt.Jenty-five systematists a nd admi n-
istrators \.,oho met to plan for the coord i na t ed management of nation al system-
atics resources .

This conunittee in their prepublica tion draft of America ' s Systemat ics
Collections : A National Plan, compiled and edited by H. s . In11n , 1:1. H. Payne ,
D. H. Bates and P. S . Humphrey (1973) p r esents a "statement of the primary
goals of the systematics collections community , " and a "desc r iption of the
specific goals of the systematics co llections commun it y with respect to
improving the condition of the col l ect i ons and the serv i ces they provide ;"
a "discussion of systemati c s in science a nd society today" and " of the
problems affecting the systematics colle ctions community ; " a list of "s pe-
cific recommendations (The National Plan) that will move to resolve t he
problems and enab l e the systematics collections community to realize i t s
goals" and a "statement of priorities and estimated costs of implement i ng
them . "

The primary goals of t he systemat i cs collections c ommunity are " t o

i mprove t he con dition of systematics collections as a National reso ur ce " and
" to improve t he efficiency of services a s sociated \~i t h systematics c ol lections
resources . Specific goals and specific ser vice-related goals include :

"1. Hanagement of the specimen inventory and asso c i ated documentation

to insure : (a) permanent conservation of t he specimens them-
selves ; (b) ready access to them and their documentation ; and
( c ) space , fa c ilities , and library reso urc es enabling sy s tema-
tists to improve the info r mation content of the collection s
756 31

through identif ication, classification, and e laboration of

the intrinsic information carried by each specimen .
2. Addition of ne'" spec i mens and associated information that: (a)
reflect the goals and priorities of science and applied
science; and (b) impr ove the quality and quantity of speci-
men and taxon related data, so that the information load
of each specimen is enhanced .
3. To make available nonproprietary, specimen- or taxon-related
information in a variety of useful configurations .
4. To enable incorpor.at ion of specimens and associated data in the
info rmat ion management system .
S. To enable ready access to the specimens themselves. and to asso-
cia ted documen ta tion and library rna terials . "

The Nat ional Plan inc ludes the follolvinB recommendations which. Ivhen
implemented, should solve many of the problems of systematics collec-
tions .

"1. Supp ort and strengthen an organization dedicated to improving

the condition of systematics collections and their services .
2. Establish a series of Councils to study specific prohlems re-
lating to systematics collections .
3. Es tabl ish Adviso ry Commi tt e es representing those elements o f
the scientific community concerned ~vith systematics collec-
tions .
4. Identify systema tics collections of importance as national
resources and designate National Resource Centers .
5. Develop standards for systematics collect ion s: physical facili-
ties , collection storage, preservation, specimen and data
acqui.si tion and documentation) collec tion growth , in ter-
institutional loans.
6. Imp lement electronic data proces sing in collection management
procedures .
7. Develop technical training programs for professional service
personnel and establish mechanisms for personnel placement.
8. Develop mechan isms whereby the resources of systematics col lec-
tions can be used more effectively in studying and resolving
problems affecting the quality of the environment and develop
programs f or improv ing public awareness of the importance of
correct identif ica tion of species to indicate environmental
changes and their effe cts on human welfare.
9. Improve the services and other contri.butions of systematics
collections to graduate education in systematic biology .
10 . Add ress the problems of libraries and publications associated
Hith systematics collections.
11. Develop and implement programs for improving public aHareness
and use of systematics collections as a national resource
important to science and to the solution of problems affect-
ing human Helfare.
12 . Study financ ial resources available and potentially available to
the systematics collections community and seek means to
implement priorities .. . "
31 757

Section B. PROC ESSING AND ACCESSIONI NG OF SPECINENS

1. THE SPECINEN

Tr adi tional l y the typical her barium specimen has been a vascular plant
aff ixed to a 11 1/2 i nch x 16 1 / 2 inch sheet , bu t recen t deve l opments neces-
s i tat e a change i n the concept of herbarium spec imens. Wood samp l es . fossils .
pollen and spores . micros l i de s, l iquid-preserved materia l s , photographs. and
drawings should be considered as specimens a nd t he r efo r e part of the herbar-
ium. These c hanges require new housing facili ties and r ecord keeping , as
well as cross ref e rencing and retr ieval sys t ems . Today most herbarium cura-
tors are urging co llectors to make a concerted effort to make their collec-
tions as complete as possible by including ma t e rial for ch r omosome counts,
wood samples. pollen s l ides . pol l i na tor s (or at l eas t observational record s) .
assoc iat ed species and detailed habitat da t a . Weeds and cultivated species
ar e a t last being considered wo rthy of in clu s i on in the collections . If t he
herba r ium is to be the source of in fo r mation and documentation it s hould be .
the materials used in biosys temat i c st ud i es (e . g ., natura l a nd artif i c ial
hybrids . ma ss coll ections . e tc . ) as \"ell as other types of studies should be
inc l uded in the herbarium . Biosystemat i sts a nd others sho uld confer \"ith
directors and curator s concerning the handling o f materials .

II . SOURCES OF MATERIALS

The infl ux of ma t e rial into an herbarium is usual ly fro m t he fol l owi ng

sources :
A. Staf f and St udent Co llections . These va r y according to staff
r esearch interes t s, spec i al i ties . and her barium goal s . Coll ec-
tion s may be for flo ri stic. monograph i c , biosystematic . or
t aximetric studies . Staff and s tudents at institutions Hhich
have no herbarium , or only teaching or r eference co llec tions,
may choose to document their st udies by sendin g specimens to
other insti tutions. These specimens may be sent eit her as
gifts o r loans and published works s hould always i nclude the
name of the herbarium where vouche r s a re depOS i ted .
B. Exchange . Dupl icates o r spec i a l col l ec tions are e xchange d among
various ins titutions. us ually on a one- fo r-one basis . This is one
of the majo r means o f adding to a co llect ion and an efficient
means of ge tting needed mate r ia l s a t minima l cost to all con-
cerned .
C. Gif ts . The se are va riou sly interpreted in different herbaria . Some
treat r eceip t of all specime ns othe r than exchange or l oans as
gifts \~hil e others include only those for which no staf f services
are required . These may range from an en t ire herbarium to a few
specimens sent to a staff specia list .
D. Loans . Loa ns are generally eithe r t et:lporar y (short term ; e .g . • a
st udy of specif ic taxa for preparation of a monograph) o r indefi-
nate (permanent; e . g . • the loan of an entire herba r ium f r om one
institutio n to anot her ) . The l atte r is a particula r ly de sirable
procedure for histo r ic and ot he r collec tions which may be for
the most part inaccessible to most bo t anists o r where proper
housing facilit i es are lacking . The s e can often be loaned to
a nother i nstitution and still have the or iginal name main tained .
E. Purchases . The purchase of an he rba rium is an un common event today .
Usually priva te or insti tu tional herbar ia are donated o r are
758 31

deposited on permanent loan at an inst i tution . Host purchase s

today involve collections from specific areas (particularly the
tropics) with prices ran ging from ten cents to over one dollar
per sheet. In some cases an institution may help to sponsor a
collecting trip in exchange for a set of specimens .
F. Identification Service . In most herbaria specialists are Hilling to
determine specimens for other \'JOrkers and interest ed amateurs .
Some herbaria have staff members I~hose specific job is service
identifications . In general the specimens sent for identifica-
tion are kept by receiving institu tions unless other arrangements
are made in advance. These specimens should be of such quality
that they are desirable as specimens (see Chapter 18 on col lect-
ing), and should not be sent I~ithout some prior agreement .
Requests for id entificat ion services should be reasonable in terms
of the time required and the number of specimens to be identified.

III. HANDLING OF I NCOHING SPECI!>1ENS

Regardless of the source , most materials are first unpacked and variously
treated to rid them of insect pests which are a serious threat to specimen
longevity . Such pests includ e the larvae of cigarette and carpet beetles ,
moths, booklice and silverfish. Procedures vary, but several methods are
commonplace:

A. Fumigation . This involves the use of such compounds as methyl bro-

mide, carbon bisulphide, carbon tetrachloride, ethylene dichlo-
ride , hydrocyanic gas , lindane, dichlorvos strips , or paradichlo-
robenzene . hlJlile it is necessary to treat all neH additions and
loans upon arrival, the entire collection should be treated on a
regular basis (see Section C--Herbarium Use, Operation and Nainte-
nance). Several drawbacks to fumigation given by Fosberg and
Sachet (1965) are :
1. Commonly used fumi gants are also harmful to humans .
2. Nany fumigants are inflammable .
3. Insect eggs and pupae are often not killed unless f umigation is
done in a vacuum chamber.
B. Heating. The use of specially constructed chambers with heating ele-
ments which can keep specimens at 44 0 C for some hours is an effec-
tive method . Hhile this is a good method for new acquisitions, it
is usually not practical in most cases f or treating entire collec-
tions (for details and references see Fosberg and Sachet, 1965) .
C. POisoning . Ther e are several methods of treating specimens making
them either permanently poisonous or unpalatable to herbarium
pests. Specimens may be poisoned by dipping or painting them I"ith
an alcoholic solution of mercuric chlo r ide. Specimens so treated
should ah~ays be clearly labelled since this compound is extremely
poisonous. It is also said to make some paper brittle and crumbly,
blacken labels, and attack aluminum .
Another method employs lauryl pentachlorphenate (a 3 . 75% solu-
tion of LPCP in pure white kerosene free fr om high boiling paraf-
fins). This solution is used at the British Nuseum Herbarium
where spec i mens are dipped in the solution in a galvanized iron
tank . The excess liquid is allowed to drain off and the speci-
mens are placed in a drying cabinet where the kerosene evaporates
31 759

l eaving the pl ants impregna ted with LPCP . For details s ee Fosberg
and Sachet (1965) and Hh itmore (1965) . Thi s technique has also
been used on mounted s p ecimens \o11th succes s. Haunted specimens
may be sprayed Hith a 5% l auryl pentachlorphenate (Mystox) in va r-
sol . An adjustabl e sp r ay gun i s used to sa turate the plant ma t e r-
i als while a cardboard shield is held over the s pecimen labe l to
preven t fadi n g o f i n ks. Dryin g time is 24 ho urs (for details of
spraying ap paratus and hand li ng of specimens see Lunde ll and
Kirkham, 1966) .
IV . ACCES SIONING OF COLLECTIONS

Accessioning is defined by Fosberg and Sa che t (1965) a s the recording of

the r eceipt a nd origin of lots of spec i mens coming into the herbarium. Each
lot is ass i gned an access ion numbe r . The minimum data that is reco rded is
name o f sender, da te of receipt, type of transaction ( g i ft, exchange , pur-
cha se) , number of sheets , place (country, province , etc .), and kinds of plants
(algae , fungi, vascul ar plants, e t c . ) . Space may be left to r eco rd the inclu-
sive sheet number s after the specimens a re mounted (Fosb er g a nd Sachet , 1965).
Other sys tems us ed include the registration of each speC imen (mos t l y abandoned
but fo r an examp l e see Hil lspaugh, 1 925 ) o r simp l y mountin g speCimens and
assigning sheet numbers . A practical and efficien t \"ay is to stamp sheets
with the offic ial herba r ium s eal and number th em before specimens are mount ed .
Since e xchan ge , loan , a nd gift r eco rds are kept, as \"ell as all co rrespon-
dence , t he separat e acces sion record may be cons idere d unn ecessary . Sheet
numbers , in addition to being us ed to assess growth a nd eff iciency in herbar-
ium operat i ons, have impo rtance di r ec tly r ela t ed to t axonomic resea r ch . They
are a n id eal means of refer r ing to a par ticul ar specimen i n a collect i on . The
exclusive use of col l ec t o rs' numbers may not be suffic i e nt, particul a rly in
designating type s .

V. SPECINEN PREPARATIO}l

Hounting proc edures vary but at present only a fe \" me thod s a r e popul ar .
It cannot be emphasized enough t hat the mounting procedure is a c ritical step
in specime n prepara tion . A care l essly mounted specimen can be essen tially
usel ess . Mounting , l ike pressing, is indeed an art .
A. Strapping. A spec imen is placed on mounting paper and the label
pasted sq uarely in the 10\"er right -hand corner . Strips of linen
tape (gumme d cloth or Holland cloth) are moistened and place d
aroun d stems , across leaves , etc. until the specimen is fi r ml y
at t ached to paper . (Cellulose o r plastic tapes which de t er i orate
or soften wi t h age or \"hich decompose when specimen s are fumi-
gated shou ld not be used . ) Care should be taken so that impor-
tan t detai ls are not covered by the strips . Length and \"idth of
strip s depend on the part to be secur ed .
Another method of strapping or stripp ing (as it is often
c alled) involves the use of liquid pl astic . Thi s method. the
Ar cher Netho d. uses a mix ture of et hyl-ce llulose a nd Dow resin in
toluene a nd met hanol , prepa red according to t he sched ule of Rollins
(19 55) given below.
Toluene - 880 cc .
Nethanol - 220 cc .
Ethocel - 10 cps . standard
Dow resin - 75 gms .
 760 31

Toluene and me t hanol are mixed first and t he resin added and dis -
so lved comple tely . The ethocel is added slo\~ ly and s tirre d until
partly d isso lved. The solution should be al lowed to s tand 24
hours befo re using . Sto re in air tight containe r s a nd use a 4:1
mixture of toluene and methanol as a solven t . The plastic is
usually dispensed f r om a pistol-grip press ure oi l can or a poly-
ethylene (mustard or cats up) sq ueeze bottle .
B. Pasting or Gluing . Specimens are often attach ed to their s hee ts by
glu ing . An efficient and effective way is to coa t t h e su rface of
a glass or coppe r p l a t e with glu e ( use a pain t brus h ) , place the
specimen on the pla t e , and t ap gent l y \<lith forceps or car ds . The
specimen is t hen l i fted . using forceps , inspected to check dis t ri-
bution of glue , and placed on the sheet . If t he gl ue is not evenly
distributed, more may be app lied using a soft brush . Tap spec i -
mens down gen tly using relatively lint-free tissue . Labels gen-
erally are app lied using the same gl ue and put on before the
specimen. I f any glue is on the exposed s ur face . t h e sheet may
be covered with waxed paper . This is a particularly good proce-
dure for gr asses and sedges . Al l f ragments , frui t s, and seeds
sho uld be plac ed i n packets or envelopes a n d t hese aff ixed to the
herbarium shee t at this time. In c ases wher e materia l s are too
bulky. the pa cket should be n umber ed \.,.ith an herbarium accession
n umbe r and/o r collector's n umber a nd set aside to be mounted later .
The specimen i s t hen covered \.1ith a sheet of newspaper ( usually
the one containing the specimen , if not t oo dirty) and placed in
a stacking box or simply placed on the table . Specimen s are so
stacked unti l 20 to 25 specimens are in a stack. A board and
Height are t hen pla ced on t he stack . If mounting is s low , t he
, l.,Ieight sho uld be applied sooner . Stacks should be al l ol.,led to dry
for 24 hours . The papers are then removed and specimen s a r e ready
fo r t he n ex t s tep. Severa l gl ues or pastes are available. The
" Special A Ti n Pas t e" is used in some herbaria, but lib r a r y pastes
such as "IPt" or gl ues s uch as " EIme r s ," " Nicobond ," o r " I?ilhold"
are a l so used . Some herba r ia use the Arche r formula (plastic) for
gluing specimens t o the sheet . Th e plastic , dispensed from a
" sq ueeze " bo ttle or pressure gun , i s appl ied to the back of a
specimen and material is placed on the sheet , weighted Hith wash-
ers or other su it able weights a nd the plastic alloHe d to dry.
All of these glues , but not the plastic , are Hater soluble.
C. Sewin g . Except where st r apping is t he only process, mos t spec i men s ,
afte r glu i ng , a re strapped in some manner--line n tape, liquid pl as-
tic. white glue, a n d/or sewing with a heavy linen t hread . A very
efficient process combines sewing and strapping (with glue o r
Archers) . Sewing is restricted to heavy stems , overlapping
leaves, rhizome s , matted bases of grasses. l arge fruits , cone s
or heads , or other places where the use of plastic or glue is
impossible or imp ractical . Threads on the back of sheets should
be covered wi th lin en or paper tape. Specimens are placed on a
sheet of cardboard and strapped . Each specimen is then stacked
one upon the other u sing 4- 5 wooden blacks to separate each
specimen. This r e qu i r es lit tle ~.,Iork space and a llows proper
ventilation.
D. Special Hand ling. One case of spec i a l hand ling is t he use of se r ial
sheets for a s i ngl e specimen, l ong a necessity i n tropi ca l a reas .
 Although several techniques are used and much depends on the notes
of the colle ctor, the series of sheets should bear a single acces-
sion number and an annotation noting on each sheet the number of
sheets in the series; e .g., sheet 1 of 5. Such a series is much
more desirable than the practice of ins i sting that regardless of
size, an entire plant should be on a sheet even though c ritical
examination of the material is impossible .
Inflorescences of ~, Cirsium, etc . o fte n shatter and may
be held intact by enclosing them in bags (cheese cloth or netting) .
Bulky f ru its , cones , et c. may be removed and stored in special
boxes or cases . Specimens should clearly indicate that both
sheets and accessory materials are present in the co llect ion .
Each sheet should also be stamped with the distinct i ve seal and/
or name of the herbarium . The use of letter s (abbreviations)
only should be avoided (e . g., specimens marked au Herbarium may
mean Unive rsity of Oxford, Oregon, Oklahoma) . Some herharia
use a stamp with the name of the institution and its official
abbreviation (see Lanjouw and Stafleu, 1964; Fosberg and Sachet,
1965).
Proper mounting depends greatly on careful collect ing. Sug-
gestions fo r collecting and pressing of herbarium specimens are
given in Chapter 18.

VI . SORTING AND FILING OF SPECINENS

Once the specimens have been glued, sewn and/or strapped and allowed to
dry (in the cases where glue and/or plastic are used), they should be sorted
for filing. Techn iques and physical arrangements vary, but ultimately speci-
mens are sorted to family , genus, species , and in fraspecific taxa. If geo-
graphic provinces are recognized in the collection, additional sorting is
required . Regardless of the filing system used, specimen s should be arranged
in the order they I.;ill occur in the herbarium. Considerable time and energy
can be saved if this procedure is folIO\"red. I n some collections, once the
specimens are sorted, they are routed to curators of special groups fo r
checking and/or filing .

Specimens are filed on shelves in herbarium cases in 24 x 16 1/2 inch

folders called genus covers. These heavy manilla or bristol folders are often
color coded . Specimens are either filed directly in these f olders or in a
th inner species f older. Today there seems to be a bre akdown in terminology
for folders. The genus cover often in rea lity serves as a specie s folder if
the r ep res enta tion of a species warrants an entire folder. In small collec-
tions the commercial "genus cover" may serve as a fam ily folder .

Specimens are placed in folders face up with labels in the lower right-
hand corner. The genus name usually appears in the lm"er lefthand corner on
the f ace o f the cover . Specific names or alphabetical sequence appe ar in the
lower righthand corner. Names may be printed or stamped on the folder or on a
label I"hich is then glued to the cover.

VII. ARRANGENENT OF SPECIHENS I N THE HERBARIUH

A. The General Co llectio n. Although ther e are numerous possible systems

of arrangement of specimens, the schemes most often encoun tered
are given belO\~:
762 31

1. Bentham and Hooker . This system LS rarely used in the United

States. Seo::! Franks (1965) fOI~ a list of larger European
herbaria using this system.
2. Dalla Torre and Harms numerical a r~ ' a!lgement of the Englerian
System . A common system i n L1h! United States and other
countries although it is oft ~ll modified so that only family
sequence is follmJed Hith genera and spo::!cies being alpha-
betically arranged . (See Dalla Torre , C. G. de, and H.
Harms, 1900-1907) .
3. Bo::!sseyan System. This system became popular in the 1920 ' sand
is still used in some herbar ia (particularly those organ-
ized after the above date) .
4. Alphabetical. This system varies fr om strictly alphabetical (by
family, then genus, species, etc .) to some modified system
(e . g., segregation o f fe rns, gy mnosperms , monocotyledons,
dicotyledons, with alphabetica] arrangement of families in
each of these groups) .

In addition to tho::!se basic systems, many herba ria also employ

a geographic arrangement. Local areas are in one color folder, \.,rith
u . S . • other Neh' \~orld countries, Old \,Iorld, etc . in other colors .
Such schemes range from fe\" geographic categories to quite elabo rat e
ones. These divisions are exceeding ly useful for floristi'c studies .
Even though colored folders are more expensive and fading colors
occasionally are confusing, the reduction of handl ing of all speci-
mens of a species to examine only those specimens from a particular
area (e . g . , county, state, or physiographic province) may justify
the added cost.

Undetermined materials (of \·,hich every herbarium has its fair

share) are generally filed at the end of genus or family and are
usually in specially marked folden; (e.g ., "undet. " ). I n addition to
specimens and special materials (discussed belm.,r) non-plant materials
are often of great value and arc included in general collections.
These include photographs (e .g ., a type speC imen, a habitat), dra\.,r-
ings, correspondence concerning a specimen , keys, notes on synonymy
and cross-reference markers ("dummy" sheet or cover) .

B. Special Collections . In addi tion to the general collection, many

herbaria maintain various special collections .
1. Type Collecti on . Type specimens s uch as holotypes , isotypes, and
co-types are often hous ed separately from the general collec-
tion, \"here unnecessary handling and risk of damage can be
greatly reduced. These are commonly filed in the same sequence
as the general collection, o r alphabetical ly . Another solution
is to place types in the general collection, but in specially
and conspicuously marked genus covers .
2 . Synoptic Collection ( Teaching or identification) . Special collec-
tions for routine identifications and student use are sometimes
separated f rom general collections . The synoptic collection
can greatly reduce time spent consulting specimens in a her-
barium as Hell as specimen wear. These collections are
usually small and may be arranged and rearranged (particu-
larly for teaching purposes) according to recent schemes
31 763

of c l ass i f icat ion, e . g ., Cronquis t . Tho r n e , Hutchinson .

3. Special Resea rch Collectio n . ~1ass collections and artifi cial
hybrids, as Hell as po llen a nd spo res , anatomical prepa r a-
tions and bud materials may often be housed separately or in
special fold e r s or cabi nets . I f t hi s is the case , some nota-
tion in the form of a dummy sheet or annotation s hould be made
in the genera l collec tion so that the existence of these mater -
ials can be readily knmm by t hose consult ing the herbarium .
The arrangement of such systems varies according to t he t ype of
materials a nd i n clination of the c urato r s . Alphabe tical, com-
munity, numerical (according to accession n umb e r) or phylo-
genetic systems a r e commonly used . ~~ere the system differs
from tha t of the general co l lection i t should be clearly
explained in the "l1erbar ium Guide " (see section C) .
4. Historical Col l ection . ~Iany ea rlier coll ec tion s , as men t ioned
above , were bound in vo lume s and these are often not amenable
to inclusion in gencra l collections . Other collcct ions include
those loans or gifts Ivith specific ation that they are to be
maintained as separate entities . The only other r eason fo r
sepa r ate housing is for protection . Some pr efe r to inc l ude
these materials in the general collec tion in speci al folder s ,
as mentioned ea r lier concerning types . I n fact many group
types and other historical mate r i als (including l et t e r s , notes.
etc . ) into one special collection . Regardless of phys ical
arrangement, a catalog of materials and t heir arrangement should
be incl uded in th e "Herbarium Guide " and/or appropri a te " dummies"
i nserted i nto the gene ral collection .

Sec tio n C . \IJo:RBARIU~1 USE , OPERATION AND HAINTENANCE

1.. IlERBARIUN USE

Herbaria exist only fo r us e regard l ess of the t yp e of collection- - r esearch ,

identification (synoptic) , tea ching , or other . Every effor t fo r maximum
efficiency in locatin g ma terial in a col l ection (information retrieva l ) should
be made Ivithout endangerin g the col lection .
A. Herbarium Guide . Every herba rium should ha ve a print e d g uide to the
collection . This gulde sho uld expl a in the systematic and geographic
ar r angement; color code of f olders; list of famili es with number s
and case nu mher (inc illding seg re gate and combined families ; e . g .,
Ericaceae vs . Pyrola cf'ae , ~Ionotropaceae); map of case arrangement;
list, location and <lr t' angement of specia l collections; and a generic
cata log . The guide ~. ;ou ld also contain he r bar ium policy stateme n ts
concerning r eshelvinr. o ( ma t e rial s , handling of spec imens , use of
e quipment and other l ;lcili ties, loan proced ures, and other perti-
nent in forma tion . Su c h g uides can save considerable t i me on th e
part of visitors as \-'e LI as the herbarium staf f .
B. Spec imen Usc . Specimens (I re prese r ved for use but should be handled
with ext r eme care sjn ce they are of scienti fic value and generally
irreplaceab le . The r o llmdng suggestions are t o a id users i n
hand l ing spec imens pro perly .
1 . Keep sheets flat , do 110t shuffl e or leaf through a fo lder like a
book. Specimens <lro bri t t le and eas ily damaged .
2 . Store properly . Preferably , specimens s hould be s t o red in he rb a r-
ium cases o r at least on shelves . Do no t c rOl.,rd t oo many
 7M II

specimens i nto a box or "pigeon hole ." Keep ma t e rials in

cove r.s when not in use .
3. Do not lay books or other heavy objects on s pecimens .
4. Pl ace loose fragments in packets or enve lop es if th e specimen to
which the fragment belongs can be asc e rtained .
5. Use materials sparingly . I t is often necessary to dissect flowers
or fruits . Use no more t han is req uir e d a nd place all fra g-
me n ts in a fragment enve l ope on the s hee t s . For dissection,
flowers and fruit s may be softe ned by boiling o r treating wi th
a softening o r wetting agent . PohI ' s sof t ening agen t (Pohl,
1965 ) i s an excellent we t t i ng material a nd may be pre pared
according to t he fol 1o\~i n g fo rmula :
17. solid "Aerosol OT " 1 .6 rol 75 % aqueo us "Aerosol or"
74% distilled water 73 . 4 ml di st illed wa ter
25% methyl alcohol 25 ml methyl al co hol
This so lution \"ill not stain or discolor sheets but i s best
used in a watch g lass . Place materials to be sof tene d i n a
watc h glass and a dd seve ral drops of the solu t ion. Hany
materia l s are sufficien tly sof ten ed in a few minut es . Fr ag-
ments may be b l otted dry and returned to the f r agmen t envel-
ope .
6 . Use a long-armed dissecting microscope. An entire spec imen can be
studied with this instrument without bending the sheet .
7. Supp ort the specimens with a v e ntilator or other stiff board when
carrying specimens, e ven fo r short distanc es .
8 . Call t o the attention of cu r ators those specimens in need of
repair.
, 9 . Do no t wri t e on the he r barium shee t unles s per miss i on to do so
has been given by someon e i n charge .
1 0 . Reshelve specimens o nly with permiss ion and always with ext r eme
care.
11. For additional i nformation on anno tat i ons , borrowing s pe cimens ,
etc ., see Sec tion C 10 (Loans) .
C. Visitors. Visiting an h erbarium is , of course, one of the best pos-
sible ways t o study material. An investigator can examine the
exact materials desired as \ofel l as search " undetermined " folders ,
con s ul t types and other special collections ( some not available
on loan), and use the library and bot anic garden as we ll as ex-
change ideas wi t h fe l low botanists . Visitors would do well to
fo llow common courtesy if they wish to have a ccess to co l lections ,
particularly a t unusual hours , s uch as evenings , weeke nds, or
holidays . A not e in advance can mean the d iffe r e n ce in having or
not having a comfortable place to wor k and access to a microscope ,
as we l l as gai ning admittAnce to special collections a n d l ibraries .
D. Loans. Most herbaria l end materials to o ther institutions (rarely to
individuals). Reciprocal agreements bet\ofeen many i nstitutions
often result in transit costs only one Hay . One sho uld, hOHever,
be sure t ha t such agreements exist before requesting a loan.
Otherwise , the borrower pays al l transit cos ts . For the most
pa rt loans should be r equested af ter some preli~inary work has
heen done so that unduly l ong- t e rm loans a r e avoided (loan periods
are genera lly for six months) . Req uests should be fo r reasonable
amounts of mate rial and fo r specific taxa r ather than communities
o r floristic p rovinces . Loans should b e r equested by the curator
or director of the herbarium fo r study by a specific person , and
 - ---- - ._- -

31 765

shou l d be addressed to the director or cur ator of the lending

institution . Special permiss i on should be obtained if fragments,
pollen, or other materials are to be removed . Even with permis-
sion the utmost discretion should be exercised. Many herbaria
ask that duplicate slides , chromatograms, and other preparations
made from their material be included when the specimens are
returned . "lfhen in doubt , the borrower should consult the curator
of the lending institution for specific instruct i ons . A commonly
accepted procedure is that all specimens be annotated whether they
are used i n ecological, floristic, monographic , anatomical , bio-
systematic , or other studies. This should include even those speci-
mens determined correct l y . A convenient way to agree is by using
an exclamation point ( 1) followed by the name of the investigator
and date (at least the year) . Printed annotation labels affixed
above the or i ginal label are the most desirable . The original
label should not be altered . Some institu t ions allow the use of
a rubber stamp with small, yet readable print (prefe r ab l y using
black ink); e . g ., "Examined in a survey of parasit i c angiosperms
of Southwestern U. S ., John Doe, 1971. " The name of the institu-
tion where wor k is being conducted is also quite desirable . Anno-
tations should reflect the history of use of an individual speci-
men . Naterials should be returned promptly on the clate due or an
extension should be requested . Nost inst i tutions honor requests
as soon as staff resources permit and at cons i derable time and
expense . Loan forms to be returned are generally sent o u t before
or at the time the loans are shipped . Upon receipt of a loan , the
borrower should e x amine , count and acknmo/ledge the shipment. Any
damage occurr i ng in transit due to mishandling or insects should
be noted on the loan acknmo/ledgment form. Each sheet shoul d be
checked for the stamp of the lendin g institution . Since sheets
may have the seal of more than one institution , careful notes
should be made, particularly if the borrmo/er has material from
several institutions. Specimens were packed by the lender in a
certain order and an effort should be made to return them the same
way.
Borrowed materia l s should be housed in "fireproof , " dust - and
pest-free cases. Naterials may be fumigated and repellents used
prov i ded they do not damage specimens , labels , and photographs.
Great care should be used in all cases . Plastic strapping is
softened by strong fumigants and repellents such as paradichloro-
benzene. This and other information i s often included on loan
forms . Most specimens are sent "lib r ary rate" by parcel post at
considerable savings to all ; hmo/ever , the lending institution
determines the mode of transit and this should be follmoJed unless
permi ssion to do otherwise is obtained.
In addit i on to the annotations, the bor r ower can perform many
other services :
(1) Determining duplication .
(2) Supplying additional data (obtained from duplicates from
other institutions .
(3) Correcting local i ty data (by annotation) .
(4) POinting out mixed labe l s .
(5) Determining spec i mens as to type--isotype. paratype , etc.
(6) Indicat i ng a mixed collection .
In most cases both parties should profit from a loan .

J
766 31

II. OPERATION AND l>L\INTENAl.~CE

An herbarium, contrary to the opinion of some, requires constant atten-

tion if materials , information and facilities sought by botanists and others
are to be available and in proper condition .
A. Specimen Care .
1. Backlog of materials . Keeping up with the influx of materials is
probably one of the g rea test problems in most herbaria . This
is often due to lack of s uppor t , staff and space and to in-
efficiency. Hhere possible , backlogs , which in turn create
storage and fumigation problems and consequently endanger
specimens , are to be avoided. Inaccessible materials are of
no value .
2 . Specimen repair. This is an end le ss prob l em . Specimens in need
of repair and r eMounting shou ld receive prompt atten tion as
soon as they are noted. Some institutions repair all mater-
ia l before i t is loaned or before reshelving i t after i t has
been used. If staff resources pe rmit, repairs should be made
as they are noted by persons fi l ing new acquisitions, insert-
ing ne\.; g enu s cove rs, etc .
3 . Fumigation. As mentioned earlier (Sect i on B) fumigation (or
other means of protection) of the ent ire collection is a
necessity in most if not all herbaria . Annua l fumigat ion
may n ot be sufficient to insu re a collection against damage ,
particularly if all new accessions are not first fumigated .
Constant alertness is mandatory . Thos e cases housing mem-
bers of the Asteraceae , Brassicaceae , Ranunculaceae and
Apiaceae should be scrutinized often s i nce members of these
families are common l y attacked by herbarium pests . Visitors
would he of great service to curators by calling attention
to signs of ins ect damage .
Fumigation of a collection may be done by a commercial
firm or by the herbarium staff. The entire collection may
be fumigated at one time or on l y part of it. The method
used will , of cou rse, vary depending upon physical arrange-
ment and size of the collection, financing , and staff pref-
e rences . A 3:1 mixture of ethylene dichloride and carbon
tetrachloride is a commonly used fumigant Hhich is placed
in each case and the case then sealed . Th is solut ion is
dangerous and should be used with extreme care . Stern et al .
(1969) and Lellinger (1972) have discussed the advantages
and disadvantages of using lindane and dichlorvos as fumi-
gants and repellants . Le1ling er (1972) indicated that
dichlorvos ("No Pest" or "Vapona " ) is probably the safer
material vhen used as a fum i gant rather than a repellent.
This method involves placing one 0.25 strip/22 .3 ft . 3 of
case space for 7-10 days with the case remaining closed .
This treatment should be used tHice each year assuming all
i ncoming materials are fumigated before insertion and that
there i s no infestation . Curators should consult both the
Stern ~~. and Lellinger papers before using this method.
Initial fum i gation of small lots of specimens, e . g ., neN
accessions or loans , may be carried out in specially con-
structed cabinets or chambers . Nethyl bromide is the fumi-
gant often used in this operation.
31 767

B. Insertion of }~terials (othe r than new accessions) .

1. Genus covers. As collections grow new species and/or genus covers
must be added. Division of mat e rial i nto smaller gr oups (sys-
tematic or geographic ) results in easier access and a r eduction
in needless handling .
2 . Dummy fol ders . As a result of changes i n nomencla ture and taxo-
nomic r evis ion, materials may in time be lost unless some pre-
cautions are taken. The probl em can eas i ly be solved by insert-
ing a dummy folder in the appropriate place in the herbarium ;
e . g ., lIymenoxys (see Tetraneuris); Calylophus serrulatus (see
Oenothera serrulata) . These may also be used to refer to addi-
tions to special collectiDns--\~ood samp l es, cones, types .
Tempora ry dummies are useful f or indica ting materials on l oan
o r those being repaired .
3. Photographs, Dr awing s , ~~ps . etc. These may be mounted on sheets
or placed in envelopes and are usually treated as specimens.
Care should be taken to make sure such materials will not be
damaged in the course of r egular fumigation .
C. Suppl ies and Equipment . All equipment , including herbarium cases,
should be checked a nd cleaned carefully . In addition to having
a c l ean and attractive place conducive to work as \.tell as func-
tional equipment, many of the sources of insect infes ta tion , dust
catchers and fire traps are eliminated . Supp l ies for mounting,
r epairs, labels , genus covers , etc. must be i nventoried and
ordered. Fortunately, mounting paper (100% rag) is used in suffi-
cient quantit i es that it is usually readily available from paper
companies and bi ological supply houses . In fact, some paper com-
panies issue spec i al catalogs and s upply lists for herbarium sup-
plies-- mounting paper, gen us cover s , pa s te , gl ue , etc . Hos t
herbaria maintain a file on sources of supplies .
D. Correspondence and Record Keeping. (See also Fosberg and Sachet ,
19115) . All rec ords and correspondence concerning exchanges ,
loans, gifts, and accessions should be kept and filed . Methods
vary , but co rrespondence is often filed a l phabetically according
to person or inst i tution . Exchange records are commonly entered
in a l edger o r journal or in a special card file . In formation
often recorded includes date , number of specimens received or
sent , origin of material , and balance . Duplicate or tripl icate
exchange forms are usually sent with an exchange lot and the
receivin g institution acknowledges receipt of shipment, verifies
specimen count and condition, and account bal an ce . One copy is
returned to the sender as a r ecord. Loan forms genera lly are
similar to exchange forms . In fac t, many herbaria use the same
form and simply check the appropriate type of transaction . Loan
agreements should be read carefully for they may specify condition s
under which specimens s hould be hous ed , type of annotation required,
and other particular requirements .
I nformat ion of considerable importance concerning specimens,
types, etc . may be in herbarium correspondence which s hould be
pr eserved . Fosberg and Sachet (1965) s ugges t binding letters
from one person or institution in chronolo gically arranged volumes .
E. Annual Report . The preparation of an annual report can be exceedingly
profitable for herbaria of n umerous kinds and sizes. It not only
provides an opportunity to reflect on progress and shortcomings of
the past year but is useful in planning for future gro~.tt h and
768 31

developme nt. Such reports usually include summaries of variou s

acc ounts and r eco r ds; e . g .• l oans and exchanges , spec ial r equests ,
and gifts . These r eports a l so often inc lude i nf ormat i on not avail-
ab l e from account l edge rs and journals and which consequently may
be of histo r ica l and othe r importa nce ; e . g .• r esea r ch in progress ,
f i eld trip reports , library and herbar ium requ isitions , new equip-
ment purchases , staff c hanges , and de velo pment of new techniques
and proced ures . This info rmation may be useful in preparing grant
prop osals a nd annual budge ts and supplying information to various
administ r a tors a nd botanical i nstit u tions . For examples of de-
tailed r eports see direc t or ' s annua l report i n c urrent vo l umes of
Al i so and Arnoldia .

Section D. liERBARIUM GLO SSARY

(Largely adapted fro m Fosb erg and Sachet, 1965)

Accession Number . The number applied to each lot of specimen s received by the
herba rium o r t he s hee t numbe r assigned to a spec ific specimen .
Accessioning . Recording the receipt and origin of l o ts o f specimens coming
into the he r barium ; also includes assigning of sheet numbers .
Annotation . A note writ ten on or attac hed to an he rbarium sheet indicating a
correction or change in identification or a point or po i nt s of interest
abou t the s pecimen ; any not e attached to a specimen.
Annotation Labe l s (annotat ion slip). A small s l ip of paper on which an anno-
tation may be wri tten and then glued to an herb arium shee t (see a l so det er-
minavi t and ap probavit s l ips).
App r obavi t Sl ip or Labe l. A special annotation l abel, indicat ing that t he
name on t he label is correct .
Arrangement (o f specimens in the herba rium). The sy stem of c l ass ification or
scheme follot.Jed i n the placement of speCimen s i n the herbarium ; e . g., Ben-
tham and Hooker , Engl e r, Cro nquis t, alphabe tical.
Ar cher Ne thod. Affixi ng specimens to mounting s heets by mea ns of sma ll strips
of liqu i d plastic ex trud ed f r om a con t ainer with a na rrow nozz le.
Boar d (o f plant press) . One of the two sti ff sheets of wallboard , cardboard
or wood between which t he blotters or ventilators , and the fol ders of
plants are laid and Hhich a r e tied to gether to form a pre ss .
Ca rpological Collection . Separate collection of fr uits and seeds .
Case. The cabinet in l.,Ihich herbarium specimens a r e s tored .
Collecting . The gather ing of spec i mens in t he fie l d .
Collection . The accumula t i on of specimens in an her barium; the s pec i mens col-
lected on a single expedition; a s ingle gat he ring of a particular species
at a given time and pl ace; a spec imen plus its replicates (many Hould pre-
fer the term " duplica te" rather than " rep licate . ")
Collection Number . The number assigned to a collection in the fiel d , when it
i s collected or Hhen the notes are ~</'titten up, t he same for all of its
replicates; it is assoc i ated wit h the specimen(s ) and identif i es them from
then on , and refer s to the data recorded i n t he collector ' s notebook.
(This is often also called the co l lector' s number and sho uld not be con -
f used Iv-ith the s heet number which is assigned to a parti c ular specimen in
t he herbarium .)
Corrugate . A sheet of pasteboard or thin metal with f lut e d duc ts extending
ac ross the sheet (no t lengthwise) used i n presses when drying plants by
means of artificial hea t; these a re ofte n calle d ventilators.
Determinat ion. Ascertaining the cor rect name fo r a s pecimen; ident ification.
31 769

De terminavit Slip . A type of annotation bearing the name of the plant and
the name of the person who identified the plant; date of identifica tion
should also be included .
Dist ribut ion. A confus ing t erm used to refer to the fi l ing of specimens as
wel l a s the sending out o f duplicates on exchan ge . l oan, etc .
Documentation. The depos ition of , or r eference to. voucher specimens i n an
herbari um .
Dryer or Plant Drye r. An appa ratu s for drying plant specimens by art i ficial
heat ; term used fo r a sheet of bl otting paper used in drying plants.
Dummy Sheet . A blank herbarium s he et or manilla paper of the same size as a
herbarium sheet or genus cover inse rte d in the herbarium for c ross ref er-
ence purposes.
Dupli cate . One of two or more specimens collected at the same time and place.
under t he same co llection number. to rep r esen t a partic ular specie s ; more
appropriately called a r eplica te if the collec tion contained more than two
sheets; an ext r a shee t of a co l lectio n .
Envelop e . Another name for packet. or pocket; a p i ece of pape r folded a nd
af fi xe d to a n he r bar ium shee t to contain fragments.
Exchange. The process of distributing duplicates (replicate s) or other
material s to o the r institution s in ret urn f or their duplica tes.
Felt Driers. Sheets of heavy blotting paper or build er ' s tl deadening felt, "
cut 12 x 17 inches , used to absorb moisture i n a press , with or without
corrugated meta l ventilators; also for tying piles of plants in folders
to make l ight packages .
Fide. Abla t ive s ingula r o f f ides , " according to or by the assurance of"--a
term used to r e f e r t o ano t her author ; e . g . , Neptunia pubescens Bentham,
fide Windler, 1966 .
Field Book. A notebook used to record data in the field a t the time of col-
lection or a book containing data collected in the field . Each collection
usually is assigned a numb er which refers to note s in the fie ld book .
Fie l d Label. A special label for recording data in the fie l d .
Field Notes. I n fo rmation recorded about a col lection or speCimen in the field
and usually inc l uding loca lity , habitat des cription , as Ivell as thos e fea -
ture s of the plant not di scernible from the dry specimen ; e . g ., height, if
entire plant i s no t col l ected; bark and branching characteristics, if a
tree o r s hrub; flower co l o r; odor ; etc .
Fie l d Press or Portfo lio. A l ight plant press carried in the fie l d when
co llecting, in to wh i ch the specimens may be placed as they are gathered .
Fil ing . The insertion of mount ed specimens , dummy sheets, and other materials
into the herbarium case s.
Fl imsies . Fo l ds of thin absorbent paper into which plant specimens are col-
lected and in which they may be dried and st ored .
Fragment. A part of a plant; a n y detached portion of a specimen ; also used
to re fe r to an incomplete o r poor s pecimen; e . g . • a sterile twig Ivith a
f ew leaves; extra materials co llected for dissection and placed in the
packet o r envel ope on a mount ed specimen .
Fumigant . A volatil e s ubs tance used to kill insect pests in the herbarium.
Fumigation. The process o f killing or getting rid of insect pests by sub-
jecting them to a l e tha l conc entration of a volatile poisonous chemical .
General Herbarium . Synonymous with general co llection ; all materials exclud-
ing special collections.
Genus Cover . A heavy manilla folder sligh tly l arger than an herbarium sheet
in Ivhich specimens belonging to one ge nus are filed; often in various
col or s in d icating geog raphic region s .
 770 II

Glue . An adhesive u sed to affix specimen s to the he r ba r ium pape r ; see also
white glue .
Gl uing . Fa sten ing s pecime ns to mount i ng sheets by means of glue or other adhe-
sive .
He r barium . A c o l l ectio n of drie d pl ants ; an inst itu t ion built around a collec-
tion of dried pl a nt s .
Herba r ium Abbre via tion . The a bb r eviat ion assigned to a n herbarium in Index
Herbariorum .
He rbarium Numb e r . The sheet number o f a sp ecime n .
Index Herbariorum . A series of i ndices to herba ria a nd collecto r s published
in Regnum Vegetab i le by the International Association for Pla n t Taxonomy ,
Utrec ht.
Man illa o r ma ni la. A coarse , un bleached paper of wh i ch fol de r s a re made .
l-Iass Colle ction . A population sample , composed of a numbe r l arge e nough to
be s tati stic ally significant, of cri tically sele cted c orrespondin g plant
fragmen t s collect ed at a particular time and place , to show the range of
variation i n selected characte ristics of the population sampled .
He rril l Case . A cardboard container 48 cm . long, 34 . 5 cm . wide and 24 cm .
high (outside dimen sions) with a doo r on one en d . Used as temporary sto r-
age fo r fil i ng of speC i men s ; develope d by E. D. Herr ill; not dust o r
in sec t proo f .
Hic rof iche . Gr eat l y reduced t r anspar en t posit i ve photographs of p r inted mate r-
ial or herbar ium specimen s des i gned for ready f ilin g and for r eading with
a spec ia l magni fyi ng projector (reade r) or wi th a binoc ular microscop e .
l>lounting . The process of affixing dr ied and pressed specimens o f plants to
he rbarium sheet s of heavy pape r.
Ne\"rs pr int . Tbe pape r on which ne\~spapers are print ed amI of ten used fo r
press ing plants; also ca ll ed s pe cimen pape r.
• Numbe r ing . The process of s tamping or printing sheet numbers ( access ion num-
bers ) on mount ing pap er or herb ar i um specimen s .
Pack Frame . A l i ght wooden fr ame Hith a shoulde r st r ap fo r back-packing .
Nounting Paper . The heavy herbarium paper to which specimens a re a ffixed .
Pa cket . An envelope fold ed and mo un ted on a herbarium s h eet to c on tain extra
mat eri als a nd f ragmen ts .
Paper Folde r . A smoo th st r ip of bone (I"rood) , r ounded at all corner s , used in
creasing paper , pressing dO\m glued paper , etc .
Para- dichl orobenzene (PDB o r moth crysta l s) . An insecticide o r repellent com-
monly used in he r ba r ia .
Pigeon - hole . Compartment in a herbar ium case in which the fold e rs of speci-
men s are ins e rted .
Pl a nt Dr yer o r Dr ye r. An appa r atus fo r drying plant specimens by artificial
heat (commonl y cons ists of a fram e to suppo r t the pres ses a bove a n e lect ric
hea t e r, se rie s of li ght bulbs , camp stove , or lan tern) .
Plant Press . An appa r atus for fla ttening and drying plant speCimens , usual ly
consisting of two light \~ eight boards or fra mes and a pair of straps or
s ash c or ds .
Pocket . An e nvelope that is pasted to an he rbarium sheet wi th the specimen to
contain ext ra pieces or detached fra gments o f the spec imen .
Po hI ' s So ft ening Agen t. An excellent de tergen t so lution f or so ften i ng f l ow-
er s and fruits for dissection .
Po rtfolio or Field Pre ss. A light plant pr ess for c arr ying in the fie ld I",hile
co l lecting , in to which the specimens may be placed immediately as they a re
gathered .
Pr epa r ation (spec i men) . The process of gett in g specimens mounte d and r e ad y
for i ns ertio n in the herbarium .
31 771

Press . An apparatus for flattening and drying plant specimens .

Protectant . See repellent.
Quire . A number of sheets of paper folded once, resulting in a number of
folds one inside another.
Registration (o f specimens) . The recording of each speci.men to be inserted in
the herbarium, a procedure now rarely follO\~ ed .
Repe llen t . A chemical used continuously in herbarium cases to prevent i nsect
in festations (also called protectant) .
Replicate . One of the specimens of a series or suite collected at the same
time and place , under a single collec tion number, to represent a parti-
cular species, often called a duplicate; an extra sheet of a collection .
Reprint . A separate copy of a journal article, specially printed for private
distribution.
Scheda , Schedula . A label; in scheda--on an herbarium label; e x confus i one
schedae - from confusion ot a label .
Secateurs (clippers) . Small hand clippers for cutting tough or woody stems .
Set (of specimens) . j.Jhen a collection is made with more than one sheet
(replicate) of each number, these are sorted into lots containing one
sheet of each number, called sets.
Selving . Reinforcing the attachment of herbarium specimens to mounting sheets
by loops of thread around the stem and through the paper .
Sheet . A standard sheet of heavy paper 11 1/2 x 16 1/2 inches (29 x til cm .)
or other standard size on I"hich a dried plant specimen is mounted; often
used colloquially to refer to the specimen that is or 11ill be attached to
a sheet.
s . n . (sine numero) . Literally \1ithout a number; a designation for citing a
collection lackin g a collection or collector ' s number; e.g . , John J . Doe,
S.n .
Sorting (of specimens). Arranging of specimens for insertion in the herbarium .
Space Treatment . See fumigant.
Special Collection . Any materials kept apart from the genera l collection ; e . g . ,
~"ood specimens, carpological collection, type collection, microslide col-
lection.
Spec i es Cover . II light manilla folder in Hhich specimens belong ing to one
species are filed 11ithin a genus cove r .
Specimen . II plant or fragment of a plant, taken and preserved to represent
a species or other taxon or a population, or to serve as a voucher to
document observations made on a plant or population of plants .
Spirit Collection ( liquid) . Separate collection of plants or plant parts pre-
serv ed in liquid preservatives .
Staff. Collective term to apply to all the employees of an herbar ium.
Sterile Specimen . II specimen lacking reproductive parts (flm"ers , fru i ts,
sp orangia ) .
St r apping . Affixing specimens to h erbarium sheets by means of strips of
gummed cloth, plastic, or glue .
Stripping . Colloquial term applied to strapping, particularly when plastic
is used .
Stripping or Strapping Plastic . A plastic used to affix specimens to herbarium
sheets; see Archer Hethod .
Type . See type specimen .
Type Specimen . The speCimen to which a name is permanently attached .
Type Collection . A special or separate collection of type specimens (holo-
types , isotypes); the type specimen (holotype) and its duplicates .
Unicate . A specimen of I.;hich there are no duplicates .
 772 31

Undetermined (undet.). Refers to unidenti f ied materials i n the herbarium;

spe cial senus covers may be so marked .
Vasculum. A col l ec ting container or can, Hhieh is usually made of sheet metal ;
standard vascula are 18-24 i nches long , 8 inches Hide and 10 inches high
and have a hinged door provided with a secure fastener on one side .
Ventilated Press. A plant pres s so made that Harm air may pass through it to
hasten the drying process.
Ventilator . A 12 x 17 inch sheet of corrugated metal, preferably aluminum, or
of corrugated cardboard, or of lattice bamboo splints, Hhich are laid be-
tHeen blotters and folders o f pl ants to facilitate the passage of air
through the leaves.
Voucher . A specimen preserved to substantiate recorded observations, and to
which reference may be made in the future to verify the identity of the
plant on Hhich studies were made.
V-shaped Rest. A f rame f or ho lding genus covers and specimens Hhen in use in
the herbarium . Large ones on Hheels are excellent for holding and moving
specimens about the herbarium \vhen filing.
\}ebbing Strap. A strap made of heavy cotton or linen fabric used on plant
presses .
Hetting Agent . A variety of solutions used to soften f low ers , fruits, etc.
for dissection.
Hhite Glue. Gelatin base or synthetic liquid used to paste specimens to her-
barium paper; e . g., Elmer's Glue, Hilhold, Fish Glue.

HERBARIUH LI TERATURE

Arber , A. 1938 . Herbals, Their Origin and Evolut i on . 2nd ed. University
Press, Cambridge .
Archer. W. A. 1950 . Ne\v plastic aid in mounting herbarium specimens .
Rhodora 52: 298-299 .
Beaman, J . H. 1965. The present status and operational aspects of univer-
sity herbaria. Taxon 14 : 127-133 .
Brenan , J . P . H. 1968. The relevance of the national herbaria to modern
taxonomic research [in Great Britain] . In : Hodern Methods in Plant
Taxonomy. V. H. HeYl-lOod (ed.) . Botanical Society of Brit i sh I sles and
Linnean Society. Academic Press. London.
Clokie, H. N. 1964 . An Account of the Herharia of the Department of Botany
in University of Oxford. Oxford University Press. London .
Cronquist, A. 1968 . The relevance of the national herbaria to modern
taxonomic research in the United States. In: Modern Hethods in Plant
Taxonomy . V. H. HeYl.ood (ed . ) . Botanical Society of British Isles and
Linnean Society . Academic Press . London .
Dalla Torre, C. G., and H. Harms . 1 900-1907 . Genera Siphonogamarum ad systema
Englerianum conscripta . Leipzig .
Dandy, J . E . , editor & compiler . 1958 . The Sloane Herbarium . British Huseum
(Natural History) . London.
Davis , P . H., and V. 11 . HeYlwod. 1963 . Principles of Angiosperm Taxonomy.
D. van Nostrand Company, Inc . Ne\v York .
DeHo1f, G. P ., Jr. 1968 . Notes on making an herbarium . Arnoldia 28 : 69-111.
Fosberg , F . R., and H. Sachet . 1965. Nanua1 for tropical herbaria. Regnum
Vegetabile 39 : 5-132.
Franks, J . l~ . 1965 . A Guide to Herbarium Practice . Handbook for Huseum
Curators , Part E, Section 3 . The Huseums Association . London.
31 773

Irwin , H. 0., W. Payne, D. Bates, /:. P . Humphrey (compilers & editors) . 1973.
America's Systematics Collections: A National Plan (Prepublication Draft).
Jones , G. N., and E. Meadows . 1948. Principal instit ut ional herbaria of the
United States . American Nidland Naturalist 40: 724-740 .
Lanjouw. J . • and F. A. Staf l eu . 1964 . I ndex lIerbariorum . Part 1. The her-
baria of the worl d. 5th ed. Regnum Vegetab11e 31: 1-251 .
Lawrence , G. H. M. 1951. Taxonomy of Vascular Plants . The Hacmillan Company.
New York .
Lellinger, D. B. 1972 . Dichlo rvos and lindane as herbarium insecticides .
Taxon 21 : 91-9 5 .
Lundell , C. L . • and R. Kirkham. 1966 . A method of applying Nystox (lauryl
pentachlorphenate) to protect mounted herbarium specimens. \-1rightia 3:
1- 7-180 .
f>lil l spaugh , C. F. 1925 . Herbarium o r gan ization. Fie l d Nuseum Technic al
Services 1 (3) : 1- 1 8.
McNeill, J . 1968. Regional and local herbaria . In : Nodern Nethods in Plant
Taxonomy . V. H. Heywood (ed . ). Botanical Society of British Isles and
Linnean Society . Academic Press . London .
Pohl, R. \01. 1965 . Dissecting equipment and materials for the study of minute
plant s tructures. Rhodora 67: 96-96.
Porter , C. L . 1959 . Taxonomy of Flower i ng Plants. t·1. H. Freeman and Com-
pany . San Francisco.
Rollins, R. C. 1955 . The Archer method for mounting herbarium specimens .
Rhodora 57: 294 - 299 .
1965 . The role o f the university herbarium in research and teach-
ing . Taxon 14: 115-120.
Shetler , S . G. 1969. The her barium: past, present and future. Proceedings
of the Biol ogical Soc iet y of Washington 86: 687-758 .
___-,._' et a1. 1973. An introduction to the botanica l type specimen
register . Smithsonian Contributions to Botany 12 : 1-184 .
Smith, D. E., Jr . 1971. Preparing herbarium specimens of vascular plants.
Agricultur e Information Bulletin No . 348 . Superintendent of Document s .
Washington . D. C.
Stearn. h' . T. 1957. An introduction to the "Species Plantarum" and cognate
botanical ,,,a rks of Carl Linnaeus. Prefixed to the Ray Society facsimile
of Linnaeus's Species Plantarum, 1 . London .
1971. Sources of information a bout botanic gardens and herbaria .
Biologic al Journal of the Linnean Soci ety 3 : 225-233 .
Stern, \.;r. 1. 1967 . Index Xylariorum . Regnum Vegetabile 49. Utrech t.
__",,,,,_' II. B. Gillenwater , G. Eason, A. Garcia-Quintana , and R. S . Cai!.
1968 . Lindane and dichl orvos for p rot ection of herbarium specimens against
insects. Taxon 17: 629-632.
Stuckey , R. L . 1971. The first public auction of an American herbarium
including an account of the fate of the Baldwin, Collins, and Rafinesque
herbaria. Taxon 20 : 443-459 .
Turrill, H. B. 1964 . Plant taxonomy, phytogeography and plant ecology . In :
Vistas in Botany . I"; . B. Turrill (ed . ) . IV : 187-22ll.
I 774

"ERHRIU," OF Til. U"'VER~IT" V I' .~"".T" CA " ()L'N~

31

TEXAS
~ee Co.
Ike
Pvrrhopappus multlcaul1s DC .

Fine har d sandy loam , grassy r oadside , eight

miles northeast of IIcevillc at Hedio Creek.

Flowers pal e sulfur yellow , tinged rose - violet ,,_ ,-";., ,i.e, ," It.~ "oo fo, "'''''..",,''
outside; anthers black (11 AM); CO=lon. """'"' """,,,J [,... ,h. 'r« ' ~'n.
C. A. Lawson 928 28 HaTch 1967

PLANTAE EXSICCATAE GI(AYANAE ANNOTA TI ON LABEL

1·127 . Sibara v;'~l n lc3 ( 1.. ) 1("..,... ,

In 1,10",10" , .Ii;;, ~ ~ I ('!HI) et in Con"ll>. Gr>f l!<l~.
01.", 1:);: (1911 ), Fom.ld ,G r3Y', ~!"". ( .. L SJ 7~3( I~.\O). Oct. R,nald L. M,G .....
S)'n. CO",",";,, ,'i'llinim L. Sp. 1'1. ti5G (I'"j) . ..I,,,.;,
';'II;.im ( L.) 1'01 •. Encr<1. S"I'I'I. i. -11:1 ('S 10): Hupkin.
in Il h.,Jo," , .. xi., 80 (19::11). C"rJm,;",I.. I~t'jcja ." !-look.
in .Ionm. 110\. i. 191 (IS;j,) . • I'obi, !,,,Iot·icia.,, ( Il ook. )
C. A . .lley. in Fl , ch. ,.. ~lcr. 1,,,1. Sem. lIort. I'drol'. ;< .
,iO (IS'~)' /'/"."'/" !,i'Ri.i<"", (1..) Green., L • .,H. Bot. I'r<&s<J by P. C. H""h;,,,,,,
Ob •. ii. ~~I (1~1~).
11",,," p),cc" Tubl" It .."I . oif l.owc', F<rry I'ike, odd ""mber
);"0,. C""n 'y. 1"",-,,,.,,,,. G,own f,om ;Olpor1<d pl."" of ,oe .J";.,,, ,011«"0<1"
Coli. A. J. SU'"" (no. ~ ".) .110,. 'i. 10' , l'otve""y of C,i;lo,llta !lou.;, Gord,n . ilr:,kJ<y.

BAJA CAl iFORN IA. MEX ICO

I'I. ASTS UF TI!~ BOCKY MUUloiTAJloi BICG!OS

FLORA "I QUElIEC, CANADA
The q",I"" P"nrn,,,ia Colorado
H"b"'u m of K«n< S,"," C"lIe.e

Coun,y

S o.

Cellwo,. I). F~
July I'>"
llcuHe,d B""d C. Hul!;",. Suo
. \11.
".

Figur e 31 - 1 , Sample herbarium and annotation labels .

(Actua l size of the UNC label is 2 3/4 x 4 1 /2 i nches.)
32 775

Chapter 32. BOTANIC GARDENS AND ARBORETM'

The purpose of the general survey of botanic gardens as presented here is

to g i ve the student of natural sciences an appreciation of the heritage of
botanic gardens, and an understanding of the important scientific and public
services t hey p r ovide today , as well as an idea of their potential for future
serv ic e . Some o f the material on the organization and developmen t of botanic
gardens may be o f litt l e value to the average student; but it i s in cluded here
to give the pro fe ssional botanist a feeling for the practical and phi lo sophi-
cal problems basic to the operation of botanic gardens . Botanic gardens may
be utilized by many d i fferent individuals and groups \~ith general or specific
interests , such as :

Botan i sts Othe r Botanic Gardens Artists and Photographers

Zoo l og i sts Industry Horticu ltu ral Soc i eties
Ornithologists Landscape Architects Garden Clubs
Arborists Teachers Home Owner s
Conservationists Students General Public
Nurseryme n

Each \.,rill be specifically con c ern ed with perhaps only one aspect t"ithin C,le
botanic garden program . The extent to \~hich a botanic garden is able to fu l-
fill each special interest ':lill depend on many factors as will be pointed out
in the fol low i ng pages .

Strict definitions of arboreta and botan ic gardens are difficult because

o f their varying degrees of d evelopment and overlapping purposes and functions .
Arboreta and botanic garden definition s in this chapter are e xtracted from the
National Recreation and Park Association Bulletin No . 90, Arboreta , Botanical
Gardens, Special Gardens (Bunc e , 1971) \.Jhich was prepared by professionals in
the fie ld of botanic garden structure and development .
1 . Arboretum . "An example area set aside f or the growing and effect ive
display o f all the different kinds of \wocly ornamenta l trees and
shrubs , vines, and other plants which may be grown in a given area,
their maintenance, proper labeling and study" (Hyman, 1959) .
2 . Botanic Garden. "An institution organized to maintain plant col l ect i ons ,
usually repr esenting a large number of genera and species ... to serve
educational, aesthetic , scientific o r economic purposes . Recent ly ,
botanic gardens offer a source of recreation" ( from Fred ~ndmoyer ,
Secretary-Treasur er , American Association of Botanical Gardens and
Arboreta contribu t ing to American Encyclopedia , 1967).
Maintained p l ant areas include many other types of developments such as pr ivate
estate gardens open to the public, recreational parks , histo r ical restorations,
public and commercial display gardens , and even landscaped high~"ay right-of-\"ay
borders and median strips. These various developments arc generally designed
to provi de aesthetically pleasing surroundings. Arboreta and botanic gardens
assume t he additional r esponsibilities of education and research. Since they
are frequent l y quite similar in design and function , they \vi ll be treated as
one and the same in this present discuss ion (\vith the exception of comments
concerning the historical development of the botanic garden) . Botanic gardens
and arboreta share the follovJing basic objectives :

"'Contributed by J . Kenneth Hoore , University of North Carolina , Chapel Hill.

 32
776

1. Display of plant material for the general public .

2. In troduction of new plan ts into cultivation .
3. Pla n t exp l or a tion of r emote nat ural regions .
4. Sci.entific i nves ti ga tion s in su ch ar eas as plant breeding , hybr id i za-
tion . a nd taxonomy .
5. Devel op ment and maintenance of an he rbari um .
6. Devel opmen t and maintenance of a lib r ary .
7. Developmen t and maintenance of var i ous r esearch l aborato r ies .
8. l'laintenan ce of representatives of the en tir e plan t kingdom f rom the
poles t o the t rop ics grown o utside or under glass .
9. General advanc ement of botanical st ud ies and horticul t ure.
10 . Conse r vat ion of natural areas for ecologic al s t udie s .
11 . Es tab l i shment and maint e nance of gene pools .
12 . Dissemination of informa t ion gained through plan t sciences .

I. HIS TO RY

\oJ . T . Stearn (1971) has summar ized the significan t sou rce s of information
r ela tin g to pl ant col l ectors , he rbaria , and botanic gardens . The general his -
t o r ical s urveys of t he development of botan ic gardens by A. '.-I . Hil l (1 915) and
St3fleu (1969) are the foundation f or th e f ollowing summary accoun t.

A. The Earliest Bot an ic Garden s . Though ga r dens exi sted in a nc ien t Egypt
and Ne sopo t amia for the g r owi ng of herbs , f ood plants , ornament al s , or for
pleasu r e and as sta tus symbols , they c an not in the p rop e r sense be de signated
as botanic garden s , that is , a ga rd en in wh i ch plants are colle c ted and main-
ta ined for scien tif i c p ur poses . Bearing in mind the p rima r y function of science
and e duca tion , th e fir s t botanic garden may be consi dered t o have been tha t of
t he " fath er of bot any ," Theophrastu s , whose garden , atta ched to hi s school , the
, "Lyceum," near Athens, was bequea th ed to him by his former teacher , Aristotle .

The Romans uti l ized small ga r dens as sources of medicine and aids to medi-
cal studies . The first such gardens In Euro pe are a ttr ibu t ed to the necessity
for medici ne a nd dr ugs . The medieval monastic gardens or i ginated in c.he late
8th century du ring th e re ign of Cha rlemagne , who aSsigned the sp eci fi c ta sk of
medic al t ra ining to the monasteries . The ty pical monastic ga rd e n included the
"ho rtus ," for vegetables and fruit , and t he "h erb ular is," for var io us herbs ,
the lat te r being the precursor of the phy s ic gardens \-Shie h were of ten es t ab-
lished in connect ion with the medical f a cu lti es o f t he universities during the
16th a nd 17th centuries .

Th e credit fo r the establishment of the first mod e rn botanic gar den belongs
to the Ita lian , Luca Ch ini (ca . 14 90 -1556) , a professo r of bo tany c alled from
Bologna to Pis a i n 15 43 or 1544 , at \"thich time he immedia t ely bega n pl a nt ing
a bo tanic garde n connec t ed with the univer s ity . Othe r un i ve r s i ty botanic gar-
dens immed i ately fol l m~ed at Padua and Florence i n 1 545 ; al l three of these
garden s wer e aided by t he pat r onage of th e Medici family . The e stablis hment
of o th er i mpor t an t gardens fo l lm"ted in succession : Bologna , I taly (1567) ;
Leyden, Netherl ands (1587); Nontpellier , Fran ce (1 593 ); Heidelb urg , Germany
(1593 ); S tra sbou r g , France (1 619); Oxfo r d , Eng land (1621) ; Paris , France (1653);
Groningen . Nethe rland s (1642) ; Be rlin , Germany (16 46) ; Uppsala , Sweden (1655) ;
Edinb urgh , Sco tl and (1670 ); Ch els ea, Englund (16 73) ; Amsterdam , Net herlands
(168 2); Vienna , Austria (1 754); Kel" . England ( 1760) ; Cambridge , Engl and ( 1762);
and Coimbra, Po rtuga l (177 3) . Luca Chin i .is ""Is o credi ted I'l ith havin g est ab-
lished the fi r st herbarium; he taught scienc.ific course s in plant t axonomy
not l imited s imply t o medicinal herbs .
32 777

Though the Pisa Botanic Garden does not ex ist t oday , records of its
design demonstrate the geometric a rrangements of plantings o r iginat ing t"ith
the monastic garde n s and characteristic of many c on t inen tal garde n s up into
the presen t century .

Th e history of botanic gardens logically fal l s into distinct pe riod s

coincident td th world e xploration and subsequen t i ntroduction in to Europe of
plants from distant lands (Stearn, 1971, Stafleu, 1969).

B. European Period (to 1560) . Plant collections of this time were prin-
cipally o f indigenous European s pecies and those collected from southern and
southeastern Europe and the adjacent Mediterranean l ands , including Egypt.
Some species had to be maintained in t ubs as pot- plants. The pot-plants wer e
kep t i n "cubicula tepida" during t he win t er ; t hese glass - houses were simp l e
r ooms with windows fac ing south, another important innovation at tributed to
Luca Ghini.

C. The Near East Period (1560- 1620). This oriental pe r iod of plant in-
troductions fro m southea stern Europe a nd adjacent Asia may be chara cteriz e d by
a few groups spectacular for th e fragrance and the brilliant colors of their
flm~ers such as hyacinths, tulips , and l ilies . The Fl emish -Aust rian botani st
and father of the b ulb i ndustry, Carolus Clusius , an inquisitive natural
scientist, botanical traveler , scient i fic hor ticu lturist. and plant taxocomist ,
was very inte r ested in t his gro up of introduced plants . His st udy and travels
carried him through the Netherl ands , Vienna , Frankfur t, Leiden , Hungary. Ita l y .
and Spain ; a ll the ,~hil e he was engaged in obta ining and describing new plants .
Clusius became a dominant fig ur e in botany during t he las t half of the 16th
century , and as dir ector of the botanic gardens at Vienna and Leiden, he
greatly in flue nced the development of botanic gardens in Europe . By the end
of the Near East period the center of plant activity had shifted f rom Italy
to Austria, Germany, and the Neth e rla nd s .

D. The Period of Canadian and Virginian Herbac eous Plants (1620- l687).
During this period the center of botanical activity turned to France , co i nci-
dent with Fr ench explora tion following t he decline of Spani sh sea pm~er .
l"hile the Dut ch t,1ere involved in the tropics and th e African Cape . and t he
English with their Virginia territories, the French exploited the Canadian
wi lderne ss . In trod uct ions i n t o the botani c garden i n PAris include d s uch
fam i liar pl a nt s as arborvitae, sumac and poison ivy. bl ack locus t. and peren-
nials such as Black-eyed Susan, Dutchman ' s Breeches , and Goldenrod . Th e
botanic garden at Paris , known as the "Ja rd in du Roi" or "Jardin des pl ant es ,"
was in no way a pa l ace garden; it was an in dependent educational and scienti-
fic institut ion with a royal endowment . originally founded as an establishment
to promot e the teaching of pharmaceutical bo tany . Pres e ntly its collections .
particul ar l y its herbarium and paleobotanical and paleontological departments .
are among t he world' s bes t; it is the oldest and most important non-unive r sity
bo tani c garden still in exi s tenc e . Some of the pri vate gardens of England
rece ived t heir first North American plants by way of French i n trod uc tions . A
few decades l ater p rivat e gardens of Engla nd, such as that of the Tradescants
at Lambeth , had early Virginian i nt roductions wh ich in cluded red map le and
tulip popla r, thus marking a second phase of this botanical period . The intro-
duction of North American speci es to t he Ne therland s from Dutch se ttlements
began a third phase .
778 32

E. Cape Period (1687~ 1772) . Dutch sea traffic around the Cape of Good
nope during the golden age of Holland is responsible f or the most character-
istic plant introductions of this period. By nm... g lass h ouses and conserva-
tories had become much more sophisticated so that the introductions o f geran-
iums and numerous succulents could be better acconnnodated. After travel in
South Africa and India, Palll Hermann, a German immigrant, ,,'as appointed pro-
fessor of botany at Leiden • \·,here he Has instrumental in the great incres.se
i n the plant collect ion at the Leiden botanic garden . I n Amsterdam at about
the same time, 1682, Jan Commelin \-las appointed to create a botanic garden .
The e fforts o f these t'lva botanists stimulated a strong interest for plants
in the Dutch sea merchants \"hich resulted in vigorous and enthusiast ic plant
collecting . 1.innaeus completed his botan ical training in Jlolland, utilizing
the latest collect ions \"hich often contained plants representative of fami-
lies previously unknmm to him. From 1735 to 1737 he \"<1S employed by George
Clifford , a wealthy Dutch banker, to describe the colle cUons in his mill
private botanic garden, De Hartecamp , H\lich included a permanent herbarium
nOl-J at the British Huseum . Ihth access to Holland's excellent botanical
libraries and print e rs, Lirmaeus \"as able to complete and publish "Systerna
naturae" (1735), "Bibliotheca" and " fundamenta botanica" (1736) . "Ge n e ra
p1antarum" (1737), and "Hor tus Cliffortia nus" (1737) . The latter desc rib ed
the collections of Clifford's garden , many of which Here introductions from
the Cape . I n 1741 Linnaeus became director of the botanic garden at Uppsala,
Sweden . Prior to this time the garden had been allO\"ed to deteriorate to
such an extent tha t plant collections I'Jere dmm to fel,'er than 300 species .
Through Linnaeus ' recommendation, the garden obtained the servi ces of the
chief gardener of Clifford' s estate, 11ith the result that by the time
Linnaeus Has stationed at Uppsala , the garden Has b eginning to i mprove.
Under Linnaeus ' sup ervision plant collections had graHn to over 3 ,000 species
Hithin seven years time (Blunt, 1971) .

The Cape plant s reached England almost e xclUSive ly through Holland, but
during the course of the 18th century, direct int roductions into England
rapid ly increased . By the end of thi s period the English gardens began to
rival those of Holland; the gardens of both nations had surpassed the impor-
tant German, French, and Italian gardens of earlier periods .

F . Period of North American Trees and Shrubs (1687-1772). Through the

efforts of Lancelot Brm-ro, English gardening exper ienced a revolution in the
development of an open landscape style. free fr om the rigi d geometrical for-
mality Hhich had persisted in the continental gardens from the monastic in fl u-
ence . Introductions of 11eH Hoody species from North America gained promi-
nence. Nany of the int roductions r eached England through the efforts of Peter
Coll inson, ",hose private gardens at Peckam and Nill Hill \~ere significant
botanic collections. Collinson corresponded extensively Hith the American
naturalist, John Bartram , Hho is credited \-Jith establishing the fi rst botanic
garden in America in 1731 on his estate on the Schuylkill River near Phila-
delphia . Bartram studied and propagated the native plants prior to send ing
them to his fr iend Collinson. Through Collinson many of the American intro-
ductions reach e d other gardens such as Philip Hiller's at Chelsea , and the nelV
gardens of Princess Augusta at Kct,] \,]hich ",e re supervised by IHlliam Aiton .

The great botanic gardens of England IVere preceded by the establishment

of several private gardens devoted to the cultivation of medicinal herbs and
other plants of botanical interest. The establishment of the botanic garden
at Ox f ord in 1621 Has specifically for the purpose o f cultivating "physic"
32 779

plants and for the demonstration of principles in the study of botany . The
Chelsea Physic Garden l"a8 founded in 1673 in London as the garden of the
Society of Apothecaries. Sir Hans Sloane, who deeded the land for the
Chelsea garden, served botanical study hle11 when he stipulated that fifty
preserved specimens of distinct plants from the garden be presented to the
Royal Society of London each year uotil tHO thousand distinctly different types
of p lants tvere attained. The <lppointment in 1723 of Phil i p Hiller as Head
Gardner \"a8 an important event for the garden . Under. his practical skill and
botanical knoHledge. many of the various plants net.,rly acquired from allover
the lvorld were be ing grmm and flm"ered for the first time in c ult ivation .
Hiller ,,,as also the teacher o f l"Til1iam Aiton, the first curator of the Roya l
Botan ic Gardens at Ke"' . Ke'" OI']es its establishment, in 1760 , to the interest
of Princess Augusta, Princess OOl']ager of Hales, "'ho set aside a portion of the
Royal Ga r den at Ke,,, !!ouse for a physic garden . Ihl1 iam Aiton "'as in charge of
this garden for thirty-four years . After the death of the P rinc ess in 1772,
George III inherited the Kew property and united the gardens ",ith those l ying
contiguously ",hich had formed the gardens of the Palace of Richmond , thus pro-
ducing the extensive area knOl-ln as the Royal Botanic Gardens , KeH . Ke'" dif-
f ers from other botanic gardens in that it has no connection '''ith any univer-
sity or educational organization as such , thus its usefulness has largely been
with the economic aspect of bo tany, assisting and encouraging botanists , trav-
elers, merchants, and manufacturers in var ious botanical investigations .

G. The Austra]jan Period ( 1772 -1820) . Ke,,, Garden ' s Si r Joseph Banks
accompanied Captain Cook on his First voyage (J768-]77l) marking the beginning
of the Australian period of dcvc']opment of hotan ·i .... al gardC'f's . Spe cies of num-
erous unfamil ia r representatives of southern heT!'j"'''Jl'~re families of plants
wer e discovered and introduced to Eng' and and fr a n there 1·0 the continent .
The tropical botani.c garden~' '''ere developed du rjng the Australian period .
Great Bri.tain is credited I"ith establishing the firs t of these economic g a r-
dens (for the cultivation of sp i ces and other commerci al plants) on the i s land
of St . Vincent, British Hest Indies .• in 1764. Th e CalClltt n Botan i c Garden ,
founded in 1786 for cu1tivfltion of spices, is no t(>d for pota to cultivation
and the introduction of tea, mA.hogany, jute , sugar cane, and the quinine-
yielding cinchonas . The Botanic Garden at Buitenzorg in .J,1V3 , I ndones ia ,
f ounded in 1817, is noted for the cultivation of rHbber ane! coffee . Other
tr.opical gardens Her e subsequently established in Halaya and Ceylon during
this p e riod .

Botanic gardens lost the restrict :ivE' character of phy~ic ga rde ns and
became more extensive in size and function , ,·lith the result tha t e v e ry I,te l l
knut·m university '''ithout a botanic garden be g an making plans fo r one . It ",as
near the ~nd of this period that true bot,mic gardens began to appear in North
America . The Elgin Botanic Garden, established in 1801 , accumulated a valu-
able plant collection and became the Botanic Garden o f the State of Ne,'] York
in 1810 . It "'as subsequently granted to Columbia Col l ege . Although i t does
not exist today, several of this garden ' s associates, Thomas Eaton , John
Torrey, and Asa Gray, Hill be remembered as SOI:le of America ' s leading bo tan-
ists .

iI. Subseguent Periods .

1. Period of tropical glasshouse plants and hardy plants fr om Japan

and North America (1820-1900) .
780 32

2. Period of \.Jest Chinese plants (1900-1930).

3. Period of hybrids (1930 to present) .

It was during the last three periods, and pr incipally the last tl"0, that
American botanic gardens \~ere established . Be f ore 1900 the Missouri Botanical
Garden or Shalv I s Ga rd en , St. Louis ( 1859). the Arno ld Arboretum, Harvard Uni-
versity (1872), and the New York Botanical Garden, New York City (1891). were
the major botanic gardens established. American botan i c gardens and
arboreta did not appear. in significant numbers until after 1920 . Presently
there are more than 15 5 such gardens throughout America. In spite of the dis-
tinction between botanic gardens and arboreta, it is necessary to us e the two
designations in terchangeably when referring to American operations, particu-
larly because the arboreta, like botanic gardens, often boast greenhouses
and displays o f he rbaceous plants .

II . ORGANIZATION AND DEVELOPNENT

This section pertaining to organization and development is reduced to

outline form because it does not bear directly upon the significance of botanic
gardens in taxonomy and related botanical research . The basic information pre-
sented here is included as an effort to demonstrate in general the total
machinery necessary for botanic garden activities . Students of botany as I~ell
as plant researchers who utilize garden facilities and materials often fail to
see beyond their own specialized interests . It is important that they be
aware that the i r individual interests are only a small part of the total pro-
gram essential to a botanic garden .

A. Personnel . Depending upon resources availab l e, botanic gardens , whether

public or p r ivate, are similarly structured as follows (Richmond, 1971) :

Botanic Garden Board - - - - - - and

Suppo rt Group
Governing Body

Director

Professional Staff Clerical Staff Labor Force

Assi stant Director Secretary Nurseryman-Foreman
Education and Program Specialists Clerks Gardeners
Horticulturists
Lib rarian

1. The Board .
a . Holds full authority for , and responsib ili ty in , all matters
pertaining to the operation .
b . Estab lishes garden policies .
c . Approves garden budgets .
d . Receives and expends all garden funds .

2. Support Group .
The support group may be parent to a trust fund establ i shed to
receive and expend money for operational purpOSes or may be
active in raising f unds fo r garden use .
a . Fiscal support for the garden.
b . Establishme nt of a trust fund for garden use.
c . Publication of newsletters and period icals.
32 781

d. Vol unteer work in the garden.

e. Public re lations .

An act ive and vigorous support group is not only one of the most
valuable assets a botanic garden or arboretum can have , i n many
cases it is essential to the f und ing necessary for survival of the
garden.

3. The Director .
a. Recommends policy to the Board.
h. Implements the policies of and is responsible to the Board .
c. Responsible f or th e staff 's day- to-day operation .
d. Plans further development fo r consideration by the Board.
e. Prepares and submits budgets to the Board for its considera-
tion.
f. Maintains professional liaison with other botanic gardens ,
arboreta, and similar institutions.
g. Promotes public relations through speaking engagemen ts , the
press , and other media .
h. Procures funds through active sol icitat i on of grants, contri-
butions, and other monies from public and p r ivate sources .

4. The Staff .
Carry out day-to- day operations and ma i ntenance under the gen-
eral supervision of t h e Di rector . Profess i onal staf f member s
also advise the Director in matters of their specialization.

B. Financing the Botanic Garden . Hith increas ing costs and responsibili-
ties, even some of the most liberally endowed organizations must seek addi-
tional operational f unds f rom both public and private sources .

1. Gove r nment Sources .

A number o f Federal agencies such as The National Science
Foundation, The Agr icultural Research Se rvice, Soil Conserva-
tion Service , and the Forest Service have funds set aside for
special projects for Hhich arboreta and botanic gardens may
qual i fy . State f unds aid through university budgets , special
boards, spec ial legislation, and in cases ,,,here i nstruction is
a significant part of the program, through the State Department
o f Education.

2. Private Sources.
a . Contributions . Varying amounts may be acquired from indivi-
dual membershi p or actively solicited from large industrial
concerns , particularly thos e based upon the extraction of
nonrenewable resources .
b . Ph ilan thropic Organizations . Societies and Foundations active
in advancing education , science, conservation, or cultural
activities are logical sources .
c . Bequests. I n f requently, substantia l sources of funds are
bequests from individuals Hho have had long and pleasant
associations ,·Iith the botanic garden .

C. Planning. Significant botanic gardens are based upon thoughtful prior

planning . Individuals qualif i ed in this area have published var i ous master
782 32

plans e mphasizing plann i ng befor e development (lolyman , 1970. and Longene cker,
1964) . The Bulletin of th e American Assoc i ation of Botanical Gardens and
Arboreta is the p r i ma r y vehicle of distribution of info rma ti on pertaining to
developing and maintaining garden facilities . In addition to fo rmula ting the
general purpo ses of a bo tanic garden , plan ning inc lud es basic matters such as
complete contour and vegetational mapping of available land areas , soil stud-
ies, initial tra i l , path , and road sys t ems , nu r sery a r eas , and designation of
existing plant groups to be set a side , undisturbed .

D. folaintenance. Maintenance , t he single mos t i mportant fac t or (aside

from a good preliminary plan ) is often ove r looked and under-budgeted during
cons iderat ions of botan ic garden fu nctions and activities . In addit i on to
the more obvious mechanical efforts invo l ving utilization of tractors , chain-
saws , and var io us gard ening tools , are s u ch indispens ib le activities as :
1. Haterin g (in cl uding i nstallati on and repair o f wate rin g s ystem) .
2 . Protec tive fencing .
3 . Plant propagatio n.
4 . Rep la cement of old or dead plants .
S . Happing and cons tructi on of ne,. trail s and mainten ance of old ones .
6. Maint ai n ing re co rds pertaining to all co l le ct ions of p lant s .
7 . Labe l ing of plants .
8 . Soil ma in tenance ; fertilizing , fu miga t ions. s teri lization . e t c .
9. Police or other. g uard force .

E . De sign . Gene ral designs of botanic gardens have been molded largely
by f ashion. The gardens o f the continent developed befo re the 19th century
mainly conformed to the geometri cal arra ngements dictated by the medieval
monastic garde n s . Later , garden design. tho ugh still confined ,.1thln a geo-
me tr ical framewo rk, began t o r eflect the growing understanding o f plant rela-
tionships . Emphasis "J 8 S placed on gro up ings of similar types of plan t s or
p l ant fami l ies s uc h as found at Kew , or of va rious phy l ogene t ic sys tems as
exempl if ied by A. L . de J ussieu in the gardens of Tr ianon , France, at the
close of th e 1 8th cent ury . This system of disp lay design was imitated by
others includ ing de Cando lIe at Geneva . As l..a rld exploration opened up dis-
tant lands , display designs ,.ere modified to exhibit plants in geographical
groups . The se general design frameworks i nc luding the open English designs,
represent a rich botanic garden heritage upon wh i ch Amer ican ga rde ns are
designed . Though small geometri c designs are still present in tribu te to
our heritage , t he general trend in the American design is an open l andscape ,
which is logica l, since until r elatively recen t l y spa ce in America was not
scarce . However, wit hin the open land scape design are to be found the more
traditional arrangements of p lant s grouped (1) as to t y pe or genus as at
t he Arnold Arbo r etum or the National Arbo r etum , \./ashington . D. C . , (2)
acco rding to geographical origin as demon strated at the Strybing Arboretum
and Botani cal Garden . San Fran ciSCO , Calif . , (3) illustratin g plant phy-
logeny from primitive to advanced fami l ies as designed by the Ar thur Hoyt
Scott Horticultural Foundation on the campus proper of Swartbmore College,
Swarthmore, 1'a ., an d (4) more r ecently lvith emph as is on plant commun it y or
hab itat arrangements s uch as developed at Rancho Santa Ana Botanic Garden,
Claremont, Ca lifornia .

The f oll owing o u tl i ne (Teusch er, 1933) demons trates not o nly t he oppor-
tuni ties fo r es t he t ic lan dscape design, but also the possib l e a r rangements
which facilit a t e various taxonomic or other botanical research bas ed on
living materia l.
32 783

1. Geographical or Ecological Groups . The emphasis i s on ecology ; an effort

is made to reproduce i n a natur al manner and on an extensive enough
scale to achi eve examples of var i ous characteristic types of vegeta-
tion.

2. The Flower Garden .

a. The perennial garden , demonstrat i ng the best selected varieties of
perennials for continuously blooming borders from spring through
fall.
b. The annual garden, demonstrating t he bes t var i eties in annual plants .
c. The water and hog garden , demonstrating water-loving plants such as
wate r l ilies gr own in basins or in natur al or arti f ic i al pools .
d. The ro c k garden , demonstrating not only col l ections typ i cal of the
local area , but possibly plantings to illustrate the flora of dis-
tant areas, as for example , Asiatic or European alpines .

3. \.Joody Plants for Special Effect.

a . The r ose garden, for beauty , hybridization , or as a testing site for
neH var i eties .
b . The spring flm... er garden , grouping the most ornamental spring-flm.;rer-
ing shrubs and trees perhaps in combination with spring- flowering
bulbs .
c . The autumn garden , grouping shrubs and trees ou tstanding for their fall
leaf color and colorful fruit in combination with fall-flowering
bulbs.
d. The color-garden , grouping the best of trees and shrubs for display of
fall foliage best exhibited against a background of evergreens .
e. The \.;rinter color-garden , a collection of trees and shrubs which dis-
play conspicuously color ed branches duri ng the winter .
f . The form or freak - garden , being a grotesque or humorous di splay of
\.;reeping, fastigiate, globe-headed, and d\.;rarf forms of trees and
shrubs .
g . Co l lection of vines, perhaps demonstrating potential as natural fe nces
or utilization with other types of plants .

4. Plants of Special Botanica l Interest.

a. The medicinal garden, featuring an ancient herb-garden or a demonstra-
tion of spice and poisonous plants .
b . The economic garden , featuring a vegetable garden and demonstrating
various grain crops , fodder plants , berry shrubs , fr uit trees,
fiber plants, and oil plants .
c. The morphological or biological garden , demonstrating any designated
plant function or design , such as vegetat i ve propagation, protec-
t i ve mechanisms, pol l inating designs, and seed dispersal .
d . The taxonomic garden, concentrating on the demonstration of particu-
lar genera or fami l ies , or groups of families in a particul ar sys-
tem of relationships . This and the preced i ng type display are
perhaps best closed to the public e xcept to students and interested
persons with permission .

5. Specia l Displays .
a . A desert garden . e. A flm·]ering Crabapple collection .
b . An evergreen 8arden . f. A fragrance garden f or the blind .
c . A he dge garden . g. Others .
d . A rhododendron dell .
784 32

I II . FUNCTIONS OF THE BOTANIC GARDEN

Activities of botanic gardens are many and variable , limited only by the
policies of the organization, the talents of the perso nnel , and the resources
of the facilities . These function s vary from general civic contributions to
disc i plined botanical research .

A. Public Service.
1. Social . Facilities of the botanic garden may be utilized for
meetings of various plant-associated groups and service organi-
zation s . l>lany such organizations make their headquarter s at
facili t i es furnished by botanic gardens .
2 . Cultural. If facil ities are adequate , various l ectures . art
exhibit s . or musical programs may be presen ted in the pleasing
surroundings of a botanic garden . As a consequence of proxim-
ity to botanic gardens, community businesses and private homes
are often more care fully landscaped and maintained.
3. Economi c . The presence of a botanic garden enhances the community
through its needs for personnel, goods, and services , and
depending upon visito r levels, it benefits local businesses .
Through cooperation with the nursery industry new ornamental
plants of possible commercia l value are introduced and through
display of new plant material avail able i n nurseries , gar dens
create a market for desirable ornamentals .
4. Recreational . Depending upon total structure of the garden pro-
gram and particularly on the space available , limited areas
may be open to the public for picnic and play areas. However,
such activities are best restricted to publi c park areas and
not included as a program of botanic ga rdens.
5. Spec ial Events . Other activities may include flower shm.,ls . garden
club projects , special or seasonal displays , and use of flowers
for hospitals or local functions .

B. Educat i on . l>lany botanic gardens are adjunct facilities to, or an actual

part of, an educational institution . As s uch , the particular garden organiza-
tion must provide courses or programs in plant related subjects , vary in g from
formal graduate level courses for college credit to popular one-day , non-credit
plant t ...orkshops. Nost significant independent botanic gardens are organized
to support their OIm plant related c ourse s serving much the same group of
peop le. Gene ral types of courses are listed beIOl....
1. Graduate and undergraduate level botany and horticulture .
2 . Short courses and Norkshops on many phas es of plant study for the
general public and professionals fr om other disciplines .
3. Natur e study programs for school groups.
4 . Environmental awareness pr ograms for pre-schoo l age groups .
5. Special workshops for teachers of natural sciences.
6 . Classes and demonstrations for professional garde ners .

C. Conserva ti on. \.Jith the increased pressure of pop ula t ion and develop-
ment upsetting or destroying natural areas, botanic gardens are in a position
of public and political influe nce not yet ful l y realiz ed .
1. Acquisition of na t ur al areas (habitat tracts) .
2. Planting of r epresentative native plants, cultivation of rare and
endan ge red species , and preserving various portions of the
botanic garden grounds in t heir natural s t ate .
 3. Various courses offe r ed to the general pub l ic \~hich str ess t he
importance of natural areas .
4. Preserva tion of var ious ornamental forms and varieties t~ h i ch may
be n eglec ted by nurseries because of cu rren t public unpopular-
ity .
5. Coop e r at i ve efforts with other conservation group s to lobby for
partic ula r legi s l ati on or governmenta l protection.

D. Research . A recent s urvey of bo tani c gardens and arboreta indicates

an extensive r ange of botanic r esearch being conducted at garden fa ci li tie s
and by garden personne l and associates (Chan, 1968) . Onl y t he mos t exten-
sive ly researched discip l ines are listed unde r each ca tegory; many o thers
exist .
1. Biochemica l Programs
Biochemical mechan isms Pho to synthesis
Amino acid s , pe ptides , pro teins \vater relations
Plant Growth Regulators 7 . Taxonomica l /Clas sifica tion Programs
2. Biophysical Programs Genera l taxonomy - c las sification
Climatolo gy (the l a r gest Si ngle discipline)
Crop-'-'ea th e r r e l at ionship Floristic Surveys
Hicrometeorology Phylogeny , evolution , adaptation
3 . Eco l ogi c a l Progr ams Experimental taxonomy
Gener al Ecology Chemo taxonomy - paleobiochemistry
Biogeogra phy - distribution Palynology
Ec osystems tJumeric al taxonomy
Cont rol (chemical and others) 8 . Fo r estry Pro grams
Community or popul ation dynamics Gene ral
4 . Genetical - Breed i ng Programs 9 . Patho l ogical Programs
Breeding , hybridiza t ion , testing Disease con t rol
Cytogenetics Host r esistance
S. Morphological Programs General Pathology
General anatomy - Horpholo gy 10. Horticultural Programs
Compara t ive Cultur al
Cell tis sue control Testing and evalua tion
Emb r yology - deve lopment Propagation - techniques
6 . Physiologi cal Progr ams Winter Hardiness
Development - grm.;rth 1 1. Field Crop Programs
Gene ral plant physiology Ce reals
Frost Hardiness Forages

The fo llowing is a li s t o f those programs which are most like l y to be

emph asized during t he next deca de :
1. Expansion of co l lec tions 5. Ana l ysis of vegetation dynamics
2 . Co n s~ rvation by comput ~r techniques
a . Preservation of na tural areas 6 . Species biolo gy studies of native
b . Conservation of biotypes or plants
gen e pools 7. Eco l ogy and ecosystems
3. Taxon omic st udi es of cultivated 8 . Native plant r esea r ch
plants (economical l y useful)
4 . Studies on deteriora t ion of t he 9 . Breeding and evalua t ion of
qua li ty of the envir onment ornamental shr ubs
a . Air 10 . Breeding and evaluation of shade
b . Hater trees
c . So il 11. Plant introduction
d . Relation of man t o hi s 12 . Propagation t echniques
environmen t
786 32

13 . Plant physiology IS . Patho l ogy of ornamental plants

a. Physiology of adventitious 16 . Plant geography
root for mation 17 . Ethnobotan y
b . Physiology of f ire-reta rd a nt 18 . Phenology
plants 19 . Cytogenetics
14. Virology - r es i sta nce t o virus
i nf ec ti ons

As applied to urban vege t at i on , the re su lts of s t udies wi th in many of the dis-

ciplines l isted above \~ill significantly benefit fut ure efforts in e stablish-
ing pol l utant r esis t an t plant material on o ur public l andscapes and highways .

E. Herbaria and Publications . Botan ic gardens throughout the world which

are activ e in re se arch usually maint ain herbarium collections or are closely
assoc i ated wi th an active he rbariuJ:1 . The major botani c garde n s distr i bute the
r e sults of plant i nvestigations at their own fa cili ties and elsewhere through
scien ti fi c publication . Representative r esearch bo tanic ga rd e n s which main-
tain h erbaria are listed be l ow Hith publications Hhere applicable .

Garden a n d Important Col l ection s Publication

Arnold Arboretum, Harva rd University ,

Nass . ; \~oody ornamentals Jou r nal o f the Arnold Arboretum
Botanic Gardens of Indonesia, Bogar,
Indones ia; Halaysian flora Annales Bogoriensis
Botanic Garde n s , Peradeniya , Ceylon; Ceylon Journal of Sc i ence ,
Flora of Ceylon Section A - Botany
Indian Botanic Ga rd e n , Calcutta ,
Ind i a; Flora of I ndia and
neighboring co un tri es Anna les of th e Indian Bo tanic Garden
Ja rdim Botanico de Rio de Janeiro,
Brazil; Bra zilian f lora Rod ri guesia
Hain Bo t anical Garden , Hoscm.,r , USSR ; Proce edings of the 1-lain Bo tanical
Native f lora of USSR Garden
Hiss ouri Botanical Garden , St . Louis;
Tempera te and tropical Amer ican Annals of the Hissollri Bo tanical
flora Garden
Nontreal Botanical Garden , Canada; rlemoires du Jardin Botanique de
Economic, Native , a nd Alpine plants Jo.lontreal
Nat ional Botanic Gardens of South
Afri ca, Capetm>JO ; Flora of South
Africa Journal of So ut h African Bo tany
Rancho Santa Ana Botanic Garden,
Cl aremont , Ca li f . ; Na tive Aliso (Journal of the Rancho Santa
flora of California Ana Botanic Gard en)
Royal Botanic Gardens , Kew , England;
He rbaceous, alpine, in sec tivorous ,
economic and s ucculent plant s Ke,v Bulletin
Singapore Botanic Garden, Singapore;
Nalaysian flora Gard en s Bulletin , Singapo re

I n addition to scholarly publications of restricted circu l ation. many botanic

ga r den s publ ish pop ular j ournals o r bulletin s such as Th e Ho r ton Arboretum
Quarterly (Hor ton Arboret um) , Garden Journa l (Th e Ne\-l York Botanica l Garden),
and Ar noldia (Arnold Ar bo r etum) which bridge the gap between the pub li c and
32 787

professional botanists and plantsmen . Complete lists of botan ic gard ens and
he rbaria and associate d research and publications are incl uded in Index
Herbariorum and the International Direc tory of Botanical Ga rdens .

IV . TilE FUTURE OF BOTIu'HC GARDE.NS

Addressing the Geneva Symposi um in 1968 , Richard A. Hm.,tard , Director ,

Arnold Arboretum, discussed t he botanic garden as an unexploited source of
information . He singularly honored th e achievements of the Jardin de Botanique
at Geneva; the work o f the three gen erations of de Candolles (Augustin-Pyramus,
Alphonse, and Casimir), al l i ntimately associa ted with the botanic garden as
perhaps unexcelled in the history of botanic achievement s . The fou nder and
first director, Augus tin-Pyramus de Can delle, \. .as much concerned that the pub-
lic should be participan t i n the developmen t of the new ga rde n. lie energe ti-
ca l ly trave l e d a nd correspon ded \.,tith botanists and traveler s , c onstantly int r o-
ducing nel'" plants to his garden for his research a nd for the e njoymen t of the
people, giving publici ty t o new and rare plants thriving at the garden . While
he apologi zed for the fact t hat the directo r was r equ ired to pay more a tt ention
to the general foundation and es tabli shment of the organization t h an to the
component plants , his mffi studies added c onsiderable in sight i nto the morphology
of plant s and t he homology o f thei r parts . His son Alphon se concentrated on
economic plan ts and I"as particu l arly interes t ed in ob serva tions on the ir ana-
tomical structure . The grand son Casimir con centrated on monograph ic works
st res sing vascul ar bund le patterns in t he stems and l eaves . All of t hese
studies util ized , in many i nstances, t he ver y same living and preser ve d
materi al maintained at the Geneva garden . The botani ca l heritage sho\oll1 i n
this single institu t ion . Th e Bo ta nic Garde n a t Geneva, is exemplar y of the
bo tanic garden of the future.

Botanica l research has been greatly aided by the estab lishment of the
Plant Re cords Center (see Appendix for historical account and exp lanation).
For example , plants from a ll locations of the world whi ch have been cultivated
in various botanic gardens and arbor eta throughout the United Sta tes fo r many
years have been negl ec ted in seve r a l monograph i c treatmen t s o r anatomica l -
morphol ogical stud ies b ecause, through no fa ult of the researcher , the loca-
tions o f particular plan ts or gr oups of plants are not generally known . The
Plan t Re co rd s Center can f unction in l ocatin g all the materia l ava ilable i n
t h is co un try for futu r e s tu dies. Oth er f unc tions of botanic gardens for f utu re
resear ch include the mainte nance in pu re fopm of primitive s trains of knolffi
provenanc e (gene pools) to be uti lized in the elucidat ion of the o rigins of
cultivate d plants . Special gene-banks and plantation s of i mportant agricul-
tural and horticu lt ura l stra ins can be maintained . The garde n operation can
continue to serve the distribution of s u itable strains of economic plant s . and
perpetuate and distribut e rare and novelty type plants. Conservation shou l d
be a maj or responsibility of botan ic gardens , whether it be the preservation
of single speci es or a lar ge scal e opera tion of ad!llin iste rin g the protection
of an extensive nat ural area containing an entire ecosystem a s a "biological
garden. "

I n addition to the purely sc i entific and conserva t ion effor ts supported

by botan ic gardens and arboreta, t here is ano ther yet not ful ly r e a lized
r espon sibility, as exp ressed by George S . Ave ry, Jr., for mer director of
the Brooklyn Botani c Garden : public e ducation . Cour ses designed to teac h
a wide ran ge of plant related subjects to t h e general public may aid in
broadening the base of American cu lture , p articu la r l y as the pub lic is
788 32

increasingly faced with more l eisure t ime. Through public co u rses , botanic
gardens and arboreta, in cooperation {.,rith the professional botanists and hor-
ticulturists associated I"ith them , are in a v i tal posi t ion to educate and
influence t he thinking of the general pop ul ation in a r eas of conservation and
environmen t a l concepts . Contemporary environmen tal necessity forces a major
duty upon bota nic gardens. Utilizing the ir staff and living plant coll ec -
tions , they must serve as interpretive centers for the mass of publ ic citi-
zens , demonstratin g the significance of our nat ur a l heri t age and of ecological
principles I"hieh have been unheeded far too l ong . Two recent publicat ions
pr esent good statements of the changing r o l e of botan i c garden s and arboreta
in our modern wor ld : The Ur ban Arboret um i n a Time of Cr isis--A summary of
Univ ersi t y of I-Iashington Arbo r et um Sympos i um in 1972 published by the Arbor etum
Foundation , Seattle . I-Iashington. and The Prospective Role of an Arhoretum pr e-
pared for Holden Arboretum . Ohio , by the Institute of the Study of Science in
Human Affairs . Columbia University .

The basic programs of botanic gar dens will differ acco rd ing to the sources
of f inancial support, and personal and professiona l empha sis of the personnel
instrume ntal i n the ope r ation . The re s ult of all t he var iables wi ll be t hat .
although all botanic ga rd ens and a r boreta may not complete l y fulfi l l all of
th eir responsibilities or function s . each should make s igni ficant contributions
through speci a li zation in one or more of i ts areas of poten tial.

APPENDIX

PLANT RECORDS CENTER (HacDonald. 19 71)

In 1964, coincid ent \"i th the estab li shment of the Unive r sity of Tennessee
Arboretum . a need \~as reali zed for the efficient retrieval of plant records
information . In 1966 a formal r esea rch project , El ect roni c Data Proc essin g of
Plant Records . was approved by the Unive r s ity of Tenness ee Ag r icultural Experi-
ment Stations . TIl e r es ults of t his research wer e r epor t ed i n the Apr i l. 1966.
iss ue of the Association of American Botan i cal Gar dens and Arbor eta (AABGA)
Newsletter by Robert D. NacDonald, \"ho had been instrumen tal i n t h e research
project and t h e devel opment of the Univers it y of Tennessee Arb oretum . At the
request of Dr. Harold R. Fletcher, President of the I nternational Ass oc iation
of Botanical Garden s (IARG) the r esearch Has reported by MacDonald be for e the
meeting of the I ABG during the XVII I nternational Horticultural Congress he l d
in Naryland in late summer of 1966 . Enth us i asm for t he potent ial of the s tudy
spread and by October . 1967 . the American Hor ticul tural Soci e t y. which had
been consid ering a data p r ocessing center of its own . and the AABGA unite d
their efforts and through the generosity of the Long\.,IQod Foundation a two-year
pilot program \~as initiated .

Success followed and the American Horticultural Soc iety Plant Records
Center is now permanen tly lo cated with the American Horticul t u ral Society a t
Ht . Vernon . Vi rginia, 22121. Plant records of the fo l lowin g ins t itu t i ons
were in data processing form by the end of 19 71 :

Denver Botanic Garden . Denver. Co lorado

Fairchild Tropical Garden , Coral Gab l es , Florida
Longwood Gardens. Kennett Square . Pennsylvania
Plant Research Institu te, Ot taN'a. On tar io
Sco tt Horticultural Fo undation, S\"ar thmore . Penn sy l vania
U. S . National Arboretum , Hashington . D. C .
32 789

Records of other institutions already completed or in various stages of prepa-

ration include:

Los Angeles State and County Arboretum , Californ i a

Rancho Santa Ana Botanic Garden , Claremont , California
Santa Barbara Botanic Garden, Santa Barbara, California
Univers i ty of Californ ia Botan ic Garden, Los Angeles
University of California Botanic Garden, Berkeley
University of Minnesota Landscape Arboretum, Chaska
Plant ing Fields Arboretum , Oyster Bay , New York
Royal Botanic Gardens . Hamilton, Ontario
South Coast Botanic Garden, Palos Verdes , California
Strybing Arboretum , San Francisco , Cali forn ia
University of California Arboretum, Davis
University of Washington Arboretum , Seattle
Mt. Cuba Botanical Park , Hilmington, Dela\"are
John J . Tyle r Arboretum , Lima , Pennsylvania

The objective of the PRC and its services centers on three areas .
1 . The PRe is developing a central data bank to contain information
about the living plant collections in arboreta , botanic gardens ,
and simi lar institutions . As such, PRe serves as a botanic garden
records-management consulting organization , aiding in the develop-
ment of ne\" record systems and improving existing systems .
2. The PRe is developing a system to provide a means to manipulate,
ana lyze , and evaluate statistical data about these collections .
3 . The PRC will disseminate information to the scientific, professional,
and amateur conununities which draw on the botanical d i sciplines
for information and materials . Special compilations \"ill be avail-
able such as checklists and indices to important horticultural
plant groups and a directory to plant sources . Presently , the PRe
has 21 computer programs which generate reports, inventories, or
data listings . As a research facility for scientific study and
a practical aid for l ocating plant source material for agencies
such as nurseries and hort i cultural research groups, the PRC will
become more and more irreplaceable as its storehouse of information
increases .

BOTANIC GARDEN AND ARBORETA LITERATURE

Avery , G. S., Jr . 1957. Botanic gardens--what role today? American Journal

of Botany 44(3): 268-271.
Blunt, W. 1971 . The Compleat Naturalist--A Life of Linnaeus . Viking Press .
New York .
Brown , R. A., and R. D. NacDonald . 1971. The A. H. S . plant reco rds center.
Arboretum and Botanical Garden Bulletin 5(4): 108-117.
Bunce , F. H. 1971. Arboreta , Botanical Gardens , Special Gardens . Bulletin
No . 90. National Recreation and Park Association . 1I1ashington, D. C.
Chan, A. (ed.) . 1968. Arboretum and Botanical Garden Bulletin 2(4) : 86-97 .
(A special issue on research programs) .
Fletcher , H. R., D. N. Henderson , and H. T. Prentice. 1969 . Internationa l
Directory of Botanica l Gardens . International Bureau fo r Plant Taxonomy
and Nomenclature. Utrecht , Netherlands .
Fogg, J . M., Jr . 1970. The nature and functions of an arboretum . Arboretum
and Botanical Garden Bulletin 4(2): 47-54 .
790
32

Hill, A. w. 1915 . The history and functions of botanic gardens. Annals of

the Hiss ou ri Botanic Garden 2: 185-223 .
Ho\"ard , R. A. 1969 . The botanical garden--an unexploited source of informa-
tion . Boissiera 14: 1 09-117 .
LanjOUlv , J ., and F . A. Stafleu. 1959. Index Herbariorum . Utrecht, Nether-
lands.
Longenecker, G. W. 1964 . Arboretum master plan . American Association of
Boranical Gardens and Arboretums Quarterly Ne\.,rsletter 61 : 6-9.
MacDonald, R. D. 1971. A history of the Plan t Records Center . Arboretum
and Botanical Garden Bul letin 5(4) : 107-108 .
Pyle , R. 1937. Report on the Committee on Botanical Gardens and Arboreta .
American Association of Nurserymen. Chicago .
Richmond, G. B. 1971. General comments on arboretum organization , f inancing,
and administration. Arboretum and Botan i cal Garden Bulletin 5(3): 75-80 .
Stafleu, F. A. 1969 . Botanical gardens before 1818 . Boissiera 14 : 31-46.
Stearn, W. T . 1971. Sources of in formation about botanic gardens and her-
baria . Biological Journal of the Linnean Society 3(3): 225-233 .
Taylor, G. 1969. Opening address . Symposium International de Geneve .
Boissiera 14: 23-29 .
Teuscher , H. 1933. The botanical garden of the future . Jou rnal of the New
York Botanic Garden 34 : 49-62 .
Thompson , P . A. 1972. The role of the botanic garden . Taxon 21(1): 115 -119.
Tsitsin, N. V. 1969. The system o f botanical gardens of the US SR and its
role in investigating vegetable resources of the country. Boiss iera 14:
135-140 .
\-lyman, D. 1947 . The Arboretums and Botanical Gardens of North America .
Chronica Botanica , Vol. 10 , No . 5/6 .
1959 . The Arboretums and Botanical Gardens of North America .
Arnold Arboretum of Harvard University. Jamaica Plain, Nassachusetts .
1970 . Hm., to establish an arboretum or botanic garden . Arboretum
and Botanical Garden Bul let in 4(52): 52-60 .
 Chapter 33 . I NFORHATION SYSTEMS AND DATA BANKING>'

Biology can a n swer many complex questions about nat ure . bu t the simple
questions o ft en confound the scientific mach ine . HO\~ many spec i es? I,There do
they grow, and what a r e their habitat preferences? l.,Then do they f l ower and
fruit? lfuich ones are rare and endangered? Such questions usua lly are unan-
swerabl e , not because the basic data never have been co llecte d, but because
the data are locked up in libraries and museums and cannot be compiled and
evaluated in a timely , economical manner . The means o f sto r ing and retrievin g
new finding s have not ke pt pace with discovery . Furthermore , man by nature
has always seemed to de rive more pleasure and challenge f r om the p rocess of
discovery itself than f r om the process of syn thesizin g the results of previous
discove r ies .

Today's biologis t, faced with t h e stagger ing overburden of historical

documentation in the wo rld's libraries and museums and with the annual ava-
lanche of new publications, specimens , and other documents, is t ending mor e
and more to ignore the existing information sto r e and instead to go back to
nature for direct answers to his questions . The risks and penalties of redis-
covery and duplication are proving more tolerable than the crush ing burdens of
dealing with the infonnation r e trieval prob lem . Yet mankind cannot afford to
keep duplicating scho l arship and documentation gene rat ion afte r generation.
O\lr libraries and museums a l ready a r e filled to over flo,.r!ng , and the world
that we seek to discover is be i ng consumed or destr oyed in the ver y pr ocess .

N O\~he r e in scie nce is the problem of redundant scholarship and documen-

tation more keenly r ea lized or the need fo r be tter means of retrieving exist-
ing informati on more e vident than in systemat ic b iology , in this c ase plant
systematics . Th e plant systemati s t must dea l Hith an enormous library of
Floras , mono graph s , bibliographies, indexes , journal articles , and other
documents \"hich often overlap to th e point of repeating large a mounts of the
same in fo rmation wo rd for '~ord . This is especi ally true of descriptive works
like Floras . lie must a l so contend with t h e million s of specimen s i n the
wor l d's herbaria. many of '''hich carry basica lly redundant information.

The computer offers a breakthrough in information tech no l ogy to rival

the invent ion of the p rinting press in the 1 5 t h cen tury . Fr om commerc ial
bank ing to law enforcemen t, the "machine" a lready has pervaded our daily
lives and begun to make an impact on scientif ic resear ch and corrununication.
Yet the computer r e volution has only begun .

Computer processing , from the appl ications point of view, is of two

t ypes : (1) numeric processing (computing) and (2) Hard (information)
processing . Broadly s peaking, the term ( elec tronic) data processing (EDP)
covers both types . ( The term automatic data processing (ADP) may r efe r to
mechanical and/or electronic da t a processing . but often i s used synonymous l y
with the term EDP .) The computer, as t he name i mp lies , achieved its widest
usage initially as a computing machine. As machine memory ca pa ci t y expanded ,
word-proce ssing capacity expanded, and the sc ience and technology of infor-
mation storag e and retrieval de v eloped. Today's big machines a r e us e d more
and mor e for information proces sing .

"'Contributed by Stam~yn G. Shetler , Department of Botany , Smith sonian

Institution.
792 33

In the l ate 19505 a feN t axonomists began to us e the comp ut er fo r numeric

processing at least in a limited way , and now it i s used widely in systematic
biology as a too l f or statis tic al and classifica tory studies . Its usefu l ness
as a n umeric processor was at on ce obvious t o biostatistically oriented tax-
onomist s . The new subdiscipline of numerical taxonomy quickly developed as a
l ogical outgrowth of the biostatistical application . Forma lly heralded in
1963 by t he appearance of the Sakal and Sneath book, Principles of Numerical
Taxonomy. this new approach offered the phil osophical basis and practical
methodology for using t h e computer to co r relate numer ical data on a la rge
scale to analyze group structure and relationships . The go al was a mo re ob -
jective means of classifying or ganisms based on actual numerical va lues r a t her
than s ub jective terminology . Numerous professional pape r s and severa l books
have been published. Today n umerical taxonomic methods, whi ch init i al l y en-
counte r e d fierce opposition at both the philosophical a nd methodological
levels , are gener ally accepte d a nd are being adopted as part of the new ortho-
doxy of systematics ; the method s now are be i ng used routinely in many s t andard
sys tematic i n vest i gations .

Information processing has developed more s lowly than n umeric proc es sing
in taxonomy desp ite the heads tart given the form er by the pioneering taxonomic
applic ati ons of Sydney W. Gould in the l ate 1940s and early 1950s when the
comp uter was brand new. The situation i n t axonomy , however , mere l y reflected
the situation in compute r pr ocessing generally. Use r s of the computer had to
reso lve major problems in designing compact fi l e structures and efficient
processing strategies . Computer technology , es pecially with re spect to memory
capaci t y . probably was less limiting than the sl ow-to-deve lop expertise of the
users. To overcome t he l imitations , the pioneers i n word pr ocessing resorted
to extensive numerical coding of information. Perhaps for thi s reason more
than any other Goul d ' s l audab l e effort s were premature a nd unconvincing to the
world\olide taxonomic fraternity as a whole . Taxonomists, no less than others,
exhib ited the typical human aversion to the subs titution of numb e r s fo r words
and generally refused to acc ep t the computer as a tool as long as this
appeared necessary .

During the mi d-1960s . David J . Rogers and his associates made the f irst
attempt to develop a generalized computer program for handl i ng taxonomic dat a .
His program package , TAXIR (TA..'{onomic I nformation ~etrieval ). is being used
for an ever-inc r easing number of applicat ions , some complete l y outside of
bio l ogy.

The most ambitious eff ort thus far to process taxonomic information by
computer has been undertaken by the Flora North America (FNA) Program, organ-
ized in la te 1966 by t he Ame ri can Socie t y of Plant Taxonomis t s under the aus-
pice s of the American I nstit u te of Biological Sciences and headquar t ered at
the Smi thsonian I nstit ut ion . Just as n umerical taxonomy was t he l ogical ,
inevitable application to systematics of the nume r ic processing capabi lity of
the computer , the current da ta-banking developments epitomized by FNA a re the
logical . inevitab le application of the i nformation processing capabi lity.

The purpose of t his chapte r is to consider t he princip les and elements

of information systems and data banking , especially compute r-based systems ,
as they apply to plan t sys tematic s . The r eader who i s in terested in f urther
information shoul d consult the general re ferences cited at the end of the
chapter and particu l arly their bibliographies which l ist numerous papers and
books .
33 793

I . DEFINITIONS

The ter m info rmation sys tem (i.e . • information storage and retrieval
system) has var ious meanings, and it s usage must be defined . Although it
has ga i ned spe cial meaning in t he context of EDP and nowadays tends to be
used e x c l usively in this conte xt , th e term does not mean implicitly that a
computer is invo l ved . In fact. it may be applied quite aptly to conventiona l
schemes fo r sto ring and r et ri eving information .

An informa t ion s ystem may be defined in general terms a s a total st rat-

~ and methodology fo r gathering, sto ring, retrieving, and exchanging (inter-
chang i ng , tra nsferring) in forma t ion to serve a spec ified pu r pose o r set of
purposes . I ncl uded i s the concrete organiza t ion of people , proced ures ,
facil i ties , e quipment , media , and data requi r ed to pe r form t he functions of
the system routinely . At the core of the system i s the data bank , the col-
lection or library of fi l es of data crea ted a nd maintained to support the
information f unctions of the sys tem . Usuall y the data bank is such an inte-
gral part of the system , fundamentally s haping a nd gove rning the system, that
it is impractical if not meaningless f rom an operational point of view to
speak o f the system or the data bank apa rt f r om one another. In fact , the
te rm data bank a s used in common parla nce today not only denotes the data but
also presumes a computerized system to process the data ; in many cor.texts,
the refore , the term has corne to mean i nfor mation syst em , as here defined .
Conceptually , nevertheless . the dat a bank should not be conf used with the
other element s of t he system.

An in fo rmation sy stem may be a s simple as a desk-top card fil e or off ice

filing cabinet , or as complex as a national library system or weather da ta
networ k. I t may be s peciali zed to serve a single user or category of users .
or generalized to ser ve many purposes and users. The mo re gene r al ized the
sys tem , the greater is the fleXibility to e xploi t it for new uses or users as
they appear, but also the l ess resp onsive it can be to the specific needs of
th e i ndividual user. Customizing a system at t he outset be fore the uses are
fully defined and the design is fixed th rough actua l experience wi t h the sys-
tem may prove unnecessarily cos tly and i ne fficient in the long run . Hhile a
system is being evolved , the bui l der us ually cannot afford to keep tailoring
it to the needs of t he moment and should be satisfied to adopt reasonably gen-
era lized procedures un ti l the sys t em's spec i fic purpose and function stabili ze.
Even in the long run he may be we ll advised to stay wi t h a general purpose
system that wi ll keep his options open and op timize t he efficiency and cos ts
of s torage and re t r ieval over several to many applications . As a general rule .
an informa t ion system s hould be de signed to answer who l e classes of questions,
not particular questions . On t he other hand, a gene r a l p urpose system can
become so generalized that it serves the purp ose o f no one in par ticu lar , and
there are many cir cums tanc es i n which the use of a system ta ilored to ans\~er
spec ifi c questions const itu tes the only practical al terna t ive . The developer
of a system a l ways is c on fr onted by a spec t rum of opportun i t ies and limitat i ons,
and cha r ting a course to f o llow is a s ub tle and comp l ex process . I n actual
p r ac tice, he strikes the best compromi se po ssible between the idea li zed goals
and the real resources and circumstances .

The process of opera ting the information sys te m to c rea t e or enhance t he

data ban k , exe rc ising all the input func tions of the system i nclud i ng r evi e\~
and editing, ma y be defined as data banking . Those who s tor e and/or retrieve
data may be defined co llectively a s the user community . Ei t her storing or
794 33

retrieving exercises some of the funct i ons o f the system and therefore cons ti-
tutes "using" the system and its data bank . [f it be necessary o r desirable
to distinguish be t \.,Ieen persons storing and persons retrieving, those who store
data , i.e ., add to tbe data bank , may be cal led contrib u tors or, co llectivel y,
t he contrib utor communi t y , and t he term user communi tv can be reserved for the
users in t he str i c t sense , i . e . , those \"ho retrieve and use th e data .

The terms "da ta" and " inf ormation" have similar connotations in many con-
texts and often are used interchangeably, but t hey are not synonymous. Let us
define the terms for our p ur poses as they frequently are used in computing cir-
cles . Data are th e indiv idual measurements , obs erva tions, et c ., e xpressibll!
ei the r as words or number s , that const itut e the factual element s of knowledge .
Informat ion i s o r ganized and e valuated data--data in context that convey a
message . In fo rma t i on i s a funct i on of context , there fore , a nd the same da ta
organi zed di ffere nt ways may convey different information . Thus , by this dis-
tinction , d ata are comp iled , Hhe r eas information must be au t hored , although
a uthorship may be a s limited as o r ganizing a set of q uestions or question-
def in i n g par ame ters to que ry the comp uter aml then ed it ing the ou tput to con-
s titute evaillated, coheren t anSl~ers to the qu e stions . From the comp u ter proc-
e ssing po in t of v iew , preci se usage of the t e nns data and information depends
on whe ther one i s foc using on the data as el~mentary fa cts or a s o r ganized and
e valuated b locks of fac ts, bu t often making 1;1 distinction is unnecessary and
und uly pedantic, either 1·/ord su f fic i ng to con v ey the practical sens e.

II . CONVENTIONAL ~ 1 STEHS

~Iode rn " computerized" (computer-based) sy stems large l y are logical ext en-
sion s of conventional systems man has develor ed and perfected t h rough long use..

The human mind with it s phenomen a l memory and powers of co rr e lat ion and
evaluation i s n atu r e ' s CIoffi, most fa ntasti c computer and infor mation system .
Since man fi rst l earn e d t o I~rite, hOh'e.ver , he has tu rned his ingenuity to
devising schemes for ex tending h is 01V!'. me nta l capacity to record and recall
information of use t o him . Hodem man is sur rounded i n every day life , pri-
vate l y a nd pub licly , by countless sys tems for storing and r e tr ievin g in fo rma-
tion . Bi rth records , health recorcl s , job records , proper ty re cords , school
records, milita ry re co rds, bankin g r ecords, shopping records, cred it reco rds ,
police reco r ds, l i bra r y records--th ese are only a few of the most important
fi l e s of vital statistic s for which he has dev i sed information sys tems .

Through t he lon g cour se of human history, civil i zed man ha s evolved and
insti tu tional ized t\~O grea t , complementary systems for storin g and r e trieving
hi s a ccumulated Idsdom--hi s re cord of hiseory and sc i e nt ific and c ultural
achievement. These two colossal systems or instit utions, \~hich e pitomize
c iviliz a tion a n d subsume all o t her lesser systems, are the l ibrary and the
museum . He has evolved the fi r st to cope with his written and, more recen tly ,
spoken record a nd the second to cope I"ith his materia l record , i ncludin g his
ar t ifacts and his voucher spec imen s of nature . Th e picto r i a l record--illu s -
tratio ns, p h otog raphs , pa i ntings --i S divided between them . Each system docu-
ments and supports the other, and each is characteri zed by a vast data bank.

To the library, in this b r oad sense, belongs a ll of man ' s I~ r i tte n archives ,
published and unpublis hed , incl udi n g the innume.rable fi l es of vital statistics
mentione d above, as I~ e ll a s his tap e a nd disk r ecord ings a nd many of his illus-
tration s , photo g raphs , and paintin gs . The library. as the rep osito r y fo r the
3 33 795

i- world's literature and the system for. exchanging it, symbolizes communication
through publication as it develo ped in the 500 years si n ce the invention of
ce
,
'e
. movable typ e . It compri ses many sys tems within the total system . Every book
or journal in itself can be viel..red as a kind of information system, containing
selected , organized, and evaluat ed data displayed and indexed in a particular
format for fast, convenient retrieva l . Customarily , the printed page is
regarded in static terms as merely a source of information , but any volume
also constitutes a means of communicating or trans ferring information and as
s such satisfies the criteria of a dynamic sys tem.
r-
Like the library, the museum as an insti tution is a hierarchy of systems .
The functions are stor ing, curating, displaying , loaning , and exchanging col-
lections of ob j ects and specimen s . Apart from mere cu riosities , these coll ec-
tions not only vouch for t he published reco rd but also provide a vast reser-
voi r of untapped data for future generations to study . Th ere are many classes
of museums, e . g ., art museums, 11is torical museums , technological museums, and
there are museums Hithin museums , s uch as herbaria (sing . , herbari um) within
na tural history museums . The system also includes countless informal and pri-
vate collections beyond the Halls o f the grea t formal institutions \·}hich con-
stitute the visible museum estabJ fshment .

The library and the museum a ' e as generalized as any systems man has been
able to conceive . They exist fOl ' t he benefit of society as a whole , and no
individual co uld ever lleed or be cap able of taking advantage of either system
in its entirety. The particular needs of individuals a r e met through speciali-
zatioll withi.n the systems . In thf' c ourse of history, numerous special- purpose
systems have evolved within the l ,:trge r sys tems to accommodate the specifi c
needs of the individual discipl in/f; and branches of human though t.

Systematic botany, among the scientific disciplines , has deve l oped its
own share of speCial -purpose information systems through the years . New ones
appear constantly as ne\·} needs a.re perceive d , but few persist for long as
measured by the cen turies . The special prominence of historical scholarship
in taxonomy gives a special prominence to the taxonomic libraries and museums
(herbaria) where the essential obj ec ts of this scholarship--the literature and
the specimens--are preserved, and to the systems devised for retrieving infor-
mation from these repositories . A few of the more durable systems or categor-
ies of systems deserve mention here.

Of the many types of taxonomic publica tions that botanists have always
produced , certain ones stand out in their sign ificanc e as information retrie-
val devices : (1) Floras and taxonomic monographs , (2) literature indexes ,
and (3) name indexes and nomenclators.

The Flora brings together selected information about the plants of a

given geographical region , while the monograph brings together the essence
of what is knm..rtl on a \wrldwide basi.s about the systematics of a given taxo-
nomic group of plants. In hoth cases, the information is organized and for-
matted to anSHer a time-tested set of predef ined questi.ons, and effective use
is made of a limited vocabuL:lrY, as s tandardized as is ever achieved in \~ritten
conununication . The binomial names are used as the ultimate address e s for the
information, and these are grouped hierarchic ally according to a systematic
classification. Diagnostic trai.ts are prese nted in the for m of d i chotomous
keys to facilitat e identification of unknown plant s . In short , as a class of
publications 'Floras and taxonomic monographs constitute one of the most
796 33

effective spec ialized information systems devised in science. The re has been
a perennial need fo r them .

Various systems for indexing and abstracting the taxonomic li te r at ure by

subject have been introduc ed th rough th e year s, and, although old syst e ms keep
giving way to newer ones , out of sheer necessity some f orm of l ite rature
retrieval con tinues t o survive . Notable among present sys t ems , fro m the view-
point of North AJile rican vascu l ar pl ant syst ematists, are the Bi bliogr ap hy of
Agricul t ure and Biological Abstracts . \.,rhich cover the broad areas of agri-
culture (including bo tany ) and biology , respec t1 vely. and have bee n r unning
for many years ; Excerpta Botaniea, ,.,.hieh abs tr acts the literat ure of p l a nt t ax-
onomy, dis tr ibu t ion , and s ociology on a worldwide basis ; a nd the various bib-
li ographic inde xes being published by the Interna t ional Associa tio n for Plant
Taxonomy.

One of the perennial prob l er:ls that the systema tic biologis t faces is to
ke e p track of all the new scientific names that are published. For many years
flowering pl ant taxonomists have been served by t wo key name-indexing systems
\~h i ch cite the original publications: Index Kel... en sis , which is pro duced in
book form at the Royal Botanic Garden, Kew , England , and covers t h e world ; and
the Gray Herbarium Index , which is produced in card form a t Harvard University ,
Cambridge , Nassachusetts, and covers only the New I,.,Torld . I n t he realm of
nomenclators, specia l ment i on should be made of J. C. Willis ' A Dictionary of
t he Flm... e ring Pl ants and Ferns , nm... in its eighth edi t ion as revised by H. K.
Airy Shaw. A nomcnclator seeks to establish the a cceptable name s , which t his
Dictionary does for family and generic names . The Dictionary al so incl ude s
synoptical de scri ption s and geographica l distributions for a l l a ccep ted names .
For a single volume , \.Jil l is is perhaps t he most valuable information retrie val
aid available to the vascular plant systema tist .

The herbarium , which traces its formal histo r y a s a s pecialized branch of

the museum back more than fo ur centurie s, has always been an i nd ispensab l e
information system of the plant systematist . Upwards of 200 mi llion specimens,
including at least 35 mi l lion in the United States alone , have accumulat ed in
the world ' s herbaria during this l ong per i od . Today , this immense storehouse
of vouchers of the plant I... orld i s managed by a worldwide network of mor e than
a thousand public herbaria, amon g t hem nearly two dozen i n stit utio ns wit h 2
million o r more specimens each and a do zen I~ith 3 million sp ecimens each .

The thou sands of transac tions that take pl ace every da y in the world ' s
herbarium system sus tain a nlultimi l lion-d ol lar en t erp rise of scient ific r e-
search and pub li c education . Every herbarium is in i t self a fi l ing system
designed for rapid retrieval o f informa t ion by kind of plant . Specimens are
filed by their taxonomic names under th e appropriate specie s , and t he sp e cies
are grouped by gene ra and fami l ies . The families are then a rranged alphab eti-
cally or, in most cases , a cco rding to some I...e l l knO\m syst ematic cl as sif i cation .
Sometimes there i s a secon dary classification by geographic r egion wi thin gen-
era or s peci es .

Finally, it should be noted that phylogen et i c and other taxonomic systems

of cl as sificati on are themselves a type of metaphy sical information system,
because they enable the systematis t to organize his knowle dge for effi c ient
re trieval , wh e ther in his mind, o n paper, o r in th e herbarium, by groupin g
like organ i sms and all information pertaining to them accord ing to their
intr insic r elationships , i. e ., their shared charac ter ist ic s . In a noth e r
33 797

sense, of co urse , the taxonomic classificat ion i s not the sys tem but a part of
the syste m, name l y , the scheme whereby the data bank is arranged o r structured .
Regard l ess of t he emphasis, a taxonomic sys t em of c l ass i fica tion is an i ntel-
lectually thrif ty devi ce for s tori ng and r e trieving information abou t plants.

II I. COHPUTERIZED SYSTEHS

A. Philo sophical Basis . For 500 years the pr in ted page has r uled as
t he medium of information exchange . Now t he hi gh-speed electroni c computer
introduces new me dia and new metho ds t o in form ation handling , making a who le
new mo de of operation possible . The computer can free th e written \.,rord from
t he physi cal limitations o f t he printed page . In t he da t a bank of the fut ure,
all data can be equal ly access ible , and vi rtually any comparison , co rrelation ,
or combination can b e made at will. Promis i ng new ways of using data are pos -
sible , a s, for example, in the use o f computer programs t o " con ve r se " directly
with the machine to identify plants by matchin g the cha ra cteristics of unknown
specimens wi th the stored values of known species . By taking advantage of the
machine ' s pm-ler to permute data and to affor d multid imensional query i ng , the
user can define a completely n ew functional r elat ionship to his data .

J ust as schola r l y man fi rst took to wri tin g and then t o printing, today ' s
scientist is communicating with the aid of ma chines. As for the systematic
botanist , s urely concern for the fut ure if not the present l-1e l fare of his
scien ce will i mpel h im to exploit the comp uter f u l l y as a tool for s t o r ing
and retrieving l i tera tu re and original data . Such exp l oi t ation must be done
in concert with devel opment in o the r scientific disciplines , of course, becaus e
the plant systema tist ' s retrieval problems are not unique .

Today ' s taxonomist can hardly cope with the new literature of his own
special t y as i t appears; yet h e continues to publish I-lith abandon, casting
his contributions int o the swe l ling sea of l iterature in t he hope t hat he has
truly add e d to the store of knowledge . With eve ry passing year, however , the
cha nces dimini sh that his co n trib utions wi l l ever be fo und by those who need
them. Likewise , no taxonomist can c laim r ealist i cally that he is able to take
ful l a dvantage of the mass of information l ocked up among the milli ons of
specimens in the 11O r ld ' s herba ri a; ye t eve ry year taxon omists go into the field
in ever gr eat er n umber s to co llec t more specimens for t he s t ockpile .

If they are to continue to discharge thei r responsibilities to science

and so c ie ty, therefore , plant systematists urgen tly need be t ter ways a n d
mean s of keeping perman e nt documen ta tion of what ha s al re ady been discovered
about o r gan isms in a fonn that permits ready compa rison with neH data so that
truly mo r e and more is l earned ahout t hem in a n efficien t manne r as discovery
continue s . Thi s calls for better t~ays and mean s of dealin g with the Horld ' s
200 million herbar i um spe cimens and de ciding realistically about the needs for
more specimens , an d vast l y i mproved means of retrieving and anal yz i ng t he lit-
erature. Using computer processin g systems now in exis ten ce , systema tic bot-
ani sts can hope to develop a botanical da ta bank and i n fo rmation sys t em to
cope wi t h these s tagger ing p r ob l ems . The r e also is hope of breaking o u t of
th e Linnaean taxonomic mold .

The present taxonomic system is an outg r owth of the Ll nnaean taxonomic

system, wh ich was an outgrowth of ea rli er folk taxonomi es, and it now is quite
i nadqua t e fo r deal in g wi t h the hundreds of thousands of organisms already
798 33

described , not to mention the millions still to be dis covered. Linnaeus knew
a much simpler \wrld than \.,re knO\o! today, and his taxonomic sys tem was des igned
to handle a memorizable number of gene r a . The human mind can cope easily \~ith
a thousand different kinds (taxa) of organisms at anyone time and tends to
deal in generics (oak , map l e, hickory), as seen in var ious folk taxonomies
existing in the wo r l d today and in historical times. (Some wou ld say that
trained taxonomist s can cope lliith upHards of ten thousand different kind s of
organisms at anyone time . ) When knOl"ledge abou t o rgani sms i s forced i nto a
system that canno t cope with large numbers, it tends to ge t los t or dealt with
artificially and sup erf ic ially , with the resul t that th e system of names
itself r e ce ives undue atten tion . The computer can extend t he taxonomist I s
capacity to remember the names and traits of organisms and can allow him,
there fo re, t o deal I.,.ith virtually any numbe r of t a xonomic units on an eq ual
and object i ve basis and to store the information so that new constructs built
from it will not des troy , di st ort, or obscure the information its elf . Accord-
ingly, the modern systema tist can begin to examine taxonomi c discontinuities
at any level he chooses and bring them into sharper fo cus using the othe r
modern methods of investigation . This s hould advance ou r very perception of
the living world.

The Flora or other taxonomi c treatis e of t he future will be a dynamic

data bank written on the "pages " of a computer memory vhich are available via
telecorrununication s to the entire Iwrld and from 1.,.llich can b e printed in hard
copy any number of subsets and permutations of data for individual or mass
cons umption . The very process of research and communic ation will be alt e red .
Today research culmi nates in publication. I nformation retrieval, if any,
comes after publication. Tomorrow the results of research viII go direct ly
to the data bank, and conventional publ ication, if any, Io}ill come as output
from the data bank. Today the Iwuld-be user of scientific information must
wait until the contributor has collected enough da t a to j ustify pUblication .
TomorrolO} the user Idll be able to ask the sy s tem at any time to compile the
late st information on a given topic , and the contributor will be ab le to sub-
mit his data a datum at a time if he so chooses .

B. EDP Terminology. Any computer sys tem has two major components : (1 )
hardHare , and (2) so ftl.are. The hardl.,rare consists of the computer itself and
all associated (p e ripheral) eq uipment . I ncluded in this equipment are data
conver sion devices and compute r termi nals .

A comp uter termin al i s a typewri t er or other device for tran smittin g da ta

and/or operating instruc tion s directly to the computer . A lo ca l terminal is
Hired to the computer and must be relatively near the computer , while a remote
termi nal is connected to the computer via te lephone or other data transmission
lines and can be located Hherever the necessary telecommunications a re ava i l-
able . Through th e us e of satellite transmi ss ion con tinents and oceans can be
spanned. The corrunon terminal is some type of teletypewriter. Io.'ith increasing
freq uency typeHriter t erminals are being linked to display systems using a
television s cr een (CRT: cathode r ay tube) and even Lo mini-computers .

Data conversion ( automation, reduction) is the p r ocess of converting

(reducing) data to machine-readable form by mechanically punching holes in
cards or pap e r tape or by elec tronically encoding on magnetic tape o r a direct
storage device (e.g., d isk) associated wi th the computer . Th e most widely
used da ta conversion device is the card punch (keypunch) , but various paper
tape and magne tic tape machines also are now l.,ridely used. Recently , optical
33 799

character reading (OCR) and other optical scanning devices have come into use .
Some of the newest data conversion devices are built to double as computer
terminals . They can be used off-line , i . e . , as self-contained units , to con-
vert data , or on- line, i . e ., connected to the computer, to convert or transmit
data . On- l ine data conversion systems , which are more costly to operate than
off- line systems, transmit data directly from the keyboard to a computer where
the data are encoded on the magnetic storage deivces .

The soft\"rare of a computer system is the tocal se t of encoded ins truc -

tions -- apart from those " hard-wired" into the circuitry of the machine--
required to run the computer and process data. Software which accomp l ishes
functions such as equipment (hardware) allocation, job scheduling , and program
compilation makes up the operating system, and individual programs of this
software are knOlffl as system programs. Soft,"are designed to process user data
is made up of application programs. Some application pr ograms are generalized
to make them usable on a wide variety of data and are controlled entirely or
in part by options specified at run time in lieu of submitting spec i all y coded
instructions . These programs are attractive because they eliminate or greatly
mLnLmLze program development for whole classes of applications . Frequently
used application programs may be held, as the operating system, on storage
devices that are connected to the computer ' s central processor at all times.
Application programs tell the computer what and how to process in order to
execute jobs . They are written in one of the standard programming languages ,
e . g. , COBOL , FORTRAN, ALGOL , that the particular machine can interpre t . Often
the programs are encoded on cards or tape in the same manner as data and sub-
mitted I"ith the data at run time.

The capabilities of the hardware are limited by the capabilities of the

software, because the softl"are controls the hardware and constitutes the means
whereby its potential is realized . Software even of a routine application
type rarely if ever 1s totally machine-independent and capable, there fore , of
running without any modification on different computer s of the same or of
different manufacturers; however , programs running on a particular manufac-
turer's computer usually are more or less compatible upward through his line ,
Le., from bis smaller to his larger machines .

Hodern third-generation computers, which can link to the latest tele-

communications to support teleprocessing (processing by remote control), offer
the user several processing options. In the multiprogramming mode, the com-
puter can execute more than one job and handle more than one processing option
simultaneously. The tradit i onal option is batch processing, i n which t he user
delivers his jobs on cards or tape to the computer center and then co l lects
his results after the jobs have been run in their turn . This is the most
economical way of processing, especially when large jobs are involved , and
much computing is still being done this way .

A variation of the batch-processing option (or mode) is called remote

batch processing or remote (job) entry (RJE) , which allows the user to submit
his jobs over a telephone line or other data-transmission system from a remote
terminal, removed perhaps by hundreds or even thousands of miles from the
processing center . The jobs enter an electronic queue at the center and are
processed in turn . Turn-around on these jobs may be very rapid, and process-
ing often is accomplished t>'ithout anyone being directly involved at the com-
puter center . Some computer systems permit the user to govern the speed of
turn-around in part by assigning a priority to the job f r om a relative scale
800 33

established for the system . To get faster turn- around he sets a higher prior-
ity but then pays a higher unit charge. The output can be directed back to
the originating remote terminal, to a printer or other output device in the
computer center itself, or to some other remote terminal or computing facility ,
as specified by the user . Bo th in batch and remote batch processing there is
a discrete , though perhaps extremely short , time lag between submitting the
job and receiving back the processed results . The computer is not executing
jobs ins tantaneou s l y, in so-called " real- time," unde r the con tro l of t he user,
but seq uent iall y or in o rder of priority as controlled by the computer opera-
tor or automatically by the system itself .

On-line or real-time processing is the ultimate option because i t puts

the computer under the direct con trol of the user himself , if only for a split
second, and enab les him to execute his job immediately . He can communicate
or "converse" Hith the computer continuously as his job is b eing executed to
guide the processing from step to step, adding or modi fying instructions as
necessary . This processing option often is called the conversational mode,
for obvious reasons. Output can be obtained in any of the ways available for
RJE.

Some times an entire compu t e r system is dedicated to continuous on-line

use by a single user, but most user s canno t justify or afford real -t i me access
to a comp ut er sys tem except on a shared- time basis. In the time- sharing mode ,
the comp uter shares its central processing unit (CPU) with several to many
users simul t aneou sly by al l ocating each user short bursts o r "sli ces " of
processing time in turn until his job is done . The process might be like ned
to juggling balls and keeping all in the air but one at anyone time . This
time juggling usually is so rapid and precise that each user gets virtually
instantaneous response to his commands and has the i llusion of exclusive a nd
con tinuous control of the entire computer system. Speed of respon se degrades
only I~hen the system becomes overloaded with users, whi c h may happen momen-
tarily or for extend ed per iods of time during the busiest part of the oper-
ating day or week . Time- sharing access to the computer ordinar ily is via
remote terminals.

The pr actical effect of the ne\~er option s that e nable mul tip r ogramming ,
real- time processing, and teleprocessing is to put distance , both figu r atively
and literally, between the user and the computer and to reduce his dependen ce
on understanding the computer ' s operation, thereby making i t pos sib le for him
to use the comput er simp ly as a utility like the telephone .

C. Literature vs . Data Retrieva l. Literature retrieval is t h e p r ocess

of identif ying and locating, by manual or a utomated means , books, journals ,
individual articles , or other documents \dthin the total mass of published
literature that are of interest to a particular user . The means may be analy-
tical reference \wr ks , b ibliographic indexes, card catalogs , or other retrieval
systems . As it i s used, especially in compu ting circles, the concept of lit-
erature r etrieva l does not necessarily include the act of physically retrieving
the items , although this obviously is implied by the term. I n the broadest
sense, however, a literature retrieval system deals with all functions and
processes required to t ransfer document s between author and user. Document
retrieval is perhaps a more apt term because the Iwrd "document" covers unpub-
li shed as \."ell as published material , including manuscripts , research notes ,
illustrations, films, tapes, and even biological specimens . Some would gen-
eralize even furt her by us i ng the term document tran sfer because the word
 ---- .. _--------- ---.-~- --=---

33 801

"r etrieval" tends to emphasize only one aspect of the transfer process . The
term information retrieval as used in most contexts refers to document (lit-
erature) retrieval. Although literature retrieval might be said to be the
original and ordinary meaning of information retrieval, the latter term has
come to have a general connotation since the advent of computers . Further-
more, this term , if taken literally, implies that the information itself is
being retrieved Hhen in fact the documents that contain the information or
references to them are what is being retrieved.

Computerized document retr i eval systems aTe now a commonplace, and they
are multiplying rapidly. Obviously. this has been one of the most urgent areas
in which to apply EDP. Anyone who has faced the task of searching the past
literature or of keeping up \"ith current literature understands the fundamental
inadequacies of the present retrieval methods and appreciates the rush to com-
puterization . The Bibliography of Agriculture is produced with a computer-
based system. Biological Abstracts has been computerizing gradually , \..rith the
resul t that a whole net" range of bibliographic services are being offered
under the umbrella of BioSciences Inf ormation Service. Only the most tenta-
tive steps have been taken in the specific area of plant systematics. The
Flora North American Program developed the "FNA Automated Bibliography" on a
pilot basis , but for lack of funds this system was never implemented . \.Jithin
the frame ...'ork of systematic botany , it Has designed to be a general purpose
system, serving all of the taxonomist ' s needs from retrieval of current titles
by several parameters, including author and subject, to retrieval by Latin
name of up-to- date lists of new names , ne ...' distribution records and maps, neH
chromosome counts , etc. Hany individual scientists have developed personal-
i zed systems, using local computing centers.

Data retrieval is the process of identifying, locating, and retrieving,

by manual or automated means, specific facts in stored documents and specimen
collections which are of interest to a particular user . Computerized data
systems a r e still in their infancy , but they hold far greater potential than
document systems as a tool for solving our modern information problem. They
firs t demonstrated their utility in the intelligence area, and today no self-
respecting intelligence or po l ice agency is without its data system . In
biology , data banking has just begun. Hhere document systems present the
user only with references to the information desired , data systems deliver
the actual data or elements of information . Obviously, the scientist \"ho is
harassed by too much literature to read ,..rill opt for a system that presents
him wi th the prec i se data he needs in preference to one that gives him only
references to these data . In an FNA survey of nearly 500 biologists, con-
ducted in 1971-7 2 , data retrieval lo,Ias favored over literature retrieval by
almost 2 : 1 as being the more urgently needed kind of in formation service
today .

The principles of machine retrieval are basically the same fo r data

systems and document systems because the elements of a document record are
the same as data to the computer, but in practice the tl"O types of systems
present quite different operational problems, from data preparation and input
to report generation . Consequently , different file designs and processing
strategies often are used , and information retrieval specialists tend to be
specialized in one or the other type of system but not both. The distinction
is far from black and \"hite , however, and many hybrid systems can be fO'llld "
In deed, the pilot FNA Automated Bibliography Has a hybrid system, pt"imarily
for document retrieval but also including actual data, e . g ., nelo,l chromosome
802 33

counts , extracted from the contents of the documen ts recorded .

The remaining sections of thi s chapter are concerned only \dth dat:a sys-
tems . The subject of document systems is heine left to the numero u s publica-
tions on literature retrieval both in and out of biology .

D. Comp onents of General ized System . A generalized representation of an

information system is illustrated i n Fi gure 33-1. This could be e i ther a data
system or a document system , depending on the contents of the data bank . It
is only one of the many schematic representations that could be dra ....'Tl. depend-
ing on the points of emphaSis, and it does not purport to show al l components ,
functions . and relationships of an operating system in their every detail. The
schematic is simplified deliberately, and in particular it does no t indicate
all of the potential feedback points nor the basically circular nature of
informa tion flOl~ through such a system . The flo\~ of da t a th r ough the system
is from the sources at the left to the data bank at the core and from there
to the users at the right who can pose direct questions to the data bank and
receive direct anSHers . [f contributors or other sources in t eract with the
system , they in effect become users . Li kewise, users I .... ho submit corrections
become contributors .

According to this schematic , the contributors are the source of ne~. .

research that cann ot be compiled more or less mechanically from existing lit-
erature, specimen collections, or other infor mation systems . The information
specialists a r e the sc i entists and technical experts I.... ho provide the critical
human i n terface between the machine and its processing system on t he one hand
and t he con t rib u tors and users on t he other hand . In this categor y there may
be several to many types of specialists , depending on the size and comp l exity
of the system-- e . g., systems analysts , programmers , computer operators , data
base specialists , data conversion operatot;"s , technical editor s , scient i fic
e di tors , managers . Nanagemen t information s p ecia lis ts , who are tr~ine d and
experienced in the whole process and know the characteristics and parameter s
o f the data base thoroughly , manage the system and interpret its operat i ons
and its products and se r vices to the " public ," the cont r ibutors and use r s .
Scientific editors are respon sible for upholding the scientific standards of
the data bank and must review and edit all da ta before t hey enter or leave
the hands of the technical specialists . All technical and s c ientific func -
tions may be perfo r med by one or wo individuals in the case of small systems .
A more deta i led representati on of the data collection process is shown in
Figure 33-3 , discussed in the n ext section .

The "informa t ion processing system" in Figure 33-1 symbolizes all aspec t s of
machine processin g, i ncluding da t a conversion as well as wha t happen s i n the
computer "black box . " Thus i n this broad sense the box in the chart repre-
s e nts both the hardware and the soft\oJare . In the stric t sense , however, the
information processing system is only t he softwa r e for perfo r ming t h e standard
functions of file creation, file maintenance and up datin g , retrieval , a n d
report generation . This i s the technical meaning of the term " in f ormation
system" in computing circles , but in contexts I .... her e there is l ikel y to be
con fusion with the more gener al meanings of t he term i t is best to use the
full expression " i nformation processing system" to denote the sof t ware .

To qualify as generalized, an information proceSSing system mu st be abl e

to pe r form t he standar d functions enumerated above and i n addition must meet
these two criteria :
 CONTRIBUTORS INFORHATION
~ ~-
SPECIALISTS
I,

\~-
QUESTIONS
NEH DATA , ANSI-lERS
INFORHATION
PROCESSING
SYSTEH
LITERATURE
I
CORRECTI ONS
DATA STANDARD

SPECIHENS
BANK

HI
REPORTS &
PUBLICATIONS

OTHER SYSTEl'lS

Figure 33-1 . Highly generalized schematic representation of an information syst~m .

00
o
w
 804 33

1. The system must be characterized by " data independence, " i.e., it

must be des ign ed \...-ithout regard to any particular type or file of
data .
2. The sys tem sh ould be controlled by a high-level language that is easy
to learn and use (i.e .• user -or iented) for defining processing
requirements and thus for invoking and directing the functions of
the systcr.J ,

Other highly desirable features of a generalized sof tware package for

information prucessing are the following :

3. The sys t em should stand alone as self-contained software, performing

a ll functions vithout dependence on a host l a nguage such as COBOL.
4. The system should be capable of p roces sing hierarch ical data st ruc-
ture, e . g . , multiple collections within a species or multiple
specimens ~"ithin a collection.
5. The system should be capable of multi- f ile search and ret r ieval,
a llowin g fo r retrieval of related information from two or more
files automatically in response to a single query.
6. The system s hould operate in both the batch and remote-entry modes
but not necessari l y on-line, although the potential for this is
desirable .

The "data bank" at th e hear t of the system may contai n not only the pri-
mary data files but also authority files and specialized application programs.
Examples of data files pertinent to plant systematics are an index of chromo-
some numbers, a r egister of type specimens , a direc tory of special i sts in th e
field, an herbarium catalog of speCimens , and a species-by-species file of
, morphological, ecological, and geographical descriptions .

An authority file is an authoritative dictionary of names or terms for

standardizing usage in the data bank . Example s are a checklist of species
names , a list of author names and abbreviat i ons , and a vocabulary of morpho-
logical or eco logical terms. These files, i n addition to providing guide-
l ines for con tr ibuto rs, editors , and users, also become tools of th e so ft-
ware which enable the system to edit automatically for conformity to stan-
dards during the "processing of input or output. Any deviation from the
standards in the dictionaries ,..rill be f lagged for the a ttention of the editor
or user . The authority file certainly is not a new invention , but the compu-
ter is a po,,,erful tool for applying such a file consistently . Hithout prop-
erly maintained authority files, data banks soon get ou t of hand .

A good data bank '''ill include a library of application programs to per-

form sp ecialized opera tions on retrieved data . Examples are programs for
computing sta tisti cs , performi ng numerical taxonomic analyses, mapping geo-
graphic distributions, const ruc t ing dicho tomous keys , and id entifying plants
on-line . These programs sho uld not be confused '''ith the information
processing system per se .

The aim of the retrieval strategy of any information system should be

maximum user-accessibility to the stored data , and a generalized system can
offer a wid e range of products and services. The numbe r of possib l e permuta-
tions of even a relatively fe'" data e l ements can be astronomica l. For every
data bank, it is necessary to define the standard reports and formal publi-
cations that are to be issued according to some schedule for the benefit of
33 805

the entire user community , but beyond this formal output the system can pro-
vide informal answers to an unlimit e d variety of personalized queries.

E. Systems Compatibility. Every effort should be made in developing a

system to achieve a high degree of compatibility \.,rith othe r systems having
similar objectives , but total compatibility is unattainab l e. To say that two
systems are "compatible" is to say nothing unless the nature of the compati-
bility is defined, because there are so many elements to make compatible in
any total system. Pushed to its extreme , compatibility would mean identical
systems with identical data banks. As sure ly as the purposes of systems are
different , the systems themselves must be d i fferent at some point. What is
more pertinent to stress than compatibility is translatability . As long as
the procedures followed allow for easy data translation from one system to
another , the conceptual ideal of compatibility is achieve d .

IV. STEPS IN DEVELOPING A DATA SYSTEH

A. Defining Purpose. The purpose(s) of a data retrieval system should be

defined ~"ithout regard for whether the system ~"ill be computerized . The compu-
ter tends to invest a system with an aura of significance and authority regard-
less of the system's intrinsic worth. It should a1\"ays be kept in mind , there-
fore , that computerization in itse l f can never give ~wrth to .wrthless data or
purpose to purposeless functions . On the other hand, a computer may spell the
difference between feasibility and unfeasibility and make a system worth
building, not because it bestows purpose, but because it allows the purpose to
be realized within the limits of the time and resources availab le.

The chief reason f or computerizing any system should be to increase the

effic iency and flexibility of access to the data, hence to enhance the effec-
tiveness of the data bank, not to reduce the absolute cost or increase the
absolute efficiency . Computerizing will not necessarily reduce the cost of
operating the system or increase the efficiency o f operation, as measured by
the conventional products and services or the f unctions of the conventional
operation . In fact, the out-of-pocket cost in all probability will increase,
perhaps many fold, because new types of personnel (e.g., systems analysts,
programmers, data conversion operators) are needed , and machine costs even if
modest represent a net,.] cash outlay. Overall system efficiency is likely to
decrease, even dramatically, at first , because ne,~ procedures must be insti-
tuted . Many would-be builders of computerized systems have been led dm..rn the
rosy path by p romises of greatly reduced costs and increased efficiency which
did not materialize. I n the final analysis , the cost and eff iciency of a
computerized system must be measured in terms of p rod ucts and services, by
comparing the new array with the conventional array. This is a critical point
especially because it means that cost-effectiveness cannot be determined before
the system has been p ut into operation and the ne,,, use pattern established,
which usually happens long after the case must be made for financial support
to build the system.

Finally , it is important to keep in mind that the main purpose for taking
a generalized approach is to have a system that will be capable of answering
whole classes o f questions of concern to many users . Systems designed to
answer specific questions or a single set of predef i ned questions seldom
justify the costs of computerization . The system should be measured by the
~ of questions it can anSHer. A well des igned system ,,,ill be capable of
responding to new types of questions that were not anticipated when the system
806 33

Has first built. The layman Hila \.Jatches retrieval from a data hank for the
first time often jumps to the conclusion that the system l.ras built to ans.Jer
the specific query he has posed. and not surprisingly he then may conc lude
that the result is not l.,ror t h the cos t. This is tantamount to concluding that
the library system was developed for the express purpose of ansl.Jering the spe-
cific question that one may be asking on a part i cular day and then judging the
Horth of the i nve stment thereby .

B. Identifying User Community. It rarely if ever is possible to knml the

exact limits of t he user communit y, because the primary users pass along in-
formation to secondary users. secondary users to tertiary users. etc. Fo r
this reason there is great danger in dra\... ing conclusions about the value of a
system from an analysis of the circle of primary users l.,rho , though more or
less visible and identifiable, may form only a small fraction of the real
user community . Thus, l.,rhile a floristic or other taxonomic data bank is
designed to se rve primarily systematic botanists , the find ings of the systema-
tist sooner o r later make their \.,ray , sometimes quite indirectly , into the
handbooks and reference l.,ro r ks of scientists and the publ i c at large . Antici-
pated direct or primary use, nevertheless, is the only concrete basis on l.,rhich
to build a system . If primary uses and users cannot be identified, then pre-
sumably there is no need for the system . Despite the difficulty, theref ore ,
of defin ing the primary us er community, especially in ad vanc e of having had
experience with an operat.ional system and real users, some effort must be made
to do this as a basis for designing the system and justifying its development .
In most cases, a previous l y existing system is being computerized , and its
pattern of use can serve as a preliminary guide to the pote ntial ne\.,r pattern
of us e .

As a general rule, the size of the user conununity will be directly pro-
portional to the size and scope of the data bank and to the degree of gener-
alization of the information service, in the same manner as t he user community
of a library or museum is gover ned .

C. Defining Data Bank . The natural tenden cy in deve loping a computerized

system is to want everything in the data bank immediately . The old adage,
"Halk before trying to run," ce rta inly applies here. The f irst task in def in-
ing a machine-readable data bank is to cut the universe of possibilities down
to a practical number. If , as is often the case, an exist ing manual sys tem is
to be converted to a comp ute r-based system , then it is advisable to stick as
closely as possible initially to the parameter s of the current data base .
resistin g the u rge to enlarge the number and variety of da t a elements . This
appr<J ach of using the old data bank as the model for the ne,.,r one takes maximum
advant age of past experience in defining us ef ul categories o f data. If a
totally new data bank is to be created , t hen it must be defined in care ful
detail at the outset. and subs equent cbanges must be made in an orderl y Hay
supported by t horough document a tion.

A taxonomic data bank dealing Idth flora (or fauna) can be represented
as a taxon X data-category ma trix of data. as depicted in Figure 33- 2 . Every datum
is keyed on one axis to a taxon . usually a species. and on the othe r a xis to a
type or category of data . The data category may be a general class of fea-
tures , e.g., morphological features, or a sp e cific characteristic. e . g . •
flm.,rer color. The data bank can grow in either direction indefinitely and
thus is open-ended . The monographic approac h to buildin g t he data bank is to
 •"

TAXONOHIC GROUPS w
w
Vascular Flora Other Flora I Fauna
I
I
1 2 n - _I __
I
l. Nomenclature I
- -1 - -
I
2. Kevs
- -l - -
1
3. Horphology I
1
I I ________L
4. GeoRraphy
- r
I I

5. Ecology II I I
.I
DATA I
CATEGORIES I
6. Cytology I
I

1
_ _ _ _ _ _ _ _ L
I
I
7. Reproductive
Biology
. I
I
L
I
8. Chemistry I I
I - - - - - - - - r
I

I
n. Other I
1 _ _ _ _ _ _ _
, .

F i gure 33- 2 . Matrix representari on of a taxonomi.c data bank 00

o
~
808 33

collect data i n many categories for a relatively few taxa , \"hile the floristic
(faunistic) approach is to col l ect data in feH categories bu t for many taxa.
It is important t o decide at the outset whether the initial effor t will be
directed t oward taxonomic or data comprehensiveness .

D. Collecting Data . The major functions of t he data collection process,

from the sources of the raw data to the publication of output ready for use ,
may be represen t ed schematica lly as in Figure 33-3 . Some data co llection systems
wi ll be more, others less complex than the system depi cted here . The major
sources of data should be identified concretely before any da ta banking is
begun. The collecting and editing procedures may differ significantly bet\.,teen
small one-man efforts , such as in the computerizing of a single university
herbarium, and l arge net\.,tork efforts involving many people and institutions,
s uch as in the FNA Program. In the first instance , one person is in total
control and may be able to ma in tain con sistency without establishing a closed
system of standards a t the outset , but in the second instance it is critical,
be cause of the different par ties involved , that all standards be fixed at the
outset and then be held to firmly . In either case , no signif icant data col-
lection should be started unti l a precise format, down to the punctuation ,
ha s been established. An example of the data col lection form bein g used for
the Typ e Specimen Register , a network system cente red at t he Smithson ian
Institution, is shown in Figure 33- 4 .

Two types of edi ting a r e always r equired -- con t ent ed iting and techni cal
editing. As in conventional publishing, ed iting for scientif i c content is
necessary to control the reliability and authority of the data . The fac ts
may be reliably collected but not au thoritatively synthesized as scientific
i n formation by an appropriate speciali st; contrarh.,tise, an authoritative
presentat ion maj be riddled with unreliable dat a . The botanical editor , who
is responsible for ove ral l quality control, must keep these t\m aspects of
the data distinct in his mind when setting up data collect i on and editing
procedur es. He \dll realize that the person who is best qualified to prepare
an autho ritative synthesi s of da ta is not necessarily the one best qualified
to collect reliable rm.,t data for synt hes i s . Thus. although on l y a speciali st
c an produce an authori t a tive taxonomic treatment, an eXperienced professional
technician or research assistant may do a more thorough and accurate job of
compiling routine spec i men data and nomenclatural data from the herbarium and
library. respectively . The botanical editor also will make s ure that an
appropriate sc ient ific revie\.,t by other special ists forms p a rt of the editorial
process in his dat a collection system .

The technical editor in the data collection system e nforces the estab-
lished f ormatting stan dard s of the conversion and processing syste ms. After
data have been processed initially. t he machine records must be recycled to
the techni cal editor and perhaps also to the botan i cal e ditor for v erification ,
to ensure that t he original data have been transcribed into the system faith-
fully, without typographi cal error s and in conformity with all standards of
the system . Several cycles of editing and updat ing may be required befo r e a
"clean" data base is produced .

Once the formats and procedures for collecting and editing data are
stabilized. then full documentation should be prepa red, including technical
specifications of the system and instruction manuals fo r the edi t ors and
operating personnel.
 SOURCES RESEARCH

...- .I BOTANICAL EDITING

I I

TECHNICAL EDITING
0;-

DATA CONVERSION

~I
DATA PROCESSING
f---

DATA BANKING USERS

J
PUBLICATI ON

Figure 33-3 . Schematic representation of a data collection system

810 33

Please type . Enter nel'; names only . "' Ess ential fields--information required.

1. FAMILy"' _ _ _ _ _ _ _ _ _ __ __ _ _ __ _ __ _ __ _ _ __ __

2. GENUS*

3. SPECIES*_ _ _ _ _ _ _ _ __ __ _ __ _ _ _ _ _ _ _ _ _ _ _ __

4. INFRASPECIFIC TAXON ______________----____- - - - - -- - --------------

( I ndicate rank: ssp , var, svr, f or, sfm)
5. AUTHOR(S)*'________ __ _ _ _ _ _ _ ______ __ __ _ _ _ ___

6. CITATION1' __~~~----_--~~-----~--------~--~~~~~._---­
(Cite periodicals and serials according to standards of B-P-H . )

7. COLLECtOR (S) t:._ _ __

8. COLLECTION NO . _______--~----~ 9. COLLECTION DATE,_____________

(Indicate whose series if not collector ' s series . )
LOCALITY :

10 . COUNTRY*'-c_-------_~~~~_~--_~~--_~~~--'"~--_ _.
(Use modern name and cite original as follows : Ethiopia ("Abyssinia") , )

11. STATE , PROVINCE, DEPARTNENT, OR

EQUIVALENT_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

12 . COUNTY OR EQUIVALENT_ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __

13 . TOt-IN OR LOCAL REFERENCE:_~--_ _ _~---------___-------

(Place important wo r ds first and omit unnecessary words.)
SHEETS : 14. HERB. ACRONYN(S)* 15 . SIlEET NO(S) . 16 . KIND(S) OF TYPE(S)
1st ______________
2nd ____________
3rd _ _____ ___

17 . RE~~KS>___~--~~----~~~~~~~~~~----~----~__-----
(If more than 2 shee t s , indicate to which sheet remarks 8IJPly . )

Note : For additional sheets . continue in "Remarks"; for additional collections

(e . g ., syntypes) , continue on back .

Source of information'_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ ___

Refe ren ces checked :
B-P-H Gray Card I ndex___ Index Kewensis Other? ________
Informa tion provided by : ___________________Datc'________

Fi gure 33- 4 . nata collection form of Botanical Type Sre cimen Register.
 -- - -------------------

33 811

E. Converting Data. Collected and edited, the data must be converted

into computer r eadab le form . I n selecting a data conversion medium and pro-
cedure, the botanist should seek the advice of a systems speci alist ",here the
data are to be processed.

Today there ar e many devices on the market for converting data , and there
are several basic types of data conversion systems :

1. Card System : The tried- aod - true , 80-column , keypunch card , c ommonly
knmm as the "IBN card," is still the best medium for many appli-
cations \·,here a limited number o f data fields are involved and a
physical record in the form of a card file also is desired . The
ca r d system \"orks best Hhen all data for a unit record can be
punched on a single ca r d . Th is permits easy filing on the one
hand and inexpensive sorting by machine on the other hand. Valuable
listings can be obtained l"ithout any computer programming by using
a carel sorter and then printing so-called 80/80 listings , i.e .,
printing column for column as given on the card. Before any com-
put er proc essing can be done, hOI,'ever, even if only to reformat
data or decode data on the printout , the cards must be read into
the computer . Often cards are used only as an interim storage
medium until the data can be copied onto magnetic tape or other
magnetic storage .
The chief limitations are the f i x ed number of 80 co l umns per
card and the bulkiness of the card decks when they become large .
Neve rth e less, the punched card should be used more o f ten by biolo-
gists. Like other scientists, biologists are accustomed to main-
taining various types of card fi les, and such files can be made
much more useful by punching them on machine-readable cards . It is
easy to learn to punch and sort cards and to design simple computer
programs for permuting the information more elaborately . Once the
data are punched on cards the door is open for any type of infor-
mation processing . Horcove:!:" , there are few data process i ng cen-
ters in the Iwr ld that cannot read punch cards .
2. Paper Tape System : For lengthy unit records that would require several
to many cards per record, 3S in most data-banking applications ,
paper tape is a relatively cheap and effective medium . Paper tape
is analogous to cards attached in an endless string and thus without
column limitation, and in fact this is the l"ay the computer "per-
ceives" a deck of cards . A specially designed type\"riter with
tape-reading and tape-punching attachments is used to encode the
data on a strip of {"hatever length is required to incl ude the
record . Hany individual records can be punched on a s i ng l e reel
o f tape, as long as they are separated fr om each other by machine-
readable codes, or individ ual records can be kept on individual
pieces of tape folded or rolled for filing . In the latter case ,
usually the pieces are spliced together into large spools or the
r e cords are copied by machine onto continuous r eels be f ore process-
ing , because generally it is more efficient to feed one long tape
to the computer than to feed many short tapes .
As with cards, the mechanical coding on paper tape is visible
and can be read by an experienced user, but unlike with cards the
codes cannot be "interpreted," i.e ., typed in black and white on
the top of the tape itself . Some paper tape devices can be pro-
grammed to punch automatically invariable data , such as the field
812 33

names of a data collection form or the fixed informat i on of a form

letter . The operato r needs only to enter the variable da t a from
the keyboard.
Paper tape systems are useful not onl y for inputting data to
the computer but also as office machines for automatically repli-
cating specimen labels , manuscripts, letters, or standard forms .
Each copy is in effect an original . Once a manuscript is on tape ,
for example , revisions can be incorporated by producing a corrected
tape and then playing i t back to get a corrected copy automatically,
HitilQut manual typing .
A paper tape system is not without drawbacks. Some tape-
punching typewr ite r s are subject to frequent mechan i cal or elec-
tronic fai l ures . In a large data operation , the tape , \\Th i ch in
quantity is bulky , may be used in such volume as to create a
serious handling and storage problem . Splicing or dup l icat i ng
smaller pieces to produce larger reels , whenever this is necessary,
can be a time-consuming nuisance in a data-banking oper ation.
Paper tape tears or mutilates more easily than cards, and such
tearing can be a frustrating problem, par t icularly if it occurs
\"hile the paper tapes are being read at high speed by the computer .
Despite its dra\"backs, paper tape offers important advantages
over cards i n many applications , and it is widely used. Numerous
examples co u ld be cited of effective applications of a paper tape
system of data convers i on. Under many circumstances , in fact, a
paper tape system is the only practical and , at least in terms of
cash outlay, economical alternative to a card system. Because
paper tape has been in use for quite a few years , industry has
been able to devel op sophisticated , p r ogrammable paper tape
machines, which offer many features , e.g ., tape-editing options ,
that can enhance a data conversion system. Programmable pap er
machines are especially adaptable for applications that require
the production of some type of finished copy , e . g . , specimen
labels, in parallel with the conversion of data to computer-
readable form . For such applications , there probably is no more
flexible data conversion device on the market short of a te r minal
supported directly by a computer of some type .
Paper tape remains, however , a relative l y cumbersome conver-
sion medium, and the trend in the information processing industry
is away f r om paper tape systems to systems of the types described
below. Furthermore, when all e l ements of the system, including
personnel, are considered , a paper tape system is not necessarily
the cheapest. \fuenever extensive editing is required during input,
a computer-linked conversion system that offers text- editing fea-
tures may prove cheaper in the long run if it allm"s for more effec-
tive personnel utilization .
3. Magnetic Tape System : Functionally, a magnetic tape system is analo-
gous to a paper tape system except for the medium of conversion .
Instead of being encoded as holes in paper tape , the data are
electronically recorded on magnetic tape . Hagnetic tape permits
enormous data compression and thus reduces the handling and stor-
age problem several orders of magnitude by comparison with paper
tape . For further ease of handling , the tapes for magnetic tape
typewrite r s are packaged in compa c t cartridges or cassettes , {"hich
can be reused many times . Thus, while cards and paper tape are
cheaper storage media , they can be used only once , whereas magnetic
33 U3

tapes , which are costly to purchase . not only allow for vastly greater
data con tent on a g i ven section of tape but al so are reusable . I n the
end, th e refo r e , t he cost of the sto rage medium per unit of r ecord con-
verted for process ing probably i s abo ut the same ,.. hether card. pape r
tape, or magnetic t ape is used .
It is not advisable or practical , however . to archive data on the
input cartridges or ca ssettes , because of the ir high unit cos t and th eir
uncertain l ife expectancy as a reliable magnetic storage medium . If t h e
data conversion medium itself must be preserved for some reason , then it
is better to use cards or paper tape, which , t h o u gh s u sc eptible to fir e
and mutilation , nonetheless preserve the data record in visible phYSical
code rather than in invisible e lectromagnetic code . If records are to
be s t ored indef i n it e l y on magnetic tape, then they shou ld be copied
promptly onto s t a n dard computer tape, wh ich is fa r more reliable as a
storage medium than the tapes of the var i ous input devices . In fact ,
a data conversion system , r egardless of the medium used , should be
designed to transfer the data from the inpu t medium to standard com-
puter storage devices in as short a time interva l as possible .
One of the chief advantages of the neHer magnetic tape machines is
that they can be used off-line as stand-a lone conversion de vices or on-
line as computer terminals . Although paper tape machines al so c an be
built to function thi s way (e . g ., the Teletype) , magnetic tape terminals
offer a faster, more flexible interface I.ith the computer . Increasingly ,
in fact, magnet i c tape devices a r e being marketed with built-in mini-
comput e r s , which pr ovide the data conve rsion system with dedicated,
programmable memor y .
Magnetic tape systems are not without their drawbacks. Chief among
these is the relatively high cost of purchasing or renting the equipment
and the tapes . Operat ions may he quite expen sive if the syst em is sup -
ported by a comp ut er link. Some magnetic tape machines are just as sus-
ceptible , if not more so , to breakdown as paper tape machin es . At least
some magnetic t ape devices on the market lose synchrony too easily , and
they may not read tapes they have produced previously, or they may not
read tapes produced on anoth er machine of the same make. This problem
adds to the hazard of storing data on t he cartridges or cassettes for
any l ength of time, because a perfec t ly good tape may read like gibberish
on a machine that has gone out of synch r on i zation . The tapes do ,.,tear out
and cannot be reused indefinitely . Finally , in some applications it is
a decided disadvan t age to use magnetic tape rather than cards or paper
tape because the coding cannot be read by eye.
All considered, magnetic tape systems almost invariably are pre-
ferred over paper tape systew.s by those 1;.,th o have used both . The best
magnetic tape systems offer most of the advantages of paper tape sys-
tems and few of the serious disadvantages , and in addition they can
furnish options that are difficult if not impossible to p r ovide \.,tith
paper tape sys tems. For many applications, the logistics of producing
and processing t apes can be simp li fied greatly by switch ing to magnetic
tape .
4. Scanning and SenSing Systems: Various machines are on the market today
that can scan typ e script or graphic mate r i al s optica lly or can sense
special markings or coordinate (X, Y) positions and then translate what
they " see " or " sen s e" into codes on a tape or other magnetic storage
device , ready for proc essing . "lost optical character readers (OCRs)
are capabl e of accurately reading onl y one to several type faces.
Usually , therefo r e , it is necessary to prepare the data by typing
814 33

them on precisely registered data forms with a special type font,

\"hich , hm.,rever, can be done \-lith an ordina r y Selectric typewriter
provided with the right head. Although OCR machines are being
improved rapidly and some now can read printed pages directly , such
as from a telephone book , in general many problems remain to be
solved. Given their present weaknesses, scann ing systems tend to
cause an extra editor i a l burden in proofreading and correcting the
initial input.
Nark-sensing has been used for years to grade examina tions in
colleges and universities and in some kinds of data capture .
Machines fo r digitizing the points in an X- Y grid are being
used to create maps and other graphics in machine- processable form .
Scanning and sensing systems clearly are gaining in i mportance
as Hays of converting data to machine-readable fo r m automati ca l ly ,
thereby eliminating the manual keystroking element of the data con-
version process; however, although they hold great promise fo r the
future , such systems are useful at present only for specialized
applications particu l arly Ruited to the scanning or sensing proced-
ures and involving very large data sets .
5. On- Line System : As exp lained above, some data conversion devices are
terminals linked directly or by telephone to mini -computers or
large computer systems to t.1hich the data can be transmitted directly
from the keyboard . I n such systems the data can be put into a proc-
essing cycle at once if desired . Often special software packages
that enable the terminal operator , at his mm command , to direct
the computer to perform various editing functions have been written
for data conversion systems of this type . Sophisticated text-
editing systems are available which , though relat i vely e xpensive
to use , constitute powerful tools for getting data into machine-
readable form accurately. Output and input may be printed on paper
o r, i n some systems, displayed on a screen .
Persons accustomed to using a good on- line system supported by
text - editing soft\"are become addicted to this method of data con-
version and are r e l uctant to return to any card or tape system
unless it is an economic necessity . Because on-line data capture
invariably is more costly than off- line data capture , at least in
terms of actual cash outlay for equipment and machine time, it is
important to develop convincing cost-effective justification before
opting for an on-line system . Cost-effectiveness in t his instance
is likely to hinge on the eff i ciency of manpm"er ut ilization .

A gene r al caution about multi - purpose data conversion systems is in order

at this point . If the main purpose is to capture data for computer process i ng,
then the system should be designed to do this in the most straightforward ,
eff i c i ent manner possible , and it shou l d not be ladened with othe r peripheral
functions . It may seem logical at first to build a system that can produce
various byproducts such as fi le cards or specimen labels in the process of
encoding input cards or tapes for the computer , or produce encoded cards or
tapes that can be used both to feed the computer and then to replicate file
cards or labels mechanically . Inevi tably, hOHever, compromises are required
when a system is designed for several purposes. Each additional function
places unnecessary constraints on the others . to some extent reducing the sys-
tem ' s efficiency in performing the other functions . Even i f the same equip -
ment is to be used, therefore , in most applications it is likely to be more
efficient in the long run to design independent systems for the separate
33 815

functions. although in practice this is a matter that the individual user will
have to determine for h i mself . In practical terms. this means that labels
sho uld be produced as one operation and input cards or tapes for the computer
as anothe r operation. Another so lution is to produce al l necessary by- products ,
e . g ., file cards , labels, as direct or indirect output from the computer. In
conclusion, the multi-purpose system often proves to be more desirab l e in theory
than in practice.

F. Processing Data . Once the data are converted into machine- readable
for m they are ready for processing and incorporation into the data bank . The
processing strategy and methods should bl~ worked out before any significant
data collecting begins . The processing step is the final an d most critical
one in creating a data bank and it must not be left to chance . The advice of
an information systems expert should be sought ;,lnd follOl.ed carefully, particu-
larly wi t h respect to choosi ng an information processing system and t o deciding
what if any new compute r programs need to be written . It is not the time to
accept the advice of an enthusiastic first - year programming student, or of
someone \.ho has made his mark in computing as opposed to information process-
ing .

The state of the art in the development of gene r alized informa tion p ro cess-
ing softl. are is highly advanced t oday . Before seriously considering the costly
course of writing his own softl. . are . anyone l>1ho contemplates buil ding a data
bank shou l d look careful l y at softl,fare packages presently on the market . and i f
at all possible he should rent or buy one of the many nOI~ availab le "off the
shelf ." The software manufacturer or vendor will maintain his package t o keep
it compatible I. . ith the current generation of hardware. and from time to time
he will upgrade the sof t~'are by adding new features . An individual or an
institutional comp ut ing center hardly can compete with indust ry in providing
s uch service . The day is past ",hen one lvould think of building one ' s own
processing hardl~are, and the day i s virtually past '. . hen one should conside r
building one ' s mm processing software.

Many comput e r serv ice bureaus exist which can rent hard\. . are and software
support on an as-needed baSis , just as , for example , large telephone systems
exist to rent telephone service. There are, o f course, weakness es to thi s
approach and certain pitfalls to avoid if possible. The ch i ef weakness is
t hat the fate of the customer ' s data processing is in the hands of the service
b ureau , which Hill be looking after i ts own interests on a commercia l bas i s .
and these interests will not necessarily coincide with those of t he customer.
Decisions \dll be taken by the firm wit h respect to hardware and softl... are with -
out consulting the c us tomer. Hhose proj ect may be af fec ted profoundly , per haps
disastrously. by the consequences . The service bureau may abruptly quit offer-
ing the softwa r e being used by the customer, for example . thereby forcing the
customer to make a costly mi d-stream conversion to another sof t wa re package or
to make a dis r uptive t r ansfer to another service b u r eau offe r ing the origin al
softwa r e . Computer cente r s , whether publiC or pr ivate . a r e notor i ous for
promising more than they can deliver at a given cost, and the \wu ld-be cus -
tomer has to select his service bureau lvisely. In particular, he must try to
avoid getting involved Idth a f l y-by-night or financially unstable firm , I.;hich
has a strong probability of going out of business or drastically curtailing
se rvi ce at a c rucial moment in the customer ' s development ef fort . This is the
major pitfall to avoid because the software vending and renting i ndustry is
stil l in a dynamically grOl. . i ng state , and the rate of business failure in
this industry is high . It should be kept in mind. however, the same \.;eaknes ses
816 33

and pi t fa lls are likely to be encountered, though perhaps in different fo rms ,

in dealing with in-house computing instal lations . In short. the manager of
a data bank , wherever he e l ec ts to process his data, must guard against the
basic capriciousness of the information p roces sin g industry at this stage in
its history .

A number of generalized information processing syst ems currently are on

the market and are being used for developing biological information systems .
These include notably TAXIR (TAXonomic Information ~etrieval) , the sys tem
developed by David J . Rogers and his associates; GIPSY (General Information
fro cessing SYs t em), developed by t he University of Oklah~ma Comp~ting Center
under the gu idance of J ames ~~ . Sweeney; SELGEH (SELf GEnerat i ng Maste r), the
Smithsonian Institution ' s general ized processing syst;'; developed under the
direction of Reginald Creighton; and GIS (~eneralized I nformation ~stem) ,
develop ed by IBN . The Flora North America Program adopted GIS as the basis
for its data banking .

V. CURRENT DATA SYSTEHS

To give more than a mention of the many data systems now operating or
under development would require another chapter. The interested student is
encouraged to consult the references cited at the end of the chapter for per-
tinent details. Most of these systems are in such a sta te of flux that to
describe thei r present charact e ristics in detail here would be pointless in
any event.

Flora North America aims to develop a truly generalized information sys-

tern to do data banking on a broad network basis . Few oth er systems of equally
generalized character appear to be underway .

Various botanists have developed systems for mapping plant distribut i ons
by computer . James II . Soper of Can ada's Na tional Huseum of Natura l Sc i ences
has pioneered in this a r ea . Recen tly , in a pro ject initiated at Colorado
State Univers ity, ~Jilliam H. Klein and Rober t P . Adams have developed some
exciting new approaches to distribution-mapping by computer in r ela tion to
critical environmental factors . Mapping by machine Has done for the entire
British flora some years ago by Franklyn H. Perring and S . M. Walters , but
this was done t"ith a mechanical tabulator ra t her than a computer. Several
other British botanists have jus t completed and published a study entitled
A Compu t e r-Napp ed Flora : A Study of the County of \.j'an"ickshire in which the
compu ter t. . as used for some of the mapping and a mechanical tabulator for the
res t of it .

In the realm of the herbarium, a number of plant taxonomists are now

using computer-based systems to store and retrieve spec imen data. Notable
herbarium projects are the Smith sonian In stitution's Type Specimen Register,
an ambitious effort to create a computerized registry of botan ical type speci-
mens on a ,,,orldwide basis, indicating ,,,here the names were published and Hhere
the specimens are deposited ; Notre Dame University's proje ct , under the direc-
tion of Theodo re J . Crovello , to computerize the specimen data in the E. L.
Greene Herbarium; and Sope r's effort at the National Museum of Nat-ural Sciences
in Ottat"a to create a specimen data bank fo r the Province of Ontario .

The Plant Records Ce n ter of the American Horticultur al Society is wel l

along in the devel opment of a data bank of pertinent botanical and horticultural
 -----------------

33 817

data concerning living plant collections in North American botanical gardens

and arboretums .

The Hunt Institute of Botanical Documentation . located at the Carnegie-

Nellan University in Pittsburgh. Pennsylvania, has concentrated on the devel-
opment of an information system to handle botanical literature, and it has
undertaken several pacesetting retrieval projects in this area . Using its
system , the Hunt Institute processes nomenclatural data for the Index Nominum
Genericorum, a project of the International Bureau for Plant Taxonomy and
Nomenclature in Utrecht. The project, \.,rhich is far advanced in the creation
of a computerized index of validly published generic names for all plant groups
of the world, is headquartered at the Smithsonian Institution. A similar Horld-
t"ide index of species names is now contemplated.

Finally, in the area of specialized application programs, mention should

be made of the promising program packages that have been developed for using
the computer to assist in plant identification . The imaginative taxonomic
matrix system of Larry E. Morse is particularly noteworthy because it enables
the botanist to sit at a remote terminal and use the computer on-line in the
conversational mode to identify unknmm plants or to construct dichotomous
identification keys on the basis of stored data matrices linked to stored
lists of taxonomic names and characters. This system can be used to great
advantage in the classroom as a teaching aid or in the laboratory as a research
tool. It is only one examp l e of the numerous application programs that are
avail able to the initiated scientist today.

VI. CONCLUSION

Of necessity botanists \"ill turn increasingly to the computer in the

years to come for help in solving their information retrieval problems, and
scientific data hanks of all sizes and descriptions ,,,ill result. It is incum-
bent upon plant systematists to make full use of the computer as a tool and to
guide the development of botanical data banks in an orderly and coordinated
way. Surely no present-day student of plant systematics can afford to omit
from his training the study of information processing technology as it pertains
to scientific data banking. He have only begun to exploit this technology,
and yet today's technology is a mere prelude to the state of the art as it \~ill
he in the future.
 Chapter 34. SOCIETIES , BIOLOGICAL STATIONS, AND RESEARCH SUPPORT*

He rbar ia , botanic gardens, arboreta , data banks, libraries (limited

sense) , record centers , taxonomic li terature, bibliographic aids , and infor-
mation systems have been treated in the previous four chapters of this divi-
sion on facilities and general information . In this chapter a miscellaneous
assemblag e of material is presented on societies, systematic centers , biologi-
cal stations and support ive organiza tions in an effo r t t o make informa tion on
taxonomic facil it ies and genera l informat i on comprehensive and comple te . The
treatment of biological stat ions and fu nding sources is made with a little
trepidation s i nce some of these facilities and agencies may be relative l y
short - lived. The facts in this chapter on biological stat i ons and research
support indicate the diversity of types of stations and financial support
at this time . An effort should always be made to procure t he most recent
information on f ield sta t ions and grant sources.

I. SOCIETIES

It is a real problem fo r any profess ional to keep abreast of the current

literature and rapid development in his specialty . The problem is compounded
in taxonomy since i t is such a synthetic field invo lving so many peripheral
disciplines --each expanding rapidly in techniques and data . The mass of cur -
r ent publications has confronted the taxonomist with an almost insurmountabl e
task of deciding what journals to get fo r his own library , what to read regu-
l arly in add it ion, and how to learn about important papers elsewhere .

One should form a habit of s urveying the current literature by regularly

and systematically checking certain abstracting journals or indices (see
Chapter 30) . The "Index to American Botan ical Literature ," published in the
Bulletin of the Torrey Botanical Club , is espeCially hel pful in this r egar d .
I n addition, make a l ist of current journals in yo ur schoo l library {">hich
regularly have articles of intere st to yo u, then survey all of these at least
once a year . Such a list could easily inc lude fifty or more journals .

Personal membership in professional and honorary societ i es is one way of

making certain that important journals are seen r egularly and also permanently
available in your olm library. However, getting the jou rnal is no t the only
reason to belong to profeSSional societies . After a ll, t h ese organizations
repr esent us and our fie ld of endeavor on a regional , national. or wo r l dwide
basis , and can sometimes be effective spokesmen in matte r s of concern to all .
\,,]e all have an obligation to support the societies or associations which
represent our area of specialty and areas of interest . Attendance at the
annual meetings of these societi es gives one an opportunity to have an input
into the b us i n ess affairs, programs, and activities of the or ganization . as
well as to knOl~ the other meJ:lbers.

Each ind i vidual must make his own choice of membersh ips, but considera-
tion should be given to at l east one or more in each of the following broad
categories .
1. General science or biology in the U. S .
(American Association for the Advancement of Science, American

*Contributed by James H. liard in , North Carol ina State University , Raleigh.

 --- -- - ---- - - -

34 823

Institute of Biological Sciences--effective "voices," hop efully ,

of science and b i ology in this country}
2. Botany in the U. S .
(Botanical Soc i ety of America--th e "mother society " for all
botanists in this country)
3. Taxonomy in the U. S . and internationally
(American Society of Plant Taxonomists, International Associa-
tion for Plant Taxonomy--the chief societies in this dis-
cipline)
4. Regional or state organizations
(This category includes numerous societies that publish such
journals as Rhodora , Bullet i n of the Torrey Botanical Club,
Castanea , Journal of the Elisha Mitchel l Scientific Society ,
Nadroiio , South~"estern Naturalist, Bulletin of the Californi a
Academy of Science ; most states have an Academy of Science . )
5. Special areas of interest
(Society for the Study of Evolution, Anerican Fern Society ,
American Bryologi cal and Liche nolog ical Society, American
Forestry Association , or thos e repres e nting per ipheral disci-
plines)
6. Honor societies (membership is by invitation only)
(e.g ., Sigma Xi , Phi Kappa Ph i, Gannna Sigma Delta, Phi Si gma)

An important organi zation for taxonomy, with institutional membersh i ps

only , is the Association of Systematics Collections . This organization was
begun in 1972 to " foster the care , management , preservation , and improvement
of systematics collections and to facilitate their utilizat ion in science . "
One signif icant result of this organization is the \-:riting and presentation
of a document entitled " America's Systematics Colle ctions : A National P l an . "
This report is addressed princ i pal l y to systematis t s concerned ~"ith the nature
and quality of America ' s systematics collections as a national resource, and
describes and reconnnends \·lays whereby the collections can be more responsive
to the needs of the basic and appl i ed sciences as well as society in general .

This organization will be an effective voice for all systematists con-

cerned with the prese r vation and utilization of biologi cal collections, and
it is therefore most important for all museums and herbaria to be represented
in the ASC .

II . BIOLOGICAL STATIONS

Most i mportant for the study of taxonomy or taxonomi c research is the

ability to get out in the field \vith the plants . Taxonomists must not los e
sight of the fact that the living plant in the field is the ultima t e source
of data . Hany colleges and universiti e s are in grOlving urban areas and field
experience for most students i s rapidly becoming limited . I t is also increas -
ingly difficult t o obtain travel money for resea r ch . Biological stations are
often the anSI"er to these problems .

Summer courses and/or research at biological stations have obvious advan-

tages over most campuses . The locations are generally in environmen ts with
unique or repres e ntative biotas , or at least within easy access to natural
areas . I t is a good \"ay to get into different areas and learn a different
flora or be in an ideal l ocation for research on a particular group . Courses
a r e generally fiel d oriented and often di ffe rent from those available on mos t
824 34

camp uses . Money i s sometimes available from the station to help defray
expenses. Students and faculty are enthusiastic biologists and a delightful
group with which to be; and, it is generally a most pleasant \vay to spend a
summer of learning a nd research .

Biologi c a l stations are easily divisible i nto either "mar i ne stations"

speciali z ing in oceanography and ma r ine species, or " inland field stations"
specializ i ng in ter r estrial or fresh - Hater biotas. Vascular plant taxonomists
shoul d become fam ilia r wi th the i nland stations and the fac i lities available .

Facil i ties, programs , and opportunities vary tremendously among stations .

Some have research facilities only, such as the Hi ghlands Biological Station ,
while others have teaching programs in addition , such as Hountain Lake Bi ologi-
cal Station . Al l have something unique to offer , either i n program or the
b i ota or ecology of the area. In 1945 there were 53 inland field stations, in
1 966 there Here 42 (Arvey and Riemer , 1966), and in 1970 there Here about 100
in the United States (Hunsaker and Dalgleish, 1970) . This growth probably
refelects the recent emphasis on environmen t al biology as I"ell as the decrease
in ava i lable na t ural areas close to university campuses .

To learn about the facilities avai l able and programs offered at a parti-
cular station , wr ite to the director of a station in the area of interest
(Figure 34-1). Addresses can be loc a ted in the pub l icat i ons cited belO\~ or
through a university in the a r ea . Hany of the stations also belong to the
OIBFS (Organization of I nland Biologi c al Field Stations) which can be con-
tacted c/o The E. N. Huyck Preserve, Inc. , P . O. Box 87 , Renssel aervi lle ,
N. Y . 12147 . Host of the larger stations have brochures I"hich are available
upon request .

References on Bio l ogical Sta t ions

Arvey , H. D., and H. J . Riemer . 1966 . I nland Biological Field Stations of

the United States . BioScience 16: 249 - 254 .
Hunsaker, D. , and R. Dalgleish . 1970 . Biological Field Stations of North
America . HSS Educational Publishing Company , Inc . New York .
Palmer , A. H. (ed.) . 1972. Research Centers Di r e ctory, 4th ed . Gale
Research Company . Detroit, Hichigan .

III. RESEARCH SUPPORT , AND i'RE- A.I\!D POST-DOCTORAL FELLOHSHIPS

\\Thether the desire or need is for a stipend , a major piece of equipment,

travel for field Hork or to e xami ne type specimens, or page cost f or a publi-
cation , the one vexing question i s ' \ lhere l2. ~ the money . " It sometimes
takes real ingenuity and aggressiveness to f i nd funds . Don ' t be afraid or
embarrassed to ask ! But, have a good plan and justification in mind first .

For extended stays in various parts of the country or I"orld , you might
consider teaching in secondary schools or colleges in the area. Often there
are American schools abroad that need teachers . Some of the addresses below
I"ill put you in contact with these . Also the Peace Corps programs I'lay be a
good I"ay to stay in an area for awhile .

Another possibility for some l imited trav el funds is th e sale of speci-

mens. This is something to consider particularly i f you are going to some
remote area \~hich has not been vJell co l lected . First , get an agreement I"ith
34 825

Figure 34-1. Inland

Biological Field Stat ions
of North America . Solid
blocks : statio ns repo rted
in 1966 ; open triangles :
a ddi tional stations
reported in 1970 .

the herbar i um curator , museum direc t o r , or researche r regarding t yp e of collec-

tions , numb er, i denti fi cation s and l ab els , methods of preservation , and cos t
per spec imen . Think beyond re gular herbarium spec imens to such th ings as
pollen samples, wood sections , r.oots or leaves fo r chemical ex tracts , photo-
g r aphs , pathogenic f un gi, pollin ating insects , or damagin g insects . The
r esearch division of some petroleum companie s may be i nterest ed i n pollen
samp l es; cer t ain chemi c al or pharmaceu t ical companies or the USDA may want
pla nt ma te r i al for ext rac ting a l kaloids , glycosides , or fats . Identifiable
voucher specimens must accompany a ny such collec t ions .

Fellm"ships or grants may be obtained from private , state or federal

so urces . Star t by as king your co lleagues Hho may have alr eady been suc cessf ul
in this venture, then the admin ist ration in yo ur OHn school (department head,
dean s o f your school , graduate school , or of research) . Someone in the admin-
i s tration usually keeps up \odt h p r ograms availab l e through the va r ious grant-
ing agencies (National Sci ence Foundation [ NSF1 . Na tional Insti t ut e of Hea lth
[NIH] , Department of He alth, Education and He lf a re [HEI-Il, Na tional Aeronauti cs
and Space Administra tion [NASA], Natio nal Defense Education Act [NDEAl , Atomic
Ene r gy Commission [AEC]) . and the various felloHship pro g r ams available. It
is always pos sib l e that you might fi nd t he money right in your OHn insti tution .

Beyond thi s , the re are nume r ous possibilit i e s as ind ica t ed be lo\" .

A. Non-state or non-federal agencies

1. Biolog ical fiel d stat ions ( see discussion in this chapte r).
2. Organiz ation of Tropi cal St udies (OTS) North Ameri can Office , P . O.
Box 1499 , South Mi ami, FL 33143).
826 ~

3. Sigma Xi Grants-in-Aid of Resea rch (Sigma Xi , 155 \-1hitney Ave ., Net..'

liaven, CT 06510) .
4. Private fou ndat ions . See : "The Foundation Directory" available in
the reserve room of most libraries; look under Life Sciences in
the Index of Fields of Interes t. Some do give grants or fellOloJ-
ships t o individuals . Some examples are :
a. The Danforth Foundation , 222 South Central Ave ., St. Louis, NO
63105 (graduate fello\is hip s , graduate fellowships for Homen,
Kent fe llOl,'sh ips) .
b. Ford Foundation Doctoral FellOloJships, 320 East 43rd St., Ne\ol
York, NY 10017 (graduate study leading to Ph . D. for Black
Americans. American Indians , Hexican Americans , and Puerto
Ricans) .
c. John Ray Hhitney Foundation Oppo rtun ity Fellowships . Hrite:
Opportunity FelloHships, Jolm Ray ~"hitney Foundation, III H.
50th St. , Nel" York, NY 10020 (for Black Ame ricans , Spanish
Americans, American Indians , residents of the southern Appa-
lachi.:m and Ozark I'lt . areas, Guam, Puerto Rico , Samoa , Pacific
Trust Territory, and Virr,in Island s) .
d . UoodroH Hilson Nat iona l Fel101"ship "I<'oundation, Box 642, Princeton,
NJ 08540 (first-year gradua te study; dissert<'ltion fellol"ship) .
e . John Simon Guggenheim I'lemorial Foundation , 90 Park Ave . , New
York, 1'!Y 10016 (fields unrestricted) .
5. National Geographic Society , Committee fo r Research and Exploration,
17th & 11 St . NH , Hashington , DC 20036 (post-doctoral travel
grants) .
6. The In ter- Unive rsity Council for Higher Education Overseas , 90/91
Totte nham Ct . Rd . , London, England (assists in recruitment of
academic staff of smaller colleges in Hritish Commom"ealth) .
7. r~ear East College Association , 305 E . 45th St ., New York , NY 10017
(teaching positions in private American colleges in Greece, Turkey,
and Lebanon) .
8. Smithsonian I nstitut ion Visiting Research Associate Program , or
Visiting Postdoctoral Research Associate Program . ~.Jrite : Director ,
Division o f Gt'aduate Studies, Of fice of Academic Prog rams , Smith-
sonian Institut ion , l·]ashington, DC 20560 (fel lolvsh ips for study
at the Smithsonian) .
9. Environme.ntal consulting fi rms (possible job or s ubcontract research
Hith various groups involved I"ith ecological studies or environ-
mental impact studies) .
10 . Conference Board of Asso ciated Research Councils, 2101 Constitution
Ave . NH , Hashington, DC 20418 (university l ecturing and resear ch;
postdoctoral felloIJships).
1 1. I nstitute of International Education , 809 United Nations Plaza , NeH
York, NY 10017 (graduate study abroad; predoctoral fell m.'ships).
12 . Council o f International Educational Exchange , 777 United Nations
Plaza, Nev' York , l\TY 10017 (aid and information on study programs
abroad) .
13. American Association of University \·,Iomen Educational Foundation
FelloHships . Hrite : Direc tor, MUI,J FelloHship Program , 2401
Virginia Ave. , NH , \.Jashington, DC 20036 (pre- and post-doctoral
fell oViship s) .
14. East-l.Jes t Center , 1777 East-Hest Hoad, Honolulu, HI 96822 (senior
specialist awards; scholarships for graduate study at the Univer-
sity of Hm"aii) .
34 827

IS . Foreign Ar ea Fe llm-lship Program, 110 E . 59 th St ., Uel" Yo rk, NY 10022

(r esearch in Africa , Near and Hiddle East , South Asia , Southeas t
Asia, East Asia , La tin America and th e Ca ribb ean , Soviet Union ,
Eastern and Hestern Eu r ope ) .
16 . Latin American Teaching Fel lowsh ips, The Fletcher School o f La\" and
Diplomacy , Tu ft s Univers ity , Medfo rd, HA 02155 (p r edoctora1
fe110\.Jships in social and natural scien ces ) .
17 . Na t iona l Hildlif e Fede r ation Fe llowshi.ps , 1412 16 th St. m.,r , Hashing-
ton, DC 20036 (predoctora1 fe 110\ls h ips in conservation and related
areas) .
18 . Oak Ridge Gradua te Fe1101.Jships . h'r i t e : Univers i ty Programs Office,
Oak Ridge Associate Unive rsities , Oak Ridge , TN 37830 (for work
at Oak Ridge or other AEC sites) .
19 . Organi za t ion of American Sta t es , 17th St . & Constitu tion Ave . NIJ,
Hash in g t on , DC 20006 (OAS Fe11o\<l'ship fo r advanced speci alized
st ud y, training , or resea rch in Latin America) .
20 . South e rn Fellol,'ship Fund, 745 Peachtr ee S t . NE , Atlanta, GA 30308
(predoctoral fellow ships for black s tudents) .
21 . tlATO fel lO\~s h ips . \-!rite : Fellm-Iship Office , National Research Coun-
cil , 210 1 Constitution Ave . NH, \/ashington , DC 201118 (postdoctoral
fe llo\,Iships f or st udy primarily , but not restric ted to , countries
of the NATO all ia nce) .
22 . Th e American Ph iloso phical Soc iet y , Independence Sq uare , Philadelphia ,
PA 19106 (postdoctoral g rants- i n- aid for res ea r c h costs ; not for
stipend s , publicat ion cos ts, or permanent equipme nt) .
23 . The Phi Kappa Phi F ellol~ships . Writ e : Phi Ka ppa Ph1 , 3001 Plymouth
Rd., Ann Arbor , HI MHOS (first a nd second yea r g raduate fellow-
sh i ps i n all a r eas) .
24. Sta te academy o f science (some have modes t g rant s- in-aid of research) .

B. State agenc ies

1. Sta te a gencies such a s museums , parks department, Hildlife agencies ,

science a nd technology hoard s , etc . (fo r summer jobs or modest
support of research.)
2. Uni versity fac ilitie s such as phytotrons , comp uting cent e rs, agric ul-
tural expe riment stations . (Often th ese have state funds or fed-
eral grants for support and can a 1lOi' use by faculty and stud e nts . )

C. Federal agen cies ( f or a comprehensive list in g se e "Catalog of Federal

Domestic Assistance," U. S . Printing Office 1 969 -0- 328-264) .
1. Grants for Imp r ovi ng Doctoral Diss e r a t ion Res earch in the Field Scien-
c e s . tlationa1 Science Foundation (funds for travel to librar ies,
museums, or field research loca tions ; spec ial equipmen t ; computer
time ; no stipends given) .
2. Institute of Inter na tional St ud ies , Office of Education , Department of
Hmo/ , \..tashington, DC 20202 (res ear ch or teaching in fore ign schools
unde r the Fulbright-Hays Act ; also , r eq uest in forma tion on foreign
travel, Iwrk, and study) .
3 . Division of Student Financial Aid , Bureau of Hi gher Ed uca tion, Office
of Ed ucation , Department of IIEI-.' , I-!ash i ng t on , DC 20202 (loans for
st udy ahroad) .
4. PellOl.ship Off ice , Office of Scientifi c Personnel , National Research
Council , 2101 Constitution Ave . N\~, \·Iashington , DC 20418 (Publica-
tion : " A Selected List of Najor FellOi~ship Opportunities and Aids
to Advanced Education for United States Citizens") .
 828 35

Chapter 35. QUESTIONS AND PROBLENS

A series of questions are presented fo r thought and discussion which wil l

provide the student with an introduction to some of the basic problems and
issues in systematics. The species biology. ec osystemat ic s . biosystematics.
and phylosystematics problems require a summary application of the various
facets of taxonomy. These problems can be used as class exercises. indivi-
dual student projects, or parts of each can be used in various comb inations
for particular systematics training emphases by the teacher. Anyone of the
problems could provide the basis for a student research project. Sample
revieHs are presented to sho\" how character or topical summaries of the fea -
tures presented in the text can be used as bases for class or individual
reviews or quizzes in the different aspects of systema tics . No effort has
been made to show test methods since systematics test methods should be simi-
lar to those found in all fie lds of ed ucation. A basic course midterm and
final examination have been included to show what can b e expected from stu-
dents at that taxonomic level rather than for quizzing methodology .

Section A. QUESTIONS FOR THOUGHT AND DISCUSSION

I. GENERAL SYSTEMATICS QUESTIONS

1. Hhat are the principles of taxonomy?

2. lo.Thy is the holistics approach in a basic course in systematics desirable?
3. Hm-l can taxonomists justify their type of work or study in this time of
environmental crisis and socia-political turmoil?
4. How does a systematics reference text or a general taxonomic text help and
hinder the development of the field of systematics?
5. '..Thy should a basic systematics course be a core course in a liberal educa-
tion curriculum? A fundamental course in botanical training?
6. l.,lhy should plant taxonomy be considered the acme and pedestal of plant
sciences?

II. HISTORY AND CLASSIFICATION QUESTIONS

1. How does a study of history of taxonomy give the student perspective in

systematics?
2. How is the organization of history into "P e riods of Development of Classi-
fication " misleading?
3. Hhy is the development of systematics dependent upon the development of
other fields of science?
4. lfuat is the practical use of a classification system ~"hich divides groups
of organisms in to hierarchies; e. g.. species. genera. fam ilies, and
higher taxa?
5. lfuy should plant taxonomists work vigorously for the production of an
effective phenetic classification of plants rather than a phylogenetic
one?
6. What are the present trends o f systematics research and ",hat is the future
of taxonomic work?

III . NOMENCLATURE AND EPITHETS QUESTIONS

1. hThat are the advantages and disadvantages o f a stable or fixed nomencla-

ture?
 n 829

2. lfuy did synonymy arise and why can it not be e liminated in t he future?
3. What are the difficulties in applying the type conc ept to all kinds of
plant s and plant remains (fos si ls)?
4. Even though there are only several hundred epithe ts \>lith their meanings
listed in the text , how can the meanings of thousands be determined
from the information in th e text?
5. Uhy should species names be conserved? Gen e ri c names? Any names?
6. \·n\at should the fundamen tal components of a speCies definition be?
7. Hhat precau tions sho uld be taken in selecting a good specific epithet?
A generic name? A higher taxon name?
8. Ifhy 1s binomial nomenc l ature conside red basic to all of taxonomy? To all
of bo tany?
9. I;lhat are the reasons for the rejection of names?

IV . EVIDENCE QUESTIONS

1. \Jhat is meant by character or evidence type, character state , ordering of

cha racters and what are some of the problems in determining type , state ,
and order?
2. Under what conditions can a meani ngful character be reliable . o r diagnos-
tic , or homologo us , or analytic, or synthetic, or two-state , etc.; or
reliable and ana l yt ic, or analytic and synthe ti c, e t c . ?
3. {·lhy should morphological evidence or characters be t he basic type used in
identification and classification?
4. Hhat are the advantages and disadvantages of a refined and definit ive
descrip tive morphological terminology based on measurements; e. g .,
angles and r a tio s?
5. Should on togeny-phylogeny be the basis for definitio ns of morphological
and anatomical terms and charac ters ? Explain.
6. What are the dangers in using one type of evidence almost exclusively;
e . g . , anatomical , morphological , or chemical in classification?
7. At what level(s) in the taxonomi c hie rarchy or system are the follmling
types of data taxonomically most useful : anatomical , embryo logical ,
palynological, cytological, chemical? Explain why for each type of
data .
8. What are some of the precau tions to take in using each of the follow ing
types of data for taxonomic purposes : anatomical , embryological , paly-
nologica l, cy t ological , chemical?
9. What are some of the fundamental problems involved in procuring data of
the fol lowing types fo r taxonomic purposes : anatomical, embryological,
pa l ynological, cytological. chemical?
10. Are al l t ypes of data of equal weight and value for c lassification?
Iden tification ? Explain .
11. How a re the fo llOt.ring features r elevant or useful in t axonomic research :
polyp loidy , karyological features , breeding patterns , meiotic pairing,
habitats , types of reproduction?
12 . Why are e cological characteristic s not used more freq uently in taxonomic
descriptions?
13. \olha t are the f undamen tal problems involved in wo r king out a satisfac tory
classification of tr ichomes. venation patterns. i nflorescence types ,
fruit t ypes?
14 . PhYSiological characteristics are seldom ever used in any phase or aspect
of systematics . Explain why not .
15. Should bas i c courses in each of the fields of evidence be required train-
ing for advanced taxonomy students?
 830 35

V. EVOLUTION AND PHYLOGENY QUESTIONS

1. Hhat is meant by evolutionary evidence and hO\'I is it used in systematics?

Phylogenetic evidence ?
2. I n the study of anatomy and morphology, hm" can phylogenetic or evolu-
tionary advancement be distinguished from ecological specialization?
3. If a p l ant possesses a single or a very few primit i ve characters , can it
be assumed to be total ly primitive? Explain .
4. Hhat is the basis for determination of "primitive or advanced " ? Cite a
few examples.
5. Hhat are some of the evolutionary "pitfalls " in morphological and ana-
tomical phylogenetic study?
6. Hhat is adaptation? Preadaptation?
7. ~'Thy do all parts of an organism not evolve at the same rate?
8. Hhat is the evolutionary role of each of the fol l owing : phenotypic var ia-
tion, fitness, competition, gene flm" barriers?
9. Is mutation or recombination the more important source of ind i vidual var-
ia tion? Explain .
10 . Hhy is evolution unid i r e ctional under strong abiotic sel ections?
11. In phylogeny should some characters be given more t"eight than others?
Explain .
12 . t-.lhat are the r el ationships between mutation rate and rate of evolution?
13 . Hhat is meant by chemical phylogeny and what is its s i gnificance to
systematics?
14 . Hhy does the search Eor evolutionary relationships inspire the majority
of taxonomists today in their studies?
15 . \<1hy are angiosperm classification systems based on phylogeny unattainable?
16 . \<1hy is vessel element evo l ut ion considered a major phylogenetic tool?

VI . GEOGRAPHY, ECOLOGY , FIELD QUESTIONS

1. Of t"hat significance is plant distribution in elucidating evolutionary

histories?
2. Hhat is meant by center of distribution? Hhy can it not be determined
readily f or most taxa?
3. \~hat are some of the chief factors that make some plants widely dispersed
and some very limited in their distribution?
4. I·lhat is meant by endemism? \ihy is there so much confusion in applying
the concept?
5. 1.Jhat are the f undamental problems in defining or classifying the follot"-
ing : dispersal types , community types, habitat types , biogeographic
regions, ecological characteristics?

VII . DATA AND l>!ETHODOLOGY QUESTIONS

1. \.Jhat is meant by documentation of taxonomic research?

2. l.Jhat are the components of good experimental design in biosystematics
research?
3. Hhy are differen t analytical procedures necessary for a comprehensive
study of local populations?
4. loJhat can be done to make breeding systems and transplant studies practi-
cal for graduate student research?
5. Hhat are the general problems associated t"ith selecting the best method
of data presentation?
 35 831

6. lfuy are experimental studies and populational analyses considered funda-

mental to modern systematics research?
7. \tIhat should the role of the botanica l ga rden and herbarium be in taxonomi c
re search , conservation , and t eaching?
8. '.</hat are some of the problems in c l assifying evidence fo r deposi t in a
da t a bank?
9. I·That is the relevance of data banking and data r e trieva l to the development
or demise of herbaria and botanical lib raries ?
10 . How Idl l t he computer reduce taxonomic labor and th e need f or taxonomists ?
How l"i11 the comput er and data bank be of service to the taxonomi st ; of
taxonomic use to the public?

Section B. PROBLEl'IS

I. SPECI ES BI OLOGY PROBLEI'!

Sel e ct one species and work ou t the characte r s t ate(s) for each of the follo\-1-
ing :

PHYTOGRAPHIC CHARACTERS
(See Chapter 6)

1. Plant Type 33 . Leaf Shape

2. Plant Duration 34 . Leaf Hargin
3. Roo t Types 35 . Leaf Apex
4. Root St ructur a l Type 36 . Leaf Base
5. Stem Type(s) 37 . Leaf Surface Configuration
6. Stem St ruc t ur a l Type 38 . Leaf Surface
7. Stem Location 39 . Leaf Vestiture
8. Branch Posi tion 40 . Leaf Vena tion
9. Branch Arrangement 41. Leaf Texture
10 . Branch Orientation 42 . Inflorescence Type
11 . Branch Transverse Posture 43 . Inflorescence Posit ion
12 . Branch Longitudinal Post ure 44 . Fl ower Type
13 . Bud Type 45 . Corolla Type
14 . Bud Structu r a l Type 46 . Calyx Type
15 . Bud Scal e Arran gement 47 . Hypan thium Type
16 . Bud Cove r i ng 48 . Per ianth Arra ngement
17 . Leaf Type 49 . Perianth Position
18. Leaf St r uctu ral Type 50 . Hypanthium Adnation
19 . Pe t iole Structural Type S1. Androecial Type
20 . Petiolule Structural Type 52 . And r oecial Position
21. Stipul e Type 53 . Stamen Structural Type
22 . Stipe1 Type 54 . Stamen Arrangement
23 . Leaf Location 55 . An th e r Type
24 . Leaf Position 56. Anther Shape
25 . Leaf Arrangemen t 57 . An t her Attachmen t
26 . Leaf Or ientation 58 . Gynoecial Type
27 . Leaf Tr ansverse Posture 59 . Carpe l Type
28 . Leaf Longitudina l Po s ture 60 . Carpel Arrangement
29 . Leaf Ptyx i s 61 . Stigma Type
30 . Leaf Number 62 . Style Typ e
31 . Leaf Division 63. Ovary Po si t ion
32 . Leaf Size 64 . Ovu l e Type
224 9

<
secondary _ endothecium

<
parietal cell
Honocotyled onou s primary
parie ta l cell middle layer
secondary
parieta1.. .cell
tapetum

<
secondary endothecium
pr i mary parie tal cell
Reduced parietal
cell secondary
parietal cell ----- tape tum
B. Quadripartition of the Hicro spore Hother Cells.
Variable characte rs include successive or simultaneous division,
tet rad formation by furrowing or cel l plate f ormation, and rnicrospores
(pollen) shed as sin gle g rains , tetrads, or polyads .

C. Development and Organizat ion of the Pollen Grain . (See Chapter 8 ,

Palynology)

D. Develooment and Structure of the Ovule .

1. Numbe r and structure of integuments .
2. He thod of vasculation (\~hen present, t-l~ere?)
3. Presence or absence of an aril. Pre- or post-
4. Presence or absence of an arillode ("false aril") . ferti li zation
5. Presence or absence of a caruncle ( s trophiole) . development
6. Presence and place of origin of an obturator .
7. Structure o f the m ic ropyl ~ .
8. Ovule type based on nature of the nucellus .
a . Crassinuc ellate ovule . An ovule ""ith a massive nucellus in
which the megaspore mother cell is deep ly embedded .
b . Tenuinucellate ovule . An ovule in l"hich there is a small amount
of nucellus with the megaspore mother cell subepidermal.
9. Presence or absence of an endothelium (integumentary tapetum) and
extent of coverage of megagametophyte .

E. Fo rm and Extent of the Nucellus .

Si ze and shape, presence or ab sence of a hypostase , ep i stase , method
of origin of integument or integuments, persistence of nucellus as peri-
sperm, presenc e or absence of a nucellar cap , and pers istence or de gen-
erat ion during development .

F. Origin and Extent of the Sporogenous Tissue in the Ovule .

Number of archesporial cells (one , f ew, many) and presence or absence
of \,ral l layers , and nature of cell divisions .

G. Negas porogenesis a nd Developme nt of the Embryo Sac .

Charact ers of special importance a r e time of \,Tall formation in mega-
sporogenesis , pO Sition of funct ion ing megaspores , and size, shape and
a rrangement of cells and nu clei composing the embryo sac (female game to-
phyte) type . Determine and record type (see Figure 9-1) .
Figure 9-1 225

EMBRYO SAC T YPES IN ANGIOSPERMS

Megasporogene sis Megogom!Iogenesls

Type
MeQospore
mot her ce ll
D, v,s, on
, D"'510n D,v'SlOn D,v,s,o n
<V ,
DMs,on Moture
emb ryo soc
" '"

@ fID ~;
~

~ (J ®
ManaSDOflc @@
a-nucleote
Po lVQonum ty p~ 0 @
@
®
®~
\@/

~
I@'
Monospo,"c
4 - roucleoTe
Oeroolhe ro type @@ @
'Q§I ~
(!)
®
®
t)
~
~
a,spo,"c
a- roucleote
AII,um lype @ @ 'vw! @ (i)~ ®@

~
(!)
@ W~ ~
;00\

®
Te T,ospofiC
16- nuCleate
@@ 0 0
o 0 0G)
Peperom,o Type @ ® ~ i(J!'
~
@ (!) ~ 0 ~
Telr05 ponc
16-nucle a te
@ ®
o 0 ~g> 00
o 0 -G 00
"enoeo typ e @ ® ' <v ~

@~ ~ ~~ 't.. ~
Teiro soo r ic
16- nucle at e ffi
Druso ty pe @ ®®
@

Tel.ospo",
fl- nucl eaTe
F"T dla' ;o TyDe @ (!) ~ @ ~ (J'" (!)''""
0 ~@
~g
[J
,4,0\
Tetrospo"c
a-nuc l eo te
P lumboQello ty pe @ (!) ~ @ ~ ..,
~ CD
@ (!) ~
,~
TeTroscoroc
8-roucleote
Ptumbo go type @
®®
®
0
0
' &v
0
0
9
Tel 'O SPO "C
a-nucleote
AdOl O t~pe
@. (!) ~ @ ®@
~

•
00

m
226 9

1. Monosporic . Embryo sac formed by the germination of one of the mega-

spores of the t etrad .
a . Polygonum type . ~hth 8-nucleate meg<Isporogenes i s, the mature
embryo sac "'ith egg, 2 synergids, J antipodals, and 2 polar
nuclei .
b . Oenothera type. Ilith 4-nucleate megaspor0genesis, the mature
embryo sac I·lith egg , 2 synergids , and a single polar nucleus .
2. Bisporic . Embryo sac formed from one of the t~lO dyad cells fonaed
after division I of me i osis .
a . Allium type . I.-lith 8 - nucleate megasporogenesis , the mature embryo
sac "'ith e gg , 2 synergids, 3 antipodals, and 2 polar nuclei.
3. Tetrasporic . Embryo sac formed by four spores resultin g from divi-
sion of the megas pore mother cell.
a . Peperomia type . 1hth 1 6-nucleate megasporogenesis, the mature
e mbryo sac "'ith egg , I synergid , 6 antipodals. and 8 polar
nuclei .
b . Penaea type . h'ith 16-nucleate megasporogenesis , the mature
embryo sac h'ith 4 "triads," one at each cardinal compass
point around the periphery , a nd 4 polar nuclei; each triad
consisting of an egg cell and 2 synergid~, only the egg cell
of the micropylar "triad" being fu n ctional.
c . Drusa type . Hith l6-nucleate megasporogenesis , the mature
embryo sac I,' ith egg, 2 synergid,s , 11 antipodals , and 2 polar
nuclei.
d . Fritillaria type . I'lith 8-nucleate megasporogenesis, the mature
embryo sac "'ith egg , 2 syn e rgids , J triploid antipodals,
and 2 polar nuclei, one haploid and one triploid .
e . Plumbagella type . l'lith 8 -nucleate megasporogenes i s , the I!lature
embryo sac Hith egg , 1 t riploid antipodal, and 2 polar
nucle i, one haploid and one triploid .
f. Plumbago t ype . I·lith 8-nucleate megasporogenesis , the mature
embryo sac Ivith egg , 4 polar nuc l e i, and 3 peripheral
nuclei , one at the chalazal end and one to each side .
g , Adoxa type . Hith 8 - nucleate megasporogenesis , t he mature embryo
sac Hith egg , 2 synergids , J antipodals, and 2 polar nuclei.

H. Fo rm and Organizat;ion of the Nature Embryo Sac .

Var i able characte rs include shape , number and arrangement of t he
nu clei and cells, nature of synergids and antipodals, presence of starch
or other food re serve , and formation of embryo sac caeca .

I. Fertilization.
Variable characters of importance include path of entry of the pol-
len tube (porogamous. chalazogamous , mesogamous). interval bet~J een polli -
nation and fertilization, and branching of pollen tub e during grQl ..,th .

J. Endosperm (Post Fertilizat i on Development) ,

1. Natur e of endosperm forma t i on : (see Figure 9-2)
a. Nuclear . Endosperm formation in \.,rhich the initial divisions are
not accompanied by "'all formation.
b . Cellular. Endosperm in Hhich the first and most of the later
divis i ons are accompanied by ,.]all formation . I f cellular,
orientation of first \,'alls should be recorded.
c . Helobial. Endosperm in I,hich the f ir st division is accompanied
by \,all formation but subsequent divisions are free nuclear,
at least in the micropylar chamber .
Figur e 9-2 227

TYPES OF ENDOSPERM FORMATION

o
, o
\ 0

'-1

if;
,,-
r

~
NUCLEAR CEL LUL AR HE LO BIA L HAUSTOR IA

[Used with permission of Dr . Barbara Palser.]

228 9

2. Presence of ruminat e endosperm .

3. Presence, absence and extent of endosperm haustoria and manner of
their formatio n.
4. tlature of endosperm food reserve, and fate o f endosperm in mature
seed .
" ,

K. Embrvogeny Type (Post Fertilization Development) .

Variable characters include relation of proemhryonal cells to body
regions of the embryo, form and organization of the mature embryo (size,
straight or curved . degree of differentiation) . and presence or absence
of suspensor haustoria . Attention should also be given to the source
and structure of the seed coat and ",hether there is persistence of
nucellus in the seed or not (perisperm) . Johansen (1950) classified the
embryos of d icoty ledons into six principal types based on their develop-
ment. A taxonomic distribution of these embryonic types and a pertinent
discussion of the relative value of these terms to systematics is given
by '..!ardlah' (lqSs).
1. Crucifer (Onagrad) type. 1·lith the terminal cell of the proembryo
dividing by a longitudinal wall , the basal cell playing only a
minor part or none in subsequent embryo development .
2 . Asterad type . \-lith the terminal cell of the proembryo dividing by
a longitudinal \"all , the basal and teTI'linal cells both contri-
hut ing in a major Hay to suhsequent embryo development .
3. Solanad type. Hith the terminal cell of the proembryo dividing by
a transverse h'all, the basal cell usually forming a 2- or more-
celled suspensor but p laying no major part i n subsequent embryo
development .
4 . Caryophyllad type. h'ith the terminal cell of the proembryo divid-
in g by a transverse ,,'all , the basal cell undergoing no division
and playing no role in subsequent embryo development, the sus-
pensor, if any, derived from the terminal cell .
5. Chenopodiad type. Hith the terminal cell of the proembryo dividing
by a transverse Hall, the basal and terminal cells both contri-
butin g in il. major "'ay to subsequent embryo development .
6. Piperad type . The zygote divides by a "'all "'hich is either vertical
or so obliquely orientated that it may he assumed to be essen-
tially longitudinal.

L. Certain Abnormalities of Development (Parthenogenesis , Apogamy, Adven-

t i ve Embrvony , Polyembryony, etc . ) .
1. Apogamy . Development of a ne,,' embryo from a cell o f the gameto
phyte other than the egg.
2 . Adventive embryony . Embryo development from dip loid cells of the
nucellus or the integument .
3. Polyembryony . The occurrence of more than one embryo in a seed .

Embryological information has proved to be of systematic value at all

taxonomic levels from the species to class . One of the outstanding examples
of a taxon heing clearly defined embryologically is the Ericales. In no
other order do the follo", ing combination of characters occur : no endothecium;
glandular, multinucleate tapetum; pollen in tetrads; pollen two - celled; uni-
tegmic ; nucel lus thin, ephemeral; endothelium present; no parietal cells in
ovule; megagametophyte monosporic and 8-nucleate, broad at micropylar end and
narro'" at chalaza I end; fluted hollm" style; cellular endosperm, first 4 cells
in a linear rOI>'; micropylar and chalazal endosperm haustoria; single- layered
9 229

seed coat; seed wi th fleshy endosperm, st raight embryo; nectary present;

anthers introTse by inversion; anther dehiscence by "apical" slit or pore;
anthers frequently app e ndaged; single archesporial cell in ovule; and zygote
elongated before division. The failure of a plant to possess a majority of
these characteristics is reason enough to cast doubt on its ericalean affini -
ties.

ENBRYOLOGICAL GLOSSARY

An ther . Pollen p roducing portion Loculus . A pollen-con taining chamb er

of stamen . of the an ther .
Antipodal. One to many cells of Negaspore . A spore formed by meiosis
the mature embryo sac loca ted of the megasporocyte and producing
at the cha l aza l end . the megagametophyte (embryo sac) .
Archesporium. One to several hypo- ~Iic ropyle. Ho l e th r ough integument(s) .
dermal ce ll s in the ovule pri- Hic rospore. A spo re formed by meio-
mordium distinguished by their sis of a microsporocytc a nd pro-
lar ger size , larger nuclei, and ducing the microgametophyte .
dense cytoplasm. Nucellar Cap. A persistent group of
Ari!. The secondary outgro\o1th of thick-\~alled and l igni fied cells
an ovule which surrounds the over the apex of the embryo sac .
ovule in pos t- fertilization Nuce llus . Female sporangium ~vit hin
stages, Hhich is derived from the ovule.
various regions but commonly Obturator. A proliferation of tissue
the funiculus or raphe . in the form of a small protuber-
Arillode . A fleshy coat derived ance Hhich is gene rally of placen-
from the micropylar rim of the tal origin and ..hich grows into
outer integument . space between nucellus and the
Caruncle . An integumentary pro- integument or also between the
tuberance restricted to the ovule and the ovary ~wll; sug-
micropylar region of a seed gested function is to aid in
and derived from cells of the pollen tube conduction to the
outer integument (characteris- micropyle .
tic of euphorbiaceous seed) . Ovule . A megasporangium enclosed
Central Cell. The cell of the by an integumentary system ; an
embryo sac in which the polar embryonic seed .
nuclei reside . Perip lasmod ium . The mass resulting
Embryo Sac . Female gametophyte from di sintegrat ion and coales-
of flowering plants. cence of tapetal cells .
Endosperm . Tissue formed from the Perisperrn . Persistent nucellus in
fusion of the t\vO polar nuclei seed; believed to be a storage
and one of the sperm cells; tissue.
nutritive in function. Polar nucle us . One of usually tHO
Endothecium. The outermost wall nuclei of the angi osperm embryo
layer of an an ther, located sac that migrates to the center
beneath the epidermis . from the poles .
Hypostase. A gr oup of thick- Ruminate endosperm . A type of endo-
HaIled, lignified cells at the sperm which exhib its some degree
base of the nucellus belO\~ the of irregularity as uneveness in
sporogenous tissue. its contour accompanied by irregu-
Integument . An enc l osing envelope larity in the inner surface of the
of an ovule ; embryonic seed seed coat .
coat .

/
230 9

Synergid . A sterile cell associated Tapetum . The inne rmost parietal l<Ia11
lvith the egg in the angiosperm layer of an anther. nutritive in
embryo sac ; a part of the egg function .
apparatus .

EHBRYOLOGI CAL LITERATURE

Bjornstad , I . N. 1970. Comparative embryology of Asparagoidead - Polygona-

teae , Liliaceae . Nytt Nagasin for Botanikk 17 : 169-207 .
Davi s, G. L . 1 966. Systematic Embryology of the Angiosperms . John Hiley
and Sons, Inc . Ne\o,' York .
Johansen , D. A. 1950 . Plant Emb r yology . Chronical Botanica Company . Hal t-
ham .
Mahes\\lari. P . 1950 . An In troduct ion to the Emhryology of Angiosperms .
HcGral.,r-Hill. New York .
1964 . Embryology in relation to taxonomy . I n : H. B. Turrill
(ed .). Vistas in Botany I V. l'-lacl>lillan Company . Ne\·] York .
Recent Advances in the Embryology of Angiosperms . 1963 . P. NahesHari (ed.).
International Society of Plant Horphologists . Delhi .
Hardla\" , C. \oJ . 1 955 . EP.lbryogenesis in P lants. John Hiley and Sons , I nc .
NeH York.

A CLEARING-SQUASH TECHNIQUE FOR THE S~UDY OF OVULE AND

HEGAGAHETOPHYTE DEVELOPl>lENT IN ANGIOSPEIU1S*

The technique outlin~d here uas devised specifically to ~xtend investiga-

tion beyond the l imits imposed by traditional sect i on methods. KnOI"n as th~
4 1/2 clearing t~chnique , this procedure incorporates features of bo th clear-
ing and squash methods and involves phase contrast optics for examination of
cell structure in optical sections through ,,'hole ovules .

1. Fixat i on. Ovularies from flOloJers in all stages of anthesis should b e

f ixed for 24 hrs . in FPA50 (formalin, propionic acid, 50% ethanol,
5 :5 : 90), Allen's modified Bouin ' s fluid , OT Randoph ' s modified Navashin
fluid (Johansen, 1940). Other fixatives may prove quite suitable, but
some , such as the various Carnoy ' s fluids, have limited value .
2. Storage . F i xed ovularies should be transferred to 70% ethanol Hhere they
may be stored indefinitely .
3. Clear ing . Th~ Hhole ovulary or ovules dissected from the ovulary in 70%
ethano l should be placed in standard 4 1/2 clearing f l uid composed of
lactic acid (85%) , chlora l h yd rate, phenol (c r ystals) , clove oil, and
xylene (2:2:2:2: 1 , by I"eight) for 2-24 hrs . (Herr , 1971) . A concave
container such as the Radnoti planchet (Radnotic Glass Techno l ogy , Inc .,
2614 Seaman Avenue, El l>lonte, California 91733) is recommended for the
clearing proc ess (Figure 9 -3) . Cleared ovularies appear nearly transparent
under the dissecting mic roscope equipped I']ith transmitted li~ht. The
ovules should be dissected f rom the cleared ovularies and by means of
a Pasteur pipette should be transferred Hith a drop of the fluid to a
Raj slide (Herr, 1971, 1973a ; Smith , 1973) for microscopic examination.
4. The Raj Slide . Two cover g lasses (No . 0-2 , 12 or 18 mm. sq . ) are affixed
Hith permount o r balsam 1-1.5 em . apart on a standard mic r oscope slide
which is then placed on a Ha r ming table at 50°C for 3 days to insure

*Contributed by J . N. Herr , Jr ., University of So uth Carolina, Columbia .

9 231

adequate hardening . The drop of clearing fl uid I"ith several ovules is

placed in the center of the 1-1 . 5 em. space and a cover glass is then
placed over the preparation so to rest on the tHO mounted cover glasses
(Figure 9-4) . The support cover glasses must be of sufficient thickness
to eliminate pressure of the top cover on the ovules.

-,, ,,-
~I
,, ,,
,,
,, ,,,

TOP
2cm

Figure 9-3 . Cross-section o f

Radnoti planchet. 5 I DE

Figure 9-4 . The Raj slide .

5. Hicroscopic Examination . The preparation should be examined Hith phase

contrast optics. Ovules in all stages of anatropous or campylotropous
development tend to lie sideHay s; that is, ,·lith the long axis of the
nucellus paralle l to the sl ide . Dowmmrd focusing, therefore , reveals
ovute structure through a series of opt ical sagittal sections (Figure
9-5). By slight shifting of th e cover glass on the support mounts,
small ovules can be reoriented for opti cal sections in the f rontal or
transverse planes .
6. Squash Procedure. After brief exposure to the clearing fluid , the ovules
become quite f ragile. If, therefore , the cover glass is lightly and
repeatedly pressed with a dissecting n eedle midway bet\~een the support
covers, the cel l s of the ovule gradually become spread apart. This
squash procedure , performed under observation with a dissecting micro -
scope , does not disrupt the struc tural integrity of the i ndividual
cells . Furthermore, it provides a method for extending the observa-
tions to the detailed features of specific structures such as mega-
sporocytes , megaspores, and developing megagametophytes previously
identified within intact ovules (Figure 9-5) .
7. Alternative and Supplementary Procedures . The clearing process in some
cases may be greatl~ enhanced if the stored ovularies or ovules are
dehydrated or dehydr'ated and pretreated prior to application of the
clearing fluid. In addition, modifications o f the standard 4 1 /2 fluid
may produce an improved clearing image.
a . Dehydration . In some cases, dehydration of the stored ovularies or
ovules to absol ute ethanol (70%, 95%, 100% ethanol) is necessary
for best results .
232 9

'- -

Figure 9-5. A. Opitcal sagittal section of an ovule f rom Glycine max

Merr. Megasporocyte during meiosis , telophase I. X 660 . (Photograp h b y
Ralph A. Geo r ge) . B, Optical sagittal section of an ovule from Phaseolus
aureus Roxb . Dyad , both cells at prophase II . X 600. (Photograph by Glenda
P . George) . C, Optical sagittal section of an ovule fr om Phaseolus aureus
Roxb . Dyad during meiosis , te l ophase II . X 600 . (Photograph by Glenda P.
George) . D, Optical sagittal section of an ovu l e from Glycine max Herr .
Preparation partially squashed to separate nucellus from integuments . 4-
nucleate megagametophyte Hith characteristic central vacuole and small
chalazal vacuole . X 660 . (Photograph by Ralph A. George). E-G , T\.,to optical
sagi ttal sections and an optica l cross section of a single ovule fr om
Phaseolus aureus Roxb . All X 380 . (Photograph by Glenda P . George) . E, Dyad
in upper focal p l ane . F . Second adjacent dyad i n l ower focal plane . G, 1\"0
dyad members , one from each dyad , in optical cross section of the ovule .
234 9

b. Clearing Pretreatment. For some species . a pretreatment procedure is

required for s uccessful clearing. The follOl"i ng methods have been
applied to \.. hole ovularies and ovules from the genera indicated .

Lactic Acid (Oenothera, LudHigia, Cornus , and Rubus)

l. Dehydrate ovularies or ovules to ab Solute ethanol.
2 . Pass the material through a g raded series of absolute
ethanol-la ctic acid (3 :1 , 2 : 2 , 1:3) I-lith 10- 15 minutes
exposure to each fluid .
3 . Transfer the material to luctic acid (85%) and incubate in
an open container at 50°C fo r 24 hrs .
4 . Replace the lactic acid ,·lith the cl earing fluid at r oom
temperature .

Lactic Acid-Phenol-benzyl benzoate , 1 : 1 : 1 by 1 (Oenothera , Lu dl<ligia,

and eornus)
1. Dehydrate ovularies or ovules to absolute ethanol .
2 . Pass the material through a graded series of absolute ethanol-
pretreatment mixture (3 : 1, 2 : 2 , 1 : 3) Hith 10- 15 minut es
exposure to each fluid .
3 . Transfer the material to the pretreatment mixture and i ncu-
bate in a closed container at 60°C for 24 hrs .
4 . Replace the pretreatment mixture Idth the clearing fluid at
room temperature .

Potassium Hydroxide (Cornus; Smith, 1973)

1. Transfer ovules from 70r. ethanol t o a 10% potassium hydroxide
solution for tHO minu t es .
2 . l~ash ovu l es in four changes of Hater .
3. Deh ydrate ovules to 95% ethanol.
4 . Replace e thanol 1·li th clear i n g fluid .
c. Alternative Cl earing Hixtures . Several modification of the standard
4 1/2 clearing fluid have applications discussed in detail else-
",here as indicated be l oH .

IKI- 4 1/2 (Herr, 1972)

Add 100 mg . iodine and 500 mg. potas sium iodide to 9 gm . of the
s tandard mixture . IKI-4 1 /2 enhances t he phase contrast image
and permits detection of accumulated starch.
PP-4 1/2 (flerr, 1973a)
Dissolve 3 mg . pota ssium permanganate in a gr am of the standard
mixture . Potassium permanganate-4 1/2 is some'<lhat unstable and
should be mixed for immediate use .
RS-4 1/2 (lierr, 1973b)
Add one part benzyl benzoate by weigh t to the standard mi xtur e .
PPflS-4 1 /2 (Herr , 1973)
Dissolve 3 mg . pota ssium permangana te in a g ram of BB-4 1/2 .
Potassium permanganate, benzyl benzoate-4 1/2 should be p re-
pa r ed for immediate use . Contrast bet\~een nuc l ei and cytopl asm
and delineation of cell layers are increased "'i th use of pp-4 1/2 ,
BB-4 1/2, and PPBB-4 1/2 beyond the leve l achieved with the stan-
dard mixture .

Pretreatment procedures and alternative clearing agents essential fo r

success in some plants produce adverse r esults in others . For example , all
9 235

pretreatment procedures listed here adversely affect the clearing image of

all legumes investigated (l'haseolus, Glycine, and Cassia) . For these genera ,
hm,lever , 8B -4 1/2 and PPHIl 4 1/2 are super ior to the standard clearing mix-
ture .

The origi nal method should be used initially in an investigation with the
alternatives and supplements incorporated later as needed . Each modification
of the original technique must be assessed for its improved or deleterious
effect on the clearing image .

Summary

Ovules or ~}ho1e ovularies to be cleared in standard 4 1/2 clear i ng fluid

or one of its modifications should be processed by the follm.,ring steps :

1. Fix material in FPA50 , Al l en ' s modified Bouin ' s fluid , or Randolph ' s modi-
fied Navashin fluid (Johansen , 1940 ) .
2. Sto re inde f init ely in 70% ethanol.
a . Dehydrate to absolute ethanol.
b. Pretreat : lactic acid at 60°C for 24 hrs.;
c. Lactic acid - phenol-benzyl benzoate (1:1 :1 hy ,,'e i ght) at 60°C for
24 hrs . (closed container); or
d . 10% KOH for 2 minutes . h1ater ; dehydrate to 95% ethanol.
3. Clear in standard 4 1/2 clearing fluid (lactic acid, chloral hydrate,
phenol crystals , clove oil , and xy len e ; 2 : 2 : 2 : 2:1 , by we i ght) .
a. IKI-4 1/2 (100 mg . iodine, 500 mg . potassium iodide to 9 gm . 4 1 /2) .
b . PP-4 1/2 (3 mg. potassium permanganate to 1 gm . 4 1/2) .
c. BB-4 1/2 (one part by I'Jeight benzl benzoate to 4 1/2) .
4. Mount ovules beneath the supported cover glass of a Raj slide and examine
in optical section l.,rith phase contrast optics.

OVULE CLEARING TECHNIQUE LITERATURE

Herr , J . M., Jr . 1971. A neH clearing-squash technique for the study of

ovule development in angiosperms . American Journal of Botany 58(8 ), 785 -
790.
1972. Applications of a new clearing technique for the investiga-
tion of vascular plant morphology . Journal of the Elisha Nitchell Scienti-
fic Society 88(3) : 137-143.
1973a . The use of Nomarski interference mic ro scopy for the study
of structural features in cleared ovules . Journal of the Society for
Advancement of Botany 1 : in press .
1973b . The cytological effects of several fixatives on the ovules
of Cassia abbreviata Oliver var . granitica Bak . NeH Botanist 1 : in press .
Johansen, D. A. 1940 . Plant Nicrotechnique . NcGraH-Hill Company . Nel,·
Yo rk and London .
Smith, B. B. 1973 . The use of a nel~ clearing technique for the study of
early ovule development, megasporogenesis , and mep,agametogenesis in five
species of Cornus L : .....American Journal of Botany 60(4) : 322-338 .
236 9

El'ffiRYOLOGICAL EXERCISE

From material on display. or preferably from specimens that have been

fixed, sectioned or cleared by the student , dete rmine the fo llmving selected
character states . Ideally, a sufficient number of floral buds should be
available so an assessment of different fixing and st?ining procedures can
be made. Accurate illustrations of embryological featu res a r e required . It
should also be remembered that a number of floral buds in various stages of
development may have to be prepared in order to give a complete embryological
description of the plant. In v ie\-l of the general paucity of accurate embryo-
logical information fo r angiosperms as a \"h0 1e, any information that can be
documented is potentially valuable and should be recorded , even though it
appears incomplete .

Spec.ies 1 Species 2

I
Hethod of Specimen Prepa ration

A. Fixative

B. Stain

c. Clearing

Character State

1. Anther

a. Endothecium Type

b. Tapetum Type

c. Type of Hall Formation

2. Ovule Type

3. Form of fluc.ellus

4. Type of Embryo Sac Development

5. Type of Endosperm Formation

6. Embryogeny

7. Abnormalities if any
 Chapter 1 0 . CYTOLOGICAL EVIDENCE*

Cytology , broadly defined , incl ude s the study of any and all charac teri s-
tics of cells and thu s embraces both mo r pho logica l and physiological wo rk.
Even the morpho l ogical aspects of cytology are eno r mous ly complex, fo r they
includ e investigations of the structure of organelles and of cells and their
contents both living and nonliving . The degree to I.hich these structural fea-
tures are faithfully represented in dead or pres erved cells depend s upon the
nature of the structure in the original living system and upon the particu lar
treatments which are employe d to preserve them fo r study . Tradit ionally , cyto-
logical studies have been made I-lit h the aid of the compound l ight microscope ;
more recently the electron microscope has provided an increasingly usef ul
means of examining the minute details of cells and organelle structure.
Clearly, the structural aspects of general cytology are closely related to
and inseparabl e from the cell ular aspects of anatomy.

The special branch of cy tology I .. hich deals with the nu cleus of the ce l l
and the hereditary material it contains, the chromosomes, is more accurately
designated as ka r yol ogy . Ever since the essential co rrec tness of the nine-
teenth century chromosome theory of heredity \..as experimen tally demonstrated
in the early decades of this century , karyology has gained i ncreasing impo r-
tance af> an aid in taxonomic \wrk . Karyology deal s Idth the chromosomes of
both plants and animals . Except for details which are peculiar to the repro-
ductive processes in each kingdom, the princ i ples of chromosome o r ganizat ion
and behavior are the same in al l eucaryotic organisms. Certain chromosome
phenomena ; e . g . , polyploidy, are more freq uent and better known in plants ;
others, e . g., duplication and inversion, have been more precisely analyzed
in animals. Fo r comprehensive treatments of chromosome cytology, \~ith illus-
trations and examples from e xtensive literat ure, and fo r clarification of
many details and discussions of controversial issues, the reader may be
referred to f>everal excellent books by distinguished studen ts of chromosomes
all of whom also happen to have been professiona l botanists (Darlington
1963, 1965; Sharp 1934, 1943; Stebbins 1971; Swanson 1957) .

This chap ter is designed to accomplish tlVO things:

(1) To provide a brief outline of karyological concepts and princip les

with emphasis on those aspec ts which have proved to be most helpful in plant
sys tematics .

(2) To outline the basic rational e and methodology for practical c hromo-
some v/Ork in the laboratory for those who wish to add the karyological approach
to their multidisciplinary attack upon the problems of vascular plant taxonomy.

The chromosomes are the physical bodies in t he ce ll which contain most but
not quite all (as modern investigations of organelles show) of th e essen tial
hered itary mate ria l, the ' DNA . This is reason enough to attach special impor-
tance to t he study of ch r omosoI:les , their numbers , form, i n ternal str ucture ,
and behavior . It also led the earlier cytotaxonomists to assume that the
characters of the chromosomes would be more reliable guides to taxonomic and

*Contributed by Ben 111 . Smith , North Carolina State Unive rsity, Raleigh .
238 10

evo l utionary relationsh i ps than many of the traditional morphological charac-

ters . This view is still \.,ridely held among chromosome cytologists and cyto-
geneticists, but i t has been categorically rejected by a number of highly com-
petent taxonomists. They maintain that the characters of the chromosomes
should be given no more (and no less) Height than any other characters \"hich
may be available for study. -.

Each of these interpretations appears to be correct in particular cases,

but ne ith er can be accepted as a general evaluat i on of the usefu l ness of
chromosome studies in resolving taxonomic quest i ons. In various genera and
sometimes higher groups , chromosome numbers, size, and form aTe monotonously
similar from species to species even though it is obvious f rom other charac-
ters of the plants that considerable evolutionary divergence has occurred .
On the other hand , there are groups such as Asplenium in temperate North
America or the Euras i an Triticum, Aegilops , Secale complex , for example , in
\~hich chromosome studies. especially Hhen combined Hith hybridization and
genetic analysis , have provided the essential clues for tracing the evolu-
tionary history of the group. Cytogenetics provides a framework in such cases
upon which the taxono~ist reevaluates the morphological evi dence of relation-
ship among the various populations of the group . Davis and HeYlvood (1963,
Chapter 6) present an excellent summary and discussion of this and other
issues in cytotaxonomy .

There is no r eliable Hay in which the relat ive usefulness of chromosome

studies in karyologically and genetically unknmm material can be pred icted
with assurance . It is necessary to examine the chromosomes . Since only a
small proportion of the total number of species among the angiosperms have
been examined , and some of the early work needs to be repeated and documented
by more reliable methods, much remains to be done . It is true, however, that
irregularities in reproduction such as infertility and ster i lity, and unusual
variation patterns such as extreme uniformity in progeny which suggests some
form of apomixis , or in contrast, unusual transgressive variants in segrega-
tion \~hich may result from irregularities in chromosome behavior often lead
the cytologist to interesting discoveries .

Section A. KARYOLOGY IN THEORY

A feH general cytological terms are defined as wel l as certain cytologi-

cal-morphological concepts which characterize the cytology of the reproductive
process in higher plants . The descriptive terminology of nuclear and chromo-
some cytology, arranged here in broad categories and \~hen possible in develop-
mental sequences, provides a conceptual outline of the karyological principles
which have developed in chromosome work. To obtain a brief characterization
of a given topic , each o f the follOt~ing subdivisions should be read in the
sequence presented . I f a novice in the subject finds that th i s raises as many
questions as it anSI"ers, he should refer at once to one of the book- length
t reatments listed in the General References at the end of this chapter. These
books also provide many detailed examples of the use of karyological results
in evolutionary and systematic studies . The material presented here also may
be utilized as a glossary by re fer ring to the general index.

I. CYTOLOGY

Cell (cyt- , - cyte). Unit of protoplasm , usually containing a single nucleus

and cytoplasm bounded by a plasma membrane and in plants usually surrounded
by a cell wall.
10 239

Cytokinesis (cell div i sion , cell reproduction). Process by \.,rhich a sin g l e

cell divides i n to two daughter cells a nd thus reproduces itself . Distinct
from but often synchronous with nuclear reproduction .
Spindle (achromatic figure). Fibrous structu re formed in the cytoplasm and
extending through the n uc lear region . Ac tive i n chromosome movement du ring
nuc l ear division and in cy tok inesis .
Ph ragmop l ast . Ex t ension of fibrous spindle-like structur e to the periphery
of the cell , active in cell plate formation .
Cel l plate . Ne\"l y formed cel l membrane transverse to the spind l e , fi r st
ob served stage i s a thickening o f the sp indle fibers .
Cytoplasm. Th e living extra-nuclear portion of the cell including its various
organelles and membranes .
Plasma membrane . The membrane ,... hich bounds the cytop l asm of the ce l l and in
plant s lies just ,... Hhin t he cell ....all.
Ce ll wall. The non-living bounding materia l .... hich is laid dm,ffi (secreted)
outside the plasma membrane and between adjacent cells . The primary t~a ll
in plants is u s ually cellulose ; other s ubs t ance s are often added which may
be var iously thicken ed , sculptur ed , or pit ted in the cells of different
t issues . Some cel l s characteri stically pr oduce a secondary ,,)'alL
Or ganelle . An y of various o rgani zed st ructures .... i thin the cel l which have
specific morphologies and functions such as plastids, mi tochondria , ch romo-
somes, nucleoli.
Heiocyte . A cell that is dest ined to undergo the process of meiosis.
Syn gamy (f erti liz a tion). The process of cellular and nu c lear fusion of the
two gametes (egg and sperm) which is one of th e two essential processes of
sexual reproduction (cf . meiosis) .
Zygote . Th e sin gle cell which re s ult s from syngamy-- t he fe rti lized egg .
Ar ti fact . An observed st ructure or apparent condition which is not a true
characteristic of the living specimen , b ut is t he resul t of post mortem
changes . cytological proce ssing techniques , or limitations of the equi pment
and the observer. Sharp distinct ions bet\o,'een artifacts and "r eality " are
oft en difficult in cytology and require good judgment as well as expe r i ence.

CYTOLOGY (Botanical Terms)

Sporocyte . A cel l which by the process of meiosis is dest ined to produc e a

set of fo ur haploid spo r es .
Hicrosporocyte (pollen mo the r cell ) . A spo r ocyte wh ich by meios is (micro -
sporogenesis) produces the micro spores o r "male spores . " The se occur i n
the anthers of seed plants .
Hegasporocyte . A sporocyte \... hich by meiosis ( megasporogenes i s) p rod uces the
usually la rger megaspores o r "female spores . " These occ ur often singl y
and mor e o r less embedded in the nucellar ti ss ue of the ovu l e in seed
plant s .
Heiospore . General t e rm for any spo re prod uced by the process of meiosis .
Secondary sporo cyte dyad . The pair of cells produced by the first division
of meiosis in tho se forms in wh i ch cell division occurs synchronous l y with
th e first meiot i c division.
Spore t e trad . The fo ur meiospor es produced f r om a single prima r y sporocy te
by meiosis . Usually applie d t.o s t ages at which t he four s po res r emain in
contac t. .
Microspore . Produ ct of mic rospo r ogenesis--"male spore . "
Hega spor e . Produc t o f megaspor ogen esis-- "female spore ."
240 10

Nicrospore mitosis (micr ospore division). Applied to the first mitosis l.;hich
occurs in the haploid mic:rospore following its fo rmation by meiosis .
Pollen grain . The male gametophyte, applied to the stages including and
follmving the microspore unti l pollen germination .
Vegetat ive (tube) cell (tub e n u cleus) . Cell ,.,rith its haploid nucleus t"hich
constitutes the bulk of the developing male gametopfly-te in seed plants .
In angiosperms, one of the products of the microspore division .
Generative cell. Cell enclosed by its membran e within the cytoplasm of the
vegetative cell \.,rhich is destined to produce the tt.JO male gametes . In
angiosperms, the other product of the microspore division .
Generative divis i on. Division of the generative cell to produce the male
gametes . Nay occur before pollen germination (3 - nucleate pollen grains),
or in the pollen tube (2-nucleate pollen grains) .
Hale gametes (sperm). Haploid sperm cells each Hith its membrane and cyto-
plasm .
Hegagametogenesis . Developmental seq uence in the female gametophyte iVhich
leads to production of the female gametes (egg cells) in the archegonium
or in the embryo sac of angiosperms .
Embryo sac. The female gametophyte of angiosperms.
Gametophyte. The haploid phase of the life cycle beginning I.;ith the meio-
spore, ending with production of the gametes.
Sporophyte. Diploid phase of the life cycle beginning with the zygote , ending
with haploid spore production by meiosis.
Alternation of generations . Alternation of diploid (2.!}) sporophyte and hap-
loid (~) gametophyte generations in the life cycles of eucaryotic plants .

II . KARYOLOGY

Nucleus (karyo-, caryo-) . Conspicuous spheroidal or ellipsoidal body within

the protoplast of a cell , enclosed by a membrane and containing the chromo -
somes (chromatin), a nucleolus, and nucleoplasm or karyolymph .
Nuclear membran e . The membrane iVhich bounds the nucleus of the cel l except
during the metaphase and anaphase stages of nuclear reproduction .
Nucleolus . Dense body within the metabolic nucleus largely consisting of RNA
which stains diff erentially . (Nucleoli usually a pp ear in telophase, dis-
appear during prophase.)
Eucaryote. An organism the cells of Hhich conta in organized nuclei \.;ith
nuclear membranes and chromosomes .
Karyokinesis. The process of nuclear divisio n or nuclear r eproduction .
Hitosis . General term for the process of nuclear reproduction \.;ith particu-
lar reference to the reproduction of the chromo somes . Hitosis in somatic
cells is used in contrast to meiosis in germ cells or sporocytes .
Prophase. The early stages of mitosis f rom the first detectable changes in
the organization of the chromatin as seen in the metabolic (resting)
nucleus to the congression of the contracted chromosomes at the equatorial
plate .
Equatorial plate . Stage in mitosis at which the contracted chromosomes come
to be arranged across the equator of the spindle wi th the centromeres of
each chromosome lying in a single plane.
Metaphase. Literally "betl,Icen the phases ," originally meaning bet\.;een pro-
phase and anaphase . By usage metaphase has become the equivalent of the
equatorial plate stage, the moment in mitosis Hhen the chromosomes are
arranged most nearly in a single plane just before anaphase disjunction
begins .
10 241

Anaphase . The stages of mitosis during which the daughter chromosomes sepa-
rate (disjoin) and move apart from the equatorial plate to the two po le s
of the spindle .
Telophase. Stages of mitosis during ,,]hich each of the newly formed daughter
nuclei become s enclosed in a new nuclear membrane, the contraction of the
chromosomes re l axes, ne\-1 nucleoli are formed, and even tual l y the chromatin
" reticulum" characteris tic of the metabolic n u c leus reappears .
Hetaholic (res t ing) phase . The condition of t he nucleus in a metabolic cell,
a nucleus which is not involved in active mitosis. The contraction of
the chromosome s has relaxed so that they form a fi n e ly divided mass of
very fine strands which often appear as a granular or ret iculate "net\wrk."
The lin ea r integrity of the i ndividual chromosomes is maintained , however,
a nd the apparen t ly anastomosing " network" is an artifact.
Neiosis. Form of nuclear reproduction in \.,.hi ch the chromosomes divide only
once while the nuclei divide twice p r oduc i ng four nuclei \.,.ith the reduced
(haploid) number of chromosomes; one of the two essentia l processes of
sexual reproduction (cf ., syngamy) .
Heterotypic division. First of the two succes sive meiotic divisions .
Leptotene (leptonema) . Earliest stage of meiotic prophase . Ind ividua l chromo-
somes are extremely long, fine threads .
Zygotene. Pairin g s t age of meiotic prophase . Homologous t hread-like chromo-
somes become int imately associated throughout their lengths .
Synizesis . "Knotted thread stage" often associated ~1ith early meiotic pro-
phase. The fi ne threaded nucleus is seen collap sed i nto a tangled mass .
This is an artifact of fixation , but represents a phys iological condi tion
in which the fin e structural details of th e living system are most suscep-
tible to distortion.
Pachytene. Thi ck t hread stage of meiotic p r ophase . Each of the complete l y
paired homolo gous chromosomes has also r ep li cated , the chromosomes a r e
four - partite .
Diplotene. The paired chromosomes separat e from each other ; a looped
structure result s because the chromosomes are held toge ther at the
chiasmata .
Di akinesis. Final s tage of meiotic prophase, t he chromosome pair s are much
contracted, but are sti ll held together by terminalized chiasmata.
He t aphase I (N I). Netaphase of the first meiotic division . Chromosomes
become cooriented at the equatorial plate with opposing centromeres on
opposite sides of the equa tori al plane .
Anaphase I (A I) . Anaphase of the first meiotic division . Chromosome pairs
diSj oin , chiasmata complete terminalization , chromosomes move to the poles .
Haximum ch r omosome cont r action .
Te l ophase I (T I) . Telophase o f the first meiotic division . Relaxation of
chromosome coiling, fonnation of nuc l ear membranes.
Interkinesis . The "resting" stage \.,.hich occurs between the two meiotic divi-
sions . In many forms there is no telophase I , interphase, prophase II
sequence; instead metaphas e II occurs immediately after t he chromosomes
r each the poles in anaphase I .
Homeotypic divis i on . The second meiotic division, so named beca use it is
" similar" to a mi t otic division.
Prophase II. Prophase of"" the second meio tic division . Chr omosomes are dyads
held together at the centromeres with the arms lying f ree .
Hetapha se II (H II). Hetaphase of the second meiot ic division . The centro-
mer es become oriented at the equatorial plate and disjoin.
242 10

Anaphase II (A II). Anaphase of the second meiotic division. Individual

chromosomes separate and move to the poles.
Telophase II (T II) . Completion of meiosis, r e laxation of chromosome coiling ,
appearance of nuclear membranes around the four haploid nuclei. The spore
tetrad results from the completion of cytokinesis . .......
Chromosome complement . The total haploid, or diploid group of chromosomes
contained within a particular cell, or ind i vidual.
Karyotype. The mitotic metaphase morphology of any given chromosome comple -
ment of an individual , race, or species .
Idiogram. Diagranunatic arrangement of photographs, dra\.Jings, or graphical rep-
resentations of all the individual chromosomes of the karyotype to show the
principal morphological features . The chromosomes are usually arranged in
a horizontal rm., of homologous pairs in order according to length and/or
centromere position.
Cytogenetics . I nvestigations \'/hich combine and synthesize the methods and
results of chromosome cytology \-lith the genetic methods and r esu lts of
breeding analysis.
Cytotaxonomy . The investigation of chromosomal relationships in populations,
races, and species in order to e lucidate taxonomic and evolutionary rela-
tionships .
Cytogeography. The geographical distribution of chromosomal differences
between various populations in relation to taxonomy and evolution .

III . C!lROHOSOHE - ABERRATIONS, REARRA.~GEHENTS

Deletion (deficiency) . The loss or elimination of a microscopically observ-

able segment of a chromosome.
Duplication . The presence of a chromosome segment which duplicates a portion
of a chromosome Hithin the same haploid chromosome complement . Duplica-
tions may occur in tandem within the same chromosome .
Inversion . Segment of a chromosome in Hhich the order of the structural ele-
ments (and included genetic materia l ) is in reverse to the order found in
otherwise homologous chromosomes.
Pericentric inversion . The i nverted segment includes the centromere region .
Paracentric inversion . The inverted segment lies entirely ,vithin a single
chromosome arm.
Inversion bridge and fragment. Characteristic anaphase configuration found in
meiosis fo llm.,ing crossing-over in a heterozygous paracentric inversion .
One of the result i ng chromosomes becomes dicentric and fo rms a bridge as
the centromeres migrate tOHard the poles at anaphase. The other cross-over
product is an acentric fragment . The relative position of the point of
crossing- over to other cross-overs determines Hhether the bridge and frag-
ment appear at the first or second anaphase (see literature ).
Dicentric chromosome. Aberrant chromosome containing t\.JO centromeres .
Translocation . Re ciprocal exchange of segments be tHeen two non-homologous
chromosomes .
Translocation ring. Configuration of 4 chromosomes formed by the meiotic
pairing of homologous segments in a heterozygo us translocation. Additional
heterozygous trans locat ions in the same complement produce additional or
larger rings.
Alternate disjunction. Centromeres of adjacent members of a translocation
ring proceed to opposite poles at anaphase I ; produces zigzag configura-
tions and balanced assortment of chromosomes.
10 243

Adjacent disjunction . Adjacent members of th e ring proceed to the same pole

at anaphase I; produce an open configuration and unbalanced ass ortme nt
with duplicated and deficient segments in both daughter nucl ei.
Structural change . General term for the various rearrangements of chromosome
segments I.;hich result from chromosome aberrations .
Structural hybridity . Hetero zygosi ty of the chromosome complement for one or
more structural changes.
Cryptic structural hybr idity . Heterozygosity of the chromosome complement
for one or more minute and microscopically undetectable struct ural changes
which presumably account for marked reductions in pairing between chromo-
somes from different races or species.
Isochromosome. A ch romosome result ing from transverse centromere misdivision
in which the two arms are identical although arranged in inverted linear
order vith respect to each other.
Chromosome fragmen t . A broken portion of a norma l chromosome.
Acentric. No centromere is present , is usually lost after one or a fev
fe\" mitotic cycles .
Centric . Cent romere i s present, may be irregularl y or regularly trans-
mitted in mitosis, is much more erratic in behavior and subject to
elimination in meiosis .
Ring chromosome . Usually ar~s~ng as a centric spontan eous or induced ch romo-
some fragment in wh ich the broken ends have r ejoine d i n a ring configura-
tion .

IV . CIIROHOSONE - HEIOTIC BEHAVIOR

Pairing . The detailed chromomere by chromomere association of homologous

chromosome threads '-lhich occurs at zygotene. Commonly also app l ied to
the continued association of homologous chromosomes obse rved in diplotene,
diakin esis , and metaphase I I"hich results from the earlier zygotene
pairing.
Synapsis . Process of chromosome pairing .
Asynapsis. Failure of the process of chromosome pairing (in presumably
homologous chromosomes).
Desynapsis . Precocious separation of previously paired chromosomes before
completion of meiotic p rophases .
Homologous . Applied to chromosomes and to segments of chromosomes ,,,hich have
the s ame chromosome ance st ry and thus contain essentially the same genetic
material.
Non-homologous . Applied to chromosomes and to segmen t s of chromosomes which
have different chromosomal ancestry and thus differ in genetic cont e nt and
arrangement.
1I0meologous. Applied to chromosomes in polyploids which are simi l ar in ori-
gin (probably onc e homologous or partially so) but which have become so
differentiated that th ey pair incompletely or not at all.
Chiasma(ta) (crossing over). Chiasma describes the cross shaped configuration
\-lhich is the ob served re sult of the eXchange of segments between homologous
chromosomes during metotic prophase . Crossing-over is the genet i c term for
the actual exchange of genetic determiners .
Chromatid. A s ingl e one of the two daughter chromosome st rand s following
chromosome reduplication in mitosis or meiosis while they r emain more or
l ess coiled about each other.
244 10

Sister chromatids. The two chromatids produced by the replication of a single

original chromosome .
Tetrad chromosome . The four chromatids of a pair of homol ogous chromosomes ,
each of which consists of two sister chromatids (late pachytene to M I).
Dyad . The individual chromosome consisting of tHO chromatids at Anaphase I
and metaphase II. .......
Terminalization (of chiasmata). Process occurring \~hen the chromosomes pro-
gressively separate as the centromeres move a\~ay from each other . This
causes the relational coils of the t\~O chromosomes to um~ind at the ends .
The nodes of the loops which originally occurred at the points of actual
exchange appear to move to\~ard the ends of the chromosomes and eventually
disappear .
Congression . Movement of the chromosomes to the equatorial region of the
cell just before metaphase .
Orientation. Arrangement of the centromeres of mitotic chromosomes so that
each lies in the plane of the equatorial plate .
Coorientation . Arrangement o f the tHO centromeres of each meiotic chromo-
some pair so that each pair is oriented Hith the cent r omeres opposite
each other and equidistant from the equatorial plane. Hare complex
coorientations result from associations of three or more chromosomes at
H I.
Disjunction . Separation of the chromosomes at anaphase .
Univalent. One unpaired chromosome in meiosis.
Bivalent. A pair of homologous chromosomes in meiosis .
Trivalent. An association of three homologous or partial ly homologous chro-
mosomes (e.g ., in triploids or trisomics) .
Quadrivalent . An association of four homologous or partially homologous
chromosomes .
Hexavalent. An association of six .
Hultivalent. An association of three or more partially paired chromosomes in
meiosis . Note: no matter hOl~ many chromosomes are involved in a multi-
valent complex. at anyone point intimate association is ah~ays that of
one to one pairing .
Autosyndesis . I n polyploids and their hybrids. the pairing of chromosomes
derived from the same parent . In A1AlA2A2 x AJAJA4A4 . pairing is AlA2
or AJA4.
Allosyndesis . In polyploids and their hybrids, the pairing of chromosomes
derived from different (opposite) parents . In AlAI BIB I x A3A3B3B3, pair-
ing is AlA3 and BlB3 .

V. CHRONOSOHE - HORPHOLOGY

Chromosome. Thread-like strand of DNA and associated nucleoproteins \~hich

replicates itself i n each complete mitotic or meiotic cycle and Hhich by
coiling appears to contract to a usually rod-like or in the smallest
chromosomes to an almost spherical form . Chromosomes and their consti-
tuent material , often called chromatin , stain deeply and differentially
in basic dyes.
Chromomere. Individual minute. more or less spherical . bead like body in the
chromosome thread .
Centromere (kinetochore, primary constriction) . A speciali zed chromomere to
\~hich the chromosomal fibers of the spindle appear attached. Hhich ini-
tiates the chromosome movement at diplotene of meiosis and as anaphase
begins . In many preparations the centromere is invisib l e and is represented
10 245

in a fully contracted chromosome by a (primary) constriction . The rela-

tive position of the centromere with respect to th e t,.,,-o chromo1:>ome arms
is one of the most reliable morphological features of the chromosome.
Secondary constriction . A constricted region ,,,ithio one of the chromosome
arms.
Satellite . A small terminal portion of a chromosome separated from the
remainder of its chromosome arm by a prominent, (often long) secondary
constriction. Often associated with a nucleolus organi zer.
Chromosome arm . The portion of a chromosome to one side or the other of the
centromere (or primary constriction). The proximal region is adjacent to
or in the direction of the centromere in contrast to the distal or terminal
region of the chromosome arm.
Metacentric (median centromere, V) . A chromosome with a median centromere has
equal arms and is V-shaped at anaphase .
Isobrachia l. Equal-a rmed chromosome .
Submetacentric (submedian centromere) . Chromosome \~ith i ts centromere near
but not exactly at the median point .
Acrocentric (subte r minal centromere , J or I ) . Chromosome with the centromere
near one end, thus ,.;rith one long and one short arm such chromosomes are J-
or I-shaped.
Heterobrachial. Any chromosome with arms of unequal length.
Telocentric . The centromere is at the end of the chromosome . Such chromosomes
are r are, most I-shaped chromosomes are acrocentric Hith a minute short arm .

VI . CHROMQSOHE - STRUCTURE, CONTENT

Chromatin . The deeply staining material (DNA and nucleoprotein) of the chro-
mosomes .
Achromatic . A region of the contracted chromosome which is not deeply staining
such as the centromere region or a secondary constriction .
Euchromatin . Those portions of the chromosomes (usually the majority) which
£0110\.;r the typical mitotic cycle of condensation by i ncreased spiralization
and consequently deeper staining through prophase to early anaphase, followed
by relaxation of coiling and progressively lighter staining which reaches its
10l"er limit in the resting stage; as opposed to heterochromatin .
Heterochromatin . Chromosome segments or chromosomes which remain contracted
and deeply staining or contract early in contrast to the behavior of the
euchromatin in the resting and prophase stages . Heterochromatin may also
despiralize and appear as lightly stained segments at metaphase I-Ihen euchro-
matin is most condensed . The genetic content or at least th e function of
the heterochromatic regions of chromosomes have b een shOlm to be qu i te dif-
ferent from that of euchromatin , and there appear to be a number of different
kinds of heterochromatin . Heterochromatic chromosomes and segments are us e -
ful in characterizing the morphology of particular c hromosomes and karyotypes .
Prochromosomes. Small condensed "chromosomes," actually heterochromatic seg-
ments of the whole chromosomes ,.;rhich appear in the rest i ng nuclei of many
plant species; the number of prochromosomes may correspond to the metaphase
chromosome number but accurate determinations are difficult .
Nucleolus organizer. Spe<;:J.fic chromomere or chromosome region associated Hith
the formation of a nu'cleolus, often but not alHays associated ,.;rith a long
secondary constriction and just proximal to a satellite.
Chromosome knob . Large chromomere, Hhich is more or less heterochromatic, \dth
a characterist i c morphology and constant position . They may be interstitial
or terminal and are best observed in early prophase .
 10

B-chromosome (accessory chromosome) . Supernumera r y chromosomes , that are in

addition to the normal diploid complement, which are primarily heterochro-
matic an d occur in varying n umb ers among different individ uals of a popu-
l ation . They may be minute (microchromosomes) or moderately large in com-
par i son \~ith the normal members of the complement.
Spi r a li zation . Process of i n ternal coiling in chromosomes \,"hich causes changes
i n appar ent chromosome length and staining inten~i~y during mitosis and
meiosis .
Allocycly . Differential cycles of coiling and condensation in different chro-
mosomes or i n different segments of the same chromosome \~hich lead to
obse r ved contrasts in the cytological appea r ance of heterochromatin and
euchromatin .
Autosomes . The typical chromosomes of the complement \~hose segregation does
not usually affect sex determination .
Sex chromosomes (allosomes) . Particular chromosomes, which may occur singly ,
as a more or less heteromorphic pair, or in multiple chromosome complexes,
whose segregation is related to sex determination .

VII . CHROHOSOME - NUMBER

Haploid (monoploid , ,!!) . One complete set of chromosomes , such as the comple-
ment in the nucleus of one of the gametes . The.!!. number of chromosomes in
the haploid", monoploid'" gametic number of chromosomes .
Dip l oid (2!:!). The chromosome complement of a zygote , two complete sets of
chromosomes one from each parental gamete - the zygotic number . The 2n
dipl oid chromosome number is the normal somatic chromosome number of any
organism which has arisen from a zygote , the organism itself is often
referred to as diploid .
Polyploid ( - ploid) . Any individual (or a cell) which contains three or more
complete sets of chromosomes. Various combinations of lvords or numbers
with " - p l o i d " indicate the number of haploid sets of chromosomes .
triplo i d'" 3n hexaplo i d'" 6g
tetraploid = 4~ octoploid '" 8~
pentaploid '" Sf!. lO-ploid '" I O,!!, etc .
Polyploidy (ploidy) . Variations in chromosome number involving more than the
di pl o i d number of complete chromosome sets .
Autopolyploid (autoploid) . A polyploid in which all of the chromosome sets
(genomes) have arisen from the same source, each chromosome is capable of
pairing (autosyndesis) \,lith a corresponding c hromosome in each of the
other sets. Easily applied to experimentally induced polyploids ; e . g . ,
a normal diploid is douhled to produce an auto tetraploid, th i s can then
be crossed with the origina l diploid to produce an autotriploid and so on .
The same events can and do occur in nature as the result of spontaneous
chromosome doubling. In natural populations , hO\~ever, both within and
between related species , every possible degree of s i milarity from essential
identity to marked genic and even structural divergence may have occurred
between the parents of the original diploid individual in \~hich the chromo-
some doubling occurred (see Stebbins , 1971 , and others) .
Allopo l yploid (alloploid) . A polyploid in which the original chromosome sets
(gen omes) have been derived from different sources as the result of hybridi-
za t ion a n d no e ff ective pairing OCCurs betHeen the chromosomes of the dif-
ferent genomes . In a strict allotetraploid only bivalents are formed at
meiosis and no multivalent associations .
Segmental allopolyploid. A polyploid in which part of the chromosome comple-
ment is essentially autopolyploid in origin , part is allopolyploid . One
 of the frequent a n d more easily recognized intermediate sit ua tions be t He en
class ical autopolyploidy and allopolyp l oidy .
Amphidip l oid (litera lly both dip l oid). The allotet r aploid formed by doub ling
the chromosomes of a steri l e Fl hybrid '''hich contains two haploid genomes
,"hich are incapab l e of effective pairing at meiosis . Ferti li ty is restored
after c hromosome doubling beca use each genome has become duplex , bi va l e n ts
a r e fo r med at me i osis , and ba l anced ch r omosome assortment e n sues .
Genome . A complete haploid se t of chromosomes; the haploid ch r omosome comple-
men t of a diploid race or species .
Euploid . A chromosome number "'hich is an exact multiple of a given haploid
numbe r .
Basic number'" x . The number of chromosomes in the o r iginal genome (haplo i d
set) from which a polyploid or a group of polyploid forms or species is
knOtm or postu l ated to have arisen .
Polyp l oid series (2~ , 3~ , 4~ , 6~ , ... ) . A group of r elated r aces or speci e s
,"ith various chromosome number s all of which are mUltiples o f a single ba sic
number ( e.g. , t h e e uploid s e r ies indicat ed) .
Homoplo i d series . A group o f r elated spec i es (e . g ., the species of Pinus , i n
a l l of them 2rr = 24) al l of ,... hich have the same ch r omosome number-.--
Dibasic po l yploidy . Al lopo l yploids ,... ith chromosome numbers ,. . hich result f rom
combinations of t wo genomcs \<Jith different basic numbers .
Aneuploid. Chromo some n umbe r s ,,,hich are no t exact mul tiples o f an original
haploid n umber, with one or more chromosomes missing (2rr - 1) or in addi-
tion (2!!. + 1) to t he e up l oid numbe r.
Aneup l oid ser i es . A series of ane uploid numbers in a group of individuals.
or species , each chromosome numher in the series differing f r om the next
by the add ition ( or subtrac t ion ) of one or two c hr omosomes (e . g . , 22!. ,
2x + 1 , 2x + 2 , 2x + 4 , 2x + 5 , etc . ) .
Dysploid . Diverse a~euploid~chromosome numbers '·Ihich do not fall into a
series o r regular pat t ern .
Honosomic . An individual in wh ich the diploid chromosome comp l ement lacks on e
membe r of one of the chromos ome pairs , 2!!. - 1. Thus one member of the com-
plemen t is monoploid or simplex . The r e are as many diffe rent monosomic
types possible as t he number of chromosome pairs .
Nul l isomic . Aneup l oid in \~hich both members of a chromosome pair are missing,
2n - 2 . This is not found at the diploid level , but can occur in poly-
ploids , in which t he gene t ic material of tbe miss i ng chromosomes may be
dupl i cat ed elsewhere in t he c hromosoJ:le complemen t so that no essen tia l
functions are lost .
Di somic . The normal diploid (2.!!.) con di t ion in which a given c hromosome is
represente d twice in the complement .
Trisomic . Aneuploid in which one of t he i ndividual chromosomes is presEon t in
trip licat e (rr + 1) . As wi th monosomics there are as many po s sible tr i s omic
types as there a r e differen t chromosomes in the haploid genome . A secon-
dary trisomic occ urs \~hen t he extra chr omosome is an isochromosome .
Tet r asomi c . An aneuploid in wh i ch a sing l e chromosome pair is duplicated
and thus is te t rapl oid, 2.!!. + 2 . This is more likely t o occur in polyp l oids .

Section B. KARYOLOGY IN l'RACTICE

-- I. CHRO}lOSONE t·mTHODS

The objective is to obtain first class microscop i c slide p r eparations of

chromos omes I~hic h may be obse rved, counted, measured , p\lOtogra phed , and o t he r -
wise stud ied 1·lith th e aid of the compound l ight microscope . Ea r ly inves tiga-
tion s '~ere based upon t he study of serial sections of appropriate meristemat i c
248 10

or sporogenous t issues which had been stained by one of t he haematoxylin pro-

cedures or \-lith an aniline dye, most often crystal violet . These techniques
are still available and valid for precise cytological investigations and when-
ever the faster and more direct squash (smear) methods do not yield satisfac-
tory results . The relatively laborious fixing , embedding , sectioning, stain-
ing procedures have a major advantag e : t.hey permit the use of numerous chemi-
cal reagents in various combinations (fixatives) in oraer to kill the living
sys t em very quickly a nd to preserve the desired features of its morphological
structure \dth a minimum of distortion (artifact) . The chief source of diffi-
culty and error i n t he method is the necessity of avoiding or possib l y of
reconstructing chromoso[fle configurations which have been cut and lie in mo re
than one section .

The aceto-carmine squash method I~as one of several such procedures in-
vented by the cytologist, John Belling, for the study of meiotic chromosomes in
the spo ro genous tissues of a variety of flo"ering plants. The basic method
has since been adapted to the study of mitotic chromosomes found in various
active l y grOt~in8 somatic meristems . A variety of stains have been substituted
for carmine by different workers and for different materials . Various combi-
nations of alcohol. acet ic acid (or propionic acid) and chloroform are employed
in the killing-fixing solution. In the acetic -orcein method , the sa turated
sol ution of orcein in 45% acetic acid is applied directly to the living root
tips and thus serves both as fixative and stain . The Feulgen nucleal staining
reaction I>'hen correctly car ried out is a specific stain for DNA . Nany workers
subs titute the Feulgen reaction for the orcein or carmine stain, or sometimes
combine it I~ith one of these stains in conjunction ,~ith the squash technique.
(Detailed proc edures and varia tions may be found in Darlington and LaCour ,
1962 , and in Sharma and Sharma . 1965 . )

The squash methods all involve spreading actively dividing cells on a

slide so t hat the cells occur loosely associated in groups in a layer on l y
one ce ll t hick . This is usually easy to accomplish with pollen mother cells
(PHC ' s) and male sporocytes in l a t er stages of meiosis because the cel l s of
the sporogenous tissue usually f l oat out separately when pressed f rom the
anther . To obtain the same sort of "mono-cellular" layer from somatic meri-
stem , a maceration process is necessary. This is usually accomplished by
placing the material (root tips , leaf buds, etc .) in IN HCl solution at 60°C
for 5-15 minutes. or by some other combination of heat and weak HCl . The
middle lamellae betl.een the cellulose cell walls a r e destroyed and the cellu-
lose itself is softened . The p r eparation is completed in each case by adding
the cover glass . after which gentle tapping fol l owed by firm p r essure is used
to flatten and spread the cells.

II . CHROHOSOHE HATERIALS

Chromosome number. size . and morphology are be st determined in somatic

(mitotic) material and may be observed in any veg eta t ive tissue in \~hich the
cells are actively dividing . Heristems from fast growing root tips are the
most convenient source "hen t he plants can be grown in culture . Root tips are
sometimes taken from recently germinated seeds , but this involves a serious
difficulty . There is no plant upon which to base identification of the species
or strain under study . Every thing depends upon the correctness of the label on
the seed packet which depends in turn upon care and accuracy of the collector
and nature of the original source . Furthermore , if different chromosome
resul t s are obtained from the root tips of different seedlings in the same lot .
 - -- - - - - - - - ----_. - -- - - - -

10 249

there is no way of associating t he chromosome differences ,... ith differ ent

plants . It is far better to gr ow plants in separa t e pots so that a one to
one c orrespondence between observations and individual p l a nts can be main-
tained . Later the plan ts may be used in cyt ogen etic and o t her experiments,
and they may become voucher herbarium spec imens Hhieh provide taxonomic docu-
mentation for the karyologieal results .

It is impracticable, oft en impossible . t o collect good root tips from a

known individual plant which is growing in it s natural hab it a t. Some othe r
tis sue must be employed if fi eld collected mat erial is necessary or desirable .
Active l y gr owing stem tips , early l eaf primordia, and i n special cases , youn g
petals. ovules , or other organs can all provid e good mito tic chromosome prepa-
rations . Herbarium spec i mens mu st be collected f r om the same plants or popu-
l ations.

Stud ies of the chromosomes i n meiosis are indispen sab l e for detennining
the presence and na ture of chromosome irregular ities . Heiosis in hybrid s
reveals the chromosomal causes of sterility Bnd the extent to which ch anges
in chromosome structure have occurred . In favorable material , e . g ., maize ,
Lilium, Solanum , detailed chromomere anal ysis at pachytene provides dire ct
morphological e v i den ce of st r uctural divergence . Analys i s of chromosome pair-
ing r elations at meiosis pennits the c l assification of the va r ious types of
polyploids , and in hybrids , provides the bas i s for deter mining the degree of
chromosome homology from which the evolutiona r y relationships of the parents
are inferred . Species vary greatly in the useful n ess of particular stages of
meio sis for de tailed study . The excellent pachy tene of Zea ~ is followed
by diakinesis and metaphase chromosomes which are so contrac t ed that structural
det ails are hidden. Diakines i s and metaphase are excell en t in many specie s
ldth relative l y l arge chromosomes , but the prec eding pachytene is so crowded
and visua lly complex that it defies a na l ys i s .

Norpho logical features of the indivi dual chromosomes are more difficult
to observe in the metaphase and anaphase s tages of meios is because the chromo-
somes are often so much cont racted that fea tures easily seen in mitosis are
obscure . The chromosomes of many species a r e small and r e l atively difficult
to anal yze even with the aid of the best oil immersion obj ec tives a t t he limit
of usefu l magn i fication. Even chromosome number determina tions based on
meiotic materia l are subject to error under these conditions unless the mat e-
rial is diploid with comple tely normal pairing and disjunction . In unknown
or cytological l y irreg ular material, these problems can be min imi z ed by corre-
lating the meiotic anal ysis with mitotic ob serva t ions .

The sporogenous cells of developing an t her s (mic r osporangia in gymno-

sperms) are t he usua l so ur ce of me i otic ma terial in seed pl ant s . (Negaspor o-
cytes are too inaccessible and too few in number to be general ly usefu l for
chromosome studie s . ) Heiosis occu rs one to t hree weeks b efor e an thesis in
many flo,,"leri ng plant s , but in various woody species, e . g ., conifer s . eornus,
and in certain herbaceous perenni als , e . g .• many bulbs . Trillium . Podophy llum;
meio s i s may be completed one t o seve r a l mon t hs before t he pol len is ac tual ly
shed. Un l ess th e time q( meios i s with r espect to other even ts in t he l ife
cyc l e is already k nown. a cons i derable amo unt of trial and erro r observation
and samp l ing may be required to determine the proper developmental stage and
the optimum time for collection . This can be critical for t he success of a
given project . par ticularly if it involves field collect ions, because the
optimum season for many spec ies is restricted t o a fe\~ weeks of the year.
250 10

III . NICROSGOPIC OBSERVATIONS

Chromosome \~ork requires good microscopy; practice and experience are the
best routes to proficiency. The equipment does not need to be elabora te or
unduly expensive . The basic req uirements are a good range of eyepieces and
objectives including at least one oil immersion lens (90-l00X) for bri ght
field which combined I.. ith the appropriate eyepieces (12 . 5-l5X) produces a
magnificat ion of 1200 to 1500X . This is near the useful limit for visual
Hork. Phase contrast optics are very useful (or difficult subjects, hut
should not be used as a subs titute for care in preparation and preCise stain-
ing . A graduated mechanical stage adds greatly to the efficiency of system-
atic slide study and the location and relocation of desired chromosome figures .
A good light source and filte rs including a green one are essential. An
attachable or built on photomicrographic camera is indispensable for recording
observations, occasionally the camera lucida drm·ling apparatus Is useful.

Chromosome number and other determinations should be based upon repli-

cated independent observations of several figures from the same slide , from
different slides , from different individuals, from different populations,
IV'henever this is possible or feasible. The most experienced observer is often
mi s l ed in his interpretation by point s of overlap between chromOSOMes, by clear
constrictions and occasional broken chromOSOMes which make one chromosome appear
as tl~O, by differences in focal plane I~hich obscure arm length s and si ze rela-
tions, and momy other details . The microscopist must often make conscious
effort to avoid subjective bias in his interpretations . This is aided by making
free hand diagrams including each chromosome! ill its proper space relation as
the figu r e is studied. Comp letion of the ske tch provides an objective chromo-
some count and aids objectivity in the evaluati.on of mo rphological details .

It is far better to make a number of slides of the same material so that

they may be quickly surveyed and only the best selected for careful study ,
than to spend much time struggling to inte rpr et mediocre preparations . Simi-
larly one passes quickly by the more difficult figures found in a given slide
and returns to study them carefully only I..hen nothing bet ter is available .
Th i s is a common and reasonable practice in most Cflses, but it can l ead to
biased results in the evaluation of meiotic pdiring relationships because the
more complex pairing configurations are also those which are mo st difficult to
interpret. Frequently , \..hen chromosomes are small and numerous, or somel.. hat
crm.. ded , it is not possible to determine whether a figure is I-lorthy of careful
s t udy uithout the enhanced reso lution provided by an oil immersion l ens . For
such nateria l a 60X oil imme rsion objective and relatively low power eyep ieces
often provide the most e f ficient combination for routine s lid e scanning .

IV . RECORDS AND DOCUHENTATtON

Accurate records are a necessary part of any scientific study; certain

complications a r ise in chromosome work Hhich need to be anticipated. A simple
numbering sys tem for ident i fy ing field col l ections and plots in cu lture with
subordinate numbers for individuals will serve to identify the plant material.
In add ition the collection vials have to be nlLmbered so that they can be corre-
lated with the pl ant s . 1\11 slides prepared from the material in a given vial;
all notes , drawi ngs, and eventually photographs derived f rom those particular
slides, also have to be identified so that they can be referred with assurance
to the particular plant or collection I~hich \-1BS the origina l source of the
material .
10 251

The plant materials are docnmented by herbarium specimens : Illhole plant

specimens for taxon identifica tto n, and if appropriate, mass collections of
particular plant parts to record individual plant variations Hhich are relevant
to the cytological results . The chromosome results are documented by photo-
micrographs (be sure to record magnif i cations) \~hich illustrate representative
observations, supplemented as needed by drm-lings and/or interpretive diagrams .
Representative slides should be made permanent as fu rther documentation of the
chromosomal results. The dry ice freezing method is excellen t (Conger and
Fairchild); various other techniques are available . Sometimes the very cells
\dlich are most interesting are destroyed in the permanizing process; the essen-
tial records and photography sho uld be completed first.

TECHNIQUES

I . OXYQUINOLINE-ACETI C- ORCEI N SQUASH HETHOD FOR ROOT-TIPS

A. Prepare a stock solution o ( 8 ·-hydroxyquinoline .

This is an almost satu ra t e d solution obtained by dissolving 0 . 3 gm
8-hydroxyquinolin~ i n 1 J.it er of distilled IJater . Prepare a day in
advance; shake occasion al ly until completely dissolved . Store in
refrigerator. It Hill remain usable [or several Heeks .
B. 1. Prepare a stock soluti oll of aceto-orcein.
Dissolve 2 . 2 grams o( orcein in 100 ml. glacial acetic. Bring the
glacial acetic ac id to a gentle boil and slowly add the orcein .
Remove from heat, c ool , and filter . (He very much prefer the
George T . Gurr, Ltd., London SH,6 , England , natuIal orcein to the
other varieties including American synthetic . )
2 . For storage : Keep as 1 00% acetic-acid stock solution .
3 . For use : (a) Aceto-orcei n s t aining solution . Dilute to 45% strength
acetic ac i d by addi ng distilled Hater . Ahmys refilter IJhen pre-
paring stain for use and also filter this L15% stain every tHO or
three days as precip itate accumulates .
(b) Orcein-HCI solution. Also prepare in a separa te dropping
bottle a solution containing : 1 pt . N HCl
9 pts . of the aceto-orcein stain
(i . e. , the 45% acetic acid solu-
tion)
C. Root-tips may be obtained:
(a) From germinating seeds-- if seeds are plentiful and individual plant
identity is not important .
(b) From the pot ball of plants grOlV\1 in cultur e . Use seedlings or older
plants I~hich have been recently repotted . Allol~ the plants to
groH undisturbed until a net\wrk of roots begins to form on the
surface of the soil inside the pot. At this stage tap out each
plant from its pot, replace immediately , and Hater thoroughly .
Hithin t\"0 to several days return to these same plants to make
root-tip collections. The "tapping-out" creates a small air
space into \Jh i ch the more active roots groH rapidly during the
next fe\~ days .
D. Procedure (Adapted frpm Tjio, J. A., and A. Levan, 1950 . The use of oxy-
quinoline in chromosome analysis . Anales de la Estacion Expt . de Aula
Dei 2 : 21-64 . )
1. Collection :
Collect fresh root-t i p s di.rectly from the plant into vials contain-
ing the aqueous oxyquinoline solution .
252 10

2. Pretreatment :
Place the vials in a cool room, T . & L. say 65 0 F. or use a 55°
room, for 4 hours . Some materials such as Pinus Idth very large ,
long chromosomes require a much longer pretreatment --as much as
24-30 hours.
3. Staining :
a . Place the root-tips in a few drops of the Orcein-HCl (9 : 1) so lu-
tion in an ordinary watch glass. Heat gently (until the fumes
of HCl begin to rise from the surface) over an alcohol lamp.
Repeat the heating at intervals of a minute or more until the
sample has been heated 3 or 4 successive times. The heating
hastens the macerating effect of the HC!. Leave the material
in this solution for 20- 30 minutes.
b. Transfer the root-tips to a large depression slide containing
aceto-orcein stain (the 45% solution) . Cover to prevent evapo-
ration and leave 20-30 minutes . Hith very small chromosomes a
much longer period in stain, as much as 12- 16 hours , e . g. ,
overnight , may be helpful.
4. Slide preparation :
a . Have clean slides and covers readily available in containers , e . g.,
keep them immersed in 95% alcohol , and I.;ipe them clean and dry
t.;ith a fine cheese cloth as needed.
h . Using a clean s l i de as a dissecting board, p l ace a stained root-
tip under alaI'; pOI.;er (lOX) stereoscopic microscope . \·lith a
needle provided ,dth a scalpel-like tip (half-spear preferred)
cut away the root-cap , and then excise a sma ll piece of meri-
stem (not more than 1-2 rom in length). P l ace this hit of
tissue in a small drop of stain at the center of a second
clean slide . Break the tissue into small pieces by quick
strokes I.;ith the flat side of the half spear needle . Spread
the tissues in a small circular area on the slide and apply
cover g l ass immediately . Observe the result and note any
bits of tissue which are still intact; give the larger of
these a quick, light tap with a blunt instrument such as the
handle of a pair of fo rceps . If the dissection and teasing
are properly done , t\Vo or three taps \Vill be sufficient. The
cover should now rest in contact lvith the drop of stain over
most of its area .
I nvert the slide on a small blotter . Use a cheesecloth
to avoi d finger prints and press moderately to hard lvith the
bal l of the thumb . The amount of pressure varies Hith the
material--size of cells, softness of the tissue , etc . It
must be learned by experience . No amount of pressure will
correct for failure to soften the tissue beforehand , or for
the pr esence of foreign matter such as sand grains under the
cover!
CAUTIONS :
1. DO NOT HEAT ORCEIN PREPARATIONS . In this respect orcein
contrasts I.;ith carmine.
2. AV.~AYS CLEA..~ THE HALF-SPEAR NEEDLE AFTER MAKING EACH
PREPARATION . NEVER HEAT ONE OF THESE NEEDLES .
3 . Label the slide I.;ith the appropriate collection number or
other symbol immediately after preparation is complete.
5. Storage and handling of slides:
a . Aceto-orcein slides may be stored in a moist c hamber , in t he re-
f rigerator overnight (or sometimes fo r tlVO or three days)
10 253

\"ithout sealin g . (Additional stain is added as necessary . )

b. They may be sealed with paraffin, with one of the various sealing
mixtures , or I"ith finger nail polish and kept (refriger ated)
for several weeks . Check at frequent intervals to be sure
seal is intact.
c. They may be made permanent by the HcCl intock method or by the Con-
ger dry ice method . (Conger, A. D., and L. ~1. Fairchild , 1953 .
A quick-freeze method for making slides permanent . Stain Tech-
nology 28 : 281- 283 . ) Alcohol destains orcein (it does not
des t a!n carmine) so the slides and covers must be moved through
the 95 % and 100% al cohol I"ith dispatch .
d. Hhen making permanent--BE SURE TO INSCRIBE PERHANENT SLIDE r-..rm-lBER
lofITH DIAl-fOND POINT PENCIL- "A!'ID HA.KE PROPER RECORD- -BEFORE
RUNNING SLIDE THROUGH PERNANIZING PROCEnURE .

II . FEULGEN TECHNIQUE1<

Th e Feulgen stain, or l euco-basic fuchsin , stains DNA specifically and

is used for cytophotometric determinations of DNA in nuclei or chromosomes or
simply as a qualitative high l y specific stain for chromosomes . Variou s t ech-
niques have been described for staining chromosomes by this p r ocedure, but the
basic principle involved in all is : liberation of the aldehyde groups of DNA
by hydrolysis , a chemical reaction binding the free aldehyde groups to the
leuco- basic fuchsin , follo~"ed by the r ~moval of unbound stain from the chromo-
somes. (See Darlington and LaCour, 1962 for further details of solutions and
schedules . )

The [01lm"ing are the essential steps of the Feulgen squash technique as
used Hith somatic tissues in Nicotiana:
1 . Fix tissue in Carnoy fixative (6 parts 95% ethanol , 3 parts chloroform,
I part Glacia l acetic acid) or 3: 1 (3 parts absolute alcohol , I part
Glacial acetic acid) for 2-24 hou rs.
2 . Replace fixative with 70% ethanol and squash or store in refrigerator .
3 . Hydrolyze tissue in 10% concent r ated HCl at 60° for 10 minutes .
4. Rinse i n several changes of distilled wa t er .
5 . Transfer to l euco-basic f u chsin for 10-15 minu tes or until desired stain-
ing intensity is reached .
6 . Rinse in 3 10-minute changes of 502 water fol l m"ed by 5 minutes in dis-
tilled water (the purpose of this is to remove the unbound stain f r om
the ch r omosomes).
7 . Transfer to 45% acetic acid .
8 . Squ ash in 45 % acetic acid on slide .
9 . Cover I"ith cover glass .
CO}IHENTS :
1. Rinsing in 50 2 water can be elimina t ed if quant itative DNA analysis is not
important . In that case rinse only 3 5-minute changes of distilled
water.
2 . The l euco-basic fuchsin and 502 ,,,ater a r e made using Dar lington and LaCour ,
schedule 4 . page 144-145, or prepared solutions can be p urchased .
3 . The most common causes of failure of this technique are poor f i xat i on.
incorr ect timing ' of hydro l YSis . imprope r ly prepared staining solu t i ons .
or poor samp l es of t he po~"dered basic fuchsin .

*Contrib u ted by Joyce Burns, North Carolina State University , Raleigh .

254 10

III. IRON ACETO-CARNINE HETHOD

(Pollen mother cells or somati c meristems)

1. Fixation (three alternatives) .

a. 3 pts 100% ethyl alcohol: 1 pt . glacial acetic acid .
b . 3 pts 100% ethyl alcohol : 1 pt. chloroforl'1 : I pt. glacial acetic acid .
c . Substitut e 100% propionic acid for acetic aC1d both in the fixative
and in the staining solu tion. Better results Hith some materials.
NIX FIXATIVES IN SHALL QUANTITIES JUST BEFORE USE!
NOTE : The volume o f fixative should alHays exceed gross volume of the plant
tissue by la-50 times. Buds tightly enclosed by protective structures
need to be clipp ed or spread open as they are dropped into the solution .
2.

in refrigerator (2-3 weeks) .

70% , or material in fixative which
freezer . It will keep f or 2 or

3.

IV . ALCOHOLIC HYDROCHLORIC ACID-CARNINE (SNOH'S STAIN)

FOR CHROHOSDHES IN SQUASH PREPARATIONS
(modified after Snow , 1963)

1. Fixation : Use fluids suitable for the aceto-carmine method discussed

above .
2. Staining .
a . Staininr, solution. Add 4 gm . of carmine and 1 ml. of concentrated Hel
to 1 5 mI. o f distilled water . Nix well and boil (GENTLY) for 10
mi nutes ,.;hile stirring . Cool the solution . Ad d 95 mI . of 85%
ethanol and f ilter. Store in stock bottles .
b . Staining proc ed ure.
(1) lI'ash out fi xative with several changes of 70 % ethanol. Allol'; an
hour f or each change. Hate r i als may be stored i n th e last
change and refrigerated if so desired .
(2) Remove buds or root tips f rom the 70% a l cohol , drain on ahsorbent
paper and place in a v ia l of stain (2-3 times as much as needed
to cover the material) . Leave materials in the stain until
they are well penetrated . The time varies and must b e deter-
mined by e xp e rience, b ut f or root-tips and anthers, which are
exposed directly to the solution , 24 hours is o f ten lon g enough;
compact tissue requi res a longer period in the stain . To has-
ten stain ing, place capped and labeled vials in a 50-60°C
 10 255

paraffin oven for 24-48 hours, or longer for some materials.

(3) Pour stain into a stock bottle for reuse (filter to remove any
debris before reusing) .
(4) Rinse material Hith I·,ater or 70% alcohol and prepare squashes as
usual substituting 45% acetic acid in place of aceto-carmine .
Excess material for future usc may be stored in 70% alcohol.
(5) Dissection o f buds and maceration of the tissue , if desirable,
should be done in 45% acetic acid .
(6) Differentiate by cautious heating over a small flame (e . g .,
alcohol burner) . (Not necessary for all materials)
(7) Squash in 45% acetic acid or for permanent slides mount in equal
parts of Hoyer's mounting medium (see Chapter 8) and 40%
acetic acid and squash .

V. QUICK FREEZ E ~!ETHOD FOR l'tAKING SLIDES PERHANENT

A. Naterials
1. A slice or small block of dry ice (solid C02)' (Obtainable from dairy
produce plants, or manufactured in the laboratory from compressed
C02) .
2 . Ti.;ro Pe tri dishes containing 95% and 100% ethyl alcohol , respectively,
each containing a short g lass rod to support the slide .
3 . Euparal or other mounting medium I"ith a solvent compatible l"rith
residual traces of l"rater (not Canada balsam) .
B. Procedure : (Adapted from Conger , A. D. , and L. N. Fairchild, 1953 . Stain
Technology 28 : 281-283).
1 . Remove temporary sealing material from slide and cover ; inscribe per-
manent slide number and make record .
2 . Place slide face up on flat sur face of dry ice for 30-60 seconds (hold
slide firmly in place by pressing it against the dry ice "'ith the
eraser of a pencil); then remove cover glass by sliding a flat
dissecting needle (such as a hal f spear) or edge of a razor blade
under a corner. \hth optimum timing the cover slip l"rill pop off
without breaking or damaging the material.
3 . Quickly place both slide and cover in the dish of 95% alcoho l . Keep
slide in original pos ition but i nvert cover so material side will
face up .
4 . Agitate slide gently for 15-30 seconds in order to disperse Hater
Hhich had condensed on the glass during the freez i ng process.
Transfer both slide and cover to 100% alcohol ; keep "I'laterial side"
up.
5 . After 30-60 seconds, remove slide from dish , drain quickly and ,,,ripe
excess alcohol from blank ends . Place drop of Euparal on the
material; re-invert and mount the cover Blass i n its original
position . Not e : The quick changes indicated here are necessary
,,,rith orcein stained mate r ial. Changes can be made more slOl"rly
after carmine or Feulgen staining .
6 . Allo\~ slides to dry on a flat surface overnight. It is best to store
recently pr?c.~.ssed slides in a horizontal position.

CHROHOSONE TECHNIQUE EXERCISE

A. Preparations :
1. Each student lvill have read and carefully considered Chapter 10 , sec-
tion B, Karyology in Practice , before attending this laboratory
session.
256 W

2. Optimum equipment and advance preparation are indicated; procedures

must be modified to fit the actual conditions . If the students
are to prepare the reagents and collect the materials this must
be done at least two or three days in advance of the present
exercise .
B. .f.quipment: .......
1. Compound microscope with oi l immersion objective and mechanical stage
(graduated if possible) .
2. High intensity light source (filters and filter holders desirable ,
not essential).
3. Stereoscopic microscope (lOX) .
4. 60°C oven and refrigerator available.
5. Dissecting instruments, alcohol lamp , vials , slides, cover glasses,
slide box , marking pencil , blotters , cheese cloth, Petrie dishes.
short glass rods .
C. Reagents:
1 . Glacial acetic acid .
2. Ethyl alcohol: 70%, 95%, 100% .
3. Stains (in dropping bottles): aceto-carmine , acetic- orcein, 9 : 1
orcein-Hel solution .
4 . IN Hel solution .
5. For fresh root tip colle c t ions : 8-hydroxyquinoline sol ution (kept
refrigerated) .
6. Supply of dry ice slices; finger nail polish, rubber cement for tem-
porary sealing .
D. Naterials :
1 . Vials containing pre-treated and pre-fixed root tips from several
d i verse plant species .
2 . If feasib l e , vials of root tips collected 3 hrs . before c l ass begins,
undergoing 8- hydroxyquinoline pretreatment in co l d room .
3 . Vials containing buds at meiotic stages , and representing diverse
species and chromosome situations . These may be collected days
to months in advance and stored in a freezer .
E. Procedure (Instructor) :
1 . Demonstrations of slide making procedures .
a . Root tips, prestained so that they are ready for use as the period
begins .
b. FIOlver bud material; illustrate dissection and sampling procedures
to locate desired meiotic stages .
2 . Demonstration of dry ice (or other) method of making squash slides
permanent . (This may be postponed unt i l later in the session if
desired . )
F. Procedure (Indivi dual Student) :
1. Each student will make root tip slides of at least 2 different species .
The pre- fixed material is to be stained in aceto - carmine; the fresh
pretreated material in acetic orcein . Slides ,viII he examined to
verify the presence of well-spread chromosome figu r es at mitotic
metaphase .
2 . Each student will find, and make slides of, as many different stages
of meiosis as possible from the materials avai l able .
G. Procedure (Group) :
3 . Each student wil l demonstrn t e his best mitotic slide(s) to the instruc-
tor and to the class. For greatest interest and variety , different
students should have made slides of different species .
10 257

4. The inst ru ctor will set up demonstration microscop es shmdng examples

of meiotic stages . Students \~ill demonstrate their own me iotic
slides to t h e instruct or and to the class and compa r e with the
stages on e xhibit.
5. Tempora r y sealing or practi ce in making slides permanent as time
permits .

NOTE: To succeed this exercise must be based upon pret ested materials \~hich
a r e known for excellence as objects of chromosome study and ease of mani-
pulat i on such as : 1. root tips of onion , hyacinth, rye , barley; 2 . buds
or inflorescences of Tradescantia pa ludosa, Rumex hastatulus , Podophyllum
peltatum , Diphyll eia cymosa , rye, ma iz e .

CYTOLOGY LITERATURE

t. General Refe r e n ces .

Darlingt on , C. D. 1963 . Chromosome Botany and the Origins of Culti-

vated P l ants . George Allen and Um-lin. London.
-=:::;:;;
_
1965 . Cytology . .1. and A. Churchill . London .
' and L . F. LaCour , Th e Handling of the Chromo somes , 4th ed .
George Allen a nd Um~in . London .
Sharma, A. Jt" and A. Sharma . 1 965 . Chromosome Techniques . Theory
and Practice . Buttenwrths . \.,tas hington.
Sharp, L . \.,t . 1934 . Introduction to Cyto l ogy . HcGraw- Hi ll. New York.
1943. Fundamentals of Cyto l ogy . I>lcGraw- Hil l. Ne\" Yor k .
Snow , R. 1963 . Alcoholic hydrochloric acid-carmine as a s t a in for
chromosomes in squa sh preparations. Sta in Technology 38 : 9-13 .
Stebbins , G. L . 1971. Chromo some Evolu t ion in Higher Plants. Addison-
Uesley . Reading , Massachuset.ts .
Swanson, C. P . 1957 . Cytology a nd Cyt.ogenetics . Prentice-Hall .
Engle\~ood Cliffs , New Jers ey .

II. Periodicals . (Note: Plant chromosome data app e ar in numerous b iologi-

cal, botanica l, and genetics journals ; also see references at end of
Chapter 11 and in sec tion II below).
Canadian Journal of Genetics and Cytology . Geneti cs Socie t y of Canada ;
Experimental Fa r m, Ottawa .
Caryo10gi a . In te rnational Journal of Cy t ology , Cytosystema t ics and
Cytogenetics . Published by the University o f Floren ce , Italy.
Chromosoma . Published by Springer Verlag , 175 Fifth Ave ., New York .
Cytogenetics . S . Karger , AG, Arnold-Boeklindstr. 25 , 4000 Basel ,
Switzerland .
Cytologia . The International Journal of Cy tology . Pub l ished i n Tokyo .
Stain Technology. A Journal for Ni c r otechni c and Histochemistry ; Offi-
c i al publication of the Biological Stain Commission .

Ill . Chr omosome Number Lists and Refe r e nces

"
Darlington , C. D. , and A. P . Ihlie . 1956 . Chromosome Atlas of FIOl~er-
ing Plant s . Geo rge Allen and Unwin . London .
Federov , A. A. (ed . ) . 1969 . Chromosome Numbers of Flm~ering Plants .
Acad eMY of Sciences of the USSR , V. I•. Komarov Botanical Instit ute .
Index to Plant Chromos ome Numh ers . 1956-1964 . Vol. 1 & 2 (Nos . 1-9)
plus a pre- 1956 suppl ement) . Universi t y of Nort h Carolina Press .
Chapel Hi l l , North Carolina .
258 10

Index to Plant Chromosome Numbers . 1965- , Annually , in Regnum Vege-

tabile v . 50 , 1965; v . 55 , 1966; v. 59, 1967; v . 68, 1969 ... Inte r -
national Bureau Plant Taxonomy and Nomenclature .
lOPB Chromosome numh er reports. A regular f eature in Taxon , publisht';d
by the International Association for Plant Taxonomy .
Love , A., and D. Love . 1961. Chromosome Numbers -.of· Central and North-
~l est
European Plant Species . Opera Botanica . Vol . 5. Almquist &
\nksel l (d is tributors) .
11 259

Chapte r 11 . GENETIC EVIDENCE*

Genetics is concerned \·lith the study of varia tion, its expression in

development, and its t r ansmissi on in inheritance . A basic undertaking of
taxonomy is the recognition, description , and use in classification, of the
variation \~hich exists bet\.'een natural populations of organisms . The analysis
of expe rimental populations provides the geneticis t Idth a pOlverful method for
inves tigatin g the underlying causes of variation and the continuin g processes
of evolutionary change . The analysis of discontinu ities in the diversity of
natural populations leads the taxonomist to classify ; i deal l y , the c lassifica-
tion evaluates the r elat ionships of existing organisms as products of these
same evolutionary causes and processes .

Nodern genetics extends from viruses to man , f rom mathematics and bio-
chemistry to psychology and medicine; many of its complexities have no direct
relevance to vascular plant systematics . The discoveries of molecular genetics,
however , have caused some major and malty subtle changes in c l assical concepts
and terminology. \,Jatson (1970) presents a gener a l account of molec ular genetics ,
I~hile Srh, DI"en, and Edgar (1965) have integrated the old \.ith the n ew. Ki ng
(1968) furnish es definitions, hThile Rieger , Hichaelis , and Green (1 968 ) provide
curren t and authoritative explanations of terms and concepts . Genetic theory
is no\" deeply ,.oven into the fabric of the modern synthe ti c theory of evolution.
Dobzhansky (1970) , Grant (1963 , 1971), l-layr (1963) , Simpson (1953) , and Stebbins
(1950 , 1967, 1970) all discuss at l ength the gene tic and cytogenetic proces ses
in evolution .

Linnaeus , by defining species and arranging them in orderly categori es ,

provided the foundntions upon which systematic biology has been bui lt . Danlin ,
a century later , I"orked muc h of his life and en titled his greatest book "On the
Or igin of Species . IT The Linnaeans considered spe ci e s to be i mmut able , divinely
created form s while variations within species I-J ere individual imperfe ctions in
the realization of the ideal form . Although the recognition of spec ies is cru-
cial to any consideration of their origin from var ieties as the res ult of varia-
tion and natural selection , Dan~in tended to consider th e species as units of
conven ience in man I s description of the genealogica l lines of descen t .

Twentieth century genetics has resolved many of the earlier contradictions

rega rding species concepts by demonstrating that species must not be c onsidered
as ideal, t ypological abstractions b u t as dynamic biological populations in
'''hich the individuals vary mo rpholo gically and physiologically and Idthin which
they interbreed. Controversy still surrounds o ur modern definitions of species
largely beCause there are many different kinds of biological populat ions Vlith
different b r eeding systems including some (agamic complexes) which have even
lost the capacity for interbr eeding . These ci r cums tan ces often c r eate serious
difficulties for the practicing taxonomist because in the absence of detailed
genetic and cytogenetic analysis he haS to make his decisions regarding the
nature of the species pop_ulations wit h which ho;! deals on t he basis of ind i rect
circumstantial evidence .

Genetic analy sis has demonstrated une quivocally that the species is a
real, probab l y the most signific ant, unit in the evolution ary differentia t ion

*Contributed by Ben \..1. Smith, North Caro lina Sta t e University, Raleigh .
260 11

of biological popul ations . The species represents the "point of no return"

in evolution , the point at Hhich the differentiation of a descendant popula-
tion becomes irreve r sib l e because pr ogr essive change prevents its return to
the ancestral condition and at which reproductive isolation prevents int er-
b ree ding with collateral speci es popul ations ,"hieh have the same ancestry .
This is the basis for the bio l ogical species concept .-. It is fundamental for
any phylogenetic system of classification .

The impact of genetics upon systemat ics i s two-fold : (1) concep tual and
general, or (2) experimental and addressed to specific problems. The experi-
mental populations of genetics usually fa ll ,,,1th10 the bounds of a single
taxonomic race or a single species. The general principles discovered in such
studies are of great sign ificance to systematic s , but the detailed results of
geneti c analyses in such popul atiol1s are of little direct value in resolving
particular taxonomic questions . On the other hand, systematists have to deal
Idth the relationships of various higher categories between Hhich divergence
has proceeded so far that genetic experimen tation is impossible. At the l ower
levels of the taxonomic hierarchy, from race or s ubspecies to species and
genus, genetic experiments of various kinds are possible in natural pop ula-
tions and the methods of genet ic s a r e directly applicab l e to the investigation
of taxonomic problems .

Geneticists have developed modes of thinking about variation , its trans-

mission, and its expression in both individuals and populations of organisms
wh ich are highly r elevant to systema t ics . The two disciplines are interre-
lated in many I-laYS , b ut only a brief summary of the mos t per tinent genet i c
concepts can be presented here . This consis ts of: (1) the basic tenets of
Hendelian heredity I~hich arise from the particulate nature of inheritance ,
(2) the features and processes which account for the occ urrence of diffe ring
genetic systems which lead in turn to the production of populations I~it h
markedly different variabilities and breeding behavi or . and (3) concepts from
genecology and population geneti cs (in words rath er than mathematical formu-
lations) which are helpfu l in dealing with populations at the subspecific ,
specific, and generic l evels of the taxonomic hierarchy, the critical stages
in evolutionary divergence. Selection and related matters have been deferred
to Chapter 26 on evolution . As in Chapter 10 , Cytological Evidence, each of
the three units which follows is designed to be read in sequence as a synopsis
of the subject , but the terms discussed may be fou nd ind i vidually by means of
the index _

I _ BASIC CONCEPTS AND CONSEQUENCES OF PARTICULATE INHERITANCE

Concepts of the ultimate physical units of inheritance have passed

through several metamorphoses from the "element s " a nd unit characters of
Hendel, to the genes of Horgan and his associates, to the nucleotide triplets
(Hhich are codons for amino aci ds, which are the buildin g blocks of po l ypep-
tide chains) , the sequences of which in turn make up the genes of molecu l ar
genetics . None of this has a l tered Hendel ' s basic conclusion that the ulti-
mate hereditary material is particulate and that it is transmitted essen tially
unchanged from cell to cell and from generation to generation. There have
been many refinements, hOl~ever , in our understanding of the process.

The genetic unit s wh i ch ac tually segregate and recombine in each cyc l e

of meiosis and syngamy are cross-over segment s, I~hich cons i st of linear l y
arranged sequences of linked genes in the chromosome, segment s of the DNA
11

macromolecule . The sizes and limits of these mendelizing chromosome segments,

and the relative degree of linkage versus disassociation between individual
genes , are functions of cross-over position and frequency which are in turn
subject to genetic control and can be profoundly modif ied by structural changes
in the chromosomes themselves.

Gene and Locus . The gene is the ultimate unit of Hendelian inher i tance , a
precise sequence of nucleotide triplets (codons) , Hhich occupies a specif i c
chromosomal locus. segreg ates a nd recombines as a unit , and which can mutate
to va r ious allelic forms I"hich have different effects in inheritance . The
locus (plural , loci) is the specific point in the linear order of genes in
the chromosome which is occupied by a particular gene .
Allele (Allelomorph). One of the alternative forms of a particular gene which
can occur at a l ocus . l.Jhen more than two alleles are known for a given
locus and gene , they are called multiple alleles .
Heterozygote . A zygote or individual carrying two different alleles o f a given
gene (e . g ., Aa) .
Homozogote . A zygote or individual formed f rom the fus ion of gametes each o f
which carr i es the same alle l e of a given gene (e . g ., AA) .
Dominance and Recess i veness. The dominance i n the express i on of particul ar
alleles and the recessiveness of others is commonly observed , but precise
investigation reveals that dominance is frequently incomplete. Although
sup e rficially similar to that of the homozygous dominant allele , character
expression in the heterozygote is usual ly to some degree i ntermediate between
the dominant and the recessive homozygote. Dominance is the general term for
inter- allelic interactions in quantitative genetics studies .
Overdominance . Overdominance occurs ,,,hen the heterozygote (Aa) exceeds both
homozygotes (AA or aa) i n the genotypic value of the character under study .
The expression of the heterozygote falls outside the range of the tlva homo -
zygotes .
Epistasis . This is the general term for the interaction of non-allelic genes
and is commonly employed in the analysis of quantitative inheritance . The
epistatic effect of an individual gene refers to the manifestation of its
effect over that of non-allel i c genes. The converse, hypostatic , refers to
the effect upon the expression of the non-allelic genes in such a relation-
ship .
Unit Characters and Pleiotropy. Specific gen e s often produce spectacular
effects upon specific characters . Further study has shOtm that most genes
affect many d iffe rent characteristics even when one o f the effects is more
obvious than the others . Also some g e nes (pleiotropic) have major effects
upon a number of seemingly unrelated characters . Since the action o f most
genes is related to enzyme systems and metabolic pathways , these results
are to be expected.
Penetrance and Expressivity . Penetrance is the frequency i n percent of domi-
nants or of recessive homo zygotes which are actually expressed phenotypi-
cally . Expressivity refers to the strength or weakness in phenotypic
expression of a g i ven gene . Both qualities depend upon the interaction
of genotype and environment .
Qualitative and Quantit~tive Characters. These terms designate the extremes o f
a continuous array of character expressions and gene effects . Qualitative
characters have been described as differences i n kind : major effects upon
colo r, shape , hyper-development or abo r tion of certain organs, etc., but
even such characters have quantitative attributes . Quantitative characters
are those which can only be descr i bed in metrical terms : numbers , lengths ,
areas, volumes, etc ., and in Hhich the net effect of a single gene among
262 11

the many affecting the same character is s mall i n r elation to the magnitude
of non-heritab le , e nvironmental ly induced modification . Quantitative char-
acters exhibit continuous variation; qualitative characters produce discon-
tinuous variation .
Oligogenes and Poly genes . Terms for the genes assumed to correspond respec-
tively to the qualitative <lnd quantitative characte"!' _e"ffects . There are
many genes Hhich are observed to have only relatively small effects. but
other genes (oligogenes) have not only major effects but also minor
effe c ts on other characters and act as "polygenes ," The polygenic inheri-
tance of recent literature is essentially equiva lent to the multiple factor
inheritance of earlier authors .
Hodif i er Genes and Genetic Background . Genes \.,rhich are known or assumed to
modify the primary effects of other genes are often called modifier genes .
The effect of any ge ne or gene complex under study all.,rays occurs in the
context of all the other genes \"hich are pres e nt in the genotype . These
other genes constitute the gene tic background or res i dual genotype .
Genotype and Phenotype. The genetic constitution of an individual organism
is the genotype . This means the total genetic endmvrnent of the organism
present in the original zygote . The phenotype is the expressed character-
istics of an individual organism . This means all the characters of the
organism Hhich result from the interaction of its genetic endmvrnent (geno-
type) Hith the environme nt during development and throughout the life of
the individual. (NOTE : Genotype and phenotype are often used in a much
restricted sense for purposes of analysis. Genotype designates the genic
or chromosomal constitution o f an individual in a particular context ;
phenotype the character expression in such an individual . )
Segregation and Independent Assortment. Segregation is the separation of
allelic pairs , the genes r eceived f rom the parents, and their distribution
to dif f erent cells, the gametes . Independent assortment describes the in-
dependence of non- allelic genes (on non homologous chromosomes) in the
segregation process; non-alleles on different chromosomes segregate ran-
doml y ,.,rith respect to each other, cf. linkage .
Transgressive Variat i on . "Then the range of variability f ound in a segre-
gating generation such as the F2 exceeds the range found in either parent
or the Fl , the variation is transgressive .
Linkage . The non-random, "linked" inheritance of groups of genes and the
characters they control Hhich happen to be spatially associated on the
same chromosome . The degree of linkage is relative to the physical dis-
tances betHeen the genes on the chromosome . These distances are inferred
from the frequencies of crossing-over bet~'een the genes as determined from
the associations of phenotypic characters in breeding experiments . The
cross - over (map) distances hetHeen genes can be affected by factors other
than physical distance, e . g . , structural modification of the chromosomes ,
or genetic effects on cross-over frequency .
Supergene. A gene complex Hhich is more or less permanently linked and trans-
mit t ed as a unit is sometimes called a supergene. It is often difficult to
distinguish such gene complexes from individual genes by experimental means .
Recombination . The constant reassortment of chromosomes , cross-over segments,
and therefore genes \.,rhich produces ne~.,r genotypes and thus new phenotypes in
each generation of sexually reproducing organisms . Recombination is the
i mmediate source o f the variation observed in populations , the raH material
upon Hhich the processes of evolution and speciation act .
Nutation . Occurrence of a heritable change in the genotype of an organism
Hhich was not inherited from its ancestors; the ultimate source of all
genetic variability (cf. recombination) . "All genetic changes, except those
11 263

due to recombinat ion , are mutations" (Dobzhansky , 1970) . A mutation may be

as minute as the substitution of a single nucleotide pair in the DNA mole-
cule , as great as a major change in chromosome structure or number .
Recurrent Nutation . Precisely the same mutation may occur again and again at
the same genetic locus . either in nature or under the influence of a given
physical or chemical agent (a mutagen). The frequency of such mutations
under a given set of conditions is the mutation rat e.
Viability and Lethality . All genes affect in some direct or indirect hTay the
survival, health, and longevity of the organism . Relative viability can
be measured by comparison ."ith a standard or "normal" population . Even
lethality, in terms of gene action, is not absolute but relative , because
death caused by the action of a specific gene or gene complex may be
haplontic in spore or gamete, or diplontic in the zygote , or delayed to a
late embryonic stage or even to early maturity.
Fitness. Success 'in producing v iable and fertile offspring .

II. THE GENETIC SYSTEN

The Genetic System refers to all the intrinsic (genetic) proce~ses and
their concerted operation \."hich affect genetic recomb i nation in a population
or species. The major components of the genetic system are the organization
of the chromosomes and their behavior in meiosis, and the breeding system .
Genetic systems range f rom that of the outbreeding diploid \."ith complete pai r-
ing and unrestr icted crossing-over of homologous chromosomes to that of the
obligate apomict Idth no sexual reproduction. Studies of the genetic system
of an organism include characterizat ion of the chromosomes and their cytogenetic
behavior (See Chapter 10 , Cytological Evidence), experimental outbreeding and
inbreeding, and an ana lysis of the reproductive bio logy o f th e species, includ-
ing isolating mechanisms. The results of such a st udy often show the taxono-
mist the most logical biological boundaries for species delimitation . Major
concepts involved in the analysis of genetic systems may be briefly charac -
terized :

Reproductive Biology. A collective term applied to all of the many mechanisms,

agents , and adaptations involved in the reproductive process of a species .
In higher plants , fo r example, reproductive biology includes floral struc-
ture, mode and/or agent of pollination, self- or cross-incompatibility ,
various types of sterility, time and season of flo\."ering. and similar char-
acteristics. Although fundamental to an understanding of the dynamiCS of
any natural population, reproductive biology is inadequately known in most
plant species .
Panmixis. The i nterbreeding of all the actively reproducing members of a
population in such a \."ay that mating is at random, i.e .• each individual
has the same probabili ty o f mating \,]ith any other individua l.
Outbreeding (Cross-breeding). A breeding system in which sex ual reproduction
involves the mating and union o f gametes of different individuals (requires
cross-po l lination i n seed plants) . The common mode of sexual reproduction
in high e r animals . I n plants outbreeding may be obligate or f acultative .
The necessity Qr.•tendency for outbreeding may be rein f orced by dioecism ,
self-in compatibility, self-ster ility , heteros ty1y and other mechanisms .
It is usually aided by \."ind, insect , or other pollinating agents. Out -
breeding may imply mating betHeen indivi duals Hhich are less closely
related than would occur lvith random mating .
Inbreeding, A breeding system in \."hich sexual reproduction involves the mat-
ing and union of gametes of individuals \."hich are more closely related than
264 11

would occur with random mating . Selfing, sib - mating, and back-c ros sing
(e.g ., parent-offsp ring matings) are examples of close inbreeding . Self-
pollination is rather frequent in plants, espec ially among annuals. It
may be obligate (e.g ., cleistogamy) or facultati ve and is often aided by
floral adaptations which enhan ce the opportunities for selfing .
Heterozygosity and Homozygosity . Applied to the pre&~nce of a pair of dis-
similar or i dentical homologous alleles (or chromosomes) respectively in
the individual, but also used to characterize the relative numbers of such
gene pairs in populations. Stri ct outbreeding produces and maintains rela-
tively high degrees of heterozygosity; strict inbreed ing produces and main-
tains homozygosity .
Inbreeding Depression and Hybrid Vigor . The inbreeding of normally crossbred
organisms usually causes depressed vigor and viability because of the
increased homozygosity of deleterious genes; outbreeding of normally inbred
organisms (and of experimental inbred lines) often r esu lt s in hybrid vigor .
Hide crosses , i . e ., bet\.;reen species or genera, sometimes produce extremely
vigorous hybrids , but more often the hybrids are relatively ,,'eak and infer-
tile .
Heterosis. Super iority of heterozygous geno types in one or more characters
over the corresponding homozygotes, the phenotypic result of gene inter-
action in the heterozygotes .
Self- Incompatibil ity . Genetically controlled physiological interactions \~hich
prevent or inhibit (partial self-incompatibility) the cOQpletion of se1f-
pollination and/or self fert ilization in certain perfe ct flowered or rnonoe-
cious plants (e . g. , species of Nicotiana , Hemerocallis, Tr ifolium) and thus
promote c ross-bree ding . The same genotypic combinations which cause self-
incompatibility also cause cross-incompatibility reactions when they occur
in separate individuals \"hich may be otherwise unrelated.
Heteros tyly. Combined morphological , physiological, and genetic mechan i sm in
c ert ain angiosperms .,'ideh promotes cross-breeding . Flol'lers of tl.;ra , some-
times three kinds \"hieh differ in the relative positions of the styles (and
anthers) are produced by separate individuals . Successfu l poll ination and
fertilization usually occurs only between flow ers which differ, e .g., long
styled by short styled or the converse , thus cross-pollination is ensured .
Apomixis . The substitution of some form of asexual reproduction for the
sexual process amphimixis in an organism derived from sexually reproducing
ancestors . Apomix is should not be confused with amixis , the asexual forms
of reproduction \"hich are common in procaryotic organisms.
Clone. All the members of a population of cells or o r ganisms derived asexually
f r om a single individual by vegetative (mitotic) reproduction .
Agamospermy. Includes those forms of apomixis in which seeds are produced by
one of a \~ide variety of asexual processes . Agamospermy does not necessar-
ily produce clones because meiotic segregation and recombination can occur
in certain kinds of agamospermy .
Agamic Complex . Comp l ex of hybrid and polyploid forms reproducing by agamo-
spermy so that each original form comes to be represented by an apomictic
population. These are extremely difficult taxonomically because each "popu-
lation " is actually the extension of a single individual of hyb rid origin .

III. GENECOLOGY AND THE GENETICS OF POPULATIONS

Geneeology . Study of the genetic basis of ecological differentiation . "A

turning- point in the study of evolu tionary taxonomy ." (See review and
def i nitions of terms in Davis and Hey\1ood , 1963, Chapter 12 .)
Deme Terminology . The suffix "-deme" as originally proposed and correctly
emp loyed is defined as "a term , always used in this terminology with a pre-
fix, denoting any group of individuals of a specified taxon ."
The nature of the association of the group of individuals or the kind of
populational unit which they may represent must ahlays be specified by the
addition of an appropriate prefix. Second order, compound prefixes may also
be used to specify relationships more precisely . TIle word deme used alone
simply means "a group of individuals of a specified taxon" and shm"s no
other relationships unless this is specified in the context (Davis and
Heywood , 1963; Heslop - Harrison, 1967). Various basic -deme terms are indi-
cated below. The deme terminology equivalents are also indicated when
appropriate in connection with other concept s described in this section.
Topodeme . A deme occurring in a specified geographical area .
Ecodeme . A deme occurring in a specified kind of habitat.
Phenodeme. A deme which differs from others phenotypically.
Genodeme . A deme which dif fers from others genetically .
Cytodeme . A deme which differs from others cytologically.
Plastodeme. A deme which does not differ from others genotypically , but does
differ phenotypically .
Gamodeme. A deme tha t is a panmictic breeding unit.
Autodeme . A deme composed of predominantly self- fe rtilizing individuals .
Endodeme. A gamodeme composed of predominantly endogamous (closely inbreeding)
dioecious plants or animals.
Clinodeme. One of a series of demes which collectively shm., a variational
trend and thus collectively form a cline .
Ecotype=Ecological Race=Ecogenodeme. A population differing genetically from
others which occupies a specific ecological habitat.
Ecospecies=Hologamodeme . All the ecotypes so related that they are able to
exchange genes freely without loss of fertility or vigor in the offspring .
In many groups this unit is approximately equivalent to the taxonomic
species.
Cenospecies=Coenogamodeme . All the ecospecies so related that they may
exchange genes among themselves to a limited extent through hybr idization.
This is equivalent in most cases to the biological species .
Biological Species. Grant (1971) : "The sum total of the r aces that interbreed
f requent ly or occasionally \"ith one anoth er, and that intergrade more or
less continuously in their phenotypic charact e rs." Hayr (1963): "Groups of
interbreeding natural populations that are reproductively isolated from
other such groups." Dobzhansky (1970): "A Nendelian population is a com-
munity of individuals of a s exually reproducing species \o1ithin which matings
take place. A biological species is the most inclusive Hendelian population;
it is integrated by the bonds of sexual reproduction and parentage." Taxo-
nomic species are sometimes equivalent to biological species but for a
variety of reasons, some good, some bad, are often less inclusive .
Compa rium=Syn gamodeme . All the cenospecies between which hybridization is
pos sible either directly or through intermediaries . Gene exchange between
cenospecies is impossible .
Hyb r id . In taxonomy, the resul t of a cross between two taxa , often inter -
specific. In geneties , the result of a cross between any two individuals
or races \.,hich differ genetically.
Hybrid SHarm . A population of interspecific or interracial hybrids and their
segregates and intercrossed derivatives. Formation of such a population
can be the first step in introgressive hybridization , but is not the actual
process of introgression which always involves successive backcrossing.
266 11

Hybrid Zone . Geographic area or region in \.Jhieh races or species meet and
hybridize .
Introgressive Hybridization . A proces s of succes s ive hybridizations in nature
which causes the mig ration of genetic material from one species into
another. An initial interspecific hybridization is followed by a series
of successive backcrosses to one of the parents (it- can occur independently
in both di r ections). The process of g(me migration is often called intro-
.&!.ession .
Clinal Variation . A s equence of morphologically interrela t ed forms \~hich have
evolved in r espon s e to an environmental gradient and form a more or less
continuous serie s , a cline . The populations in the series are clinodemes .
eltoal variation i s in contrast to introgresston but easily confounded ~ith
it .
Isolating Hechanism. Any of a number of intrinsic (genet i c or cytogenetic) or
extrinsic (geogr.aphic , ecological environmental) mechanisms or circumstances
which prevent interbreeding and thus isolate two parts of a once continuous
population .
Pre zygotic Isola tion. This results from any geographica l , ecological , or tem-
poral circ umstance or behavioral , structur al, or genetic incompatib il ity
which preven t s th e formation of hybrid zygotes .
Postzygotic Isolation . This results from mechanisms which reduce the v i ability
or fertility of hybrid zygotes or individuals .
Effective Popul ation Size . The number of individuals of a population which are
actually reproducing in any generation , the breeding population.
Nendelian Population . An in t erbreeding group of individual s sharing a conunon
gene pool.
Panmictic Unit=Gamodeme . A local population \"hose individual members have the
same chance of mating and producing progeny--the smallest Hendelian popula-
tion .
Gene Pool . The total genet i c potentialities in the sum total of the genes in
a breeding population from \"hich the genes of the next generation are
derived .
Gene Freq uency. The proportion of a given allele at a specific locus 14hich
occurs \,,1th reference to all other alleles at this locus i n a given breed-
ing population . Usually expressed in values from 0 to 1. 0 . A gene fre-
quency of 0 . 5 indicates that h3lf the loci in the popula tion carry the
specified a l lele.
Gene Flow. Changes in gene frequency I"hich result from the migration of genes
or gametes into a breeding population from a different population .
Har dy-\<Jeinberg Law . The principle that both gene and genotype frequencies are
constant from generation t o generation p r ovided that there is no migration,
mutat i on , or selection \dth respect to the gene in question , i.e . , the popu-
lation is in lIardy-t.Jeinberg equilibrium . The principle is based upon a
large random mating population .
Genetic Drif t. This is the loss or the f i xation of a gene \4i thout or in spite
of selection because of the "accidents of sampling . II Genetic drift has
been too often invoked to account for circumstances for which the investi-
gator had no ready explanation. The random fixation (gene frequency = 1. 0)
or loss (f = 0 . 0) is most likely to occur when one of two conditions exists :
(1) the pop ulation sixe is very small, 10-30 individuals , or (2) the effec t
of the given gene is very small , in which case the effect of fixation either
~ay \IIould be difficult to detect .
Founder Principle (Ernst I>layr) . If a small population invades a ne\" area and
becomes the basis of a new evolutionary line , divergence from the original
11 267

ancestral population will be accelerated because the founder population

includes (as the result of sampling) only a limited portion of the ances-
tral gene pool.
Selection . " Anything tending to produce systematic, her.itable change betl"een
one generation and the next" - G. G, Simpson .
Fitness versus Flexibility . Fitnes s is the degree of adaptation of an indivi-
dual or a population to the environment in which it lives . The ultimate
measure of fitness in terms of population genetics is the relative ability
to reproduce over generations . Flexibility is the capability of response
to change , i . e ., the adjustment of adaptation to environmental change . Fit -
ness places a premium on homogeneity with minimal expressed variability;
flexibility depends upon a wide range of free or potent i al genetic varia-
tion. The best compromise between these tlofO conflicting demands upon a pop-
ulation appears to be a ,.;ide range of variation coupled with a large modal
class of individuals t-thich are r e latively uniform and near the opt imum in
adaptive characters .
Natural Selection . "Th is preservation of favorable individua l differences and
variations , and the destruction of those "rhich are injurious. I have called
Natural Selection or the Survival of the Fittest" - Darwin . Evolutionists
now state this in genetic terms : "Natural selection is the differential
reproduction of genotypes" - Lerner .

GENETICS LITERATURE

I. Periodicals .

American Naturalist . 1867+ . Published for the American Society of

Naturalists . University of Chicago Press. Chicago .
Canadian Journal of Genetics and Cytology. 1959+. Pub lished by the
Genetics Society of Canada. Otta,.;a .
Evolution . 1947+. International journal of orgclllic evo lu tion . Published
by the Society for the Study of Evolution.
Genetical Research . 1960+ . Published by Cambridge Uni versity Press .
London .
Genetics . 1916+. Official publication of the Genetics Society of
America .
Genetics Abstracts . 1968+ . Published by Information Retrieval Ltd .
London .
Genetika . 1965 . Published by Akademiia Nauk SSSR . Hoscol'; .
Hereditas . 1920t . Published for the l-1endelian Society of Lund for
Scandinavian Association of Geneticists .
Heredity . 1947+ . An international journal o f genetics . Longman Group
Ltd . Edinbur gh .
Journal of Heredity . 1910+ . Publication o f the American Genetic Asso-
cia tion. \']ashington , D. C.
P lant Bre eding Abstract s . 1930+ . Published by the Commom.;ealth Agri-
cultural Bureau . Farnham Royal , England .
Silvae Genetica. 1951+ . Published by J . D. Sauerlandcr . Frankfurt an
Hain . '''''

II. General References.

Davis, P . H., and V. H. He)l1.;Qod . 1963 . Principles of Angiosperm Taxon-

omy . D. Van Nostrand. Princeton , Ne'IV Jersey .
268 11

Dobzhansky , Th . 1970 . Genetics of the Evolu tionar y Process . Col umbia

University Press . New Yo r k.
Gran t , V. 1963 . The Origin of Adaptations . Columbia Universit.y Press .
Ne\.,I York .
1971 . Plant Speciation . Columbia University Press . New York.
Heslop- Harrison, J . 1967 . New Con cept s in FlmoJe'r-ing Pl ant Taxonomy .
Harvard Uni versity Press . Cambridge, Nass.
King , R. C. 1968 . A Dictionary of Genetics . Oxford University Press .
New York .
Mayr . E. 1963 . Animal Species and Evolution . Belknap Press. Harvard.
Cambridge. Hass .
Rieger, R . • A. Hichaelis, and H. N. Gr een. 1968 . A Glossary of Genetics
and Cytogenetics . 3rd ed . Springer Ver l ag . New York .
Simpson, G. G. 1953 . The Major Feat ur es of Evolution. Columbia Univer-
sity Press . New York.
Sr b . A. 1-1 • • R . D. Owen. and R. S. Edgar . 1965 . General Genetics . 2nd
ed. IJ. H. Freeman & Co . San Francisco.
S tebbins, G. L . 1950 . Variation and Evolutio~ in Plan t s . Columbia
University Press. Net., York .
1967 . Adaptiv e radiation and trends of evolution in higher
plants. Evolutionary Biology 1 : 101-142 .
1970. Variation and evo l ution in plants : progress during the
past twen ty years. In : Essays in Evolutionary Biology, Hecht, H. K.•
and \oJ. C. Steere (eds . ) . Appleton-Centur y- Croft s . New York.
Ha ts on, J . D. 1 970 . No l ecular Biology of the Gene . 2nd ed. \~. A.
Benjamin , Inc . New York .
12 269

Chapter 12. REPRODUCTIVE BIOLOGY A.t~D SYSTEHATICS

Despite th e vast array of individual variation and the taxonomic problems

such variation may cause in some cases , the actual processes of evo lution and
speciation occur at the population level rather than at the level of the indi-
vidual organi sm . This is so because evo lut ion , broadly defined, is a direc-
tional change in gene frequency through time , and change s in gene frequency
only occur in populations of an organism. (Individuals do not change their
gene frequencies, thus they do not adapt in the evolutionary sense, they merely
adjust during their life span, within their fixed genetic limits.) Further-
more , all populations. even of a singl e kind of organism, do not evolve at the
same rate because different populations are under different select ive pressures
that affect, in different ways , the actual or potential e volutionary history of
the population. Accordingly , knowledge of population dynamics and biological
interrelationships often aids in a better understanding of the evo lutionary
results of importance to systematics. Two of the most obvious population char-
acteristics of such importance are population size and population breeding
structure .

Section A. POPULATION SIZE AND DENSITY

A large interbreeding population, contain ing many hundreds or thousands

of individuals , \.,/ould normally have a much larger gene pool than a sma ll popu-
lation of perhaps on ly a dozen or so individual s . A large gene pool provides
a vital storehouse of genetic variation which in turn provides a population with
the capability of adapting , over a period of time , to gradual changes in the
selective fo rces of the environment . The gene pool of a small population has
proportionally fewer different alleles and, consequently , lacks the adaptive
capabilities of the large population under changing environmental conditions .
However, just on the basis of chan ce, a new mutant or a particular ne\·] gene
combination is much more likely to become estab lished in a small population
than in a large population . A large gene pool thus has a s tronjl; buffering
effect that prevents rapid shifts in ge ne frequency and possible overspec i ali-
zation.

The chance (as opposed to selective) fl uctuati on of a gene ' s frequency in

a population is called drift . If the frequency of a particular gene "drifts"
to either 0 or 100% , the gene f r e que n cy is fixed and. since selection has pre-
sumably played no important role in the process, it is called random f i xation.
In small populations , random fixation is much more likely, and more frequent ,
than in very large populations . In deed, it is very diffi cult, even with strong
selection. to raise the frequency of a particular gene to 100% or to lower it
to zero in a large population .

The population st ructure that provides maximum evolutionary potential is

thus neither very large nor very sma ll, but is generally considered to be a
series of interbreeding populations of intermediate size. The re c an be a
relatively rapid development of slight genet ic differences between some of
the populations and then reb lending of the slightly divergent gene pools .
Such a reticulate breeding structure has the equivalent gene pool of a large
population, yet has much more adaptive flexibility \"ithout the disadvantages
of the small population.
270 12

In the establishment of a population at a new site . population si ze will

most likely be small for the fir st few generations . Thus . regardless of the
ul timate size of the population, the gene pool may show some of the effects
of drift and random fixation that were involved in the ear lier stages of popu-
lation estab lishment. Furthermore , the ent ire fate of a population may depend
on the genes b rought into the population when it Has s.tarted . If. fo r exam-
ple, a new population is sta rted by a single seed that was somehow transported
to the nel'; site, the very limited gene pool \.;i11 produce a population of
rather uniform indiv iduals ,,,ith, presumably . a relatively low evolutionary
potential. The reduction in population variability caused by the small size
of the origina l gene pool provided by the organisms that started the popula-
tion is called founder effect .

Population size, bm"ever, involves more than the gene pool : it involves
some form of competition bett.,reen the individuals of the population . Since
populations of organisms tend to occur in relatively specif ic ecologica l sit-
uations which have finite spatial limits, a second important aspect of popu-
lation size is population density. In plant populations an increase in den-
sity means more direct compe t ition between individuals for moisture, nutrients
and light. The l arger the population in a given area (i . e ., the mor e dense
the popul ation) the more seve re the competition between its members and the
more harsh the natu ral selection .

In plants , population size in the spat ial sense i s established by certain

environmental limits which define the ecologically suitable habitats of each
species. Some populations , limited to a very specific soi l type, type of r ock
outcrop, pollinator range , or moisture condition may be only a few squar e
meters in area; populations of plants with Hider physiological tolerances may
occupy many square miles . Hany populations of certain plants are more or less
linear and follOl'; stream banks. roadsides, coas tal dunes and similar linear
habitats for considerable distances. However , populations of a given species
are never uniformly distribu ted within the geographic range of the species,
nor are the individual plants usually uniformly spaced within the population .
Both populat i ons and individuals tend to occur in "clumps " or " patches " \.;rithi.n
their respective areas. The proximity of one population to another , and the
maintenance of a particular plant density uithin each population, becomes
extremely important not only in the matter of pollination and seed production
but a l so in r egard to gene £101" and attendant individual and population varia-
bility which is of evolutionary and systematic importance .

For some plants, such as t r ees and shrubs that live for many year s and
get larger each year, population density actually decreases as the population
gets older and individua l plants continue to die as they get shaded out by
the more robust or faster growing individuals in the population. This "self
thinning" has also been ob~erved in some annuals that maintain a rather uni-
form density at maturity despite varied densities of the seed at planting time .
This comes about in tt.;o ways . In the first case , the first seedlings to germi-
nate produce a wat er-solub l e chemical substance that inhibits the germination
of the other seeds in the immediate area . The first seed lin gs to start gro\"th
thus have a strong competitive advantage over the later seedlings that must
compete with a larger , estab lished , seedling for light , water , and so il
nutrients. The root s of some plants, such as those of the black walnut tree
(Juglans nigra) are suspected of giving off substances ,.;hich are toxic to many
plants, incl uding their own seedling, thus providing life- long protection
against c.ompeti tion. In the second case , germination is not inhibited , dense
stands of seedlin gs are produced, and disease (such as damping off ) quickly
removes many of the \~eak seedlings and population density is lowered .

Although an individual plant obviously cannot migrate in response to

increa s in g population density, migration is still an important fac tor in rela-
tion to competition and density in plant populations. There are numerous
adaptations of fruits and seeds which promote migration a\./ay from the pare nt
plant and often 8\~ay f r om the population . For example , t he seeds of the mag-
nolia (Hagnolia grandif10ra) have a bright red fleshy covering, called an ari! .
Hhen the seeds are ripe, they hang expo sed outside the follicle s that make up
the cone-like fruit . The fleshy ari1 contains a water-soluble chemical which
inhibits or prevents seed germination; only when the aril is removed \"ill the
se ed spro ul:. If a seed falls to the ground under the parent tree , it will not
germinate. If a seed is eaten by a bird , however, the fleshy ari l and its
chemical inhibitor are removed as the seed passes through the digestive tract
of the bird. A very hard seed coat prevents the digestion of the seed itself,
therefore , when the bird excrete s the seed , pre sumably at some distance away
from the parent tree, it Idll be ready to germinate and the seedling \"ill not
be in fa tal competition with the parent tree.

A high reproductive rate, if actually translated into high population

densit y rather than into an expanded range, often ceases to be an adaptive
advantage and becomes an adaptive liabi lity that 101"ers the biological fitnes s
of the organism involv e d. (Also see Chapter 16 on Geography and Plant Distri-
but ion . )

Section U. BREEDI NG SYSTEHS

Hany of the evo lutionary properties of populations depend upon the hetero-
zygosity maintained by a relatively steady rate of recombination '"ithin the
population . HO\~ever, in outbreeding populations chromosomal differences (e . g .
inversion, translocation), assortive mating , and, occasionally, varying amounts
of intrapopulation ste rility provide a certain degree of restri ction to com-
plete or random recombination. Nany f lo\"ers are normally bisexual or hermaphro-
ditic , thus inbreeding at the level of the individual is possible for most
higher plants . Conversely, in these plants capable of se l f-pollination there
are found a number of mechanisms that provide for some recombination through
either frequent or occasional olltcrossing . Since both inbreeding and outcross-
ing each have survival and evolutionary value und er diffe r ent conditions of
natural selecti on , it is not surprising to fin d that many plants have evolved
slightly amb i valent r eproductive systems that allo\~ for the effect ive operation
of both of these divergent breeding mechanisms within a single population or,
in some cases , a sin gle individual .

In general three different systems, based on the degree of inbreeding

versus the degree of outbreeding, can be reco gnized in plants : (1) predominant
outcrossing, (2) predominant selfing, and (3) mixed selfing and outcrossing .
In addition, several highly successful methods of asexual reproduction have
evolved among some of the higher plants and the combination of sexua l and
asexual reproduction adds to the evolutiona r y flexibility (and often the
range of phenotypic variation and taxonomic confusion) of those plant groups
in which it occurs . (See Chapter 6- VI, A and B for terms relating to pollina-
tion types and pollination . )
272 12

In dioecious plants, \,'here one plant bears only male or stami nate flm~ers
and another plant bears only female or pistillate f lo\vers, \"e find maximum
obligatory cross-ferti l ization. However, the enforced immobi l ity o f plants
makes strict dioecism of dub i ous survi va l value, especially in a small popula-
tion \,fhich might consist of plants of only one sex . Se l ection might there-
fore f avor the production and survival o f plants that ~~re functional l y dioe-
cious but I"hich may either normally, or under certai n environmental conditions,
produce a f e\-l perfect flowers . Suc h plants '"'QuId be polvgamodioecious . If
some perfect flower s oc c ur on an othenlise female plant the plant l.JQuld be
gynodioecious. The perfect flOl>'ers (as He ll as some of the pistillate flOl>'ers)
of this gynodioecious plan t could b e self-pollinated, but most of the seed set
\>'Ould be the result of normal cross pollination from another, staminate, plant.
The gynodioec i ous condition, Hhich occurs in some species of the p i nk, mint
and aster families (Caryophyllaceae, Lamiaceae, Asteraceae) allO\ols f or a maxi-
mum of recomb i nation and a minimum o f homozygosity .

In populations of s e xual ly reproducing organisms maximum g enetic recom-

b i nation, or g ene flow, Hould occur if the populat i on t>'ere panmictic, that is,
if all individuals in the population could mate at random . This means that
anyone indi vidual is equal ly l ikely to mate Hith any other individual of the
opposite sex in that particular population. Although panmixis is a useful
evolutionary conce pt f or theoretical calcul ations , or as a reference point in
expe rimental Hork, t rue panmixis most likely does not occur, or if so, only
rare ly, in natural populat i ons of higher organisms . Among plant populations
panmixi s might be most c l osely approached in a population of wind pollinated
plants . Hm>'ever , interp l ant distances , \,dnd velocity, and the prevailing wind
direct i on \>'Ould all b e f actors that Iwu l d reduce the randomne ss of wind pol-
linat i on and there f ore the randomness o f interbreeding among the individuals
comprising the population.

I . OUT BREEDING SYSTENS

Once an organ i sm b e comes evolutionari ly adapted to a particular environ-

ment, or set o f selective forc e s, it can usually maintain itself indefinitely
i n a state of eco l ogica l e quilibrium with the other biotic elements to I"hich
it is adapt e d as lon g as the tota l environme nt itself remains essent i ally
stable. Under such conditions , selection Hou1d be for continued stabilization
of the organism; increased uniformity rather than increased variability 1:>'Quld
have the most select i ve value; homozygosity ,>'ould tend to increase and hetero-
zygosity twuld tend t o d e cr e ase. I n plants, Hil i ch have evolved a much more
variable ser i es o f reproductive methods than have animals , strong stabilizing
selection might a l s o lead to changes in th e breed i ng system itself--there
might be a shift to more i nb reeding (self-pol lination) and possihly to some
form of apomixis or as e xual reproduct i on.

Among obligate outbreeding plants it is possible that they might be highly

heterozygous gene tically even though they are quite uniform phenotypically.
Also, h e terozygosit y i tsel f may increase individual survival values and also
population f itness through het e rosis and homeostatic effects, Hhile the increased
gene pool possible Id th heterozygosity provides a relatively high degree of
evolutionary potential under changing environmental conditions or changes in
the environmental selective f orces that might be incurred by migration .

Hith enforced outbre e d i ng, th e accumulation of particular alleles, or

groups of alleles , can most easily take place only in relatively small groups
12 273

of in terbreeding individuals that are in sorn.e way separated from other g r ours
or populations of the same species with \~ hich they \-lOuld norma ll y interbreed .
TIle organisms be longin g to separated , or allopatric , populations therefore
tend to different iate more readily from each other than do the sympatric
organisms of a single population . If tHO or more populations of a particular
organism are sepa rated or isolated geog raphi cally for a long enough time (i.e.,
for e n ough genera tion s) it is possible that, in add ition to any morpho logical
differences that might develop , certain physiological or genet i c changes wo uld
evolve that ,,,auld prevent effecti,ve interbreeding herl·men members of the tl>'O
oopulations should they nOH again come into con tact at some point . If this
occurs , the tHO for ms of the organism are said to be reproductively isolated
(see Chapter 26 , II) and one of the first evolutionary steps toward the possi-
ble formati on of a ne\.j' species has been completed (note Figure 21-15).

For many years , it Has believed that because of the strong recombination
effect of obligate outhr eedlng , geographic isolation \Jas a necessary first
step in the differenti ation of races and the development of other mor e effec-
tive isolating mechanisms. On the o the r hand , there is nOI"/ some e}(perimental
evidence to indicate that s trong disruptive selec tion, even Hith ITk'lximum
crossing or interbreeding of th e tHO divergent forms of a polymorphic pop u la-
tion, can be suffi cient to maintain, a nd presumably to isolate , two diver gent
groups of individuals l.Jithin th e same population . Such sympatric speciation
is most likely a rare occurrenc e in nature , but it sh.ould no longer be con-
sidered completely impossible . Given a parti.cular combination of generation
time, population s ize and polymo rph ism and a strong disruptive sele ctive force,
sympatric speciation could be r ealized .

Many of the higher plants are bisexual and capable of se lf-poll inat ion
and self-fertilizat ion . If a single floHer bears bo th male and female repro-
ductive structures , that is, both stamens and pistils , it is said to be a per-
fect flOHer . An imper fect flO\"er Iwuld be unisexual , bearing either stamens or
pistils but not both . If a single plant produces both male and female flowers ,
as is the case in many wind pollinated species such as corn (Zea mav~), oak
trees (!tuercus ~. ), and alder (Alnus) , the plant i s said to be m~cious.
In a modification of the monoecious condit i on , staminate and perfect flO\"ers
may occur on one plant (andromonoecism)or pistillate and perfect flOHers may
be f o und on a plant ( gynomonoecism) . Heterozyp.;osity and evolutionary potential
are even more strong ly assured if the unisexual flowers are bo rne on separat e
plan ts . Such plants, as indicated previously, are said to be dioecious.

In the case of plants wi th perfect floHers , se lf-pollination is not only

possible but is, in many case s , the primary type of pollination . Such close
and con tinued inb reedin g tends to reduce recombina tion, reduce heterozygosity,
reduce variability, and reduce evolutionary po t entia l. On the other hand ,
se l f-po l lination makes it possible for a single pl ant to reproduce and star t
a population ; it can also insure adequate seed set in some plant populations
in the absen ce of suffi cient or appropriate specialized pollinators. In
mono ecious plants , s e l f-pollination is certainly possible bu t not probable ,
since the pollen is usually distributed by the wind and the female flowers of
one plant are much more likely to receive pollen from t he male flowe r s of a
neighbor i ng plant than from the male flOl"ers on their own plan t. Here , out-
breeding is predominant, and recombination , heterozygosity, and evo lutionary
potential are accordingly increased .
274 12

Outcrossing is obviously obligatory for dioe cious plants , <)nd this raises
several p r oblems in terms of the optimum sexual composition of a population in
terms of the ratio of male and female p l ants . Very often, the first problem
is "solved " by the fact that , as previously mentioned, many dioecious plants
do actually produce a £e\-] perfect flm.Jers capable of self-pollination . Also ,
many o f t he species of dioecious plant s are perennial.; ·annual seed production
is not essential to survival of the population . I n the second case , selective
pressures, possibly related to some aspect of conservation of energy within
the population , have caused a shift to more female plants and felVer male plan ts
i n some spe c ies o f dioecious plants .

If the male and female parts of a per fect flot~er, or the male and female
fl Ol,'ers o f a given monoecious plant , mature a few days apart, the plants can-
not self - pollinate an d must outcross . If the pistillate fl.ol.Jers (or the pis-
tillate portion of perfect floHers) ITI.1.ture first the plant (or flol>le1.') is said
to be p r otogvnous_. In a protand1.'ous plant (or flol-ler.) the reverse is true :
the stamens of a given floHer mature and shed their. pollen before the stigmas
of the flm~er are receptive .

St ron g protandry has apparently produced a shift tOl-lard the andromonoec i ous
condition in several species of the carr.ot family (Apiaceae) . Here the numer-
ous small flo\o,'ers are horne in aggregations called umbels. Large compound um-
be ls, as in Queen -Anne ' s lace or Hild carrot (Dat1...cus carota), are made up of
many small umbellets . If the indivi dual perfect flOHers are protandrotls , the
firs t f l O\~ers to open in the umbel may be pollinated by flm"er8 nearer the
center of th e umbel that open later. As this process continues , it is evident
that the stigmas of the central flm.Jers of the terminal umbel will be recep-
tive after all of the pollen from the flo\~ers of this umbel is shed . HOI-leVer ,
stamens from the outer flQ10lers of the lateral umbels (\oIhich begin to open as
the central floi"fers of the terminal umbel finis!1 blooming) furnish the pollen
necessary to pollinate these central f 10\ole1:"s of the primary umb el.

In a number of other plants , self-pollination is still possible. despite

strong protandry . In many members of the Asteraceae, fo r example, the pol len
i s shed as the stigma , still morphologically and physio log ically immature ,
pushes up through the ring of anthers . The ",ass of sticky 90llen is l i terally
pushed Otlt o f the floHer , and most of i t is usually soon picked up hy a polli -
nator . As the stigma matures , it separates, exposing its pollen-free recept i ve
i nn er surface . At this stage, pollen from another source, when placed on t he
st igma, Hould effect cross-pollination . lioh'ever, if cross-polli nation does
not occur l-.'ithin a certain t ime, the tips of the stigma begin to re f lex until
the receptive inner surfaces come in contact Idth the style \~hich may still
be covered Hith pollen that Has not picked up by the pollinator . In this \~ay ,
self-pollination is accomp l ished and seed production assured even if cross-
pollination has not tak en place .

In s ome plants I'lith pe r fect flOl·/ers self-pollination is unlikely because

of the arran gement of the anthers and stigmas Hithin the flower . If intra-
specific differences in floral structure are responsible for the enforced out-
crossin g , the inCO Mpatibility system is said to be !J.eteromorphic . Heterostyly
is a good example o f heteromorphic incompatibility and is found in Oxalis
( Oxalidaceae ), l!oustonia (Ruhiaceae) and a n umber of other genera . As the
name i mplies , the flowers of one species are of t\~O forms : some flO\Jers have
l ong s t yles and some flol.Jers have s hort styles . In addition, the stamens of
the long-styled (or "pin") flol..Jcrs are Hell beloH the stigma \~hile the stamens
12 275

of the short-s tyled (or " thr um") flOl.J ers are above the sti gma . In this \.,Iay.
cro ss-pollin at ion bet\.,Ieen "pin " and "thrum" flal-Jers is made more likely than
self - pollination . As usual. the system is not absolute , and sometimes "ille-
gi timate cross-pollination" may occu r; in this situation, pollen from the 1 01"
stamens of one long-styled flOl"er reaches the stigma of another long-styled
flO1"er rathe r than the stigma o f a shor t- styled flO1.;er , or vice versa . HOI.,I-
ever , even I.hen il l egitimate pollinations a r e made , ferti l ization may not occur
because of secondary morphologica l or physiological adaptations cor r elated with
the heterostylous condition . "Thrum" pollen is often larger than " pin" pollen,
the papillae on the stigmatic surfaces are different in the two flower forms ,
and there are often differential rates of gr m.;th of the pollen tubes, the best
grm.;th occurring I.;hen th e pollen type i s the opposite of the style type (Le.
I.;hen the pol linations are "legitimate"). To function effective l y as a unit ,
the various genes controlling all of the correlated characteris tic s asso ciated
Id th the tl';O flm.;er types , " pin" and "t hrum ," must be closely associated , per-
haps as a supergene or block of genes , on a single ch romosome . But even such
linkage can be variously modified under different selective forces . If the
survival value of self-pollination is great enough , heterostyly may be
mod i fied to homostyly , in which the styles a re of constant length and the
stamens are attached to the corolla tube in such a pos ition tha t self-po llina-
tion is both possible and probable . In another modification, t r isty l y , the
an thers of a single flower are at tHO levels and there are three style lengths
cor re lated ,.;itlt specific stamen positions .

In other plants, which produce perfect flm.;e r s of only o ne form in I.;hich

self- pollination canno t be prevented, genetically contro lled phySiologica l
fact ors may prevent s elf-fer tili zation . This condition , called homomorphic
self - incompatibility , effectively blocks inbreeding and may be either of tl';O
types : gametophytic self-incompatibility in which the inhibit ion of fe rti li-
zat i on is the result of gene action in the po llen grain (the male gametophyte)
itself as the pollen tube grol';s through the sty l e ; and sporophytic self-
incompatibility in which the inhibition of pollen germination or pollen tub e
growth is imposed by the gene action of the sporophytic tissue of the st i gma.
In those cases that have been investiga t ed, there is a correlation between
binucleate polle n and gametophytic self - incompatibility and the formation of
trinucl eate pollen and the s porophytic type of self-incompatibility . In both
forms , hO\~ever, only one or t l.;o genes , and their alleles , seem to be involved.
Although found in about a dozen unrela t ed plant families , the genetic bas i s
of the self-sterility mechanism is remarkably similar in all cases: the
pollen tube tvil1 not grow on a stigma or through a style if the pollen and
the stigma or style have a particular allele in conunon . Thus . a plant o f the
genotype 5152 would produce t wo kinds of pollen, 51 a nd 52, neither of I~hich
would gro\~ through the 51 52 sporophyt i c tissue of the pistil of the flower of
the pl ant that produced the pollen . 5uch pollen grains I~ould , however, be
effective on a stigma or in a style with another genotype such as S353 or
S455 .

In the ground cherry , PhysaliS (Solanaceae) , self - incompatibility is due

to t wo independently ass.erting genes, Sand 2 and the ir alleles . This impo ses
an even more stringent selection for outcrossing: in the cross between two
plants of the genotypes S4552l26 and Sl5422Z6 for exampl e, eight genetically
different types of pollen wou ld be produced but only 1/4 of the pollen pro-
duced by plants of the first genotype I'lould be effective on the stigmas of the
plants with the other genotype . The I.;asted pollen is part of the biological
"cost" of maintaining heterozygosity and a greate r evo lutionary potentia l in
these particular plants .
276 12

Al though sexual reproduction is the mechanism which insures variabili ty

and evolutionary potential through recombination , asexual reproduction has
strong survival value under certain conditions in I ..hieh the production of
uniform (but not necessarily homozygous !) preadapted genotypes can maintain.
or even increase , a population in a given environment . Or. in cases where
various forms of ste rility prevent effective sexual reproduction, asexual
reproduction is obv i ously the only method of s urvival of the organism, and
in the h i gher plants several secondary types of asexual reproduction have
evo lved that provide for reproduction Id tho ut the fusion of male and female
gametes .

II . ASEXUAL REPRODUCTION

Vegetative propagation or reproduction is quite common among the peren-

nial f lowering plants and can take place by the format ion of nel" p l a nts from
rhizomes, tubers . stolons a nd root divisions which become separated from the
parent plant . Layering . the rooting o f detached branches , propagation from
individual bulb scales , and the formati on of vegetative propagules of various
kinds on the l eaves, ste ms, or in the inf l orescences of certain plants can
also prod uce large numbers of genet i cal l y identical i ndividuals called a
c l one . I f environmental conditions remain stable , t he members of a clone may
maintain a populat ion for many years . Some clones, such as the long-lived
coast redwood (Sequoia sempervirens) which reproduces by stump sprou ts from
old root crowns , and some of the rhizomatous grasses of the American prairies ,
provide examples of c lones that have survived for tho usands of years .

Because of the uniform geneti c background of the members of a clone, any

variability be tl"een the members, even wi thin a particular population, I"ill
probably be phenotypic variation cau sed by slight environmental differences .
Clone material, the re fore , is ideal for studying the morphologica l and physio-
logical effe c t s of different environments on a part icular plant or series of
plants from one or more popu l ation s from differen t areas of habitats .

Another form of asexual reproduction , cal l ed agamosp ermy, involves the

production of viable seeds \o,fhich have been formed without fertilization ,
al though in some cases pollination is required to trigger the agamospe rmous
process . The embryo in such seeds may develop from an unred uced egg (this is
parthenogenesis) or some other unreduced cell in the embryo sac , or it may
develop from a somatic cell in the ovule . In all cases, the embryo is geneti-
cally identical with the seed-producing parent. If there are several different
gen otypes i n th e population , these same genotypes \. . 111 be represented unchanged
in the next generation , but possibly in different p r oportions. Although r ecom-
binat ion is not possib l e , the series of populations maintains a smal l amount of
f l exib ility through time as selection operates to increase or decrease the per-
centage of each genotype in each generation as conditions change . Such a situa-
tion occurs in the common dandelion (Taraxacum officinale) and numerous other ,
though usually perennial, members of the aster family (Asteraceae) , grass fami ly
(Poaceae), and rose family (Rosaceae).

However, despi te the immediate fitness exhibi ted by some asexually repro-
ducing organisms. unless some sexual r eproduction is possible, the members of
a c l one , as are most other organisms that result from asexual reprodu ctive
processes , are at an evo lutionary dead end .
12 277

I n thos e organisms that combine some degree of sexual reproduction , which

insures recombination and evolutionary potential . '<lith various fo rms of asexual
reproduct i on, which provides for the rapid mult iplica tion of adapted genotypes.
we see excellent examples of evolutiona r y buffer ing. Such organisms have gen-
e rally had lon g evolutionar y histories and \.;ri l 1 most likely continue to s urvi ve
under \.,.hat ever drastic loca l or \.Jorld-wide environmental changes that man causes .

III. INBREEDING SYSTENS

De spit e the long-term e volutionary advantages of cross-fertilization , a

high degree of homozygosity and a positive pollination mechanism that regularly
insu r es optimal seed production, even in the absence of pollinators , is of con-
siderab l e selective advantage i n some plant popUlations such as those of many
annual p l ants or colonizing species in new o r ephe meral habitats . Here the
advantages of immediate f itness (i . e ., survival ) and the rapid production of
numerous offsp ring preadapted to the environmen tal conditions of the nel-J o r
temporary habitat far o utwe i gh t he long- term benefits to be derived from o u t-
crossing . It is not sur prising that many of our weedy annua l s, under the
strong selection for inunediate fitness. are found to be autogamous . that is ,
they are self-pollinating , self-fertile, inbreeders .

The value of autogamy in either annual or perennial plants of colonizing

speci es is also obvious . A single seed of a self-p ollinating plant is capable
of giving rise , in seve ral generations, to an en tire co l ony of plants . In
many instan ces of di s pe r sal, even long distance di spersal, it is not unreali s-
tic to a ss ume multiple i n troduc tions of a species into a given area ove r a
period of time . The es t ab l ishment of pioneer pop ulations from different seed
sour ces provides for some gene t ic variability in th ese populations that are
disjunct from the main area of dist r ibution of the species .

Since autogamy occ ur s in several un related taxa that are predomi n ant ly
outcrossing , and since self -pollination is more likely to be present in dis -
junc t pop ulations of a given sp ecies , autogamy is considered to be an advanced
or derived condition tha t has arisen indepen den tly i n different groups of
plants . The str ong corre l a tion between sel f-fe rtil ization and coloniz ati on
has been referred to as " Baker' s La\oI ."

A special si t uation ha s bee n described for seve r al plant spec i es which

have dimorphic flowe r s and are regularly cross- poll inated near the ir center of
dist ribut ion but which have homomorphic self-poll ina ted flowers in t he peri-
pheral portions of their ranges . This occurren ce of popul ations of self-
pollinating plants in an otherwise outcrossing species can sometimes be corr e-
lated with environmental factors that great l y reduce the number of pollinators .
The \~eather of the Faeroe Islands. i n the north Atlant ic off the coast of
Norway , is often cloudy and there is much rain . As a consequence the numb er
of ins ects i s q ui te low and s everal widesp r ead species of plants that are
normall y cross-pol l i n ated by insects in con tinen ta l Europe are adapted to
self-pollinat i on (by r aindrops ! ) i n the Faeroes . Autogamy is also f requent
in pl ants t hat inhabit des er t or semi-desert areas \oIhere flowering may be
periodic and pollinators may be scar ce. Other no rmally outcrossing plants
may adapt to occasional unfavorable seasons or periodic fluctuations of tem-
perature or moist ur e by \~hat amount s to facu ltative autogamy. Under unfavor-
able environmental conditions only self - pollinated flow ers are produced but
under more n earl y optimum condit ions cross-po llination takes place . An
278 12

interesting extension of this shift from self-pollination to cr oss-pollination

in an ind ividual plant in a single season can be seen in the mous e - tail (Hyo-
surus 2.£ .; Ranunculaceae) i n which the flowers are normally self-pollinated .
Here the carp e ls are separate on an elongated receptacle in the center of the
flm.,rer and the carpels present ,,]hen the flm.,rer opens are pollinated with pol-
len from the same flower . If the environmental concl~tions remain static or
get ,verse (e.g . , if the habitat gets too dry) no further carpels are produced .
However , if the plants remain healthy and have abundant ,,'ater the receptacle
may continue to elongate and produce additional carpels above the already
deve loping achenes formed earlier. The newer carpels \"i11 necessarily be
cross-pollinated since no more pollen is available from that particular flm,'er.

I1hen we find both inbreeding and outcrossing in t\.;o closely related spe-
cies, in a single species, or even in a sin gle plant , it is quite likely that
some fl oral dimorphism will be correlated with the two pollination types.
Under reduced pollinator selection the self-pollinated floh'ers may be smaller,
less showy, and have smaller anthers. The ultimate in this type of dimorphism
occurs in those plants, such as violets and henbit, which normally produce
both open , showy, cross--pollinated chasmogamous fl O1,'ers and on the same plant
also produce small, often drab cleistogamous f lowers that never open hut se1£-
pollinate with i n the closed floral envelope . The cleistogamous flowers of
henbit (Lamium amplexicaule ; Lamiaceae) , a small winter annual weed of la\offis
and \"aste places, are usual l y produced in the late \"inter or early sp r ing and
are folloHed short ly by, or even intermixed \"ith, th e more s ho\")' chasmogamous
flm.;ers . In the common purple violet (Vio la papilionacea; Violaceae) , an
herbaceous perennial of woodland clearings and open areas, the fragrant and
showy chasmogamous flo\-!ers appear early in the spring when the days are short
and the cleistogamous fl m.;ers fo11m" \-lith the longer days of sunnner as the new
crop of leaves mature. Such plants as the violet and henbit thus have very
effect i ve mechanisms to insure both outbreeding and i nbreeding as a normal
part of each rep r oductive cycle.

However, even i n species that are predominantly self-pollinated, there

usually remains the capacity for occasional outcrossing and a rene\.Jal of
heterozygosity. A series of population studies of predominantly self-polli-
nated species of cultivated plants such as wheat , oats , and lima beans , has
shown that normal outcrossing in these closely inbred plants may ran ge from
1 - 12% . In addition , the heterozygotes appear to be more vigorous and fertile
than the homozygotes . This further indicates that self-pollination is a
derived type of reproductive process that, under certain current conditions ,
has a higher selective val ue than outcrossing despite demonstrated heterosis _
(Also see Chapter lIon Genetics and Chapter 26 on Evolution . )

Section C. REPRODUCTIVE BI OLOGY IN RELATION TO SYSTE~~TICS

The di rect relationship between the total reproductive biology of a group

of related species and their taxonomic treatment can most easily be seen by a
brief comparison of the "key " characters used in the separation of species of
oaks \"ith the "key" characters used in the separation of species of orchids
from one another ; in the wind-pollinated oaks , fruits , leaf and bark charac-
ters are used in both identification and classification ; in the orch i ds, in
which there is often an extremely close morphological r elationship between
flower structure and a specific pollinator, flower structure is the primary
basis for both classification and identification. Even ~"ithin a group of
rather closely related species the selective pressures associated \"ith the
vital function of reproduction are such that different methods of reproduction,
different pollinator s , and different population structures may all playa role
in the size of the s pec ies gene pool , the total amount of recombination (and
variability) present and , ultimately, in the evolutionary production of a num-
ber of important biological differences that can be of taxonomic value. By way
of a sununary , Table 12-1 lists a number of general cha r acter differences which
may exist between two closely related s pecies , one of which normally ou t crosses
and the other of which is normally self-pol linated in some fashion .

Table 12-1. General character differences bet\.,reen related xenogamous

and autogamous species of pl ants . (From Ornduff , 1969 . Reproductive biology
in relation to systematics . Taxon 18: 121-133 . Used Idth permission)

Characters of xenogamous plants Characters of autogamous plants

Outcrossing Autogamous
Self- incompa tible Self- compatibl e
Diploid Polyp lo i d
Low recombination index High recombina tion index
Flo\o.'ers many FIOl.,rers fewer
Pedicel s or peduncles long Pe dicels or peduncles short
Sepals large Sepals smaller
Corollas rotate Corollas funnelform , cylindric , or
closed
Petals large Petals smaller
Petals emarginate Petals less emarginate or en t ire
Floral co l or pattern contrasting; Floral color pattern less contrast-
bicolored; polycolored ing; greenish, l"h1tish, or
monocolored
Nectaries present Nectaries reduced or absent
Flm.,rers scented F101"ers scentless
Nectar guides present Nectar guides absent
Anthers long Anthers shorter
Anthers extrorse Anthers introrse
Anthers distant from stigma Anthers adjacent to stigma
Pollen grains many Pollen grains fewer
Anther appendages ,.,rell developed Anther tips reduced (Asteraceae)
Pollen presented Po l l en not presented
Pistil long Pist i l shorter
Stamens longer or shor ter than pistil Stamens equal to pistil
Style exserted 5 tyle included
Stigmatic area Hell defined , Stigmatic area poorly defined , less
pubescent pubescent
Stigma receptivity and anther Stigma receptivity and anther
dehiscence asynchronous dehiscence synchronous
Nany ovules per flow e r (inflores- Fewer ovules per f low er (inflores-
cence) cence)
Many ovu l es not maturing to seed All ovules maturing to seed
Some f r uits not maturing All fr ui ts maturing
,-
Ray florets many Ray f l orets fewer (Asteraceae)
Ray florets conspicuous Ray florets inconspicuous or absent
(Asteraceae)
Disc corollas pentamerous Disc co r ollas tetramerous (Aste r aceae)
Narrow distribution Hide distribution
 Table 12-2 . Insect Po llination Flm..rer Syndromes (from Baker and Hu rd , 1968) .
N

""
0

Nectar
Pollination Cla ss Predominant (honey)
and Ins ect Type Anthesis Colo r s Odor FloHer Shape Flo\Jer Dep t h Guides Nectar Other

1. Can tharophily Day and Va riable , Strong, Actinomorphic Flat to bowl- None None Pollen or
(Beetles) n i ght usually fruity (haplomorphic shaped (rare - or food bodies .
dul1 or and actinomor- ly closed) open Flower parts
aminoid phic acc. to in large
Leppik) numbers

2 . Sapromyophily Day aud Purple- Strong , Usually None or deep None Open Of ten no food
(Carrion and night brmffi or often of actinomorphic i f of trap or provided .
dung flies) greenish decaying type none; Transparent
p rotein sorne- w1ndOHS or
times other fea-
pseudo - tures con-
nectar-tribute to
ies tempo r ary
traps.
Hobile appen-
dages to
flowers or
';c.ai l s" to
pe tals often
present .

3. HyophUy Day and Variable Variable Usually None to None None or Bee - flies
(Syrphids an d night actinomorphic moderate present have tongues
bee-flies) to 50 mm;
syrphids can
chew pollen .

~
N
 ~
4. Mel ittop hily Day and Variable. Present; Aetinomorphie None t o mod- Presen t None La r ger be.es N
(Bees) night except usua lly to zygomor- e ra te ; o f ten or ean open
or pure red sweet phie; mor- a b road tube pres- "closed"
diurnal phology of (gu llet ) en t flowers. The
flowers can (us u- most var ia-
be recog- al) ; ble of the
nized ope n syndromes
to
con-
cealed

5. Sphingophily Usually Usually Strong ; Us ually actin- Deep narrow Usually Pres- Rim of corol-
(Hawkmoths ) no cturn- t·,hite or usually omorp hic ; he ld tube to corol- none ent; 18 may be re-
a l or pale (or Sl-lee t horizontal or 1a (or con- flexed and
c r ep u s - green) pendent spur) cea1- dissected .
c u1ar ed; often no
a mple a l igh ting
pl atfo r m.
Anthe rs
versatile.

6 . Phalaenoph ily As in 4 As in 4 Hay be As in 4 Less deep None Pre s - Le ss extreme

(Small moths) less t han in 4 en t; modifications
st r ong con- than i n 4
than ceal-
in 4 ed

7. Psychophily Day a nd Var i abl e Hode r- Usual ly actin- Deep ; narrow Often Pr es- Usua lly flat
(Butterflies) night or (pink ate l y omorphic; usu- tube to cor- pres- ent ; wide margin
diurnal b eing strong; ally held up- ol1a (o r ent can- to corolla
very sweet right spur) cealed; tube; not
common) ample disse cted

N
00
~
282 12

The extent to I'Jhich pollinators and reproductive methods may modify

floral structures ~lithin a single family is Hell illustrated by Verne Grant's
studies (1965) on pollination in the Polemoniaceae (,,,here plants in indivi-
dual species in various genera either self-pollinate or are pollinated by a
series of flies, bees , butterflies, birds and hats ) and the \.Jork of Leppik
(1964) on floral evolution in the Ranunculac eae . In each instance floral
mod i fications can be correlated Idth the type of pollinator, thus indicating
something of the role of pollinating me chanisms , and pollinators , in both
inte rs pecific isolat ion and speciation . Some of the more common insect
pollinator-flol"er relationships pointed out by van der Pi j 1 (1960-61) and by
Faegri and van der Pij 1 (1966) have been s ummarized by Bak er and Hurd (1968)
and are g iven in Table 12-2 .

The evolutionary, and taxonomic consequences of the various floral adap-

tations or modificat ion s associated \-lith xenogamy, autogamy , dioec ism , hetero-
styly , apomixis , cleistogamy and other aspects of sexual or asexual reproduc-
tion provide an interesting, and almost unlimited , field of biological study
fo r those interested in "getting behind the scenes" in systeMat ic work IVhere
data are accumulated and attempts are made to anSI·le r the question "\-Ihy " in
r egard to certain types and patterns o f plant variation upon "'hich I~e base
lIk'toy of our taxonomic conclusions .

Section D. REPRODUCTIVE BIOLOGY STUDY HETHOnS

Careful field observations Hill usually indicate pollinator type and

specificity as I,'ell as the sequence of anthesis within a single floHer , an
inflorescence, or for an entire plant . The funct ional details of flOl~er
structure are also often best studied in t he field although laboratory obser-
vations are obviously required for the deta ils of f loral morphology "'hi c h can
be observed fresh (for ultraviolet patterns n ot other\~ise apparent), cleared
(for venation patterns and other anatomical detail) or otherwise prepared
(for chemical analysis or electron mic roscope ohservation) . Pollen studies
(size, viability, morphology--see Chapter 8) a r e easily organized into blocks
of laboratory activity, as are the cytological studies (see Chapter 10) that
are a necessary part of any detailed reproductive biology research. J\OI-Jever ,
compatability studies, tests for apomixis, and experimenta l hybridization
require experimental field plots or suitable g reenhouse space .

A series of variously treated caged and llllcaged flOl"ers I~ill usually give
a prelimi nary picture of the reproductive patterns Idthin a population of
plants . A practical preliminary program involves a total of 25 flo\~ers for
each "treatment " in each popul ation . A practic.al min imum of 5 plants should
he involved in each treatment . If each plant bas many flol-Je rs, then all five
treatments (5 f lol~er s each) could be done on one plant and only 5 plants per
population Hould be needed.

25 flmolers tagged but othen:ise left alon e as a test f or cross -pollina-

tion and as a control to determin e normal seed set.

25 flOl~ ers caged, tied in bud , or othenllse protected , and then self-
pollinated to test for self-compatibi lity , which \.,ould be indicated
by full seed set .

25 floHers caged and left alone to te!'lt for self-pollination ; normal

seed set indicat es self-pollination and self-fertility (or apomixis) .
12 283

25 flowers caged and emasculated to test for apomixis; seed set under
these conditions \-lDuld indicate apomixis .

25 floh'ers caged (and emasculated if they are self-fertile) and crossed

\.,ith plants from another population ; reduced seed set could indicate
de v eloping sterility beth'een t he t\.ro populations heing tested .

Nylon netting (or nylon stockings) over Hire f rames (or f rames made by
cutting the sides out of 1/2 gallon cardboard milk cartons) eas ily s e rves to
exclude pollinators from individual flOl.Jers or inflorescences . The "cage "
should be tied to a stake for sU!J p ort and the nylon netting tied around the
st em of the plant belm" the ( IOI-ler or inflorescence .

In nk,king the various pollinations , pollen from r.ipe anthers can be

tapped onto a glass slide and the stigma rubbed in the pollen or , for smaller
floHers, the pol l en must be transferred from the sl i de (or anther ) to the
stigma Hi th a fine br.ush . The tools us ed should be cleaned in 70% alcohol
from one pol lina tion operation to the n e xt to prevent the accidental transfer
of unwanted pollen .

Actually proving the mechanism of apomictic seed pr.oduct ion involves con-
siderable eMbr.yological work , but apomixis ~~ , can be detected readily if
self-fertile flm,'ers set s eed after e masculation and isolation from pollina-
to r s . In the case o f self-st e rile flowers only caging is necessary in the
first stages of the experiment but, if seed forms , it may be the result of a
101~ percent of self-fertility instead of apomixis . Emascu lation in subsequent
trials would indicate \·Ihich process \,'<1S responsible for seed set . Needless to
say , emasculat ion must be done prior to the dehiscenc e of any of the anthers
in a floHer and must be done carefully Hith fine forceps or a small hooked
needle to prevent damage to oth er fl oral structures . If any pollen has been
shed in a particular floHer that flower can either he removed f r om the in-
florescence or tagged Hith a color-coded string around the pedicel to indicate
that it may have self ·-pollinated.

Additional caged amI emasculated flmlers can he used for attempts at

hybridization I~ith related species. Although ease of hybridization may indi-
cate a fa i rly close relationship the reverse is not necessari.ly true.: hm"ever ,
the production of Fl hybrids and a study of their morpho logy and of their
meiotic chromosome pair ing seldom fails to provide valuable backg r ound
ma t e r ial to any evolutionary study .

LITERATlffiE ON REPRODUCTIVE BIOLOGY

Baker, H. G. 1963 . Evolutionary mechanisms in pollination biology . Science

139 , 877-882 .
--,,--c-' and P . Hurd, Jr . 1968 . Intrafloral ecology . Annual RevieH of
Entomology 13 : 385-414.
Ehrli ch , P . R., and P . H. Raven . 1965 . Butterfl ies and plants : a s t udy in
co - evolution . Evolution 18: 586-608 .
Faegri , K., and L. van d~r Pijl. 1966 . The Principles of Pollination Ecology .
Pergamon Press . Oxford .
Gr ant, V. 1905 . Pollination in the Phlox Family . Columbia Un iversity Press .
Nev York .
Grun . P ., and /\. Rad low. 1961 . Evolution of barriers to crossing of self-
compatible I~ith self-incompatib le species of Solanum. Heredity 16 : 137-143.
284 12

Leppik, E. E. 1957 . Evolu tionary relationship between entomophilous plants

and anthrophilous insects . Evolution 11 : 461-481 .
1964 . Floral evolution in the Ranullculaceae. Iowa State Journal
of Science 39 : 1-10 1.
Levin, D. A. , and H. H. Kerster . 1967 . Natural selection for reproductive
isolation in Ph l ox. Evolution 21: 679 - 687. ....,
Lewis, H. 1963. The taxonomic problem of inbreeders . Regnum Vegetabile 27 :
37-44 .
Love. A. 1963. Biosystematics and classi fication of apomicts . Feddes
repertorium specierum novarum regni vegetabilis 63: 1 36 - 149. Berlin .
Ornduff , R. 1969 . Reproduc t i ve biology in re l ation to systematics . Taxon
18: 121- 133.
Pandey. K. K. 1960 . Evolution of gametophytic and sporophytic systems of
self-incompatibility in angiosperms. Evolution 14: 98-115 .
Ray, P . H., and H. F . Chisaki. 1957 . Studies on Amsinck i a . 1. A synopsis
of the genus , \.Jith a study of hete r ostyly in it . American Journal of
Botany 44 : 529-536.
Solb r ig , O. 1971 . The population biology of dandelions . American Scientist
59 , 6B6-694 .
Van der Pijl , L . 1960-1961. Ecological aspects of floHer evolut i on I , II.
Evolution 14 : 403-416; 15: 44-59 .
Vuilleumier . B. S . 1967. The o rigin and evolutionary development of hetero-
sty1y in the angiosperms . Evolution 21 : 210-226 .
13 285

Chapter 13 . CHEHICAL EVIDENCE1,

The potential importance of chemical evidence i n taxonomy was suggested

by a number o f early taxonomists. DeCandolle, Hoffmann, Hallier and Holisch
(Hegnauer, 1969, see : Harhorne and Swain) are among those who anticipated
this innovation, but it is only in recent decades that rapid methods of phys i -
cal analysis and structural elucidation of organic compounds have been devel-
oped to the degree that reasonably rapid surveys of plant extractives are
feasible . As more comprehensive surve ys o f plant chemistry have become available,
taxonomists have s hown a growing interest in t he appl i cation of chemical char-
acters to taxonomic problems.

Virtually every chemical component found in plants, including both the

primary metabolites (invo l ved in vital functions in the cell) and the secon-
dary products o f me tabolism (accumulat ed in ce lls but not cons i dered to be
involved in vital processes) , has been used in chemical comparisons of taxa
at various taxonomic levels. This chapter is a brie f survey o f the applica-
tions of chemical evidence to taxonomy . Some specif ic examples of the taxonomic
use of chemical evidence are discussed to i llustrate this approach . The scope
of chemical evidence ranges from data on the distribution of a single compound
to a tabulation of a wide range of chemical constituents; and thus, chemical
evidence can range from a single taxonomi c character to a system of characters
which can rather thoroughly define the group under study.

Section A. HISTORICAL PERSPECTIVE

tillile there are many pre- twentieth century references to the possible role
of chemistry in taxonomy , the work of Abbott (1886) represents one of the ear-
liest attempts to correlate chemistry with the phylogenetic level of develop -
ment. She found the saponin-containing plants occupied the middle level of
Heckel!s scheme of plant evolution (multiplicity of f loral elements) . She
concluded that all saponin-containing groups ar e closely allied and possess,
in addition, some other structural similarities. The following quote from
Abbott seems much more contemporary than 1886:

The f acts obtained from these studies tend to show a chemical

progression in plants, and a mutual dependence between chemical
constituents and change in vegetab l e form... The evolution of
chemical constituents in which they f ollow parall el lines with
the evolutionary course of plant forms, the one being intimate l y
connected with the other. and consequently that chemical constit-
uents are indicative of the height of the scale of progression,
and are essentially appropriate for a basis of botanical classifi-
cation . In other words, that the theory of evolution in plant
life is best illustrated by the chemical constituents of vegetable
form.

In 1909, Greshoff (cited by Gibhs 1958) suggested that chemical charac-

ters should be included in a natural classi f ication, and he proposed that

*Contributed by C. R. Parks. University of North Carolina, Chapel Hill. This

work was partia l ly supported by National Science Foundation Grant GB30780.
The author would like to express his appreciation to Miss E. Dale Johnson for
her aid in preparing the manuscript.
286 13

every description of a ne\~ genus or species should include a short chemical

description of that plant taxon .

One of the first successful attempts to combine chemica l and morphologi-

cal evidence in the study of a single genus \.;ras the work of Baker and Smith
(1920, cited by Gibbs 1958) on the essential oils of ~Eucalyptus. They sug-
gest ed that the level of relation ship should be r eflect ed in chemical similari-
ties (primitive plant = chemically primitive) . According to the morphological
and chemical data collected on 176 Eucalyptus species , they divided the genus
into three groups differing in both morphologica l struc ture and chemical con-
st ituents . They concluded that primitive species are those which have feather-
veined leaves and high pinene (I) content in their essential oils (terpenes),
while more advanced types have intermediate venation and contain pinene and
cineole (II) . According to their system, the most advanced t axa show butter-
fly wing venation and contain oils with phellandrene ( III ), piperitone (IV)
and geranyl acetate (V). Despite the fact that more recent research (cited
by Al ston and Turner 1963) on essential oils in Eucalyp tus has prOVided evi-
dence that more intraspecific variation occurs than was shown by the early
wor kers, the fact remains that the work of Baker and Smith was the first com-
prehensive chemotaxonomic-morphological study of a complex genus, and in gen-
eral their concl usions have been a significant contribution to the taxonomy
of Eucalyptus .

CH,
.... CH ...... I
CH~ I C~CHJ,.,
I CH~ I or
CH~ CH
"'C/
,
I. CH l
Cineole .l- PheUandrene Pi perilane G efao ~ 1 Acetate
"'·Pinen e

As a result of his studies, HcNair (1935) set up four quan tifiable chemical
crit eria . His premises \~ere that more closely related taxa produce more
similar chemical products, and that more highly evolved member s produce larger
molecules (alkaloids. vo latile oils and glycerides) . He furth er assumed that
unsaturation in oils i s an advanced cha racter, a nd that the degree of increase
in molecular weight and unsaturation is proport ional to phylogenetic advance-
ment . Applying these assumptions he concluded , for example . that the Hagnolia-
ceae is more primitive than the Berberidaceae or the Ranunculaceae; that the
mono co ts are more primitive than th e dicots ; and that the polyp etalous dicots
are less advan ced than their gamopetalous relatives . His data supported the
phylogenetic hierarchy according to Engler and Gil g (1919, cited by McNair

*The abbrevia ted method of presenting str ucture will be follO\~ed in the
r emain der of the structures diagrammed in this paper .
1935) in some cases, but in others it I.;as more applicable to the Bessey system .
According to McNair ' s evidence , hQl.;ever, the Ranales I.ere placed too low in
both phylogenetic sy::;tems . Hany reviewers (see Gibbs 1958 ; and Alston and
Turner 1963) doubt NcNair ' s criteria of chemical advancement .

In a later paper ~lcNair (19 45) attempted to shQl~ that chemical ontogeny
can be evidence for chemical phylogeny . One example he used cor r elated p r o-
gressive changes in terpene con t ent and aging in Eucalyptus . According t o
his criteria of chemical advancement of terpen es , more advanced compo unds
were more typically c haracterist ic of mature trees . He also attempted com-
parisons involving seed chemical constituents .

The difficulty in obtaining comprehensive chemical information of the

type necessary for phylogenetic comparisons between large numbers of plant
species proved to be almost impossible until the development of modern
physical methods in organic chemistry . Ye t despite the great difficulty in
chemical l y characterizing taxa in the early chemotaxonomic sur veys , these
early contributors must be credited with providing a philosophical founda ti on
for the appl i ca tion of modern chemistry to taxonomy .

Chemica l surveys of taxonomi cal ly more r es t ricted g r oupin gs of p lant

species have been mo r e successful i n providin g data for the working phyloge-
nist . The dis tr ibution of be talain s [formerly known as nitrogen-containing
antho cyanins (see XI a nd XII for basic structur es )], along with other evi-
dence, led Lawrence et a !. (1939) to sugges t th at the Cactaceae should be
aligned with t he Centrospe r mae; and this disposition is now general l y accepted
(Hab r y and Dreiding 1968 , see : Nabry , Alston, and Runeckles) . From an exten-
sive survey of anthocyanin con t ent in plant species , Beale et a1. (1941)
observed that the incidence of pelargonidin pigmentation is more common in
tropi cal spec ies th an in those from temperate regions . They al so noted that
the flowers of the maj ority of t ropical or subtropical species contain ing
cyan idin (VII) or delphinidin (VIII) were appreciably redder
than temperate species with the same anthocyanins in their flowers . They
concl uded tha t natural sel ec tion favors t he survival of r ed - f l owered fo r ms
in the tropics . It is now wel l known that pollination vectors which favor
red are common in tropica l regions.

OH
HO OH

OH OH OH
VI. Pelor90ni din VII. Cyonidin VIII. De l phin idin

Beale and his collabora tors (1941) also surveyed 32 species of Tulipa
for anthocyanin content . This rep resen ted one of the fi rst efforts to use
the di st ribution of a gro uping of compounds in a genus as a taxonomic char-
ac t er . They fo und t ha t the anthocyanin dist r ibution fully supported the
288 13

generic structure as determined by morphologists. The chemical constituents

sele cted fo r analysis , as morphological traits selected, must be chosen in
light of the knowledge concerning the broader distribution and selective
forces affecting particular classes of compounds.

In the 1940 ' s Nirov (1967) began his detailed 'analysis of conifer terpen-
oids. His work, along with that of others , has led to the elucidation of many
taxonomi c problems i n the conifers .

Serological methods have been applied to taxonomy since the turn of the
century, hut over-all these have contributed relatively little to systematics .
Study of the results of the earliest serological investigations proved that
serology was a potentially valuable instrument of taxonomic analysis ( Ches ter
1937) . The exhaustive serological survey of the plant kingdom led by Mez and
his associates (cited by Chester 1937) culminated in the construction of a
phylogenetic tree, the "Stanunbaum." Though Nez ' s work stimulat ed much contro-
versy, relatively little of it has been incorporated into the mainline of
systematics. Gell et a1. (1960) speculated that the comparatively crude sero -
logical tools of that day , coupled ~"ith the absence of an additional reliable
criterion to aid in eva lua ting conflicts of interpretation between serologi-
cal results and morphologically-based taxonomy. led the taxonomis ts to choose
their own time-tested te chniques .

Paper chromatography was developed by Consden et al . in 1944 . Bate- Smith

(948) showed that the methods of paper chromatography could be applied to
plant pigments in his analYSis of petal-p i gment mutants in Dahlia variabilis.
These techniques are now widely applied in plant systematic studies.

Section B . THE APPLICATION OF NATURAL PRODUCTS CHENISTRY

TO TAXONONY AND PHYLOGENY

Different groupings of natural products may have a narro~" or wide botani-

cal distribution. The distribution of a compound, or groups of similar com-
pounds with a limited occurrence , may be correlated ~"ith a taxa system for
~"hich relationships have been proposed . The chemica l data may strengthen the
hypothesis or provide clues for an alternative interpretation . If correlations
are not fo und, it is necessary to collect additional chemical or botanical
data. Extractives which are lim ited to a few families or orders are proving
to be particularly useful in this regard . Certain compounds are limited in
this distribution to one or a few species , and these are of litt le taxonomic
value.

Compound class£s such as the monoterpenes (see I-V) and the flavonoids
(see I X and X) are ubiquitous in their distribut ion. Hhile the distribution
of an individual flavonoid or monoterpero id may have little o r no significance
above the generic level, the general groupings of terpenoids or flavonoids
found in families or even in higher categories may be of considerable signifi-
cance.

The natural products surveyed in taxonomic studies are usually secondary

metabol ite s . Alston (1966 , see : Swain) , as well as a number of others , has
pointed out that secondary metabolites are subject to low selection p res sures,
and thus would be conservative characters . Hhile this argument has some
obvious validity, stud ie s of plant biology and biochemist r y often show that
13 289

OH

HO HO P' o OH
OH
OH

IX. OH
Apigenin (0 common flavon e)
x. OH "o
Quercetin (0 common flavonol)

the secondary constituents are very much an integra ted part of the re la tion-
ship of the plant to its environment and to its metabo lism . In fact. secon-
dary constituents may be under high selection pressures . The anthocyanins .
for example, which are secondary constituents, are correlated with the type
of pol lination vector mentioned earlier; therefore, the floral anthocyanins
would be expected to be under strong selection pressures .

It is beyond the scope of this brief chapter to completely revieH the

literature . Hegnauer (1962, 1963, 1964 , 1966, 1969) has compiled five vol-
umes on chemotaxonomy, and primary papers and revie"s continue to be produced
at a very rap id rate. Nevertheless, some selected examp les are pertinent to
our discussion.

Cons id ering the taxonomic importance of instances of compounds with unique

distributions in plants, the study of the " nitrogenous anthocyanins" is perhaps
the best known example . These red pigments were first recognized as different
from the anthocyan ins by Schudel in 1918 (see ~~bry and Dreiding 1 968, see :
Mabry et al . ), but the structures were not fully resolved until 1962 . The
betalains , as they are now known , include the red to violet betacyanins (XI)
and the yellow betaxanthins (XII) . These compounds are distributed only in
certain plant families, and this has been recognized for some years as being
of importance in establishing the natural relationships of families in the
Centrospermae. Ten betalain-containing families are now recognized in this
order (Mabry and Dreiding 1968 , see: l'labry et al . ) . One family , the Cary-
ophy11aceae (once thought to be a central member) , lacks these compounds .
Recent anatomical evidence, coupled with this chemical evidence, supports the
removal of this family from the Centrospermae, and' provides for its subsequent
placement in a separate order, the Caryophyllales (Mabry 1966 , see : Swain) .

Hegnauer (1 969 , see : Harborne and S~']ain) has revie,.;red phytochemical

evidence ,,,hich can be used in phylogenetic placement of the Cornaceae and its
al lies . He notes that Takhtajan allies the Cornales with the Araliales in the
Umbelliflorae and also considers these to be derived from Rosaceous stocks .
He states that Cronquist , on the other hand (unlike most othe r treatments) ,
allies the Araliales ~71th Rutaceous stocks . Hegnauer fo und that the rather
specific iridoid glycosides (XIII) and the more specific forms of widely dis-
tributed compounds found in species of Comus are very different from the
chemical composition of species in the Araliaceae. A wide array of cornalian
genera seem to be linked by the occurrence of iridoid glycosides in their
290 13

tissues. The particular composition of phenolic acids (I·ddely distr ib uted

comp ounds , see XIV) in the Cornales is more like that of the Rosales and less
like the Umbellif lorae. Eyde (cited by Hegnauer 1969 , see : Harhorne and
Swain) agrees that flor al anatomy does not favor the linking of the Cornaceae
and the Nyssaceae in the Umbelli f lorae . From a consideration of the chemical
evidence, it can be inferred that the Cornales should.,.. pe excluded from the
Umhelliflorae and, instead, should be affiliated lvith some members of the
Rosales . On the other hand , hO\.Jever. there are striking chemical rese~lblances
betl.Jeen the Rutales and the Umbel1iflorae. On the basis of the chemical evi-
dence, Hegnauer Houle! link the Corna1es ,~ith Rosaljan stocks and the Umbe 11i-
florae ,,,ith Ruta1ian-like ancestors . Hegnauer concludes that comprehensive
comparative studies of other types of characters of the taxa concerned should
be undertaken.
HO~

HOVN~COOH HO

~ HO-(, }COOH
HO

Hooc /0cOOH
I XIII Corn in XIV. Gallic ocid
H
(on iridaid glycos i de-) (a pheno l ic acid)
Betanidin XI I. Ind icaxanthin
(a belocyonin) (0 betoxanthinJ

In Table 13-1 some major groupings of plant natural products are listed
along "'ith some comment on their botanical distribution . This brief summary
is only intended to give some insights into the scope of "natural products
chemotaxonomy" and some starting points in the literature. At the end of
Table 13-1 is an annotated list o f general references which will be of value
to the student Hho ,dshes to probe further into the area of chemotaxonomy .

Table 13-1. A summary of some groupings of natural products I"hich have been
used in chemotaxonomic correlations including some comments
on the botanical distribution of these constituents .

Compound grouping Comments on botanical distribution Citation

Terpenoids Hidely distributed \,Teissrnann (1966.

see: Swain)
Honoterpenes \.,ide1y distributed, of taxonomic numerous reports,
value primarily below the generic see Nirov (1967)
level or others

Sesquiterpenoids rather ,.,ide distribution, but Herz (1968 , see :

particularly important and useful Habry et a1.)
in the taxonomy of the Compositae Herout and Sorm
(1969, see : Har-
bOTne and St,fain)
13 291

Table 13- 1 continued .

Compound grouping Comments on botanical distribution Citation

Diterpenes rather wide distribution i n the Erdtman ( 1968 , see :

seed plan ts. taxonomically useful Nabry et a1.)
in specific groups such as the Ponsinet et al . (1968,
conifers see : Habry et a 1. )

Triterpenes \Vide distribution in living organ- Ponsinet et a1. (1968 ,

isms, taxonomically useful in see : Nabry et a1.)
several angiosperm families such
as the Cucurbitaceae, and in the
fungi

Carotenoids the universally d istributed photo- Goodwin (1966 , see:

synthetic carotenoids useful in SHain)
algal class ification ; the noo-
photosynthet ic carotenoids in
f ruits of possible limited value
as taxonomic markers in the
angiosperms

Flavono i ds very large number of diverse com- Bate- Smith (1963 , see :
pounds foun d throughout the vas- Slllain) ; Harborne (1966,
cular plants and in nearly , if see : SHain ); Hagner
not all, angiosperms ; of great (1966 , see : SHa i n) ;
taxonomic value belOlIl the genus HilUams (1966 , see :
level and of possible use in the Slllain) ; Alston (1968 ,
classification of highe r cate- see : Habry et a1. )
gories (extensive literature)

Lignins and useful in the classification of Erdtman (1968 , see :

lignans higher categories Nabry et a1.)

Quinones \~idely distributed among living Hathis (1966, see :

organisms, but compounds of this Swain)
type I.;ith limited distribution
have potential value in the
class ificati on of a number of
angiosperm families

Polysaccharides universally distributed , probably Percival (1966, see :

useful in the classification of SHain)
higher categories - particularly
among the algae, comparative data
mos t1y lacking

Plant glycosides great chemical variation in the Par i s (1963 , see :

non-sugar portion of the molecule , SHain)
varyingly of great usefulness in
taxonomy
292 13

Table 13-1 continued .

Compoun d grouping Comments on botanical distribution Citation

Asperul osides and Rubiaceae, Scrophulariaceae , an d Bate- Smith and Swain

Aucuhins (Iri - related families, Plantaginaceae, (1966 , see : Swain)
doid glycosides) Cornaceae , and others Kooiman (1969, 1970)

Ranunculins found only in the Ranunculaceae Ruijgrok (1966 , see :

Swain)

Cyanogenetic Ranunculaceae and other plant Ruijgrok (1966 , see :

compounds families Swain)

Polyalcohols widely distributed but have taxo- Plouvier (1963 , see :

nomic potential Swa i n)

Sulfur compounds a chemically diverse grouping of Kjaer (1966, see :

compounds which i n their various Swain)
forms are videly distributed, the
isoth i ocyanate-producing glycosides
are characteristic of families in
the Rhoeadales

Amino acids (non- Liliaceae and related families, Bell (1966, see:
protein) Leguminosae (particularly Papi1- SI'Jain)
i onatae , and other groups)

Alkaloids a chemi cally and biosynthetical ly Hegnauer (1966 , see :

diverse group of compounds rarely Swain)
found i n the lower vascular p l ants Schafer (1964)
and irregularly distributed among Price (1963 , see :
the angiosperms; highly useful in Swain)
the taxonomy of some groups

Betacyanins and Centrospermae ( " Betanales " ) Habry (1966, see :

betaxan thins Swain)

Alkanes (fatty widely distributed , possibly of Shorland (1963 , see:

acids and wax es) use in classification be10\. the Swain)
genus level , and of use in organic Douglas and Eglinton
geochemistry (1966, see : Swain)

Fatty acid found in seven families Bu'Lock (1966 , see :

epox ides Swain)

Acetylenes d i stributed in the basidiomycetes Bu ' Lock (1966, see :

and at least 13 angiosperm fami - SHa i n)
lies, of particular use in the S~rensen (1968, see :
link between the Compositae and Habry et al . )
the Umbellif10rae
Assorted compounds ferns Berti and Bottari
(1 968 , see : Reinhold
and Liwschitz)
13 293

Table 13-1 continued.

Comments on botanical
Compound grouping distribution Citation

Assorted compo und s lichens Huneck (1968, s ee : Rein-

hold and Liwschitz)

to,lide range of chemi cal Umbell ife rae Heywood (1971)

approaches

tiide range of chemical Leguminosae Ha t borne et a1. (1971)

approaches

General References :
1. Adams , R. P . 1972. Numerical analysis of some common errors in chemo-
systematics . Brittonia 24: 9-21. [Errors typical of chemotaxonomi c
ana l ysis are discussed and analytical procedures are presented which
will help to minimize these problems.]
2. Alston, R. E . 1967 . Biochemical Systematics , p . 1 97 - 305. See : T .
Dobzhansky , H. K. Hecht , and \.J . C . Steere (eds . ), Evoluti~ry Biology,
Vol. 1. Appleton-Centur y Crofts . New York . [ This is a crit i ca l and
detail ed review of chemotaxonomy , with particular reference to the use
of secondary consti tuents . The art i cle is very readab le and contains
a n excell ent bibliography . ]
3. 1965. Comparisons of the importance of basic metabolites,
secondary compounds and macromolecules in systematic studies. Lloyd ia
28 : 300-312 . [lfuile secondary constituents have been mo st useful in
chemosystematic studies , the ana l ysis of the macr omolecule holds great
promise for the future.]
4. , and B. L . Turner. 1963. Biochemical Systematics. Prentice-
Ha ll. Engl ewood Cliffs, New Jersey . [The first general textbook to
appear in this area of taxonomy includes a discuss i on of the application
and use of chemical data in the field and a review of chemotaxonomic
studies arranged by compound groupings . ]
5. Fairbro t hers , D. E. 1968 . Che mosyst ematics, with emphasis on sys t ematic
sero l ogy , p . 141-174 . See : V. H. Heywood (ed . ). Hodern Methods in
Plant Taxonomy . Academic Press . New York . [This u sefu l review places
particular emphas i s on serological and electrophoretic methods and
i n c ludes an excellent bibliography . 1
6. Gibbs, R. D. 1965 . A classical taxonomist ' s view of chemistry in taxonomy
of h i gher plants . Lloydia 28 : 279- 299 . (The results of a wid e range
of Simp le chemical t ests on a spectrum of plant material are provided
as e x amples for the application of chemi cal da t a to the placemen t of
taxa in various taxonomic categories.]
7. Harborne . J . B. 1 968 . Biochemical systematics . The use of chemistry in
plant c l ass ification , p . 545-588. See: L . Reinho l d and Y. Liwschitz
(eds . ). Progress in Phyto chemi st ry . -----Vol 1 . lnterscience Pub l ishers .
New Yo r k . (This paper is a general introduction to the application of
chemical character s t o taxonomy with a review of chemotaxonomic st udies
arrange d by compound groupings . ]
8. Hawkes, J . G. (ed.). 1968 , Chemotaxonomy and Serotaxonomy . Academic
Press . New York . [The text contains a compendium of papers in a
var i e t y of areas in chemotaxonomy with emphasis on macromolecul ar
approaches . 1
 294 13

9. Hegnauer, R. 1962 , 1963, 1964, 1966, 1969 . Chemotaxonomie der Pflanzen.

Vo ls . 1-5. Birkhauser Verlag . Basel. [These volumes offer an
exhaustive treatment of plant chemistry arranged by taxonomic group-
ings . ]
10. Levin. D. A. 1971. Plan t phenolics: an ecological perspective. The
American Naturalist lOS : 157-181. [The article discusses phenolic
compounds as defensive constituents in plants . with ecological con-
siderations and evaluations. 1

Additional books of interest :

Harborne, J , B. (ed.). 1972 . Phytochemical Ecology . Academic Press. New

York . [A collection of papers is presented in Hhich the ecological roles
of a wide array of phytochemicals are discussed.)
1970. Phy tochemical Phylogeny. Academic Press. New Yo rk. [A
collection of papers is presented shoHing contributions of phytochemistry
to the study of phylogeny . ]

The extensive survey of a very large number of dicotyledonous and mono-

cotyled onous species for flavonoid and cinnami c acid content by Bate-Smith
and his associates (summarized by Bate-Smith , 1969, see: Harborne and Swain)
represents a good e xample of a survey fo r the distribution of a widely occur-
ring compound grouping for phylogenetiC comparison purposes. (The classes of
flavonoid s include the catechins, XV ; 1eucoanthocyanidins, XVI; flavones, IX;
f1avonols, X; anthocyanidins , VI-VIII; flavanones, XVII; flavanonols. XVIII;
chalcones, XIX; dihydrochalcones, XX; aurones, XXI; and isoflavones, XXII.
Two common cinnamic acids are p-coumaric acid, XXIII, and caffeic acid , XXIV.)
Some phytochemical correlat ions may be noted from Bate-Smith t s s u rvey o f the
angiosperm po1yphenols. In the dicots, nearly all of the predominately woody
families from Casuarinaceae to Ebenaceae contain leucoanthocyanidins. while
nearly all of the predominately herbaceous families from the Aristolochiaceae

OH ((W0H
OHHO v OH H0O ) - 0 0
OHOH H 0 0 ) 9 0
O:H HO v OH'I~OH
I ,,;, I ,,;, I '" I
OH
"-
OH OH
OH
II
o
"0 II
0
OH "- ,
o
H
xv. Catechin XVI. Leucocyanidin XVII. Butin XVIII. Tnxifolin XIX. Dahlia chalcone
(a flavan-3-al) (a fiavan-3,4-dio l ) fa f lava none) (0 chalcone)
(a flavononol)

HOWOH
I 0-
'I \:
-
OH H0O)0
I 'CH 'I
-0~
OH
OH H°GOo°
I - do OH
OH

xx.
~
OH
>I
0

Phloretin
~-
OH
I>
0

XXI . Aureusidin
"
OH 0

XXII. Genis tein

" HOOCH=CH-COOH HOQ-CH'CH-COOH

XXIII. p-Coumaric ocid XXIV. Caffeic ocid

(0 d ihydrochalcone) (on ourone) (on isofJovone)
 13 295

to Compositae are l acking in these . The trihydroxylated substituted flavo-

ooids (see XXV for myricetin . a flavonol \.lith this type of B ring substit u -
tion) follow a s i milar, but less lTk1.rked. distribution. ifhole families in t he
Tuhiflorae lack , or essential l y lack, the f l avonols which are replaced by
flavanes, f l avanones or chalcones . Vario us of the c innamic acids show similar
d i stributions and correlations .

The polyphenols of monocotyledons are not different from those of the

dicotyledons e x cept that ellagie acid (XXVI) , \.,ridely distr i buted in the woody
dicotyledons . is lacking in the monocotyledon s ; and the flav onols are gen e r -
ally less frequent in the monocotyledons . Some monocotyledonous families ,
considered morphologically advanced . have primitive phenolics by analogy t:l
the dicotyledons; a nd some monocotyledonous families . which show general l y

HO n, - CO,H HO
~OH
OH

~
OH OH CO,H

xxv. My r ic(> l i n XXVI. Ellogic acid XXV II . He~ollydro~y dj pllenic oc i d

advanced phenolic characteristics. have individual genera with phenolics like

the primitive dicotyledons . The most advanced monocotyledonou s families , such
as the orchids and the grasses, are advanced in their polyphenols in a manner
similar to the more advanced dicotyledonous families . The ellagitannins , com-
p l ex derivatives of hexahydr oxydiphenic acid (XXVI) (a ring opened fo r m of
eHagic acid) . are centrally distributed in dicotyledonous orders allied with
t he Rosales . They are absent i n the monocotyledons a nd i n the Ranalian d i coty-
ledons . It can be suggested from this compound distrib u t i on that the monocoty -
ledons diverged from ancestral dicotyledons related to the Rana1ian complex .
rather than f r om Rosa1ian ancestors. Some genera, containing primitive pheno-
l i c characteristics , belong to families that are otherwise advanced in t h i s
re spect , thus i n dicating that the ancestry of these families is traceable
t hrough these ge nera to the primitive angiospcrmous stock .

Erdtman (1968. see: Mabry et a1.) notes that patterns of chemical con-
st i tuents are more usef ul to the taxonomis t t han single chemical trai t s , and
that it is impor tant to inves t igate in each taxon as many constituents of
dive r gent biosynthetic origin as possible . He and his associates inves tigated
s i mple phenols , flavonoids , tropolones (XXVIII) and terpenoids in conifers .
It has been possible to chemically characterize , to a degree , the conife r
families and show many generic relationships . On the basis of the chemi cal
eviden ce . Erdtman concludes that it is reasonable to merge Ducampopinus wi t h
Pinus and to remove Sciadopi t ys from the Taxodiaceae . Nicotine (XXIX) is found
in the Equisetaceae , Lycopodiaceae, Rosaceae , Leguminosae, Asclepiadaceae ,
Solanaceae and Compositae (Erdtman 1968 , see : Mabry et a1.) . It is not known
i f this compoun d is synthesized by the same rou te in each of these plant fami-
l i es . There is no reason to attach particular importance to the nicotine
 296 13

distribution, however . since other compounds specific to one ot' a few nico-
tine - 'containing families have no parallel distribut ion to i t.

Information concerning the biosynthesis of extractives being used fo r a

phytochemical correlation is i mportant sinc e simi lar or identical molecules
synthesized by different biochemical pathways should' not be utilized as evi-
dence of genet ic r elationships. Erdtman (1968. see: Nabry et al . ) points out
that anthraquinon es occur in both fungi and higher plants . One cons tituent of
lichens a nd molds . endocroc in (XXX). is known to have a bios ynthesis of poly-
keticle origin (acetate pathway); but a simil ar compound, tectoqu i none (XXXI),
found in teak wood, Tecton a grandis , is der ived from an isoprenoid napthalene .
Both the polyketide and the meva l onic acid pathway are involved i n t h e natural
synthesis of the teak constituent . It is clear that the need to collect data

y
Q
HO 0
o-Q
N
~ I
CH 3
HO'$CH'

0
.&
0

,
0
.# COOH
ceo
'"
I
0

II
0
I
CH
,

H
XVIII. ~_ Thujoplicin XXIX. Nicotin e XXX. Endocrocin XXXI. Tectoq<.linone

(o tropolone) (on olkoloid) (on onlhroquinone) (on onlhroQuinone)

on biosynthetic routes does not make the job of the chemotaxonomist any
easier, but n umerous chemotaxonomists (Alston 1966, see : SHain; Erdtman 1968 ,
see : Mabry et a1.; Herout and Sorm 1969 , see: Harborne and $\"ain; Mentzer
1 966, see : S\Jain; S0rensen 1963, see : Swain; and othe r s) have stressed the
importance of obtaining this information.

Hegnauer (1969, see : Harborne and Swa in) points out that t he ability of
a taxon to accumulate a particular natural product in measurable quantities
is more taxonomi cally important than the mere presence of that cons tituent in
trace amo unts . His opinion is contrary to the working assumpt ion of many
chemotaxonomists who labor to show quali tative presence or absence of a con-
stituent. Hore data and analysis is necessary t o determi ne if t h is interesting
observation can be r einforced .
v
Herout and Sorm (1969 . see : Harborne and S\Jain) have discussed the prob-
lem of negative evidence in che motaxonomy . Since most phytochemists are not
oriented toward ta xonomy . they tend not to publi sh negative evidence which
could be useful for taxonomic correl a tion purposes. Another hindrance , geneti-
cally controlled and environmentally induced variation \Jithin a nd between spe-
cie s, can also be confo unding in the u se of phytochemical data. There are
other limitat ions to the use of natural products chernotaxonomy; but, despite
all of them , it promises to become a sourc e fo r much new information for th e
taxonomist .
13 297

Section C. CHEHICAL EVIDENCE AS A BIOSYSTEHATIC TOOL

While most of the taxonomic conclusions resulting from studies of natural

products chemistry have been concerned with the relationships of higher cate-
gories , biosystematists us ing chemical techniques are i nterested in relation-
ships at or bel ow the generic level. In most cases the chemical. data has been
treated as a cha r acter or g r ouping of characters and , in the final summation,
is used in concert with info r mation fr om all available sources . This is be·-
cause the chemical evidence obtained in such studies usually invo l ves the
analysis of a single gro up of related compounds such as the flavono ls or the
monoterpenes ; and while the chemical determinations may be quite detai l ed and
quantitative , a relatively small amount of actual genetic variation may be
involved . The addition of chemotaxonomic information t o data from other areas ,
such as cytology , morphology , genetics , et c . can provi de a solid foundation for
systematic decisions .

I. STEPS HHICH SHOULD BE FOLLOWED IN TilE SOLUTION OF A CHEMOTAXONOHIC PROBLEH

A. Selection of chemical proced ures .

1. The facilities available and the investigator ' s experience in
chemistry \"i11 determine the amount and kind of chemical
analysis which is reasonable.
2 . The pertinent chemical and biosystematic literature should be
checked to determine which procedures and compound groupings
may be profitably studied . (If adequate work is reported in
the literature, the preliminary survey may be abbreviated or
omitted entirely .)
B. Preliminary survey .
1 . Run pilot separations on the most distantly related taxa involved
in the proposed study to determine if adequate chemical varia-
tion exists in the group .
2. Continue the pilot separations on more closely related taxa in
order to make a crude approximation of the l eve l of chemi cal
variat ion . I f a la rge amount of variation is encountered within
a species, it \dll be clear that extensive sampling of that spe-
cies will be necessary to define the parameters of chemical
variation in that species .
3 . Carry out simple chromatographic and spray reagent tests to det er-
mine the general types of compo unds \"hich are being iso l ated.
C. Environmental and physiological variation .
1 . Ninor differences in the concentration of constituents are common,
but major quantitative or even qualitative differences are some-
times observed . The stability of occur rence of constituents in
the same and different organs should be determine d from plants
which have been grown under the most uniform cond it ions poss i-
ble . Organs to be extracted should be collected at a standard
physiological stage .
D. Chemical analysis .
1. Advantage& ~and d i sadvantages of limiting the chemotaxonomic work
to chromatographic separations and chromatographic spray tests.
a. Ben efit s: Such a proce:iure i s rapid , and thus allows for the
survey of a large number o f taxa. The investment i n equip -
men t and time is minima l .
b . Problems : Characterization of structures is minimal; and
therefore, matching of constituents isolated from different
298 13

taxa may become little more than an educated guess . Lack

of information on chemical structure makes it impossible for
the investigator to estimate the gene tic bases involved in
the var iation he is measuring .
2. Identification of major constituents by chromatographic and spec-
tr.al methods increases the reliability ~ .of matching chemical
constitu ents in different taxa and provides so me information
concerning the genetic bases o f thl~ variation being studied .
Information concerning the structure and distribution of minor
constituents may be incomplete or lacking .
3 . hTith full stru ctura) determinations of e xt ractives , the signifi-
cance and distribution of these constituents can best be deter-
mined . Of course, the investment in time and equipment by the
investigator is much greater.
E. References on techniques of analysis .
1. Geissman, T. A. (ed . ) . 1962 . The Chemis t ry of Flavonoid Compounds .
The Nacmi.l1an Company . Ne\v York .
2 . Harborne, J . B. 1967. Comparative Biochemistry of the F1avonoids .
Academic Press , NeH York .
3 . Nabry, T . J ., K. R. Harkham , and N. B . Thomas . 1970 . The Systema-
tic Identification of Flavonoids . Springer-Ve rlag . Ne\v York .
4. Ribexeau-Gayo n, P . 1972 . Plant Phenolics . Hafner Publishing
Company . Nev York.
5. Stuessy, T . F . (ed . ). 1972. Ref erences and Lists of Equipment
for Systematic Studies with Flavonoids, Mustard Oils, Proteins
(Electrophoresis and Serology), and Terpenoids. (Prepared for
the A. S . P.T . symposium, " Biochemistry and the Taxonomist , " held
in August, 1972 , in Ninneapo1is, Minnesota) .
6 . von Rudlo f f, E . 1969 . Scope and limitations of gas chromatography
of terpenes in chemosy st ematic studies . See : H. K. Seikel and
V . C. Runeckles ( eds. ), Recent Advances in Phytochemistry , Vol.
2 . Appleton-Centu ry-Crofts. New York .

II . AN ANNOTATED LIST OF PAPERS RECONMENDED AS EXAHPLES OF THE APPLICATION OF

CHEHICAL DATA IN BIOSYSTEHATIC STUDIES

A. Comparison s at the generic level.

1. Arditti, J . 1969. Floral anthocyan ins in species and hybrids of
flroyghtonia , Brassavola, and Cattleyopsis COrchid aceae). Ameri-
can Journal o f Botany 56 : 59 68 . [Floral anthocyanin distribu-
t i on is shOlm to be correla t ed I.;ith generic groupings , and,
based on this and other evidence , suggestions are made vhich
\olOuld correct doubtful generic placement of some species . ]
2 . Bate-Smith, E. C. , and T. C. l.Jhitmore . 1959 . Chemistry and tax-
onomy in the Dipterocarpaceae . Nature 184 : 795-796 . [Genera
in this family are ShO\VTI to have a character i stic chemistry,
and on this basis the authors suggest c er tain species should
be moved to different genera \vith Ilihich they have a greater
chemical similarity . ]
3 . Griffiths , L. A. 1960 . A comparative study of the seed polyphe-
nols of the genus Theobroma . Biochemical Jo urnal 74 : 302-30 5.
[Theobroma species can be readily distinguished from Herrania
species by a gener i C character i stic polyphenol distrib u tion . ]
B. Comparisons at the species level.
1 . Adesida, G. A. , E . K. Adesogan, D. A. Okorie , D. A. H. Taylor,
and B. T . Styles . The 1 imonoid chemis try 0 f the genus Khaya
13 299

(Neliaceae) , Phytochemistry 10 : 1845-1853 , [Limonoid consti-

tuents are i dentified from six species . and are used to charac-
terize the species i n this morphologically poorly defined
genus , 1
2, Albach , R, F ., and G. H, Redman , 1969 , Composition and inhe ri-
tance of flavanones in citrus f ruit . Phytochemistry 8 : 127-
143 , [Flavanone distribution in Ci trus taxa is used to aid
in the definition of species groups and boundaries , as \,fell as
to verify putative hyhridity , ]
3, Conklin , H. E " and H, H, Smith . 1971. Peroxidase isozymes : a
measure of molecular variation in ten het'baceo us s pecies o f
Datura. American Journal of Botany 58 : 688-696 . [Th e taxo-
nomic affinities \Vithin this genus have been reconsidered i n
the light of peroxidase distt'ibution determined by electro-
phoretic separations of leaf proteins . 1
4. El li son, 11. L ., R. E . Alston , and B. L . Turner . 1 96 2. Nethods
of presenta tion of crude biochemical data for systematic pur-
poses, Idth particular refer ence to the gen us Bahia (Composi-
tae). Amer i can Journa l of Botany 49 : 599 - 604 . ~romographic
profiles for species of Bahia and related genera a re compared
by polygonal diagrams, affinity values , and synthet i c n umeri-
cal i ndices . ]
5. Emboden, 11 . A. , Jr . , and II . Lewis . 1967 . Terpenes as taxonomic
characters in Salvia section Audibertia . Bri ttonia 19 : 152-
160 . [Nine t een species of Salvia can be recogn ized by t he
characteristic terpene compositiol~, and this characte r i s also
a good indicator of introgression in one case studied . ]
6. Gell , p , G. H., J , G, Ha\~kes , and S . T . C. t1ri ght . 1960 . The
appl ic ation of immunological methods to the taxonomy of species
within the genus Solanum , Proceeding::; of the Royal Society of
London ( B) 151 : 364 383 . [Immunuelectrophoretic techn iques were
applied to ext ra cts from 37 Solanum species . Inte rpre tations o f
the sero log ical data tend to suppor.t conclusions from other bio -
systematic anal yses of Solanum . )
7. Giannasi , D. E. , and C. tl. Rogers. 1970 . Taxonomic signif i cance
of floral pigments in Linum (Linaceae) . I3rittonia 22 : 163-
174 . [The division of thirty Linum species into five subgen-
eric gro upings is supported by the distribution of petal caro -
tenoi ds and flavonoid s . ]
8. Johnson , B. L . 1972 . Seed protein profi l es an d th e or igin of the
hexaploid whea t s . Anerican Journal of Botany 59 : 952-960 .
[Seed protein profiles were developed e l ectropho retica lly for
wheat spec i es a t three ploid levels, and a monophy l e t ic mod e l
for the origin of the hexaploid wheats best fits t his data . ]
9. , and 11. N. Thein. 1970. Assessment of evo lutionar y
affinities in Gossypium by protein e lectrophoresis . American
Journa l of Botany 57 : 1081-1092 . [l-lith the e xception of the
placement of a few species, the accepted sub gen e ric treatment
of Gossypium is supported by electrophoretic profiles o f seed
proteins generated for twenty- five species . ]
10 . NcClure , J . t~ . , and R. Eo Alston . 1966 . A chemotaxonomic s tudy
of Lemnaceae. American Journal of Botany 53 : 8 49- 860. [Twenty-
two taxa could be identified by their specific flav onoid con-
stit uents e ve n when morphological intergradation obscured con-
ventional means of determination . Only minor variations in
f lavonoid glycosides resulted when the plants were grown und e r
various c~ltu ra l conditions . ]
300 13

11. Parks, C. R. 1965. Fl oral pigmentation studies in t he genus

Gossypium . I . Species specific pigmenta t ion patte r ns.
American Journal of Botany 52 : 309-316 . [Flower color mutants
in three spec i es of Gossypium do not obscure , beyond recogni-
tion , the basic pattern of petal pigmentation characteristic
of each species . ] .~.
12 . Payne, H. H. , R. \.,1 . Sco r a , and J. Kumamoto . 1 972 . The volatile
oils of Ambrosia (Compositae:Ambrosieae) . Brittonia 24 : 189-
198 . [T\.Jenty species from Ambrosia and two other genera con-
tain the same essential oils i n dif f erent concent r ations .
This data provided a method for disting u ishing s pecies , but
little evidence for evolutionary aff i n i ties could be gleaned
from i t. ]
13 . Roberts , E. A. H., H. Hight , and D. J . Hood. 1958 . Paper chro-
matography as an aid to the taxonomy of Thea Camell ias. The
Ne\.J Phyto l ogist 57 : 211-225 . [The polyphenol const i tuents of
a ,.;ide range of tea cu l tivars are determined , and the taxonomic
complexity of this group is underlined ; otherwise, the accepted
s ubgener i c treatment of the genus Camellia i s generally sup -
ported by the results of this study . ]
14 . Stone , D. E . • G. A. Adrouny, and R. H. Flake . 1969 . New world
Juglandaceae . II . Hi ckory nut oil s, phenetic simila r ities ,
and evolutionary i mplications in t he genus Carya . Amer ican
Journal of Botany 56 : 928-935 . [All species conta i n the same
unsaturated and saturated fatty acids, bu t in quite different
relative quantities ; and the quantitative data supports the
morphological division of the genus into two sections . ]
15. Stuessy, T. F . 1971. Systematic rel ationships in t he \.Jhi t e-rayed
species of Helampodium (Compositae) . Brittonia 23 : 177-190.
[An e x tensive cytological and chromatographic survey conducted
\.Jithin this gr oup provides additional evidence for species
delimi tations . 1
16 . Vaughan , J . G., and K. E. Denford. 1968 . An ac r ylamide gel
electrophoretic study of the seed proteins of Brassica and
Sinapis species, \.Jith special reference to t hei r taxonomic
value . Journal of Experimental Botany 19 : 72 4- 732 . [The tax-
onomic affinities of species i n these t\.JO genera are recon-
sidered in light of the results of the comparative study of
seed proteins . ]
17 . Zavarin , E., L. La\.Jr ence , and H. C. Thomas . 1971. Compositional
variations of l eaf monoterpenes in Cupressus macrocarpa, f .
pvgmaea , f . goveniana. f . abramsiana and f . sargentii. Phyto-
chemist r y 10 : 379-393. [The popu l ation structure of the
Cup res sus spec i es \.Jas determined by GLC analysis of foliar
monoterpenes.]
C. Comparisons below the species level (including hybrids) .
1 . Alston, R. E., and B. L. Turner . 1963 . Natural hybridi zat i on
among four species of Baptisia (Leguminosae) . Amer ican Jo urnal
of Botany 50 : 159-173 . [The parentage of putative natural
hybrids in Baptisia was de t ermined by chromatographic analysis .
(The Baptisia story is presented in detail i n a series of
related papers including one by T. J. Mab r y , J. Kagan , and H.
RosIer. 1965 . Phytochemistry 4 : 487 - 493 , in which the flavo-
noid constituents referred to in earl ier papers are fully iden-
tified) . ]
13 301

2. As h takala , S . S. , a n d D. F . Forward. 1971. Pi gmentation in ir i s

hybrids: the nature and development of flavonoid pigments and
t heir effect on flOl"er col or in cul t i vated hybrids of Iris ger-
manica . Can ad i an Journal of Botany 49: 1965-1973 .
[Six strikingly d i ffe r ent co l or varia n ts of Iris germanica were
shOl,," to have the same basic flower pigments; however , diffe r ences
in r elative concent rations of pigments were found to be a major
factor in deter mining flm"er color . ]
3. Belz e r, N. F., and H. Ownbey . 1971. Chromatographic comparison o f
Tr agopogon species and hybrids . American J ournal of Botany 58 :
791-802 . [Flavonoids were determined fo r species, species hybrid s,
and second generation segregates of Tragopogon. From the resul ting
segregation data , the genetic bases for some of the flavonoid e x-
t ractives were deter mined.]
4. Ca rter. L . C .• and B. G. Brehm. 1969 . Chemical and morphological
analysis of introg r essive hybridization between Iris ten ax and I .
chrysophylla . Br ittonia 21: 44 - 54. {Although n~ully obviou~
from previous stud i es , i t was determined from this analysis that
hybridization and backcrossing are occurring in sympatric pop u la-
tions of these Iris s pecies .
5. Che r ry . J. P . , F . R. II. Katterman , and J . E. Endrizzi. 1970 . Compara -
tive studies of seed proteins of species of Gossypium by gel e l ec -
trophoresis . Evolution 24 : 431-447 . [The sectional stat us of t he
genus Gossypium and t he origin of the amphidiploid species a r e
reconsidered i n the l ight of data from a n e l ectrophoretic sur vey
of seed proteins . ]
6. Craw f ord, D. J . 1970 . Systematic studies on Nexican Coreopsis ( Com-
positae) . Coreopsis mutica : flavonoid chemistry, chromosome num-
bers , morphology , and hybridization . Brittonia 22 : 93- 111. {On
the basis of these fo ur biosystematic app r oaches , Crat~ford divid e d
and defined seven var ieties of Coreopsis mutica . ]
7. 1971. Syst ematics of the Coreopsis petrophiloides -J ucid a-
teotepecensis complex. American Journa l of Botany 58: 361-36 7 .
lUsing the same methods as in his 1970 paper , he demonst r ated t hat
those Coreopsis spec i es are all part of one variable taxon with no
discernible subspeci f ic units . ]
8. Gl ennie, C. W., J. B. Har borne , G. D. Rowley, a nd C. J. Harchant .
1 971. Corre l ations between flavonoid chemistry and plant geog-
raphy in the Senec i o r adicans complex . Phytochemistry 10 : 2413-
2417 . [Flavonoids determined for forty - fo u r clones from twenty -
tHO species were no t correlated with kn own morphological or cy t o-
logical pattern s i n this group , hut rath er t~ere correlated with
the plant geography of these species . ]
9. Levin , D. A., and B. A. Schaal . 1970. Ret iculate evolution in Ph l ox
as seen t h rough protein electrophoresis . American Journal of Bo t any
57 : 977 - 987. [ Electr ophoretic profiles of seed proteins we r e suc-
cessfully used to verify the pu tative hybr i dity of some stabili zed
t axa in Phl ox. ]
10. 1972 . , .... Seed protein polymorphism i n Ph l ox pilosa
(Polemoniaceae). Br itton i a 24 : 46-56 . [Evi dence is
provided to show t ha t seed protein polymorphism in 1:. pilosa is
caused by introgression with f . glaberrima . ]
11. Le vy , H. , a nd D. A. Levin . 1 971. The origin of novel flavono i ds i n
Phlox allotetraploid s . Proceedings of the National Academy of
Sciences 68 : 1627-1 630 . [Flavonoids not f ound in diploid progene -
tors were identified from their tetrap l oid products , and a me chan i sm
302 13

is presented to explai n the origin of these hybrid spec i f ic com-

pounds. ]
12. Mirov , N. T . 1956. Composition o f turpentine of lodgepole X jack
pine hybrids . Canadian Journal of Botany 34: 443-457. [The com-
position of terpenoid constituents in pare~t trees, putative natural
hybrids, and a synthetic hybrid are determined . The putative hy-
bri ds are shown to be chemically similar to the synthetic hybrid on
the basis of these constituents . ]
13. Mooney, H. A. , and I.J. A. Emboden, Jr . 1968. The relationship of ter-
pene composition , morphology and distribution of populations of
Bursera microphylla (Burseraceae) . Brittonia 20: 44 - 51. [The
variation in morphology and chemistry is explained by the distri-
bution pattern and the presence or absence of adjacent populations
of other species of Bursera.)
14 . See1igmann , P . , and R. E. Alston . 1967 . Complex chemi cal variation
and the taxonomy of Hymenoxys scaposa (Compositae) . Brittoni a 19 :
205-211. [Chemical patterns \,'ere found to be geographically con-
sistent , but not related to the morphology or taxonomy of this
group of closely related species. Thus , the authors conclude that
one variable species is involved . ]
15. Hall, J . R., and T . H. Ifhitaker . 1971. Genetic control of leucine
aminopeptidase and esterase isozymes in the interspecific cross
Cucurbita ecuado r ensis X C. maxima . Biochemical Genetics 5 : 223-
229 . [Similar , but not identical, proteins \olere found in each spe-
cies , and each enzyme Has found to be under the control of a co-
dom i nant allele . ]
D. Studies concerning the influence o f environment on phytochemicals .
1. Abrahamson , 1-,1. G., and O. T. Solbrig . 1970 . Soil preferences and
v a r iat ion in flavonoid pigments in species of asters . Rhodora 72 :
251-263 . [Environmentally induced variation i n alcohol so l ub l e
secondary compounds was considerably greater than that attributed
to genetic differences in asters Hh i ch showed similar soil preferen-
ces . ]
2 . Adams , R. P . 1970 . Seasonal variation of terpenoi d constituents in
natural populations of Junipe.rus pinchot ii Sudw . Phytochemistry 9:
397-4 02 . [Summer to winter variations in the relative composition
of volatile terpenoids was observed . Summer collections were found
to be more variable than lolinter collections . }
3. Ball , G. A. , Jr . , E. O. Beal , and E . A. Flecker . 1967. Variation of
chromatographic spot patte.rns of two species o f clonal plants grown
under controlled environmental conditions . Brittonia 19 : 273-279 .
[Chromatograph i c spot variat i on observed in clones of Spirode1a
gr own under a variety of cultural condition s are ascribed to en
vironmental variation . ]
4 . Parks, C. R., S . S. Sandhu , and K. R. Nontgomery . 1972 . Floral p i g-
mentation studies in the genus Gossypium . IV . Effects of differ-
ent growing environments on flavonoid pigmentation . American
J ournal of Botany 59 : 158-164 . [Different growing regimes had
l i ttle effect on floral flavonols in several lines from each of
the four species studied , but evidence is presented that leaf
constituents may be less stable . 1
13 303

Sect i on D. CHRONATOGRAPIIY--BASICS AND DEFINITIONS

Chroma tograph y'~ is a fa mily of separation techniques of great resolving

pm.er , and of ten of considerabl e complexi ty, us ed to sepa r ate a wide assort-
ment of molec ules .

A. Some physical prop e rties of molecules which make ch r omatogr aphy possible
are primaril y the fo llowing.
1 . Adso rption is the tendency for a molecule and a finely divided solid
to become at t ache d .
2 . Sol ubil ity i s the tenden cy of a molecule to dissolve in a liquid .
3 . Volatility is the tendency of a molecule to evaporate .
Two of th ese propert ies of mol ecules (or one repeated twice) are
exh i bited when the chromatographic mixture is placed in a movin g
o r dynamic experimental sit uation .
B. Inte r actions fo und i n the chromatographic separation .
1. Partition i s the tendency of a s ubstanc e to dis tribut e itself between
two non-mixing liquids, a liqui d and a so lid , a liquid and a gas ,
etc .
2 . Phases are the comb ina t ions of the solids, liquids and gases between
IVhich the substance being separated is distributed .
In a ctua l practice a moving phase pass es over a stationary phase
a nd is, in as far as it is poss i b le, in equilibrium with it . The
phas es are chosen so that the compon ents i n the mix t ure move at
varying rate s . Chromatographic separations are possibl e because
of t hese di fferences in migration r ates .
C. Major ch romatograph ic systems.
1. Principles .
a . Pa rtition chromatography involves a moving l iquid phase over a
stationary liquid film on a suitable suppor t.
b . Adsorption chroma tography involves a moving liquid and a solid
ad sorbing surfa ce (n ot a s upport for a liquid film) .
c . Gas -liquid- partition chromatography (gas chromatography or GLC)
involves a moving gas and a stationary l iquid film.
Note: In man y cases i nvolving the common l y used methods of
chromatography (pape r, l'LC o r co lumn) , it i s not readi l y
possible to de termine \~hethe r partition or adsorption phenomena
are t aking place . In fact, partition and adsorption e vents are
oft en co-occu rring .
2 . Pract ices.
a . Paper chromatography involves a movin g liquid a nd paper as a n
adsorb e nt or support .
b . Thin-laye r chromatography involves a moving liquid and a fin e ly
d ivided solid as th e adsorbing or suppor t sur face . Structural
s upport is provided by a gla ss plate to I.hi c h the finely divided
solid is bound .
c. Co lumn chroma t og raphy involves a movin g liquid which passes through
a gl ass column packed with a finely divided solid a dsorben t.
D. Nor e definitions associa t ed with chromatographic separations .
1. Adso r ben t . S tationar y phase in adsorp t ion chroma tography .

*Discussion of chromatography is taken from Bobb itt et al . (19 68) , a nd it is

recommended that the student refer to that readable and concise treatment.
 304 13

2. Bands . Substances separ ating and moving through a column in column

chromatography .
3. Developmen t. Hovement of the moving phase over the adsorbent or
support.
4. Elu tion . Washing the sepa ra ted substances from the adsorbent or
support.
5. Elution analysis . Analysis of a mixture by progressive elution from
a column.
6. Effluent. The l iqui d which flows out of a co lumn .
7. Gel electrophoresis . A type of chroma tography in which the solutes
migrate in an electrical field .
8. Gradien t e l ution analysis. As in elution analysis , but solvents are
changed during development .
9. Irrigant (solvent) . The moving phas e in paper, TLC or column chro-
matography.
10. Rf value. The ratio derived in paper and TLC chromatography by
dividing the movement of the separated substance by the total
movement of the solvent (solvent front) .
11 . Solutes (collectively-sample). Substances being separated .
12. Spots . Separated subs t ances appearing on paper or TLC ch roma tograms .
13 . Spray rea gent . Reagent sprayed on a paper or TLC chromatogram to aid
in visualization.
14. Visualization. Treatment of a chromatogram which l ends spots visible .

Section E. SONE COM}!ENTS ON PROTEIN ANALYSIS

l,.,rhile the importance of protein analyses in chemotaxonomy has been pointed

, out , a discussion of methods of protein ana l ys i s is beyond the scope of this
gen eral review . The data obta ined by electrophoretic separations of crude seed
or other protein ex tracts are used as supportive evidence in systematic inter-
pretations, much as paper chromatograms of unknown phenolic mixtures are us ed .
Various degrees of analytical r efinements may be ca rried o u t in these studies ,
ranging from simple purifications of the crude extract to enzyme purifications
or even " finge rpr inting " of a specific pro t ein from different t axa .

Another major approach to pro t ein ana l ysis i s serology. Serological

reactions occur ",hen foreign proteinaceous (usually, but not necess ar ily) sub-
stances , antigens, are injected in to an animal , and these antigens elicit the
formation of other presumably proteinaceous s ubstances , antibodies , in that
host animal . The antibodies precipitate (agglutinate) or otherwise affect the
ant i gen . The host is usually a domestic anima l, of ten a rabbit . In the mo s t
common serological test , the quantitative p recipit in test, aliquots of the
antigen ext r act are mixed in varying dilutions with the an tibody preparation
(antiserum) . The amount of precipitate produced is proportional to the
" strength" of the reaction; that is , the greater the simi la rity of the test
substance to the antigen , the stronger (more p r ecipitate produced) is the
r eaction . Immunoelectrophoresis is a r efinement of the precipitin test , and
allows for the generation of qualitative data . Ext racts are prepared and
subjected to agar-gel electrophoresis. Antiserum is added to a trough paral-
lel to the electrical tract of antigen separation, and the ant i gens and anti-
bodies diffuse into the agar. l"There the appropriate antigens and antibodies
mee t, stabilized precipitates are formed which are detectable by various
means. The precipitates become visible as arcs . By this means the reactions
of more than one protein in the antigen mixture may be studied . This is not pos-
sible by the s traigh t pr eci pitin test . Recently. data from this method have
13 305

been applied increasingly to the solution of taxonomic problems .

CHEMICAL EVIDENCE LITERATURE

Abbo tt, H. C. de S . 1886. Certain chemical const ituents of plants considered

in relation to their morpho l ogy and evo l u tion. The Botanical Gazette 11:
270- 272 •
Alston . R. E. , and B. L. Turner. 1963. Biochemical Systematics . Prentice-
Hall , Englewood Cliffs . New Jersey .
Bate- Smith , E. C. 1948. Paper chromatography of anthocyanins and related
substances in petal extracts. Nature 161 : 835-838 .
Beale , C. Ho, J. R. Price • . and V. C. Sturgess . 1941. A survey of anthocyanins
VII. The natural selection of flo\"er color. Pr oceedings of the Royal
Socie t y of London (B) 130: 113- 126 .
Bobbit, J. No, A. E. Scht"arting, and R. J . Gr itter. 1968 . Introduction to
Chromatography. Reinhold Book Corporation . New York .
Chester, K. S. 1937 . A c r itique of plant sero logy (continued) Part II.
Application of serology to the classification of plants and the identifi-
cation of plant products. Quarterly Review of Biology 12 : 165-190.
Consden , R., A. H. Gordon, and A. J . P . Nartin . 1944 . Qualitative analysis
of proteins : A partition chromatographic method using paper . The Bio -
chemical Journal 38 : 224 - 232.
Ge11 , P . G. H., J. G. Hawkes , and S. T. C. Wrigh t. 1960 . The application of
immunological method s to the taxonomy of spec i es within the genus Solanum .
Proceedings of t he Royal Society of London (B) 151: 364-383.
Gibbs, R. D. 1958. Chemical evolution in plants . The Journal of the Linnean
Society of London . Botany 56 : 49 - 57 .
Harhorne, J ., B., and T. SI.ain (eds . ) . 1969. Perspectives in Phytochemistry.
Academic Press. New York.
_--;-==.' D. BOulter, and B. L. Turner (eds . ) . 1971. Chernotaxonomy of the
Leguminosae. Academic Press. New York .
Hegnauer, R. 1962, 1963 , 1964, 1966 , 1969 . Chemotaxonomie del.' Pflanzen.
Vols . 1- 5 . Birkhauser Verlag. Basel.
Heywood , V. H. (ed . ) . 1971. The Biology and the Chemistry of the Umbelliferae.
Academic Press . New Yor k .
Kooiman , P . 1969 . The occurrence of asper u10sidic glycosides in the Rubia-
ceae. Acta Botanica Neer1andica 18: 124-137.
1970. The occurrence of iridoid glycosides in the Sc rophu1aria-
ceae . Acta Botanica Neerlandica 19: 329 - 340 .
Lawrence , \1'. J . C. , J . R. Price, G. 1'1 . Robinson , and R. Robinson . 1939. The
distribution of anthocyanins in flowers , f r uits and leaves . Philosophical
Tra nsactions of the Royal Society of London (B) 230: 149-178 .
Mabry , T. J . , R. E. Alston, and V. C. Runeckles (eds . ) . 1968 . Recent
Advances in Phytochemistry . Vol . 1 . Appleton- Century-Crofts . New York .
HcNair, J. B. 1935. Angiosperm phylogeny on a chemical basis . Bulletin of
the Torrey Botanical Club 62: 515-532.
1945. Some comparisons of chemical ontogeny with chemical phy-
l ogeny in vascular plants . L10ydia 8: 145-169 .
Mirov, N. T. 1967 . • ~The Genus Pinus. The Ronald Press Company . New York .
Reinhold , L ., and Y. Liwschitz (eds .). 1968. Progress in Phytochemistry .
Vol. 1 . Interscience Publishers. New York.
Schafer. C. 1964 . Biogenetic and chemotaxonomic considerations about bisbenzy-
lisoquinoline alkaloids, p . 4-7 . See: H. R. Arthur (ed . ). Symposium on
Phytochemistry . Hong Kong University Press.
Swain, T . (ed.) . 1963. Chemical Plant Taxonomy . Academic Press . New York .
1966 . Compara tive Phytochemistry. Academic Press . New York.
306 14

Chap ter 14 . PHYS IOLOGICAL A.1'm ULTRASTRUCTURAL EVI DENCE

Physiological evidence, including the biochemical , developmental and bio-

physical aspects, provides a basic set of data Hhich ~are of increasing sig-
ni f icance to taxonomy today . This is particularly true for those metabolic
characters related to chemical systems, biosynthetic patIH~ays, and primary
and secondary products . Chemical characters are used to distinguish hybrids
and species as Hell as to provide evidence in the characterization of l arger
taxa; e . g ., Hatch-Slack patll\~ay of malic acid carboxylation is possessed by
many tropical species of Panicoid grasses , and is f ound in a number of tropi-
cal species in the Cyperaceae, Arna ranthaceae , Chenopodiaceae, and Portulaca-
ceae . According to Laetsch (1969) these species are also characterized by
having a bundl e sheath and dimorphic chloroplasts . (See Chapter 13 on chemi-
cal evidence . )

As more i s learned about plant development, grm!th features should be of

greater value to systematics, particularly in t ropica l species in \~hich many
types o f groHth patterns and mechanisms occur (see Chapter 6). According to
Toml i nson and Gill (1973) "detai led groHth analysis is scarcely begun but
offers unlimited scope for future Iwrk." An adequate and applicable classi-
fication of these patterns t/Ould inuneasurably aid our present understanding
of tropical taxonomy. Features of dormancy and germination have been used
in identification and characterization of taxa, but a systematic study of
these c haracteristics should result in a more effective and efficient means
of identification of seeds and seedlings, as \·;ell as provide a sounder b asis
for classification (see Chapter 6) . Development of individual flOl~ers and
inflorescences is used in delimitation of larger taxa , e.g., the centrifugal
stamens in the DillQniidae and centripetal stamens in the Rosidae (see Chap -
ter 6) . The ovular developmental patterns , gametogenesis, sporogenesis ,
embryogeny , embryogenesis , and endosperm characters have been used as ciiag-
nostic characters in distinguishing larger taxa , e . g . , monosporic P.1egasporo-
genesis in the Onagraceae and helobial endosperm development in the AlisMa-
tidae (see Chapters 8 and 9) .

The physical processes of diffusion , osmosis, plasmolysis , and imbihi-

tion are so ,dde-spread that they are of no apparent significance to taxonomy .
The follOl·,ing biophys ical features, hO\Jever . might be of some taxonomic sig-
nificance primarily as structural or cheMical evidence : uptake against a
concentration grad i en t in many species; selective absorption , as selenium by
species of Astragalus ; accumulation of bases , as in the leaves of most mem-
be rs of the Nyrtaceae; or gas exchanp.;e features , as through the pneuma to-
phores of some species of Lud~:igia or through the epide rmis of submerged
species of Potamogeton . The types of conduction mechanisms or transport
f10H systems, Hhen worked out, should 5hOl" taxonomic r elationships .

Nechanical functions are so closely related to form ( " structure is func -

tion " ) that morphological rather than physiological features have long b een
used by systematists. The amount and distribution of sclerenchyma and co1-
lenchyma are definitely related to supportive systems. Anchorage is related
to type of root system , as tap, fibrous , or fascicled . Protection i s afforded
by stinging trichomes , spines , thorns, prickles , and silicic and ox alic acid
crystals . All of these characters are treated as structural (see Chapter 6) .
14 3D)

Reproduction, involving reprodu ctive features and pollinatio n- fertiliza-

tion types, has been used taxon omically from .J. broad b iological standpoint
(see Chapters 6 and 12) . In general, physiological adaptations are usually
expressed as structural characters in taxonomic ,,,ark .

Even though most physiological evidence is treated by the chemotaxonomist

as chemical chari"lcters and by the traditional systematist as morphological ,
anatOl!lical, or cytological, the taxonomist should be 31;'are of nel'] developments
in plant physiology . Particularly they should have an al"areness of research
in the field of physiological mechanisms such as those of. or in , organ initia -
tion and development, mitotic and meiotic control . active ahsorption . mineral
accumulation, and regulatory controls . These mechan i sms, ~-Ihich Ivill usually
be expressed as chemical or physico-chemical systems , can he of taxonomic
value in the de f inition and delimitation of family and highe r taxa as Hell as
providing evidence of fundamental relationships. Developmental response stud-
ies I-Jill determine the reliability and meanin f,fulness of many of the diagnost ic
characters used in classification and identificat ion .

Since phys iological evidence has been so disjoined and dismembered in its
treatment by taxonomists (and by the authors of this text ! ) the functio nal
features described in this book are listed here Hith the chapters in Hhich
they occur :

Structural expressions of phys iology in ChaptE'_r (i on morphology and

Chapter 7 on anatomy .
Developmental aspects of physiology in Chapter 8 on palynology and
Chapter 9 on emhryology .
Biophysical and reproductive aspects of p hysiology in Chapters 12 and
15 on reproductive biology and ecology, res p ectively .
Chemical aspects of physiology in Chapter 13 on chemotaxonomy .

Ultimately. physiological conditions and processes Hill be directly cor-

related Hith structural characteristics at the cellular level. Although cyto-
logical evidences have historically been derived mostly from the structure and
behavior of chromosomes. i. e ., karyology (see Chapter 10) . the I~idespread use
of the electron mic roscop e has shOlm the cell cytoplasm to be a highly organ-
ized region composed of many structurally complex and interrelated membranous
organelles . Broadly based comparative ultrastrllctural information on the
plastids, endoplas1'lic reticulum, Golgi apparatus , mitochondria . and vacuoles
is available for very Ie\-! species . In addition, the fine structure of plant
cell \Valls (including pitting) and pollen grain exines (Chapter 8) are areas
of increasing importance .

An example of the significance of the electron microscope in providing

additional characters to angiosperm classification is illustrated by the
investiRat ion of Behnke (1971 , 1972 , 1973) on cells of the phloem tissue . It
has been shown that sieve tube elements can be separated on the basis of the
contents of their plas tids as observed Hith the EM . Sieve tube elements con-
tain plastids of either .the S-type or th e P-type . S-type plastids store
starch only I~hereas plastids of the P-type contain protein alone or protein
and starch . It has been demonstrated, furthermore . that there may be uniform-
ity of plastid type Ivithin taxonomic groups as high as the rank of class.
HOl1ocotyledons are thus far knOlm to possess only P-type plastids . The taxo-
nomic placement of a plant that varies in plastid type from remaining member s
of the taxon to ",hich it has heen assigned is sllspect . As more plants are
studied the phylogenetic trends in plastid morphology may be clarified .
308 14

Although the physiol ogical and ultrastructural characteristics of vascu-

lar plants have not proven, and may never prove, to be of the same taxonomic
and evo lutionary significance as in the Thallophyta (see Klein and Cronquist ,
1967). data and info r mation derived from these fields should be critically
evaluated in making taxonomic judgments . The synthesis of data f rom all
fields represents one of the real challenges to the systema tic botanist.

PHYSIOLOGICAL AND ULTRASTRUCTURAL LITERATURE

Behnke , H. D. 1971 . Sieve-tube plastids of Nagnoliidae and Ranunculidae in

relat ion to systematics . Taxon 20 : 723-730 .
1972 . Sieve-tube plastids in relation to angiosperm systematics--
An attempt tm"ards a classification by ultrastructural analysis . Botanical
Revi eH 38: 155-197 .
1973 . Sieve-tube plastids of Hamamelidae . El ectron microscope
investigations Hith special reference to Urt icales . Taxon 22 : 205 - 210 .
---;-c' and B. L . Turner . 1971. On specif i c sieve- tube plastids in Cary-
ophyllales . Further investigations wi th special reference to the Bataceae .
Taxon 20 : 731-737 .
Johnson , Sister Clement, and H. V. Brm-m . 1973 . Grass leaf ultrastructural
variations . Araerican Journal of Botany 60 : 727 - 735 .
Klein , R. N. , and A. Cronquist . 1967 . A consideration of the evolutionary
and taxonomic significance of some biochemical , micromorphologica l , and
physiological characters in the thallophytes. Part I. Chemicitl , micromor-
phological and functional phylogenetic criteria . The Quarterly Review of
Biology 42 : 108-210 .
Laet s ch, \oJ . H. 196 9 . Relat i onship bethTeen chloroplast structure and photo-
synthetic carbon-fixation patllh'ays. Sc ience Progress Oxfo rd 57 : 323-351.
Levitt, J . 1969 . Introduction to Plant Phys i ology . C. V. Mosby Company .
St . Louis , Missouri .
Plant Physiology . 1926+. Baltimore : Official publication of American Society
of Plant Physiologists .
Steward. F. C. (ed . ) . 1959. Plant Physio l ogy . (Severa l vo l umes) . Academi c
Press , In c. Net., York .
Tomlinson , P. B . • and A. N. Gill. 1973 . Growth habits of tropical trees :
some guiding principles . In : Tropical Forest Ecos ystems in Africa and
South America : A Comparative RevieH. Heggers. B. J . , E. S. Ayensll , and
\-1 . D. Duckworth (eds . ) . Smithsonian Institution Press . Hashington, n . C.
 Chapter 15. ECOLOGICAL EVIDENCE

Ecology is basic to taxonomy in understanding (1) the distribution of

taxa and composition of floras, (2) the genetic and phylogenetic relations
of taxa, and (3 ) the variation in populations and evolutionary adaptations .
The three major sections in this chapter are: A. HABI TATS wh i ch are classi-
fied primarily according to edaphic and vegetation r elations ; B. ENVIRON-
MENTAL RELATIONSHIPS which are treated as biotic, abiotic, spatial, and tem-
poral ; and C. ADAPTI VE FEATURES \"hich presumably are or have been related
to environmental composites or mosaics of factors . Distribution principles,
ranges , and biomes are described i n Chapter 16 on geography; communities are
treated in Chapter 17 on fie l d studie s .

Sec tion A. HABITATS

Habitat is the " home " of the individual organism or its native environ-
ment . Habitat descript ions should be brie f but as comprehensive as possible.
For pioneer and early successional species the edaphic (A- F and M- P be-
low) and the geomorphic features (Chapter 17) shoul d be indicated. For late
successional and climatic species the community or vegetation type (G-L)
should be identif i ed s i nce the vegetation represents the mosaic of environ-
mental factors at a given time in a given place . Reg ional climatic data can
usually be obtained from local weather stations, if the locality is Riven for
a collection . Locality data is a basic part of all collection in formation.
The geomorphic feature will give clues to microclimatic conditions. This
classification system provides one word adjectives for the major habitats in
North America north of Mexico . The terms ,"ere selected or adapted from
Clements (1902) , Gray (1967), and Dansereau (1957) .

A. "Tater - Hydrophytes (hydor: water)

~enthophilous . (benthos : sea-bottom) Dwelling submerged in and
attached to the bottom of estuaries , sounds , seas .
Crenophilous . (krene: spring) D,,,el l ing in springs and seepages.
Hal oplanktophilous. (hals , halos: salt + planktos: free -floating)
Dwelling on the surface of salt water .
Linmophilous . (limne: lake , pool) Dwe lling in lakes .
Li mnoplanktophilous . (limne : lake , pool + planktos : free - floating)
Dwelling on the surface of fresh water .
Oceanophilous . (oceanus"': ocean) Dwelling in the ocean or marine
'~aters.
Potamophilous . (potamos: river) Dwelling in r ivers .
Rheophilous . (rheos : stream, current) DHelling in running ,. . ater .
Stasophilous . (statos : standing, fixed) Dwelling in stagnant water.
Tipho ph i lous . (tiphos: standing ,vater , pool) Dwelling in ponds or
pools .

B. Clay or Nud - At.:gi-llophytes (argilla*: white clay, potter's clay)

Limiphilous. (limus* : mud) Dwelling on mud .
Pelophilous . (pelos: clay) Dwelling on c.lay .
Pe l ochthophilous . (pelos: clay + chthon: soil , earth) DHell i ng on
mud banks .

*Latin, others are Greek.

310 15

C. Sand - Psammophytes (psammos : sand)

Aigialophilous . (aigialos : strand, sea-shore) Owelling on beach
strands or sand flats .
Amathophilous . (amathos: sand, sandy soil) JlI-le lling on sand hills ,
plains , or scarps.
Anemodophilous . (aoemos : Hind) Dwelling in . bIOI-wuts.
Cheradophilous . (alluvium, silt, detritis) Dwelling on wet sandbars.
Syrtidophilous . (syrtos: \~ashed along by a stream, alluvial)
Ot'Jelling on dry sandbars .
Thinophilous. (this, thinos : beach, shore. sand-heap ) Dwelling on
sand dunes .

D. Rock - Petrophytes (p etra : rock)

Actophilous. (acta* : seashore, promontory) Dwell ing on rocky sea-
shores .
Calciphilous. (calyx , calcis* : lime) D\~elling on l i mestone .
Chasrnophi lo us . (chasma: chasm , gul f) Dwelling in rock crevices or
joints .
Cremnophilous. (kremnos : precipice , cliff) [K.,,-ell i ng on cliffs .
Granitophilous . (graniticus : pertaining to plants grOl.,,-ing on gran-
ite . Hod . Latin) Dlvelling on granite .
Gypsophilous . (gypsos: ,.,,-hite plaster , gypsum) Th.,,-elling on chalk,
o r alkali or limestone.
Halophilous. (hals, halos : sea, salt) Th.,,-elling on salt flats.
Lithop hilo us. (lithos : stone) Dwelling on flat rocks .
Petrodophilous . (petra: rock) DHelling in bould er f ie lds or flats.
Phellophilous . ( phel leus : stony ground) DHelling on stony g r ound .
Siliciphilous . (silex, silicis* : any hard stone, f l int) Dwelling
on qua r tz or sandstone .

E. Humus - Saprophytes (sapros : decayed, rotten)

Oxygeophytes (oxy: sour + geo: earth)
Coprophilous. (kopro s : dung) D\~elling in or on dung .
Eurotophilous. (euros, eurotos : mold , decay) Th.,,-el1ing in duff or
humus .

F. ~ - Oxylophytes (oxys: sour)

Sphagnophilous . (sphagnos : a kind of moss) DHelling in peat or
sp hagnum .

G. Harsh - Helophytes (helos : marsh)

Helophil ous. (helos : marsh) Dwel l ing i n fresh water marshes .
Limnodophilous . (limnodes : marshy) Dwelling in salt marshes .

H. Grass l and or Prairie - Psilophytes (psilos : bare, naked)

Coryphophilous . (koryphe : head, top) DI.,,-elling in alpine meadows .
Leimophilous . (le imon : meadow) DHelling in 10\~ g rasslands. mea-
dows , or coastal prairies .
Pedion ophilous. (pedion : flat , open country) Dwelling in the
plains , or short grass l ands .
Psilophilo us . (spilos: bare, naked) Th.,,-elling in prairies .

I. Savannahs and Open \Joodlands - Pyrophytes (pyr, pyros : f ire) and

Orgadophytes (orgas , orgados : field , meadow)
 - .
15 311

Orgadophilous . (orgas, orgado : field , meadO\ol) Tholelling i n open

wood lands .
Pyrophilous . (pyr , py ros : fire) Dwelling in burned-over areas or
soils . savannahs .

J. Scrub and Shrub - Fruticiphytes (frutex, f r u ti c i s : shrub)

Aitha l oph1 l ous . (aithales : everg r een) Ol..re lling in e verg r een
thickets .
Lochmodoph ilo us . (lochme: thicket) D\<Jelling in dry thickets .
Ptenothalophilous . (ptenothales : " de ciduous " ) Dwelling in deciduous
thicket s .
Sclerophilou s . ( sk leros: tough , hard) D\<Je lling in chapar r al.
Xeropooph i lou s . (xeros : dry + poa : g rass) DI~elling in heath balds.
thickets , o r pocosins.

K. Forests - Dendrophytes (dend r on : tree )

Ai phy llophilous. (aiphyllos: evergreen le af) DI'} elling i n broad-
l ea ved everg reen forest s .
Ancophilous . (ankos : mountain glen . valley) Dwe l ling in canyon
or cove forests .
Conophorophilou s . (konos: coen + phoras : bearin g ) DHelling in
coniferous forests .
Hel ogadophilous. (he los : marsh , swamp + orgas . orgados: meadm,',
fie l d) Dt..relling in swamp forests.
Hy l ophilo us . (hyle : t..rood , forest ) D\~elling in forests.
Ombrohyl ophi l o us . ( ombro s : rainstorm + hyle: ,,,ood, forest)
Dwe l ling in rain forests.
Xerohylophilous. (xer os : dry + hyle : wood . fo r es t ) Dwel l ing in
dry , deciduous forests .

L. Tundra - Crymophytes (crymos: icy cold , c hill)

Crymophilou s . Dwelling in polar bar r e ns .

>! . Ice or Snow - Psych rophy tes (psychros: cold , frigid)

Psychrophi lo us . (psychro s : cold , frig i d) Dwelling in cold pl aces .

N. Desert - Eremophytes (eremi s : dese rt)

Chersophilous . ( che rsos: dryland) DI..rell in g in dry waste pl aces .
Eremoph ilous . ( e remis: desert) D~Jelling in deserts .

O. Hill or Hount ain - Lophophytes (lophos : crest , ridge) or Orophytes

- -(oros: mountain)
Acrophilou s . (akron : top, summit) DI.;elling on peaks .
Ancophtlous . (ankos : mountain glen, valley) Dwelling in canyons.
Lophophilous . (lophos : c r est , ridge) Dwelling on hill s or slopes.
Orophil ous . (oros : mount ain ) Dwelling on mountains.
Pagophi l ous. (pagos : hill) Dwelling in foothills .

P. Distu rbed - An throPQphy t es (an th ropos : man )

Agrophilous . (agros : field) Dwellin g in cultivated fields .
Aletophilous. (aletes: wanderer) Dwelling on roadsides, waysides,
railroads .
Ch ledophilous. ( c hledos: mud, dirt , debris) Dwel l ing in waste
place s or rubbish heaps .
312 15

Section B. ENVIRONNENTAL RELATIONSHIPS

Ecological characteristics as expressions of the environmental r elat ion-

ships of species should be determined or o b served . Populations have tole r-
ance limits for each of the factors ; e.g. , l ight , heat, moisture, nutrients .
Even though it i s practically impo ss ible to determine·· experimentally the
regime for any factor , i. e ., temperature amplitude or t olerance limits, the
environmental cond i t ions for the best grm.,rth and reproduction shoul d be noted
by every obse rvant field worke r. The acuity of observation of plant behavior
under d i f ferent environmental conditions \.,rill determine , to a great extent ,
the degree of success in t r ansplant and field variation studies and in the
understanding of clinal relationships and flo r a or commun i ty dynamics; e.g . ,
phenology , periodicity. Species should be characterized ecologically con-
sciously and intelligent l y , if impo ssible experimentally, rather than in-
stinctively or intuitively (see Ecology Exercise for ecological characteris-
tics summary) . Th is treatment is basically a classification of factors \·,ith
definitions of the characteristics .

I. BIOTIC. Pertaining to the living environment.

A. Genet i c. Relations pertaining to hereditary rela t i onships .
1. Genetic. Pertaining to immediate gene relationships of organisms .
2 . Phenetic . Pertaining to the observable characterist ic s of organ-
isms, not necessarily closely related genetica lly .
3 . Phyletic . Pertaining to racia l relationships of organisms .
4 . Phylogenet ic . Pertaining to racia l history relationships of
organisms.
B. Nutri tiona l Relations. Pertaining to nour i shment of organisms .
1 . Autoparasite . Paras ite living on a parasite ; organism depen-
dent upon a parasitic host fo r its nourishment; e . g ., mis tle-
toe on mistletoe.
2 . Autophyte (autotrophe). Plant that makes its own foo d; a self -
nour ishing plant .
3. Hemiparasite . A partial parasite, usually chlorophyll bearing,
o ne that obtains part of its nourishment from its host ; e . g .,
Comandra , Euphras i a .
4 . Parasite . Plant that obtains its nourishment from a living
plant , its host .
5. Saprophyte. Plant that obtains i ts nourishment from dead organic
matte r.
C. Social Relations. Pertaining to the grega r io usness of individual
organisms within populations ; intrapopul at i onal relationships;
sociability . This classification of sociability is based on
that of Braun- Blanquet (1932) Hhich is difficult to apply i.,rith-
out tra i ning by an individual who has had exper ience with the
system .
Soc . 5 Plants occurring in great crowds ( pur e populations) .
Soc. 4 Plants i n small colonies , i n extensive patches , or forming
carpets .
Soc . 3 Plants in smal l patches or cushions.
Soc . 2 Plants grouped or tufted .
Soc . I Plant s growing singly .
0 . Symbiotic Relations . Pertaining to the environmental relationships
between two or more species ; interpop ulational r e l ationships .
15 313

1. Nutualism . Si tua tion in which two species a r e dependent upon

one another for survival or perpetuation of each; e . g. ,
alga--fungus in the lichen, yucca-pronuba moth .
2 . Commensalism . Sit uation in \~hich one species helps ano ther
without harm or benefit to itself; e . g . , heliophyte provid-
ing the shade for a sciophyte .
3 . Neutralism . Seemingly no relationships between two or mo re
species within a community.
4 . Competition . The struggle or rivalry betl. . een tIm or mo r e species
for a common requirement or resource ; e.g ., moisture , light .
5. Amensalism . Situation in which one species definitely inhibits
another wi thou t harm to itself; a lle l opathy; e.g. , Juglan s
secretes substance that inhibits development of many species .
6 . Parasitism. Situat ion in Iv-hich on e spec ie s definitely harms
anoth er by obta i ning its no urishmen t directly from it .
E. Reprodu c t ive Relations . Perta i ning to the i ncrease in popul ation .
1 . Apomixi s . Asexual types of reproduction involving either
agamospermy or vegetative propagation .
2 . Amphimixis . Sexua l reproduction involving union of tIm gametes .

II. ABIOTIC . Pertaining to the physical environment .

(Relative degree of tolerance or amplitude in ecology is e xpressed
by the prefixes " steno- It , mea ni ng narrow ; "meso-It, mean i ng middle ;
and " e ury- It, meaning wide; e .g . , stenothermic - meso thermic -
eurythermic refers to temp erature, st enohyd ric - meso hydric -
euryhydric refers to water . stenophotic - mesophotic - euryphotic
refers to light . )
A. Climatic Relation s.
1 . Temperat ure Relations.
Cryotherm or Hekistotherm. Plant growing in minimum heat. as
in arctic-alpine zones; tundra plants .
Ni crotherm . Plan t gro\. . ing i n little heat. as in the cold
temperate zon es; boreal fo r est pl ants.
Hesotherm. Plant grOl~ in g in modera te heat, as in t he warm
temperat e zones ; deciduou s hardwood plants .
Ne gathe rm. Plant growi ng in gr eat heat, as in the tr opical
zone ; trop i cal r ain forest plants .
2 . Hoisture Re l ations .
Hydrophyte . Plant gr owing i n water either as free floating ,
submerged , f l oating and anchored, or emer gen t and
anchored .
Hygrophyte . Plant grm. . ing under highly humid conditions .
Hesophyte . Plant growing under medium moisture condit ions .
Xerophyte . Plant gr owing under dry conditions .
3. Light Relations.
Heliophyte. Plant grovin g in full sunlight .
Sc i ophyte . Pl ant grmdng in the shade .
B. Edaphic Relations .
1. Soil Texture R~lations .
Psammophyte . Plant that grol~s in the sand .
Pelosarnrnophyte . Plant that grows i n loam, sand and c l ay .
Pelophyte . Plant that grows in c l ay .
Saprophy te. Plan t tha t grows on dead organic matter .
2 . Soil Aeration Relat ions fo r Vascular Pl ants .
 314 15

Aerobe . Organism \'lho$e roots requ ire free oxygen .

Anaerobe . Organi sm Hhose roots live j n absence of free
oxygen .
3. Soil Nutr ition Relations.
Oligo t rophe. Organism that lives in soil poor in mineral
nutrients . '.
Hesotrophe . Organism that lives in soil moderately supplied
with mineral nutrients .
Eutrophe. Organism that lives in soil abundantly suppl i ed
l1ith mineral nutrients .
{, . Soil pI! Relations .
Acidophyte . Plant living in acid soil, beIO\,' pll 6.5 .
Basophyte . Plant living in basic soil , above pH 7 . 5 .
Neutrophyte. Plant l iving in circumneutral soil , pH 6 . 5-7 . 5 .
Calciphyte . Plant living on basic sailor rock abundan tly
supplied with calcium ions.
Halophyte. Plant living on basic soil abundantly supplied
Hith chloride ions .

III . SPATIAL RELATIONS .

A. Numerical Relations.
1. Abu nd ance . An est imation of the plentifulness of individua l s of
a species \"ithin a region . One scale of abundance is 1. very
rare, 2 . Rare, 3 . Inf r equent, 4 . Abundant , 5 . Very abun -
dant .
2 . Density . An actual count and determination o f the number of indi-
viduals on a unit area basis; average number of individuals per
area sampled .
3. Frequency . The percentage of sample plots in Hhich a spec ies
occurs . Frequencies can be grouped into classes based on per-
centages of sample plots in Hhich species occur such as :
A 1-20%, B 21-40%, C 41-60% , D 61-80% , E 81-100% .
B. Distrib ut ional Relations .
1. Random Individuals . Disper sed irregularly; Hithout definite pat-
tern .
2. Clumped Random . Aggregates o f individuals dispersed irregularly
lvithout definite pattern .
3. Unifo rm . I n dividuals dispersed in a regular and definite pattern .
C. Areal Rela tions .
1 . Cover . Amount of horizontal space shaded or occupie d by an indi-
vidual. The fol l oHing cover classes \,'ere adapted from Braun-
B1anquet (1932) by lIelmut Lieth:
R - one spec imen in a r ea, fe\" if any nearby
+ - Cover less than 1%
Cov . 1 - Cover 1-5%
Cov . 2 - Cover 5 -1 2 . 5%
Cov . 3 - Cover 12 . 5-25%
Cov . 4 - Cover 25-50%
Cov . 5 - Cover 50-100%
2. Basal Area . Number of square feet of cross-sect i onal surface of
in dividual tree species determined at breast height (4.5 ' ) per
unit area such as an acre . Foresters use th is method of deter-
mining the amount o f timber per acre of land and usually mea-
sure the d.b.h . (diameter breast height) and then convert the
 15 315

d . b . h . IS to area using standard tables . Basal area is also

the amount of ground cover of bunch grasses or rosette plants .
3. Size classe5 . Refers to the numher of individuals of certain
diameters \~ithin an area or stand. l:Jithin a stand of trees ,
i . e ., there mi gh t be a group of individuals uith a d . h . h . of
4-6", another group \-lith a d . b . h . at 12-15" , and still another
group ,dth a d.h.h . of 22 -24" \~hich Hould be t hree size classes
Hithin that stand.
4. Stratification. l-lithin most \'lOady conununities t h e species occur
in layers Hith those at the top f orming the 1. Canopy , those
i n a layer just belcl-! the canopy forming the 2. Suhcanopy ,
and just beloN tha t is a 3 . Shrub layer, and j ust above the
ground is a 4. Herb l aye r, and frequently beneath the herbs
is a 5 . }1oss lichen laver .

IV . TEl'lPORAL (TItle) RELATIONS .

A . Life (Life cyclic; sexual)

Adult Sporophyte
Embryo Sporangium
Zyg ot e Sporocyte
syngamy'cccccc:-____________________________c=~cc--c~ leiosis
Gamete Spo r e
Sex Orga n
Adult Gametophyte

B. Phenology (Annual cyclic) . St udy of periodicity in plants as related

to climatic events; e.g ., time of leafing , flmlering , fruiting ,
budding.
1 . Vernal . Phenomena that occur in spring .
2 . Aestival . Phenomena that occur i n summer .
3. Autumnal. Phenomena that occur in fall.
4 . llibernal . Phenomena that occur in Hinter .
C. Periodicity (Dailv or seasonal cyclic) . Diurnal or seasonal rhythms
or fluctuations; e . g . , temperature, light, moist ure regimes or
amoutl ts .
1. Thermoperiodism. Responses of plants to periodic changes i n tem-
per ature , usually considered on a diurna l basis .
2 . Photoperiodism. Responses of plants to the length of the diurnal
light and dark periods , as regards fl m-ler initiation , t uberiza-
tion, leaf abscission, etc . Long-day plants ini tiate flm~ers
,,,hen the day length is greater than a critical minimum (usually
11 - 13 hours); short-day plants do so Hhen the day length i s
less than a critical max imum (usually 10-14 hours; day-neutral
plants flm"er independently of day length .
3 . Hydroperiod i sm . l:esponses of plants to per jodic changes of com-
plete soil saturation by Hater .
D. Succession (Generation and Conununity cyclic) . Changes in t i me of com-
mun i ties in a ..given a r ea from pioneer to cl i max conunu nities .
Invaders .
1 . Pioneer (Species or Communities) . Plant invaders of an area
previously uninhabited by plants or into an area d isturbed
by man .
2 . Transient (Spec ies or Communities) . Later invaders into
pioneer and s ubsequent conununities prior to climax .
316 15

Sere . A group of plant conununities that successively occupy the

same area f r om the pioneer to a mesic climax.
1. Hydrosere . A sere in \~hich the pioneers invade Ivate r .
2. Halosere . A se r e in which the pioneers i nvade salt water or
salt-flats.
3. Psanunosere. A sere in \~hich the pioneers i nvade sand.
4. Pel osere . A sere in which the pioneers invade clay o r mud .
S. Lithosere . A sere in which the pioneers invade rock .
6. Subsere . A sere in which the pioneers invade an area dis-
turbed by man; e . g ., pasture , old field .

Section C. ADAPTIVE FEATURES

Adapt i ve features wi thin a flora or community can give clues as to the

(1) origin of the flora or community , (2) migration patterns in the area ,
(3) evolutionary trends \~ithin populations, (4) and indications of present
or past climatic trends . Life form and leaf size analyses , t"hen compared
with norms determined for similar communities elsewhere , can indicate cli-
matic shifts or stabili ty ; diaspore and pollination analytical studies can
give hints as to origin and evolution of the flora or community; and fidel-
ity and vitality percentages can indicate the youth or maturity and migra-
tional stability within a flora or community . Studies of other adaptive
features within a f l ora or community; e . g ., those associated wi th floral
morphology could be of equal evolutionary significance .

I. LIFE FORMS (Lf ) . Physiognomic characteristics . Based on Raunkiaer

(1934) .

Life Form Symbol Description

Therophytes Th Annuals : f r om seed germination to seeds in

one year.
Geophytes G Perennials : perennating organs (bulbs ,
tubers , rhizomes , corms) underground and
protected; plants die above ground
Hemicryptophytes 11 Perennials or bienn i als : perennating buds
very close to ground surface (tussock
and rosette plants)
Chamaephytes eH Perennials : perennating bodies above ground
(coarse herbs or shr ubs Hith buds less than
0 . 5 m above ground) .
Phanerophytes P Perennials: perennating buds more than 0 . 5 m
above ground ; trees (t), shrubs (s), woody
vines (v) .
Epiphytes E Perennia l s : borne on other plants or struc-
tures high above ground (no r oot contact
with soil).
Succulents S Perennials, r arely annuals : either s t ems ,
leaves or both very fleshy .
Paras i tes Pa Perennials or annuals : non- chlorophyl lous
plants dependent upon other plants for all
nourishment or chlorophyl lous plants de -
pendent upon other plants for t"ate r and
mineral salts.
15 317

Life Form Symbol De script ion

Sap rop hyt es Sa An nuals or perennials: Living on dead

organ ic ma tte r.

II . DIASPORE TYPES (Dia) . Di s persal mechanisms . Based on Van de r Pi jl

(1969) a nd Ridley (1930) .

Diaspore Type
Gr Di sseminul e Symbol Dis persal Agent - Adaptation

Sporochore Spa Hind - -minute or dust seeds, fr uits . or

spores .
Pteroch or e Pte \Und--win ged seeds or fruits .
Pogonocho r e Pog Wincl--plumed seeds or fruits .
Cyclochore eye Wind-- tumb ling i n fructescences o r plants .

Epizoochore Epz Animal- -barbed, hooked or viscid seeds or

f r uits .
Synzoochore Syz AnimaI-- inten tionally buried or stored
seeds or f r uits .
Endozoochore Enz Animal--visceral (non-digestible) seeds
or fruit s .

Ba llochore Bal ~Iechan i c al -- expulsio n or jactitation seeds

or fruits .

Barochore Bar Cravity--heavy s eed s o r f ruit s .

Hydro chor e Hyd Water--floating seeds, fr u i t s , or vegeta-

t ive parts .

Anthro pochore Ant Nan--none .

Atel echo r e Ate No spec ial adaptation- -on the spot

d ispersa l .

Polychore Po l Nore than one agent--coul d be wind and

anima l adapted fr u its on same plants .

Note: \.jinel means total Innd Dispersed and Animal means to tal Animal
Disper sed Diaspores i n Spectrum .

III . FIDELITY. I ndi c ates the degree to which a specie s is restricted to a

part i cular type o f community. In the fo l l owing c l asses Fid . 3 anel
4 wou l d i ndicate that a species is a characteris tic of that community
and Fid . 5 ,,! ould indicate that the spec ies is an indicator of that
commun i t y .
Fid . 1. St ranger s, appearing accidentally.
Fid. 2 . 1.~differen t s, witho u t a decided affin ity for any type
of community .
Fid . 3 . Pr eferents , pr esen t in s everal type s of communities
but predominantly in one type .
Fid . 4 . Se lect i ves , usually found in one typ e of community but
occasion ally found i n ot he r s .
318 15

Fid. 5. Exclusives, found in one type community only, rarely in

another .

IV . VITALITY . Indicates the vigor and reproductive capacity of a species

l·dUnn a community. The following classes have been recogni7.ed by
most ecologists: ~.

Vit o 1 . Ephemeral, advent ive and not reproducing .

Vit o 2 . Vegetative apomicts , reproducing asexually by vegetative
propagules but not completing a sexual cycle of any sort .
Vito 3 . Amphimicts, regularly reproduc i ng sexually or occasionally
by agamospermy .

V. LEAF SIZE. Leaf size characteristics seem to be related to different

environments . The fo11m... i ng classification is adapted from Raunkiaer
(1934): (Note : these terms have no morphological significance except
size . )
L. S. l. Leptophyll, to 25 'q. mm.
L. S . 2 . Nanophyll, to 225 'q . mm . 25 x 9
L. s . 3 . Hicrophyll, to 2,025 'q . rom. 25 x 9 2
L. S . 4. Nesophyll, to 18,222 sq . mm . 25 x 9 3
L. s. 5. Nacrophyll , to 164,025 'q. mm. 25 x 9 4
L. S. 6. Negaphyll, larger than 164,025 'q . mm. 25 x 9 5

VI. POLLINATION.
A. Pollination Types . Classification based on agents by which pollen
is transferred from the pollen sac or anther to the ovule or stigma.
1. Anemoph i ly . Pollinated by wind .
2 . Anthropophily. Pollinated by man.
3 . Cantharophily . Pollinated by beetles .
4. Cheiropterophily . Pollinated by bats .
5. Entomophily . Pollinated by insects .
6 . Hydrophily. Pollinated by \~ater .
7 . Hymenopterophily . Pollinated by bees .
8. l>lalacophily . Po llinated by snails or slu gs .
9. l>licrome1ittophily . Pollinated by small bees.
10 . Hyiophily. Pollinated by diptera.
11 . Myrmecophi1y. Pollinated by ants.
12 . Necroco1eopterophily . Pollinated by carrion beetles .
13 . Ornithophily . Poll ina ted by birds .
14. Phalaenophily . Pollinated by moths.
15. Psychophily. Pollinated by butterflies.
16 . Sapromyiophily . Pollinated by carrion or dung flies .
17 . Sphingophily . Pollinated by hawk moths and nocturnal lepidoptera.
B. Flot... er - Po11inator (Visitor) Relationsh ip s. Adapted from Fae gr i and van
der Pij 1 (1966)
1. A11ophily. F1ot...er with no morphological adaptations for guiding
pollinators, can be utilized by non-adapted pollinators.
2. Euphi1y . Flower strongly adapted for utiliza tion by specialized
pollinators.
3 . Hemiphi 1y . Flower imperfectly adapted fo r utilization by inter-
mediately specialized pollinators .
4 . Nonophily. Euphilic flotoJer utilized or visited by a single or
closely related species of pollinator.
5 . Oligophily . lIemiphi1ic flot'ler utilized or visited by some relat ed
taxa of pollinators .
 15 319

6. Po l yphily . Hel'liphil i c flower utilized or visited by many dif -

fer ent taxa of pollinat ors .
C. Pollinator (Vi sito r)-FlO\~crin g Species Relationships . Adapte d from
Faegri and van der Pijl (1966) .
1. Al l o t ropy . Poll inator poorly adap t ed for utilization of flowers;
food obtained f r om fl O\,Iers form pa rt of a mi xed diet.
2. Dy s tropy . Pol linator (visitor) unadap ted or shows no relation-
ship to flower organization, frequently destructive, but ma y
cause pollinat ion .
3 . Eut r opy . Po ll i na t or fully adapted for utilization of flower ,
taking main food from flO\~e r .
4 . lIemitropy . Pol linator has intermediate degree of specia lization
for uti lizat ion of flO I"er .
S . Honotropy . Pollinator strongly adapted fo r utilization of one
flow er o r flot-le r from closely related speci e s .
6 . Oligotropy . Po l linator, with intermediate degree of spec ia l iza-
tion for utilization of flOl.er, that v isi ts flowers of
r elated taxa ( species).
7 . Po lytropy . Po l l inator, I·li th in termed iate degree of specializa-
tion fo r uti lization of floHer, that visits raany dif fe ren t
taxa of flowe r s ( species) .
D. Cleistogamy. Cla ssif ication ba sed p r imarily on environmental
c1eisto gamy . Adapted f rom Uphof (1938) .
1 . Cleistogamy . Se lf -pollination t-lithin c l ose d flot-lers , as in
~.
2. Edaphoc l e i s t ogamy . Self-pol lination in s ubterranean f lowers , as
in Commelina .
3. Hydrocleistogamy. Self-pollination in c l osed fl owers , t-lhich are
norma l l y expanded, due t o submer sion , as in Hayac a .
4. Hygr ocle i stogamy . Self-pollination i n c lose d flowers . I~hich ar e
normally expan ded , due to great fog den s ity , as in Liparis .
5. Oligo tropi c Cleistogamy . Self-pollination in closed fl ol"ers ,
I"hich are normally expanded , due to low soil nutri ents , as
in Specu la ria .
6. Psyehro e l eist ogamy . Self-pollination in c l osed flowers , ",hieh
are normally expand ed , due to 10l~ temperatures, as in
I mpatie n s .
7. Seioeleistogamy . Self - pollinat i on in c losed Elot"er s , t-lhich a r e
no r mally e xpanded , due to shad i ng , as in Viola arvensis .
8. Thermocleistogamy . Self -pol lination in closed flOl"e r s , I.hich
are normally expande d , due to high temperatures , a s i n
Eranthemum.
9. Xeroe l eistogamy. Self-pollinat i on i n closed flower s . which a r e
normally expanded, due to drough t , as in Dicl i ptera .

VII . ENVIRONHENTAL EFFECTS UPON HORPHOLOGICAL FEATURES (Dev e l opmental Res pon ses
and Adaptations)
The taxonomist should note the morphological features that may be
correlated wi th di ffe r en t environmen tal fact o r s s uch as light , mois ture,
and soil aer at ion . primarily in order to better und erst and s truc tura l
var iation in the field and laboratory. Secondarily , he should incor-
porate range measurement ext r em ~s due to environmental extremes in his
description o f a particular taxon and a lso be aware of t he fact that i n
identificat i on , meas urement extremes are not ah.ays given in manuals due
to insuf fic i ent observation . Developmental or behavioral r espons e s should
320
15

be distinguished from genetically fixed adaptations. A few morphological

features have been selected to shaH developmental or adaptive responses to
intense light, Im~ moisture, and poor soil aeration as examples of struc -
tural-environmental interactions . These observations have been made by
many experienced fi e ld and laboratory Horkers , sometimes with l ittle
understanding of the real significance or cause oJ the response. For
specific references to experimental and analytical evidence for these and
other interactions, consult the ecological texts and periodicals cited in
the literature section of this chapter, particularly Daubenmire (1959).

A. Norphological Features Associated t"ith Intense Light (Heliophytic

reactions or adaptations)
I ntense light results, developmentally, in
1. Thicker stems
2 . Reduced internodes
3. Reduced and thicker leaf blades
4 . Nore trichomes per unit area
5 . Thicker cuticle
6. Smaller stomata and blade cells
7. Better developed palisade layers
8 . More prolif i c branching

Adaptively, species living in full sunlight exhibit the developmental

characteristic s cited above .

B. Morphological Featu r es Associated with Low Hoi sture (Xerophytic

reactions or adaptations)
LOIv moisture results , developmentally , in
1. Reduced s i ze of shoot
2. Occasionally increased size of root system
3. Smaller leaf blade or absence of blade
4 . Smaller and fewer stomata that may be conf ined to grooves or
crypts
5 . Hare heavily lignified tissues in blade
6. Thicker cuticle
7 . Nore numerous trichomes per unit area
8. Better developed palisade layer
9 . I ncrease in vascular tissue per unit area
10. Reduction in cell s ize and increase in thickness of \va11
11. Decrease in volume of inte rc ellula r space

Adaptively , 101.; moisture has resulted in the development of

a. Ephemeral annuals
b. Non-succulent perennials with storage taproots and scl erotic
shoots
c. Non-succulent perennials with extensive fib rous root systems
and reduced shoot-transpiring surfaces
d. Succulent perennials

C. Norphological Features Associated with Poor Soil Aeration

(Anaerobic reactions or adaptations)
Poor soil aeration results, developmentally, in
1 . Roots usually less numerous
2 . Root branching less complex
3 . Roots usually shorter and occupying less space
 15 321

4. Roots and underground stems thicker with the deve lopment of more
lacunar or aerenchyma tissue
5. Root systems shallow \Vith the pres ence of numero us pneumatophores
6. Shoots smaller or stems are buttressed and with lacunar or aeren-
chyma tiss ue

Adaptively, species with roots living under anaerobic conditions

exhi b i t the developmental or behavio r al characteristics li sted above .

VIII . HETEROTROPHY IN VASCULAR PLANTS*

A. Nutritional Re la tionships .
1 . Parasite . A plant dependent on a host plant for food and/or \Vater .
Parasitism has apparently arisen independently at least six
times i n angiosperms in the following orders of dicotyledons :
Laurales (Cassytha of the Lauraceae), Santalales (Santalaceae ,
Loranthaceae , Olacaceae and others) , Rafflesiales (Raffles ia-
ceae , Hydnoraceae , Balanophoraceae), Scrophularia l es (Scrophu-
lar i aceae, Orobanchaceae), Polemoniales (Lennoaceae , Cuscuta
of the Convolvulaceae) , and Polygalales (Krameriaceae) . No
parasitic monocotyledons are known . A single report exists
of a parasitic gymnosperm, Podocarpus ustus, from New Caledonia .
The f l owers of parasitic angiosperms are among the largest
(Raff l esia sp.) and smallest (Pilostyles sp ., Arceuthobium
sp . ) of all known flm.,rers.
a . Auto- parasite . A plant parasitizing itself as in some Santala -
ceae; sometimes called sel f - parasitism or root grafting .
Most auto- parasites are hemi-parasites.
b . Holo- parasite . A plant lacking chlorophyll and thus entire l y
dependent on its host, as all Orobanchaceae and Cuscuta .
Halo-parasites are termed stem parasites or root parasites
depending on the portion of the host they attack.
c . Hyper-parasite . A plant that parasitizes other parasites as
some Loranthaceae. Hyper-parasitism has sometimes been
referred to as ep i-parasitism.
d . Hemi-paras i te . A chlorophyll-containing parasite superfi-
c ially appearing completely autotrophic, as in many members
of the Scrophulariaceae . Hemi- parasites may be stem or
root parasites . Sometimes herni-parasites are called serni-
or demi- parasites.
2 . Saprophyte. A plant lacking chlorophyll and obtaining nourishment
from decay i ng organic matter . All saprophytes Hhich have been
carefully studied have been shown to be epi- parasites .
3 . Fungal r elationships .
a . Epi- parasite . An achlorophyllous plant con taining a fungus
continuous \.,rith the mycorrhizal associate of surrounding
trees as in Monotropa of the Pyrolaceae (or Ericaceae)
and plants belonging to thi rteen or so genera of the
Orchidaceae .
b . Mycorrhiza : -' The symbiotic relationship bet\Veen a fungus and
the roots of vascular pl ants. Mycorrhizae are widespread
in ang i osperms \.,rith only a fel'] fami lies (Brassicaceae ,
Chenopodiaceae , Cyperaceae) and aquat i cs thought to be

*Contributed by Lytton J . Musselman , University of North Carolina at Chapel

Hill.

j
322
15

entirely non-mycorrhizal.
Endomycorrhiza', A fungus Ivith i ntracellular hyphal
infections in the host root cortex; the most common
type of mychorrhiza .
Ectomycorrhi za. A fungus \-lith intercellular infections
in the host cortex . All members.. 9f t he Pinaceae are
ectomycorrhizal ; in angiosperms ectornycorrhizac are
usually limited to amentiferous families. Truffles are
the fruiting bodies of an ectomycorrhizal fungus.
Ectendomycorrhiza . Fung us l"rith both inter- and intra-
cellular infection in the host cortex . Known only in
tn'!e species Hhich are usually ec t ornycorrh iza l.
4 . Bacterial relationships. Bacterial root nodules are best komJn
in the Fabaceae but also occur in many other families of angio-
sperms and Podocarpaceae of the gymnosperms .
5 . Algal r elationships. Blue green algae are knmm to inhabit the
roots of the cycad genus Cycas . Algae are also foun d in the
roots of some angiosperms .
6 . Root grafting . The fusion of tHO or more contiguous roots , usually
t r ee roots, involving the morphological and anatomical union of
cambium , ph l oem, and xylem of the two roots . Only a feH re-
ports of roo t g ra fting in herbaceous plants exist. This term
h as also been applied to auto-parasitism in angiosperm root
par asites .
B. Morphological Considerations of Heterotrophy .
Heterotrophy, especially parasitism , has resulted in extreme speciali-
zati on and reduction in the plant body of the parasite . Diffi-
culty arises Hhen t r ying to define the highly specialized organs
of parasites i n the usual morphological context.
1. Haustorium . The o rgan Hh ic h forms the morphological and physio-
logi cal bridge between host and parasite, usually considered
to be a highly modified root.
a . Primary haustorium . A haustorium \.,rhich arises directly from
a root apex , considered mo r e advanced than a secondary
haustorium .
b . Secondary haustorium . A hausto r ium which arises laterally
from any root or stem and not from a root apex.
c. Endophytic system . The greatly reduced vegetative plant body
of a holo-parasite which is entirely contained \,,;rithin the
host plant. In Pilostyle s and other members of the
Rafflesiaceae the only indica tion of the presence of the
parasite is \"hen the flQlver buds burst through the host
bark.
2 . Roots .
a. Pilot roo t. A root \.,rhich gives rise to haustorial roots but
does not form h austoria itself. as in Pholisma of the
Lennoaceae .
b . Haus to rial root . A root producing haustoria. usually applied
to parasites Hhich e x hibit root dimorphism .
3. Host responses .
a . Corallo id root. A short, s1;vollen , dichotomously branched
mycorrhi zal root .
b . Isopbasic growth. Integrated host-parasite growth where the
cel l s of the parasite are carried along by the apical
meristem of the host.
15 323

c. hloodrose . Placenta-like structure comp osed of host wood

in response to infection by parasite.
d. Nodule. Globose s\,Jelling in root Hhich conta ins bacteria .
e. Hitches broom . A malfo rma tion of a host by parasite in-
fection resl'lting in shortened in ternodes and a pro-
liferation of branches , as Arceuthobium on Pinus .

ECOLOGICAL LITERATURE

I. Periodicals
Ecology. Official publication of Ecological Society of America .
1920+ . Duke University Press . Durham .
Ecological Nonographs . Official publication of Ecological Society of
America. 1931+ . Duke University Press . Durham .
Journal of Ecology . Official publication of British Ecological
Society . 1913+ . Cambridge Univers ity Press . Cambridge .
Vegetatio . Official publicat ion of International Association of
Plant Sociologists . 1948+ . D. "T. Junk, Publishers . Hague.

II. Special References.

Braun-B1anquet, J . 1932 . Plant Sociology . (transl . by H . S . Conard
and G. D . Fuller) . NcGraw-lIill Book Company . New York.
Clemen ts, F . E . 1902. A Sys tern of Nomenclature for Phytography .
Botanische Jahrubucher fur Systemat ik, Pf1anzengeschichte und
Pflanzengeographie 31, beiblatt Nr . 70 .
Dansereau, P . 1957 . Biogeography : An Ecological Perspec tive. The
Ronald Press Company . New York .
Daubenmire, R. F. 1959 . Plants and Environment . 2nd ed . John Hiley
and Sons, Inc. Ne\" York.
Faegri, K., and L. van der PijI. 1966 . The Princip l es of Pollina-
tion Ecology . Pergamon Press . New York .
Gray, P . 1967 . The Dict ionary of the Biological Sciences . Reinhold
Publishing Corporation . New York .
Jackson, B. D. 1928 . A Glossary of Botanic Terms . J . P . Lippincott
Company . Philadelphia.
Kruckeberg , A . 1969 . The implications of ecology for plant sys -
tematics. Taxon 18 : 92-120.
adurn , E . P . 1971. Fundamentals of Ecology . 3rd ed . H. B . Saunders
Company . Philadelphia .
Percival, Nary S. 1965. Floral Biology . Pergamon Press . New York.
Pijl, L. van der . 1969. Principle s of Dispersal in Higher Plants .
Springer-Verlag. Berlin .
Raunkiaer, C. 1934 . The Life Forms of Plants and Statistical Plant
Geography . Clarendon Press. Oxford .
Ridley, H . N . 1930 . The Dispersal o f Plants throughout the \.,Iorld .
L . Reeve and Company, Ltd. Ashfo rd.
Stearn, H . T . 1966. Botanical Latin . Thomas Nelson and Sons , Ltd .
Edinburgb .
S\"artz, D . 1971. Collegiate Dictionary o f Botany . Ronald Press
Company . Ne~~ York .
Uphof, J . C. Th . 1938 . Cleistogamic flowers . Botanical Revie,~ 4 (1):
21-/,9 .
 324 15

ECOLOGICAL CHARACTERISTICS EXERCISE

Select one or two species and work out the character states for each of the
follm..ring characters :

CHARACTER Species III Species 112

Habitat

Biotic Relations:
Nutritional

Sociability

Symbiotic

Climat i c Relations :
Temperature Regime

Air Moisture Regime

Light

Hind
Edaphic Relations:
Soil Texture Regime

Soil Aeration Regime

Soil Nutrients
Regime

Soil pH Regime

Spatial Relations :
Numerical

Stratificational
Temporal Relations:
Phenological

Successional
Adaptive Features :
Life Form

Leaf Size
Diaspore Type

Fidelity

Vitality

Pollinating Agent
16 DS

Chapter 16. GEOCRAPIlIC EVIDENCE AND DISTRIBUTION

An understanding of geographic evidence and principle s of distribution is

basic to inte rpre ting t he origin , migration, and evol ution of species and
flo ras . Distribution ranges correlated \dth differences or similarit ies i n
structural features have been and are the bases for the de termination of r ela-
tionships of taxa a t various levels. The fo llowing quotations show the roles
of geographic evidence and distribution in the determination of taxonomic and
evolutionary relationships .

"The maps (on distribution of morphological fea tures in Aconit um)

show that certain featu r es seem to vary independently though l oosely
cor r elated Hith each other and with geographical a r eas . Geographi -
cally plac ed symbols shoH the corre l a t ions of these features . Three
groups become identifiab l e from the maps: (1) 3-lobed leaf and
limited pube scence , at lOH elevations, Maryland to s outhHestern
Georgia and Hest of the Appalachians in Tennessee and West Virginia;
(2) 5- lobed l eaf, gr ea t er pubescence , in the Appalach i ans , from
western Pennsylvania to eas t e r n Tennessee and northl..estern South
Carolina and along the outer Piedmon t of North Car olin a and Vir-
ginia ; and (3) 5- to 7-10bed leaf, ex treme pubescence , and ' muticum '
helmet in populations north of the glacial boundary in New Yor k ,
Ohio , i.Jisconsin and Iowa . The taxonomic interpretat ion of these
three en t ities must take into account three charact eristics shown
by the maps." (llardin, 1964, Variation in Aconitum of eastern
United S t a t es . pp . 88- 89).

" Th e f l ora of the northern part of North Amer i ca and that of

the northeastern part of Asia are, because of geographical contin-
uity, closely related. There have e vident ly been exchanges of
species and extensions of ranges of species betHeen the two areas
until the ve r y recent geological past , s uch processes probably
continuing at the present time. The rel ationships of t hese con-
tinuous flo r as have been i nvestigated especially by Hul ten in his
many excel l ent papers .

A study of the habit and habitat of the temperate disjunct

genera of eastern Asia and eastern North American Hill a l so reveal
the na ture of the vegetation when these genera and t heir rel atives
and associ ates , both ex tinct and ex tant , were once the dominant
elemen t s o f the floras in the geological past. The fo llowing point s
can be especially noted .

In the first place , th e greate r number of gene ra is composed

of woody plants, only a few consisting exclusively of herbaceous
plants . Among the woody plants are trees like Carya , Liriodendron .
Sassaf ras, Gymnocladus, Cladrastis, Stewartia , Halesia, Nagnolia ,
Gordonia . an d Catalpa , shr ubs like Hamame lis , Pachysandra , Chiogenes .
Chionanthus . Hi tchella . I llicium . I tea . Pier is , and Lyonia, and woody
vines like Decumaria, Schisandra, Wisteria, Parthenoc i ss us , and
Campsis . " (Li , 1952 , Floristic relation ships between eastern Asia and
eastern North America . p. 404).
 326 16

The comprehensive field of plant geography includes s tudies of origin ,

distribution , adaptation, and associations or gro up ings of plants . The treat-
me nt in Section II of this chapter is a syno p tic descriptive sUJlUnary of the
types of associations such as biogeographic regions , biomes, and flora
provinces. The fundamental principles of distribution and definitions of
classified geographic terms pe rtinent to taxonomy are.. presented in Section B.
In Sect i ons C and D major and minor disjunct ranges are discussed . Th e ranges
pre s ented in these sections are used on l y as examples to shm., some of the
major disjunct or discontinuity ranges in the 'h'or ld and some of the distribu-
tion patterns in North America . Th is , in no sense, is intended to he an
exhaustive summary of the subject (consult the references cited for more
detailed information) . Th e dynamic aspects of b i ogeography and the subject
of adaptations are not descri bed in this text except in a limited Hay in
Chapter 15 . The plant conmtun ities in eastern North America are summarized
in Chapter 17 .

Section A. BIOGEOGRAPHIC REGIONS , llIO~mS , FLORA PROVINCES

No hard and fast definitions have been devised for the distinctions
between region , biome , and province but for the sense in which the terms are
used here the following definitions are applicable : (1) Biogeographic Region _
maj or , more or less continuous land mass I"ith major geologic or ecologic
barrie r s to migration bet\veen adjacent regions , such as the Atlantic and
Pacific Oceans between the Nearctic and the Palearctic Regions and the Sahara
Desert bet\.,een the eastern Pal earctic and Ethiopian Regions ; (2) Biome - a
group of associations or organisms ~"ithin a region with a definite physiog-
nomy , such as the needle-leaved trees (conifers) of the Taiga ; a n d (3) Flora
Province - region which i nc ludes the total range of the dominants of several
related types o f vegetation or communities , such as the southeaste r n swamp
fores t s I"ith cypresses , gums, and cottomJoods .

1. BIOGEOGRAPHIC REGIONS
Based on Polunin (1960)

1. Nearct ic . North Ameri ca north of Nexico.

2. Palearctic . Euras i a - North Africa north of the Saha ra and e - \., mountains
such as the Himalayas .
3. Ne0tropical. So u th America and Caribh ean Regions .
4. Ethiopian . Africa south of the Sahara Desert .
5. Oriental . Asia south of the Himalayas and a ma j or part of the East In dies.
6. Aus t ralian . Australia , Ne\" Zealand, and South Pacific Islands.
7. Antarctic . Antarctica, southern South Amer ica.

II . HAJOR BIOHES OF THE \1ORLD

1. Tundra and Ice

2. Tai ga (Boreal Coniferous Forest)
3. Temperate Deciduous Forest
4. Scrub and Sclerophyl l Forest
5. Grassland
6. Desert
7. Tropical Rain Forest
8. Temperate Rain Forest
 327
16

Biome Distribution in Relation to Altitude and Latitude

in
Eastern North America
(Temperature Relations)

High I ce and Soo,"

(Tundra)

Taiga

Temperate Deciduous
Forest
Temperate Ice
Tropical Deciduous and
Lm.,r Rain Forest Forest Taiga Snow

South -.~------------~.~ North

Latitude

Biome Distribution in Relation to Moisture

in
Temperate North America
(SH summer wet, winter wet; Sw = summe r wet, ,,,inter dry;
s~.;r '" summer dry, winter wet; SW'" summer dry, Hinter dry)

SW Sw
Temperate Deciduous Grassland
Forest

"Coniferous Forests"

sW sw
Sclerophyll Forest Desert
or Scrub

III . MAJOR FLORA PROVINCES OF NORTH AMERICA NORTH OF NEXICO

Based on Shantz and Zon (1935) , Shreve and
Higgins (1964), ,\.;reaver and Albertson (1956)

Province Dominant or Characteristic Plants

Tundra - Arctic and Alpine "(Sedges, Grasses , Saxifrages, Lichens, Dwarf ~.Jillows,
Gentians, etc . )
Coniferous Forests
Boreal Forests (Spruce, Fir, Tamarack)
Northeastern Pine Forests (Jack, Red, Hhite Pine)
328 16

\.Jestern
Subalpine (Engelmann Spruce, Alpine Fir , Limber & Lodgepole Pines)
Montane (Ponderosa Pine, Douglas Fir)
Northl"restern Nixed Conifer Forests (Larch, Redlwod , Western Cedar .
Western \<ihite Pine, Western Hemlock, Sitka Spruce)
Southwestern Pinyon-Juniper Forests (Pinyon Pine; Juniper)
Southeastern Pine Forests (Longleaf, Loblolly, Slash Pines)
Hardwood Forests
Northern Aspen (Aspen, Birch)
Northeastern Hardwood and Southern Appalachian Nixed Nesophytic (Haple,
Beech, Birch. Hemlock)
Eastern Oak-Hickory Forests (Many species of Oaks and Hickories)
Southeastern Broadleaved Evergreen-Hammock and Bay (Live Oak, Red Bay,
American Olive, Holly, Hagnolia)
Southeastern S\vamp Forests (Bald Cypress, Gums, Cottom"roods, Ashes)
Subtropical Nangrove Forests (Buttonwood, Red, Rlack and l.Jhite Nangroves)
Western Broad Sclerophyll Forests (Oaks. Incense- cedar. California Bay)
Western Hoodlands (Oaks, Pines, Redbud, Gooseberry)
Chappara l (Chamiso, Manzanita, Hountain I'Jahogany, Wild-lilac, etc.)
Southwestern Savanna (Mesquite. Desert Grasses)
Grasslands
Prairie (Tall Grasses: Bluestem, Indian, Needle)
Mixed Prairie (Hid and Short Grasses : Bunch, Needle , Slender l.Jlleat)
Pla ins (Short Grasses: Grama, Calleta , l-lire, Western Hheat)
Pacific Prairie (Bunch Grasses: Wire, Hheat , Poa)
Desert
Northern Desert Scrub (Salt Sage, Sagebrush, Rabbit Brush, Hinter Fat)
Salt or Alkali Desert Scrub (Greasel"rood, Pickleweed, Samphire, Salt Sage)
Southern Desert Scrub (Creosote, Bur Sage, Ocotillo)
Arboreal Desert (Cercidium, 01neya, Jatropha , Bursera)
Succulent Desert (Pachycereus, Carnegia, !:£p.llOcereus, Opuntia-Cacti)

Section B. DISTRIBUTIONAL PRINCIPLES AND TERMINOLOGY

General agreement on the principles o f plant distribution was reached many

years ago by geographers, ecologists, and taxonomists. The concepts dealing
with origin, establishment, dispersal, barriers to mig ration, and continuous
and discontinuous ranges are accepted by individuals working with distribution
problems . The criteria for place of origin and center of distribution, however,
are debatable ; continuous range is an impossibility but is accepted and under-
stood as a concept in the sense of general distribution throughout a climatic
regime under the proper edaphic conditions l"rhich are always disjunct; the
agents of dispersal: wind, water , and animals, are agreed upon but determin-
ing the actual effectiv e agent for a given species in a local area or r egion
occasionally presents a dilemma; barriers to migration: climatic, edaphic, and
biotic, are the broad general classes of obstacles to dispersal but determining
the actual barrier to migration of a species in a limited area can be a problem .
Determining the limiting facto r(s) for establishment of a species Hithin an
area is a traditional problem f or experimental e cologists .

I. DISTRIBUTION PRINCIPLES
Based on Good (1964)

1. Plant distribution is primarily controlled by distribution of climatic

conditions .
 2. Plant distribution i s secon darily controlled by distribution of edaphic
factors.
3. Plant migration is brought about by transport of dispersa l phases of
individual plants.
4. Species ranges are limited by genet ically controlled tolerances.
5. Flora movements have occurred in the past and a pparen tly are still occur-
ring .
6. Flora movements have occurred a t higher latitude s and altitudes due to
variation and oscillation in climate .
7. Flora composition has varied with change in the relative distribut:lon of
land and sea .
8. Barriers t o migration of plants are phySiographic , climatic, edaphic. or
biotic.

II . AREA AND DISTRIBUTION DEFINITIONS

Based on Cain (1944)

1. Allopatr i c Distribu t ion. Distribu tion in which closely related spt~cies or

subspecies occupy different range or area .
2. Center of Area or Place of Origin . Area with greatest differentiation
within a taxon;
area with greatest abundance of individual organisms ;
area t.;rith maximum size of ind iv iduals;
area with greatest numb er of convergent lines of dispersal ;
area with l east dependence upon a restricted habitat;
area with the greatest number of dominant genes .
3. Continuous Distribution . Areas with ,.;ride-spread distribution of one or
more species.
4. Discontinuous Distribution. Areas with disjunctions in th e ranges of one
or more species .
S. Dispersal . The spreading abroad of an organism from one part of an area
to ano ther ; in plant s by means of wind , water, animels, or mechanical
devices.
6. Disseminule or Diaspore . The agents of dispersal characteris tic of dif -
ferent spe ci es , such as plumed seeds. wi nged fruits.
7. Endemic Species. Sp ec ies limited to a local area or restricted geographi-
cally .
8. Establishment . The successful germination . development. and reproduction
of an i ndividual plan t or population within an a rea .
9. Origin. The source from which anything ar i ses , such as species, popula-
tion, community .
10. Relict Area. Area occupied by remnant of an earlier flora .
11 . Sympatric Distribution . Distribution in which closely related species or
subspecies do occupy the same r ange or area .

III . RANGES ACCORDING TO SIZE OF AREA

1. Local. Very limited range. usually applied to endemics within a

small area. . ...
2. Regional . Range covers part of a vegetation cl imax region or physiographic
province.
3. Continental . Range usually covers more than one c l imax region or two or
more major physiographic provinces.
330 16

IV . GENERAL TYPES OF DISCONTINUITY DISTRIBUTION

1. Alt i tudinal. Distribution pattern broken into two or more non-adjoining

elevation parts .
2. Bipartite. Distribution pattern broken into two parts, a large and a
small wit hin the same general region or area .
3. Bipolar . Distribu t ion pattern broken into t\W parcts, one at each pole .
4. Diffuse . Distribution pattern broken into many small more or less equal
parts, usually corresponding to local discont inuous climatic patterns
or edaphic conditions .
5. Disjunct . Distribution pattern broken into two or more parts; a large and
one or more s mall parts not in the same region or area .

V. \.JORLD-\HDE TYPES OF CONTINUOUS DISTRIBUTION (CENERIC LEVEL)

Based on Good (1964)

l. Cosmopolitan . l.Jorld -lvide. 4. Circumaustral. South Temperate

2. Circumpolar. Frigid Zone. Zone .
3. Circumboreal. North Temperate Zone . 5. Pantropical . Tropical Zone .

VI. ORIGIN AND ESTABLISH~mNT OF PLANTS

1. Adventive . Species that is imperfect l y naturalized ; a species that repro-

duces spontaneously for a while and then disappea rs.
2. Allochthonous. Native species that has not originated l,Jithin the area ; a
past immigrant that l.S now reproducing spon taneous ly l..rithin the area.
3. Autochthonous. Native species that has originated within the area and is
reproducing spon taneously.
4. Domesticated . Species planted by man but not reproducing spontaneously .
5. Ind igenous or Native. Spec i es occurring within an area and not propagated
or introduced ~ccidentally by man .
6. Introduced . Species accidentally or purposely propagated in an area by man .
7. Naturalized . Introduced species that is now permanently established and
reproducing spontaneously .
8. Sporadic . Repeatedly introduced species that never becomes permanently
established.

VII . DISSEHINULES OR DIASPORES AND DISPERSAL

(See Chapter 15 for definitions and classification)

A. DIASPORE TYPES

1. Atelechore 5. Cyclochore 10 . Polychore

2. Anthropochore 6. Endozoochore 1l. Pterochore
3. Ballochore 7. Epizoochore 12 . Sporochore
4. Barochore 8. Hydrochore 13 . Synzoochore
9. Pogonochore

B. CLASSIFICATION OF Pl.. ANTS ACCORDING TO PARTS NORHALLY DISPERSED

1. Carpophyllophyte . Fruit and bract or accessory flm.Jer part of plant .

(Iromwod, clover, linden)
2. Carpophyte . Fruit part o f plant . (Oats)
3. Embryophyte . Embryo or plantlet part of plant . (Hangrove , duckweed)
4. Holophyte. Entire, above-ground part of plant . (Russian thistle)
16 331

5. Inflophyte . Infructescence (old inflorescence) part of plant. (Tumble

grass)
6. Spermophyte. Seed part of plant . (J eweh:eed , larkspur)
7. Sporophyte. Spore part of plant . (Club-moss, Christmas fern)

Section C. RANGES AND DISTRIBUTIONAL PATTERNS

The two basic problems in all of plant distribution are (1) the explana-
tion for the distribution of closely related but disjunct taxa in the \<lOrld
and (2) the explanation for local distri.bution patterns . "'hat is tbe evidence
for a GondHana Land and Laurasia? Did closely related species that are not~
disjunct have a continuous distribution at one time? Have the present major
barriers to mi gration such as the Polar Ice Cap , the Sahara Desert, and the
Atlantic and Pacific Oceans been in existence since the advent of the angio-
sperms? Are refugia such as the Southern Appalachians and Southeastern Asia
real or figments of the imagination? lias there been parallel evolution in the
same taxa in various parts of the Horld? From a local distributional pattern
standpoint , for example , in the flora of the Carolinas, hOH can a prairie ele-
ment, a boreal element, a subtropical element, a montane element in the outer
piedmont, a sand hill refugium element, a coastal pla in element in the lower
mountains , and southwestern Georgia-nortln·,estern Florida l ime sink element be
explained? The anSHer to these and similar questions are basic to an under-
standing of the ori gin, migration, and evolution of species , floras and com-
munities.

Local, r eg ional, int racont inent.:"tl , and intercontinental distributional

patterns and ranges of species and genera provide basic clues to the comple x
geographical and historical relationships of floras . Floristic components
are f rom many genetic sources fro m many directions over a long period of time .
An analysis of the distributional patterns Hithin a region can lead to an
understanding of the floristic r elationships of t.he vegetation of a g iven
area Hith that of contiguous as Hell as in disjunct regions of the earth . A
knD\o,ledge of floristic relationships can lead to an understanding of the evo-
lutionary and taxonomic relationships of similar taxa . Geographic evidence
is fundamental to the determination of systematic affinities of taxa .

As an example, the analysis of the distributional patterns of genera of

seed plants in eastern Nort.h America is presented in Table 16-1 to show the
floris tic relationships bet,,,een the Southern Appalachians and ,,,estern North
Ame ri ca ( from Hood, 1971) . Some intracontinental and endemic ranges on the
generic level involved in the study by '''ood (1971) are given in Table 16- 2 .

Detailed mapping of nmges of species is necessary in geographic work .

Napping by physiographic provjnce, rock type , soil type , or cl i matic regimes
Hill provide explanations for local distributional patterns Hhich t.Jill give
insight into taxonomic , floristic and evolutionary relationships of various
taxa in contiguous and disj unct regions.

In Table 16-3 an analysis of distributional patterns within some Blue

Ridge escarpment gorges in the Southern Appalachians is given to show the
variety of relationships that can be derived from a local study " ithin a
restricted area (from Cooper and Hardin , 1971).

The major intercontinental disjunctions in the geographic ranges are

presented in Section D.
332 16

Table 16- 1 . Partial analysis of distributions of genera of seed plants

occurring in eastern North America (from :<1ood, 1971).

Genera occurring in southern Appalachians 557

' ..
Genera represented in both southern Appalachians and
,.;estern North America 365*

Genera ± continuous l y represented betHeen southern

Appalachians and ,,,estern North America 237

Genera wi th taxa ,,,ith dis_;unctions between southern

Appalachians and Hestern No rth America 158*

Genera \.;ith disjunction(s) in range of one or

more species : 48
Genera of various distributions Hith one or
more taxa in southern Appalachians and dis-
junct taxa of same rank in western North
America : 46
Genera restricted to North America (or nearly
so) with related taxa in southern Appalachians
and t"estern North America : 20
Genera "Tith r e lict Tertiary distribution , \"i th
related taxa in southern Appalachians and
Hestern North America: 46

Gener a not represented in both southern Appalachians

and western North America 192

Endemi c genera, restricted to eastern North America

or nearly so 52

Te rtiary relict genera , most restricted to eastern

North Amer i ca and eastern Asia 57

Genera >-lith other distributions 83

Genera not occurring in southern Appalachians 475

Gene r a with one or more representatives in hoth east-

ern and western North America 144

Gene r a Hith other dist r ibutions 331

Total indigenous genera 1032

Nonindigenous naturalized genera 384

Total genera of seed plant s 1416

*Because some genera appear in more than one category , the subtotals will
add up to more than these figures.
16 333

Table 16-2 . Some intracontinental and endemic range s in North America (from
Hood , 1971) .

1. Genera represented in both Southern Appalachian and l,~estern North America .

(Partial List)

Abies Orchis Salvia Ambrosia

JuniEcrus Oxalis Stachys Antennaria
~ GeraniuT'1 Vaccinium Aster
Potamogeton Verbena Asclepias Bidens

2. Endemic Genera represented in Southern Appalachians but not in Hestern

North America . (Partial List)

Orontium* Sanguinaria Galax Pe ltandra

Nedeola* Leiophyllurn* Comptonia* AmianthiuI:l*
Nestronia* Dalibarda'~ Xanthorrhiza* Epifagus*
Hvdras tis'" Neviusia'~ Honot ropsis", Diervilla

*Honotypic

3. Relict Tertia ry Genera. most restricted to eastern North Ame rica and
eastern Asia b ut not in Hestern North t\.l'1erica . (Partial List)

Svmplocarpus Saururus Astilbe Shortia

Aletris Lirioclenclron Decumaria Pieris
Convallaria Hepatica Apios Ph r)'Iil§.
Tipula ria Pvrularia Panax Tr io steum

4. Genera restr ict ed to North America (or nearly so) Hith related taxa in
Southern Appalachians and Heste rn North America . (Partial List)

Thuja Robinia Dode catheon Pycnanthemum

Camassia Ceanothus Frasera Trichos tema
Xerop\wllum Dirca Phlox Collinsia
Eriogonum Oxypoli~ !!ydrophvllum Sericocarpus

5. Gene ra either knm·m to be Relicts of Arcto- tertiary Floras or showing

Relict Patterns of Distribution Hith survivors in Eastern and \')estern
North America and in one or more areas notable as Refugia for Arcto-
tertiary Relicts . (Partial List)

Taxus Disporum Tofieldia Asa rum

Tsuga Ervthronium Jr illium Berberis
lIystrix Narthecium Ostrya Calycanthus
Clintonia Stenanthium Aristolochia Dicentra

.-.
334 16

Tab l e 16- 3 . Some important distribut ional patterns of vascular plants in

escarpment gorges (from Cooper and Hardin, 1971) .

1- Species reaching their upper elevationa! limits

'.
A. Species fIlOS t common on coastal plain

Itea virginica Callicarr~a americana Viburnum nudum

Decumaria ba rbara Lorinseria arealata ealopagon pulcheUus

B. Species chiefly on the piedmont and coastal plain

Platanus occidental is Asimina triloba 2uercus stel la t a

Fraxinus pennsylvanica ~ pallicla Quercus marilandica
Ho ru s rubra Quercus falcata Diospyros virginiana
-------
Carya cordiformis
Liquidambar sty raciflua

II . Species reach i n g their 10lver elevational limits

Pinus pungens Comptonia peregrina Leucothoe recurva

III. Appalachian endemics

A. Present

Shortia galacifolia Diervilla sessilifo lia

Clethra acuminata Kr i gta montana
Stuart ia ovata Leucothoe recurV8
Hydrangea arbor escens ssp. radiata

B. Absent

Cladrastis lutea

IV . Characteristic species of mixed mesophytic forest

A. Absent , hut to be expected

Cercis canadens is Os t r ya virginiana

Acer pensvlvanicum Hagnolia tripe t a l a

B. Present, but of very rest ri cted dist r ibut i on

Betula lutea (above gorges) Hagnolia acuminata

Acer saccharum Halesia carolina
Aesculus octandra

V. Problematic distributional patterns

Fagus srandifolia Quercus rubra

Tilia hete rophylla Carya glabra a nd ovalis
16 335

Section D. HAJOR DISJUNCTIONS IN THE GEOGRAPHIC RANGES OF SEED PLANTS",

Among the most fascinating aspects of plant geography are the very Hi de ,
intercontinental disjunctions in the natural geographic ranges of many seed-
plant groups . Most intriguing to the geographer are the possible explanations
for these wide gaps in range, e.g ., the floristic links betHeen Africa and
America , and their possible bearing upon the theory of continental displace-
ment. Most enlightening to the systematists are the evidences of taxonomic
relati onships shm.,rn in the analyses and explanations for these distributional
patterns .

This treatment is an attempt to review and classify the major intercon-

tinental d i sjunctions in the geographic ranges of seed plants ; six guiding
principles are follm.,1ed Hhere possible . Sixteen categories and 34 subcate-
gories of disjunct ranges of intercontinental magnitude (Table 16- 1) are
classified according to continents and latitudinal bands invo l ved. \,Jidely
distributed taxa are, by their very nature, disjunct; they are therefore
classified along Hi th the more traditionally discontinuous types . Thus, for
the most part , on l y endemics and taxa Hi th smaller intracontinental d i sjunc-
tions are omitted from this revie\~ .

Generalized distribution maps are presented for selected taxa characteris -

tic of some of the discontinuous types. Other taxa are list ed for each of the
categories and subcategories , and compendia of good distribution maps are cited
for ready reference to additional examples . Possible e xp lanations fo r the major
discontinuities are discussed .

Finally, a statistical summary is given of the numbers of disjunct taxa

and the largest disjunc t categories . Among the Angiospermae (the flm~eri ng
plants), app r oximately 78 per cent of the 324 families, 24 per cent of the
genera , and 1 per cent of the species are Hidely disjunct. The largest num-
ber of disjunct gene r a are found among the African-Eurasian- (Pacific) (600),
Asian- Pacific (460) , Pacific (370), North American-South American (ca. 360) ,
Pantropical (334). North Temperate (316) , and Subcosmopolitan (125) categor-
ies; the largest number of disjunct fami l ies among the Subcosmopolitan (90),
Pantropical (59) , North Temperate (20), African-Eurasian-(Pacific) (ll) , and
Nor t h American-South American (13) categories .

To achieve the goals in this section of (I) revieHing Hhat is knOl.ffi about
the major distributional disjunctions, (2) classifying their patterns, (3)
mapping or listing examples of each major type, (4) citing use ful compendia
of range maps of seed plant taxa, and (5) discussing the possib l e explanations
for the major discontinuities , I have been guided by several principles , or
general considerations, \o.'hich are hereHith briefly stated. All should be
self - evident but a r e all too often ignored . Some have recently been stated
by Wood (1971) in his excel lent paper on floristic relationships bet\,;een the
southern Appalachians and western North America, an intracontinental range
disjunction not considered here .

*Contributed by Robert F . Thorne , Rancho Santa Ana Botanic Garden, Claremont ,

California . This is a condensed version of " l>lajor Disjunctions in the
Geographic Ranges of Seed Plants " in The Quarterly Revie\~ of Biology
47(4): 365 - 411. Used \~ith permission .
336 16

GUIDING PRI NCIPLES

1. Only taxa that have been reliably revised should be serious ly

considered .

2. Only accurate distributional data should be used in constructing

distribution maps .

J. Almost all taxa have discontinuous ranges.

4. The current distribution of a taxon may not indica te its past

distribution .

5 . Coincidence i n range disjunction, although certainly suggestive,

does not mean that t\W taxa have similar dispersal h i stories .

6. The smalle r the rank of a taxon, the more instructive is it s

disjunct range.

Classification of In tercontinental Disjunctions in Range

In classi f ying the major types of intercont inental discontinuities in the

ranges of seed plants (Table 16- 4), I have had to make rather arbitrary deci-
sions to limit the number of categories. Eurasia, Afri ca , and Australia are
considered to be the Old Horld continents, although Australia is occasionally
treated as a very large island . North and South America , the New h1orld, are
mostly treated as separate continen t s , but sometimes it is useful to treat
the two as a unit. The Pacific-Indian Ocean basins and the Atlantic-Caribbean
basins are both considered of continental rank with respect to marine seed
plants . Australasia, as a phytogeographical area, here includes Australia
and Tasmania, NehT Zealand , New Caledonia , and usually Ne,~ Guinea, all with
their satellite islands.

I. EURASIAN-NORTH ANERICAN

Because Eurasia and North America lie across several major latitudinal
belts, it seems best to subdivide this areal type into Arctic, Boreal, and
Temperate .

1. Arctic .

Hide-ranging plants that display this far-nor thern distribution pattern,

by geography, are necessarily disjunct betl~een Eurasia and North America across
the Arctic Sea, and often also across the North Atlantic between Europe-Iceland
and Greenland-North America and across the Bering Sea bet,~een Alaska and Asia .
These are plants which are believed by Hu1ten (1937) and Love (1962) to have
survived maximum Pleistocene glaciation in refugia mostly north of the ice
sheets. Hith the melting back of the maximum ice, they have quickly spread
into their present far northern ranges.
 TABLE 16-4 ....
~

MAJOR DISJUNCTIONS IN THE.. GEOGRAPHIC RANGES OF SEED PLANTS

1. Eurasian-North American I V. African-Eurasian (-Pacific)

1. Arctic 1. African- Mediterranean
la o Circum- Arctic 2. African- Eurasian
lb . Beringian- Arctic 3. African- Eurasian-Ma1esian
Ie. Amphi - Atlantic- Arctic 4 . African-Eurasian-Pacific
2. Boreal 5. African- Eurasian-Australasian
2a. Circum-Boreal 6. Indian Ocean- Eurasian (- Pacific)
2,b. Beringian-Boreal V. Amphi-Indian Ocean
te. Arnphi-Atlantic-Boreal VI . Asian-Pacific
3. Temperate 1. Asian- Papuan
3a. Circum- North Temperate 2. ASian-Papuan- Melanesian
3b . North and South Temperate 3. Asian-Papuan - PaciEic Basin
3c. Fragmentary North Temperate 4. Asian-Papuan-Australasian
3c-l. Amphi-Atlantic Temperate VII. Pacific Ocean
3c-2. Mediterranean-American VIII . Pacific-Indian-Atlantic Oceans
3c- 3 . Eurasian- Eastern and Hestern IX . American-African
American X. North American- South American
3c-4 . Eurasian- Eastern American- 1. Tropical
Nexican 2. Temperate
3c-S. Asian-Eastern and {"estern 3. Bipolar
American XI . South American-Australasian
3c-6 . Eurasian-Eastern American 1. South American-Australasian
3c-7 . Asian-Eastern American 2 . South American-Aus tra'l asian- Asian
3c-S. Eurasian- I.;restern American 3. South American - Australasian-Madagascan
3c-9 . Asian-Hexican Highland XII. Temperate South American-Asian
3d . Wide Intracontinental Disjuncts XIII. Circum-South Temperate
XIV. Circum-Antarctic
II . Amphi-Pacific Tropical XV. Subcosmopolitan
III. Pan tropical XV! . Anomalous

w
w
~

"
338 16

1a. Circum-Ar ctic . The circum-Arctic type of disjunction comprises the

Arctic circumpolar and the circumpolar, Arctic-montane plants of Hulten (1937) .
The circum-Arctic species are found in northern Alaska and Canada, the Cana-
dian archipelago , Greenland, Iceland , northern Scandinavia , Spitzbergen, Franz
Josef Land , Novaya Zemlya, the Siberian Arctic, and often farther south at
alpine l evels in the Eurasian and North American mOlln t;ains. a1 though each is
usually lacking from one or more of these areas . Few species are continuously
circumpolar. Hulten (1963) counts 352 species as circumpolar or more or less
circumpolar . He counts another 107 taxa as having a more or less circumpolar
range but represented by different races or slightly differe.nt species on
both sides of the Atlantic . Be considers this flora an old one, and thinks
it probable that it "may have been spread by wind blo'h'ing over the frozen
Polar Sea or by floating on ice" (p . 72) .

Only a relatively feH small genera are largely restricted to c ircum-

Arctic (and alpine) ranges, namely Arctagrostis , Arctous , Braya, T>iapensia ,
Dryas , Dupontia , Loiseleuria, and Oxyr ia . Honkenya , Koenigia , and Phippsia
are also essentially circum-Arctic , but they reappear i n southern South Amer -
ica and \"il 1 be discussed beloH . The species Diapensia 1apponica L. illus-
trates this circum-Arctic type of distribution.

lb. Beringian-Arctic . This sub~roup consists o f those plants \1hich are

ma inly distributed on the Arctic shores of North America and Eurasia on either
side of the Bering Sea but Hhich do not reach both sides of the North Atlan-
tic . Included a r e the northern Beringia radiants and Arctic-Pacific plants
of Hulten (1937) . Hu1ten s uggests that these plants probably survived the
Pleistocene glaciations in refugia in Alaska- Yukon, the northern part of the
present Bering Sea , and in northeastern Siberia . Some formerly may have had
more complete circum-Arctic ranges that \"ere much reduced by glaciation in
eastern North America , Greenland , and the European Arctic .

One plant that sho,,]s the Beri.ngian-Arctic type of disjunction is Diapen-

sia lappon i ca L . subsp . obovata (Fr . Schm . ) Hult ., separated by considerable
gaps in northern Canada and Siberi.a from the amphi-Atlantic Q. .!iP.ponica L.
subsp . lapponica.

lc . Amphi-Atlantic-Arctic . This group is the reverse of the Beringian

group of Arctic disjuncts, for the species ar e mainly distributed on both
sides of the North Atlantic . They have received much critical attention from
Hulten (1958, 1963) . Hulten (1958) maps 278 species , but these include Arc-
tic, boreal , and temperate species, 129 of the Arctic and 148 of boreal or
temperate distribution . Hulten (1963) explains many of these amphi-Atlantic
plants as fo rme r circum-Arctic species \1hose ra nges have been much r educed by
changing c limatic conditions. Some may have spread, hm1ever, f rom centers on
one side or the other of the Atlantic . Ce rtainly some of the maritime and
aquatic species can be accounted for by long-distance dispersal by sea cur-
rents o r by birds . There seems little need, if any at all, for a Late Ter-
tiary or Quaternary land-bridge across the North Atlantic to account for
plants having this type of di sjunc t range .

Diapensia lapponica L . subsp . lapponica and Salix herbacea L. illustrate

this type of range discontinuity .
16 339

2. Boreal.

Plants cons idere d to have a disrupted boreal range in North America and
Eurasia are those ,~hose nor t hern limits generally lie sou t h o f the Arctic
shores and whose upper altitudinal limits do not reach the alpine levels of
the Arctic-alpine species . They are absent usually from Greenland and often
from I celand. Hany have been mapped by "ulten (1958, 1968) and by Heusel,
Jager , and \~inert (1965) . Unlike the Arctic disjuncts , these plants appar-
ently survived maximum glaciation in refugia south of the ice (Hulten, 1937) .

2a. Circumboreal. Plants having this wide distribution in the boreal

belt of Eurasia and North America have been called boreal circumpolar plants
by Hulten (1937), and they are very well knotm to temperate botanists . They
are especially abundant in the coniferous forests , bogs, marshes , and oligo-
trophic lakes of this zone .

Some of the more familiar species of this circumboreal disjunct group are
Adoxa moschatellina 1.. . , Calla palustris L ., Caltha palustris L . (s. l. ), Calypso
bulbosa (1.) Rchb . f . , Carex chordorrhiza Ehrh . • .f. disperma Dew .• .f. rostrata
Stokes , Cvpripedium calceolus 1.. Drosera rotundifolia 1. , Juniperus communis
1., Henyanthes trifoliata 1., l'loneses uniflora (1.) Fray, Monotropa hypopitys
L . , l>lyrica gale L . , Ranunculus reptans L., Rhynchospora alba (1.) Vahl, Rubus
idaeus L., Sparganium angustifolium Hichx . , Subularia aguatica L . , Veronica
scutellata L . , and Viburnum opulus L . (5 . 1 . ) .

2b. Beringian-Boreal. Taxa o f this subgroup have much less extensive

ranges than the circumboreal species . They are primarily distributed on e ither
side of the Bering Sea in eastern I-:urasia and l~estern North America, but they
do not generally reach the Arctic shores of Beringia and they do extend far-
ther south on ei ther side of the Pacific Ocean . Oplopanax , l~ith tHO or three
closely related species in North America , Japan, and Korea , is repres entative
of this discontinuous type . In North America O. horridus (Sm.) Hiq . • the
infamous Devil '5 Club of the Het coniferous forests o f the Pac ific NorthHest,
illust rates a Hell-knmm intraco nt inental disjunction by its jump from Hon -
tana to Isle Royale and adjacent islands on the north side of Lake Superior .
At least eight additional genera, Achlys , Chamaerhodos, Glehnia, Leptarrhena ,
Lysichitum, Fauria, Smelm,'skia, and Thellungiella, have a Beringian-boreal
tyue of discontinuous distrihution .

Some o f the boreal species or species-pairs ~1ith ranges in eastern Asia

and ",estern America, a fe'" reaching across boreal America to the Atlantic
Coast, are Ach!E triphylla (Smith) DC.-.6. . japonica Haxim . , Achillea sibirica
Ledeb., Aruncus sylvester Kostel . , Carex macrocephala Hilld ., Cassiope lyco
podioides (Pall.) D. Don, and Cornus canadensis L .

2c . Amphi-Atlantic-Boreal. This subgroup, along with the more northerly

amphi-Atlantic-Arctic group, has been thoroughly discussed by Hulten (1958 ,
1963) . Because of their occurrence on both sides of the Atlantic Ocean , spe-
cies having this type of disjunction are Hell known to European and American
botanists. Rhynchospora'" fusca (L.) Ait . is just one of some 1 48 species ,
most of which belong to this group, that have been mapp ed by Hulten (1958) .

3. Temperate.

This very large group of disjunct plants is certainly the best knmm and
340 16

most frequently studied of all, as might be expected from the heavy concen tra-
tion of the world ' s botanists in the North Temperate zone . To this group be-
l ong the many genera that dominate the temper ate deciduous forests of Eur asia
and North America . It is an ancient group , mostly origina t ing no later than
the widespread early Tertiary forests of the northern hemisphere . Many genera
have retained their wide ranges of Tertiary time s ; others have become reduced
to fragments of their former vast ranges. These latter have received much
more attention because of the i r rather romantic, highl y discontinuous ranges ,
but surely they are no more informative to the geographer than the genera that
retain most of their former ranges. Bo th kinds of genera , however , together
with the fairly extensive fossil re cord for the woody re l icts , tell us much
about the vicissitudes the Tertiary forests must have experienced to the
present time .

The more fragmented North Temperate d i sjuncts are today preserved in the
scattered deciduous forests of the North Temperate zone, primarily in eastern
Asia (Japan, Korea, Taiwan , and China), th e lm"er Himalayan s l opes, the Near
Eastern Casp ian- Caucas us - Black Sea areas, Europ e (including the Nediterranean
regio n ), easter n North America , the Nexican and Centra l American highl ands,
and western North America .

33. Circum-North Temperate. By analyz i ng various r egional temperate

flora s . \-,1i11is I Die tionary (1966), and Engler I s Syllabus (1964), I have been
able t o list 118 wide- ranging, primarily tempe r ate genera, 62 that are t reated
here as circum- North Temperate, and 56 that are equal ly ,,,idely distributed in
the North Temperate zone but are also r epresented in the temperate areas of
one or more of the southe rn continents . Most of the 118 possess re l atively
uninterrupted ranges across North America and across Eurasia . The 62 assigned
to the circum-North Temperate group mostly do not cross t h e equator . The few
that do , like Acer , Alnus , Arabis, Campanula , Corydalis, Draba , Juniperus,
Pinus, Oue r cus , Rosa , and Sedum, are confined in the southe rn hemisphere to
the tropical mountains. Man y of these gene ra are by no mean s restricted to
the t emperate zone but have one or mo r e species represen ted in the Arc tic or
boreal zones (as Arenaria., Armeria, Betula , Chimaphila, Draba, Juniperus ,
Hertensia, Orchis. Parnassia , Pedicularis , and Silene) or in the tropics
-(as Acer . Lonicera, Pinus, and Quercus) .

3b . North and South Temperate . Presumably most of the preceding circum-

North-Temperate genera have achieved their far-r eaching ranges through the
normal dispersal capacities of their member species. Their species, a t l east
in total, must have a wide amplitud e of tolerance to often severe environmen-
tal conditions in the continental heartlands . Many of the 56 widely distri-
buted North Temperate genera that have suc cessfully migrated across the
tropics into temperate a reas of one or more of the three southern continents
have likewise probably Moved by normal short-range dispersal along relat i vely
continuous mountain chains , as Descu r ainia, Epilobi um, Gentiana, Lotus. Ribes,
Salix , Saxif raga , and Trifolium. On the other hand, other genera s hoH very
large discontinuities across the tropics , as in Anemone , Euphrasia. Geum, Pha-
lari s , Primula , and Spa rganium . These amphitropical genera may have achieved
such large disjunctions by the extinction of montane species formerly s p an-
n ing the gaps, or perhaps more likely. by having disseminules transporte d from
one temperate zone to the other by migrating birds.

3c . Fragmentary North Temperate . Organized under this subheading i s a

variety of ,,,idely discontinuous groupings that have conside r ably l ess exten sive
16 341

ranges in Eurasia and North America than the groups discussed above . Their
disjunctions in one or both continents probably represent s ever e restriction
of ranges that were relatively continuous during more favorable Tertiary cli-
mates. Most of the taxa thus are Tertiary relict s . Lon g-d is tance dispersal,
however, is more than likely for some of the maritime, marsh, and aquat ic
plants .

3c-1. Amphi-Atlantic Temperate . Since the \.,tarmer parts o f Europe and

North America lie widely separated by the Atlantic Ocean, I'lith fe\V islands
of any consequence between the continents to serve as stepping-stones, it is
not surprising that this type of discontinuity is very rare in comparison Hith
the relatively frequent Arctic and boreal amphi-Atlantic disjuncts . It is best
represented by Corema, with f. alba (L.) D. Don along the western coast of the
Iberian Peninsul a and on the Azores and f. . conradii Torr . , the only other spe-
cies, along the Atlantic coast of North America from Newfoundland to NeH Jersey .

Of all the amphi-Atlantic species mapped by Hulten (1958). only 6 angio-

sperms are primarily temperate amphi-Atlantic disjuncts . They are Drosera
intermedia Hayne, Eriocaulon septangulare Inth ., Juncus tenuis Hilld .• Limo-
sella subulata Ives, Ranunculus hederaceus 1., and Spar tina patens (Ait .) NuhI.
It is surely no coincidence that these are all aquatic, maritime , or rudera l
species .

3c-2. Nediterranean-American. Approximately 35 genera have their maxi-

mum development in the xerothermic, Hediterranean climates o f the Old Horld
Hediterranean region and southt.,testern North America o f the New \\lor l d. some of
them also ranging disjunctly to other areas o f Nediterranean climate i n South
Africa. Australia , or Chile. This kind of discontinuity has been of special
interest to Californians because the Hediterranean climate in North America is
centered about southern California . Four families , the Cistaceae, Cneoraceae.
Ephedraceae , and Resedaceae; 18 groups of vicarious species or speCies-pairs,
listed in Stebbins and Day (1967); and one species, Styrax officinalis L. ,
also possess this kind of major disjunction .

3c-3 . Euras ian-Eastern and Hestern American. Of the North Temperate

genera with fragmentary ranges this category of td de disjuncts has ranges
most similar to the nearly continuous circum-North Temperate genera . At
least 24 genera , although with rather large gaps on one or the other conti-
nent, and o ften on both, are found in Europe, one or more areas of Asia, and
in both eastern and t.,testern North America . Among these relict s are such t.,tell-
knOlm temperate genera as Aesculus, Amelanchier, Cercis , Corylus, Juglans,
Ostrya , Platanus , Staphylea, Styrax , and Taxus .

3c-4. Eurasian-Eastern American-Hex ican. Hard l y distinct fr om the

above group, the genera in this subcategory are not found in the western
United States and Canada but are found in the highlands o f Nexico (and often
in Guatemala or farther south) . The 6 genera that have this special type of
disjunct range are Carpinus , Clethra (this genus is slightly aberrant since
it is absent from Europe". but is found in Hadeira), Fagus , Halenia, Tilia ,
and Ulmus . Some of the Eurasian -East ern and ~.Jestern American disj unct genera
discussed above , li ke Amelanchier, Juglans , Ostrya, Platanus, Styrax, and
Taxus , are also represented in the Nexican highlands and often farther south.

3c- 5. Asian-Eastern and Western American . Another large category of

Tertiary relicts are those genera , at least 38, that are r epresented i n both
342 16

east ern and west ern North America , but are nOH absent from Europe and mostly
also from the Near East . The y usually have their cente rs of development in
easter n a n d southeastern As i a . Among these genera are Aralia , Calycanthus ,
Chamaecyparis . Cl intonia . Dicen t ra , Disporum. Leucochoe . Phvsocarpus . Smi la-
cina , Thuja . Torreya, Toxicodend ron , J rillium . a nd Tsuga . Excellent maps
showing the disjunct American ranges of some of these . and some other genera
have recently been published by \\Iood (1971) .

3c-6 . Eurasian- East ern North American . There seem to be only four gen-
era , Castanea, Convallaria . Epigaea, and Po l ygonatum, that occur both in
Euro pe and As ia and on l y in the eastern pa rt of North Ame rica . Presumably
t he east-\Y'est orientation of mountain chains and seas in Eu r ope . wh ich pre-
vented the escape of tempe rate species southward befo r e the Pl eistocene gla-
ciers , was a lmost as disastro us to the s urvival of Tertiary genera in Europe
as was the severe deteriora tion of climate in western North America .

3c- 7 . As i an-Eastern Amer ican . This is the largest category of Te rt iary

relict temperate genera, the genera having found haven only in eastern No r th
Ame r ica and in ASia , most o f them only in eastern and so utheastern Asia . I
have l isted 74 genera in th i s much - studied group, early b rought to o ur atten-
tion by Asa Gray (1846, 1859) . and more recently discussed by Li (1952) and
by Hood (1971) . Among the more conspicuous or better knOl-JU of these numerous
gen e r a are : Arisaema , ~ . Catalpa, Caulophyllum, Cladrastis . Croomia,
Gordonia. Hamamelis . Il l icium . Liguidamba r. Liriodelldron, Nagnol ia. Ne l umbo .
Nyssa . Panax . Podophyllum , Sassafra s , Sauru r us . Schisandra, Shortia, S t ewartia.
Symplocarpus , \-li steria , and Zizania. Some of these are as correct l y treated
in the amphi- Pac i fic tropical dis junct category . fo r their ranges in either
America or As ia or in both are as tropical as they a r e wa r m temperate . Among
s u ch genera are Berchemia , Cle thra , Gordonia , Illicium, and Schisandra . Two
ge nera , Arundinaria and Symplocos , ~lere placed in the amphi Pacific tropical
disjunct categor y because all but one of t heir American species a r e in t he
tropics, although often at mon t ane elevations .

Because so many of the Asian-Eastern American disjuncts have preservable

parts. they have left a rather r ich fossil record in areas ,,,here they a r e now
ex tinct . The di stribution of fossils in su ch genera as Liriodendron, Magno-
lia (s ee Meusel , 1969). Nelumbo , and NYssa (Eyde and Barghoorn, 1963) , in di-
cates that these highly discontinuous tempe ra te-to-tropical genera once had a
nearly continuous distribut i on across Eurasi a and North Ame r ica in the Ter-
tiary . Presumably the same is true for t he herbaceous dis j uncts, \"hose
chances of being preserved as fossi l s are much less .

3c- 8 . Eur a sian-\~estern North American . A small group of primarily tem-

perate genera are the nine restricted to I.estern North America and Eura sia ,
mostly eastern Asia . Because they are few and the disjunction is rela tively
les s spectacular, this group has undescrvedly received little attention .
Heterocodon , Paeonia, a nd seve r al generic-p air s have been discussed by Stebbins
(1940) . According to Stebb ins (1938) the two I"estern American species of
Paeonia are closer to t. delavayi Franch. of southwestern China than to the
herbaceous species of Europe . The other genera 11ith this t ype of disj unct
ran ge are Adenocaulon, Boschn i akia , Calocedrus , l'-lahonia, Phot inia , and Pseu-
dotsuga . They are little differet1t from the nin e genera earlier listed as
having a Beringi an-boreal distribution. In fact, this Eurasian-Hestern North
American group could just as well be called Beringian-temperate or even amphi-
Pacific Nor t h Tempe r ate .
16 343

3c - 9 . Asian- Nexic8n Hi gh l and . Hardly distinct f r om the preceding group

of genera a r e the six gene ra fo und only in eastern As ia and the Hexican high-
lands or fa rther south . Several of the preceding group , such as Adenoc au lon ,
Mahonia , Photinia , and Pseudotsuga. occur also in the cool He xi can or Guat e
malan highlands . The six that do not occu r in North America north of Nexi co
are Abelia, Deutzia. Langsdorffia. Leibn itz i a, Hitrastemon . and Sa r cococca .
These could also be absorbed . perhaps as well , in the amphi-Pacific trop i cal
group of discon t inuo us genera di scussed nex t be l m... .

In addit ion to the numerous genera, many li sted abo ve, that have a fragmen-
tary North Te mperat e distribution , there are at least 18 f ami lies and three s ub -
families so restrict ed : Calycanthaceae , Cistaceae, Cneoracea e , Datiscaceae,
Ephedraceae , Hippocastanaceae, Hydrastoideae, Illi ciaceae , Juglandaceae ,
Hi tras t emon oideae, Nyssaceae , Paeoniaceae, Platanaceae, Podophylloideae .
Resedaceae , Saururaceae , Schi sandraceae, Stemonaceae , S tyracaceae . Ta xa ceae ,
and Taxodiaceae .

3d . I~ide I ntracontinental Disjuncts . Hith in the temperate 7.one

of North America on the one hand and of Eurasia on the other there
are taxa with spectacular disjunctio~s that are not considered her e because
they are intracontinenta l. In distances involved, ho,,,ever , they a r e often
mo r e widely discontin uous than those tha t we are attemp t i ng t o class i fy here .
Such North American endemic disjun cts as Camassia, Dirca , Dulichium (wi th
fossil recor d in Europe) , Il iamna, Sullivan tia , and Xerophy llum, are mapped
and many others are d iscuss ed by \~ood (1971) .

II . AHPJlI-PACIFIC TROPICAL

This discon tinuous g r o up includes all those taxa that are fo und bo t h i n
tropi ca l Amer ica and the t r opical lands on t he '~estern borders of the Pacif i c
Basin . A t o tal of 89 genera , 4 t ribes o r subtribes, J or 4 s ubfamilies , and
8 to 11 families of flower i ng p lant s are amphi-Pacific t ropical in their dis-
tribut ion . The number of taxa is inexac t because of the overlap with cate-
gories disc ussed above . Clethra , Hag(1o l1a , Hi trastemon, and Nelumbo are
particularly troublesome , and probably rightly should be included here . Van
Steenis (1962) ha s presented a useful list of "amphi-transpacific genera and
other affinities" organized in four l atit udinal groups . Some of the mos t
note,~ ort h y arnp hi-Pacific tropical genera are : Anaxagorea , Cleyera , Endian-
dra, Hedyosmum, Nelio sma , Passiflora, Pentapanax , Perrott etia , Persea , Phoebe,
Picra sma , Rhynchoglossum , Sau r a u ia, Sloanea , Sp athiphyl lum, Svmplocos , Talauma ,
Turpin ia , and Xy l osma . Al l of these and 27 more reach southeastern ASia , in
clud ing the Nales ian islands Hest of New Guinea . Six t een additional gene ra
range muc h far ther to the Mascarene Islands, Nadagascar , or even easte r n
tropical Africa . Among them are Aphananthe , Calliandra, Callicarpa , Ca lophyl -
lum , Clu sia , Elaeocarpus , Glochidion , Protium , and Suriana . Sixteen genera
reach only to Austra lasia ; these inc lud e Batis , Br edeme yera, Epistephium ,
Iresine , Licania, Lindenia, Huehlenbeckia , Nicotiana, a nd Sicyos. Fou r gen-
era , All agoptera, Br achist us, Leucaena , and Pritchardia , r e a ch neithe r Aus -
tra lasia nor Asia but do reach Pol ynesia o r as far wes t as the Fi ji and
Solomon Islands .

Of the larger ca t egories , th e families Actinidiaceae (including Sa urau1 a) ,

Clethraceae , Magnoliaceae , Nelumbonaceae , Sabiaceae (including l'!eliosoma) ,
Staphy l eaceae , and Symplocaceae, although each has some ,"arm temperate repre-
sentatives , are essentially amphi-Pacific tropical in dist r ibution . The
 344 16

:-j
. '-J.
.~ .

C~NF.JAlIU1> DI.ITlu .UTlOS 01 m E C,.cUM·BoHAL Boc Sr lc' .... Sch~u<h,eTid fMiuJiTis

The 1)'P'C<liluiMp. palwlr;, occuro in Lun .... and the Jub.p. dm.,.icdna in 1'0<1 1> America. Modified
from Meu.d, Jagtr, a n,l Winert (1%5) and Huh~n (1968) ; b. ", m.p a. in Fig. 2.

< .. ~-
T fi E WOR LO -......:...:._ ....
.. ",,, ""''' ." . " "'.<ToO.

G DfnALIZm DISnlBI!I10N 0 7 nu: MIAN·I.I.nDM AWD.JCAN GENU. Ham<:melu, THE WrTal.H.uru:

Bucd in part on U (1952); twc IIl<Ip as in Fig. 1.

Figure 16-1 . Generalized distribution o f Scheuchzeria and Hamamel is. Used

\,'ith permission .
16 345

T HE WORle
'- "','"-'-
"., ' ;y;'~~
\

."0" ,.....c .... . 00"' ''0'

PAST ... " .. Pusi:I'IT KNOWN DISTI.1BUTLON OJ' TilE MIAN-E.uTnN ANU.lCAN, 0 .. haHAP.I Bl.TTD
AM"m-PACIl'IC T~P[CAL, CENU5 Ndumbo, WAD.1-LOTlIS
Modi6ro from Good (1964); b;uc m!lp aI in Fig. I. The blilCk drdes rcpraent fouil nrords.

CU<nALlttJ> DJSn.lBUTION or mE P"N"BOPICAL GIN~'$ DiOJ/ryro, (•. L.) or nIl EaUlAaAl.

Much modified from Fem;l1d (1951); blle map ;u in Fig. I.

Figure 1 6-2. Distribution of Ne l umbo and Diospyros. Used \·.rith permission.

-.
 346 16

Bataceae reach only Australasia, but the Trigon iaceae , Elaeocarpaceae, and
Chloranthaceae are represented not only in As ia but in Madagascar as \VeIl.

III . PANTROPICAL

The samE! rather conservative approach to genera .... .as ,.;ras used else,vhere
in this survey ,dth other taxa , has been used to drm" up a list of 231 genera
of seed p lan ts that are trul y pantropical, i . e ., are represented by indigenous
species in all the major tropical areas of the lvorld. These major tropica l
areas are defined as including tropical America, tropical Africa, tropical
As ia inc luding Malesia, and tropical Australia . An additional 103 genera are
not believed to have indigenous species in Australia (Burbidge, 1963) but occur
elsewhere in all the major tropical areas. Thus , we can accept a total of 334
widely distributed tropical genera (many with some temperate species) , a rather
tiny total out of possibly 12 ,500 currently accept e d genera of seed plants.
Fifty-nine angiospermous families are also essentially pan t ropical .

I V. AFRICAN-EURASIAN (-PACIF I C)

An analysis of Hillis ' Di ctionary (1966) p roduced 555 currently accepted

genera that range from mainland Africa to Eurasia or beyond . They are readily
divisible into six subcategorie s .

1. African-Mediterranean .

Only about 9 genera are restric ted essentially to Africa south of the
Sahara and to the Nediterranean region, a fet~ extending north into Europe or
eas t into the Orient . These are Althenia, Capnophyll~.!!!. , Caris , Cytinus,
Desmazeria , ~, Hyacinthus, Romulea , and Tolpis.

2. African-Eurasian .

Thirty-eight genera, including such t~ell-kno\m plan t s as Adenium, Aeonium,

Aloe, Catha, Cotyledon , Huernia, llyphaene , Fissenia, a nd Sansevcrinia, and the
family Barbeyaceae, extend from Af.rica only to Ar abia. At least 200 Eenera
and 3 families, Trapaceae , Tamaricaceae, and Moringaceae, reach f arther into
Asia, usually at least to India, but do not extend in to Nalesia . These include
Balanites, Borassus, Bost.Je l lia, ficer , Commipho ra , Gladiolus , r1oringa, Ochna,
Pedalium, Phoen i~ , Salvadora, SCilla , Sesamum, Tamarindus, and Trapa .

3. African-Eurasian-l·lale sian .

Another 106 genera have repres enta tives in Hales ia, which here is defined
to include all the island groups fro m l>!alaya to the Nel·} !lebrides . A few o f
these are An8raecum, Asparagus , Azima, CnestiE_, Combretodendron , Ellepanthus ,
Ensete, Irvingia , Phrynium , Pothos, Quisgualis, Sauromatium, Stromboisia , and
thightia . Six families have a similar disrup ted rang e : Ancistro cladaceae ,
Ctenolophonaceae, Dipsacaceae, Dipterocarpaceae, Pandaceae, and Salvadoraceae .

4. African-Eurasian-Pacific .

Another 100 discontinuous genera have representatives also in Fiji, Tonga ,

Samoa, or mo re distant islands in the Paci f ic BaSin , as Alangium, Barringtonia,
Bruguiera.• Canarium , Cerbera, Claoxylon , Elatostema, Flagellaria, Gardenia ,
16 347

Gre\"ia. Jasminum, Korthalsella, Leea. Lumnitzera. Nyri stica , Olea, Pandanus,

Pittosporum, Premoa t Syzygium, Taeniop hyllum, .exlocarp us , and _Zeuxine . Six
families extend disjunctly from Africa to Fiji or the Pacific Basin: Alangia-
ceae, Casuarinaceae, Cycadaceae. Flage ll ariaceae , Pandanaceae, and Pittospora-
ceae.

5. African-Eurasian-Australasian.

Ninety-six additional genera extend so uth into Australasia, some \dth

representatives on New Caledonia (Acanthus, Acridocarpus . Aponogeton, Cer iop s .
Olax , $onneratia , and Tristellateia) or New Zealand (Ga s trodia. Glossostigma.
Iphigenia , Pelargonium, and _Sebaea) . Actually 205 of t he 555 African-Eurasian-
Pacific disjunct genera have representatives on the continent of Australia .
al t hough many I.ere included in other totals above because they e xtend fa rther
east into the Paci fic .

6. Indian Ocean-Eurasian (-Pacific).

Forty-seven genera t hat do not reach mainland Africa do have discont inu-
ous ranges from I-ladagascar, the Nascarene, Cornaro , or Seychelle Islands to
Eurasia, mostly to southeastern and eastern Asia and to Hal esia. and often
to Australia o r the oceanic islands of the Pacific Basin . Some of the better
known of these genera are Allantospermum . Alyxia, Balanophora, Carallia,
peeringia, Dillenia, Elaeocarpus. Erythrosperrnum , Foetidia, Geniostoma, Hal
oragis, Hedvchium, lIiptage , ~oporum , Nepenthes, Ochrosia , Pipturus , San
doricum , Timonius , and Vateria . When added to the mainland African Eurasian
(-Pacific) disjunct genera, slightly mor e than 600 genera share this common
pattern of disjuction .

In viel~ of the r e lat ively large number of dis cont inuous fami l ies and
genera reaching Eura si a and beyond from Africa and adjoining islands, one
must seek an exp l anation other than long-ran ge di spersal to account for this
very common type of disjunction . Some of the maritime, aquatic, and f l eshy-
fruited taxa on islands most likely are dispersed over large distances by
sea currents and bird s . HOI"ever, many of the wide-ranging taxa have probably
migrated over land or over small water gaps by normal methods of short-distance
dispersal at a time when much of northern Africa, Arabia, and southern As i a
were much moister than they are at present. In add iti on , there may have been
much more land between the continenta l Seychelle Islands and Ceylon and India
in the past, ove r which some of the taxa might have migrated . The us ual
Gondwanaland , can tinental-drift, tectonic-plate movement , and similar exp lana-
tions have little if any bearing here because such geologica l phenomena must
have taken place , if ever, long before even these relict distributions were
initiated .

V. AHPllI-INDIAN OCEAN

Very few taxa are restric ted j ust to Africa and Australasia, thus dis-
junct across the Indian dcean . Only 18 genera apparently are so discontinu-
o us between mainland Africa and Australasia . mostly Australia, with another
11 having representation in Australasia and only I>ladagascar, the }iascarene , or
the Seyche l le Islands. This total is likely to be reduced as species are found
to be naturalized in one area or the other or as careful revision s shm~ the
representation on either side of the Indian Ocean to be generically dist inct
or to belong to larger , wider-ranging genera . This rath e r rare type of major
348 16

discontinuity direct l y contradict s the rathe r loose speculation of some

authors that ther e is a c lose relationsh ip between t he fl oras of Austra lia
and Af r ica . Inasmuch as 1 5 of the 29 genera have Madagascan . Hascarene , or
Seychel1e re presenta tives a nd 13 have insu lar pop ulations east of the I n di an
Ocean other than t hose in Australia-Tasmani a , long-d istance dispersal would
seem t o be the l ogical explanat ion fo r th ese dis j uncts . Adansonia , Anacamp-
se r as , Ch rys i thrix , Die tes , and Triraphis a re prima rily African o r Hadaga scan ;
wher eas Arthropodium, Bubbia , Caesta, Cassin i a , Cunania, Hclipter um, Hibbert ia ,
Humea , Kerau drenia . Hacadamia. Rulingia, Soulamea, and Vi llarsia are primarily
Australian , Papuan , or Ne\" Caledo n ian . Othe r gene r a are mor e evenly divided
(Australina , Brachyachne, Bul binella , Costularia , Entolasia , Granunatotheca,
Po t amophi la, and \o,Iur mb ea) , o r a r e pr i mar ily oceanic (Astelia , Cohn ia , a nd
Cossinia) .

VI . ASIAN-PACIFI C

Anothe r l arge discontinuous ca t ego r y , b ut one somewha t l ess wide- r anging

usually than the African-Asian ( - Pac ific) t ype of dis trib ut ion, is r ep resente d
by about 460 genera and five families . These taxa have species in main land
Asia and others in easter n Ha l esia t o t he Bisma r cks . i n the Solomons , i n t he
New Hebrides , in Fiji-Tonga- Samoa , or on mo re distan t islands in t he Pac i fic
Bas in. These disjuncts are best trea t ed in four subc ategories .

1. Asian-Papuan .

Nearly 200 genera r ange from Asia to New Guinea or t he Bismarck Archip e l -
ago : Anisoptera . Hopea , I xonanthes , Kadsura , Lithoca rpus, Pentaphragma , Ploi-
~ , Pome tia, Sar candra , Sho r ea , Svcopsis, Wa l sura, and Zanonia
among oth er genera , a n d t he two fami lies Crypter on iacea e and Pentaphragmata -
ceae .

2. Asian-Papuan-Helanesian.

Another 130 gener a reach beyond the Bismar cks to t he Solomon I slands (46) ,
the Santa Cruz and Ne\-! Hebrides I slands (15) , Fiji (29) , Tonga (19), or Samoa
(21) , inc luding Aeg iceras , Ai lanthus , Alpinia , Amoo ra, Artocarpus. Carvota ,
Disch i dia , Dysoxy1um . Hanguana , Hoya , Hydnophytum , Koe l reuter ia , Livis tona ,
Nadh uca , Hangifera , Helicope , }Ie troxy I on , ~ , Pal aq uium, Semecarp us , Tris-
tan i a , and Xanthophyl lum and t he fami l y Daphn iphyll acea e .

3. Asian-Papuan - Paci f ic Bas i n .

\..rell beyond the Andes i te Line in the Pacific BaSin , about 40 genera have
representation on the islands of Po lynesia (many on the distant i slands of
the Hawaiian chain) : c . g . , Alectryon , Alstonia, Berrya , Cyrtandra , Dendro-
bium , Fagraea, Freyc i netia , ,9a hnia. Nelastoma , Pl anchonella, and l..rikst r o e mia .

4. Asian-Papuan-Australasian.

To the south 92 additional genera have ranges t e r mina ting i n Aust ralia
(62 ), New Cal edonia (23) , or New Zeala nd (6) , i nc l ud ing Aegialitis, Apos tas i a,
Bassi a, Cardiopteris , Curcuma , Helicia , l'lelaleuca , Neolitsea, Philydr um , Po ly o-
sma , Siphonodon, Stemona. S tyl idium , Yanda, Tet r ame les a nd Zingiber , and th e tl'lO
families Cardiopter idaceae a nd Philyd ra ceae . Actually , i f one counts those
genera credited to t he So l omons and islands farther eas t i n the Pacific , 195
16 349

of the Asian-Pac i fic disjunct catego r y are believed to have ind igenous spe-
cies in Australia.

For many of the se taxa , reaching only to eastern "!alesia, Australia , or

the other con tinental i s lands, normal methods of short-distance dispersal are
probab l y adequa te to explain the currently dis junct ranges . It is assumed
that 1n th e pas t . particularly durin g periods of g r eat tectoni c activity and
during the glacial lowering of sea- levels, much more land a t time s connected
Nalesia to mainland Asia , and Australia to Halesia . For various good biogeo-
graphical considerations. s hort over-I-meer hops may have been necessary for
disseminules to r each such presently isolated continental islands as New Cale-
don ia, NeH Zea l and , th e Solomons, New Hebr ides, Fiji , and Tonga (Thorne, 1965),
1969) . Beyond the And esite Line lie only islands of ocean i c, volcanic o r igin.
These must have been r eached by disse minules carrie d over long distances by
bi r ds, sea or ai r curre nt s , or by man, altho ugh numerous atolls and guyots ,
f la t-topped sea-mount s . in the Nestern Pacific Basin do ind icate that once
there were more hi gh islands available there for stepping-stones (Thorne ,
1963) .

VII . PAC I FIC OCEAN

Approximately 370 gene ra and 7 families that r each neither the Asiatic
no r the American mainland have discon tinuous distributions betHeen at least
tHO Pacific isl and groups (Australia here being treated as a very large Pacifi c
island) . These are truly Pacific taxa . Some have relatively restricted ranges
t hat hardly merit comparison l-li th taxa shot'ling i ntercont i nen t al di sj unct i ons .
Such are the 44 genera disjunct be t Heen I-/este rn Nalesia (east to the No luccas)
and eastern l1alesia (New Guinea and the Bismarcks); 19 genera betHeen Austral ia
and Hel.,. Ca l edonia ; 48 bett.,.een Australia and New Zealand ; 5 between Austra l ia
and the New Heb rides or Fijis ; 16 bett. . een New Guinea and ~ew Caledonia (mostly
also on intermedia te ~lelanesi an islands or i n Queensland) ; one , Carpodetus ,
between New Guinea and New Zealand; 43 between ''''estern Malesia or NeH Guin e a
and the Fiji, Tonga , or Samoan Islands . Si milar t-Iestern Pacific disjuncts are
the Stackhous iaceae a nd the 5 genera that each represent monogeneric fami l ies:
Ea lanop s, Byb lis, Corynocarpus, ~omatia, and Galbul imima . Somewhat ,dder
gaps in range are shmm by the 39 gener a distributed from western Ha l es ia to
Australia (IS), to New Caledonia (11), and to NeN Zealand (10).

Hore Nide-ran ging in the Pacific are at least 37 genera that have repre-
sentation from Nestern Halesia or New Guinea to the Hawaiian or other Poly-
nesian is l ands (22), including Ascarina, Astronia , £oprosma, Cyathodes, Fara-
daya . Lepin ia. Neubergia, Olearia, Pseudomorus, San ta lurn, and Tr imen ia (Tri-
meniaceae); from Au stra l ia to the Ha,.,.aiian o r other Polynesian islands (Neste-
gis and Corokia ); or are disjunct within the Pacific Bas in (14) : Apetahia ,
Charpentiera , Cheirodendron , Cros sostylis , Earina, Fitchia , Heryta , Neso luma,
Oparanthus , Pelea , Pelasodoxa. Phyllostegia , Reynoldsia , and Sclerotheca .

Some of the widely distributed genera of the Pacific-Ind ian Ocean areas
discussed under categories IV and V could as wel l be treated he re as a s ub-
catego ry, Pacifi c-Ind ian Ocean disjuncts . At least 14 genera range from the
Indian Ocean islands, including Nadagascar , to Australia o r islands beyond:
Arth r opodium , Astelia , Bleeke ria, Bubbia , Cohnia, Cossin ia , Geniostoma ,
Hibbertia. ~~ , Kerauclrenia , Hacadamia, Pipturus , Rulingia , and Soulamea .
350 16

Because its species are entirely restricted to the cool , shallow coastal
waters of the Pacific Ocean and its tributary seas and gulfs along both the
North American and Asiatic continents, Phvl10spadix of the Zosteraceae must b e
considered a Pacific genus . It CQuld, hO\~ever, \'lith some justification have
been included under Category I as a Eurasian-\~estern North American (Berin-
gian-Temperate or Amphi-Pacific North Temperate) d is:funct . Similarly , the
more tropical sea-grasses, Enhalus and Thalassodendron, of sha l101" l"Isters of
the Ind ian and I<le stern Pacific Oceans pelong here in the Pacific-Indian Oceans
subcategory of wide disj uncts.

VIII . PAC IFIC-INDIAN- ATLANTIC OCEANS

Instead of being intercontinental disjuncts separated by the oceans , the

taxa of this category are interoceanic disjuncts separat ed by the continents.
By definition, relatively fe\v taxa belong to this group . Host notable are the
tddely distribut ed genera of marine phanerogams . Zostera ( s .l.) forms marine
meadows in shallow, coo l Ivaters off the coasts of all the con tin ents on either
side of the equator to the subarctic o r s ubantarct i c. The more tropica l Ha10-
dule, Halophila , Syringodium , and Thalassia have extraord inarily large and
discontinuous ran ges in the shallow waters of the Indian Ocean-Red Sea-Hestern
Pacific Ocean and the Car i bbean Sea and Gulf of Nex ico, all ex t end ing v i a the
Gulf Stream to Bermuda in the Atlantic . lIalodule and Halophila exten d south
to Brazilian coasts (den Ha rtog , 19 72) and one species of Ilalophila entered
the eas tern l·lediterranean Sea from the Red Sea via the Suez Canal. The eastern
Atlanti c Ocean seems to be strangely lacking in these genera except for records
of I!alodule from Na uritania, Senegal, and Angola (den Hartog, 1970) . Cymodo-
cea (s . s . ) and Posidonia , the sole genus of the Posidoniaceae, have rather
different disjunct patte rns. Neith er 1s foun d in tropi cal American Haters,
but each has a species tha t ranges from the Atlantic coasts of No rth Africa
or of the I berian Penin sul a t hrough the Nediterranean Sea and one or more
species in Australian ,.,,-aters . Posidonia is extratropical, b ut the mo re tropi-
cal Cymodocea ranges \~id e ly in the Indian Ocean-Red Sea areas and into the
western Pacific as far as the Ry ukyu Islands and NeH Caledonia .

Perhaps the only terrestrial angiosperm posseSSing a some,,,hat similar

pattern of di sj unct ion is the palm genus Pritchardia , assuming that the Cuban
genus Colpothrinax is i ndeed congeneric "dth it. The combined range is Fiji ,
Tonga , Tuamotu s , Hal"aii, and Cuba (van Balgooy, 1971), a truly remarkable, if
not anomalous , ran ge . Host of the circumt r op i cal strand plants reach conti-
nental as \~ell as island shores , hence are technically and j ust ba r ely excluded
f rom this category.

IX . Ar-!ERICAN-AFRICAN

Since I have published a l engthy discussion o f this disjunct category

(Thorne, 1973), I shall give just a brief sununary here. THelve angiospermous
famil i es , most of them largely American , are limited essentially to America and
Africa (including Ha dagasca r): Bromeliaceae, Cactaceae , Canellaceae, Car icaceae,
Humiriaceae , Hydnoraceae, Loasaceae, Nayacaceae, Rapateaceae, Tur neraceae, (Fig-
ure 16-6), Velloziaceae, and Vochysiaceae. Seven subfamilies, includ ing Napo-
1eonoideae of the Lecythidaceae , Siparunoideae of the Nonimiaceae, and Stre-
litzioideae of the Nusaceae; 10 tribes and subtribes, including the rafflesia-
ceous Cytineae and the leguminous S'''artzieae; and 111 genera, as Annona , Bar-
bacenia, Chrysobalanus, Elaei~, Guarea , Hyptis , Henodora, Ocotea , Pitcairnia,
16 351

• -
L!-. _
L • • -
,

,---~--. " /"

--:-- .- : .' j '
-:.J0.-?"
OUGINIJ. GtNUAI.I1:W MAl' OF THE fioPlCIJ. An,rCAN· AslAN-MAU:S1AN FAMlLY DIl'TD.OCAJlPAo:AE

Drawn by Chris Davidson; base map as in Fig. 2.

MUCH GL'<I;RALIUD DUT1lBUTION OF 111" C'''CUM_MCTIC, CrkcuM- Bou.AL, AND B.""" .... Gl:NUI
Emp~lr!J.m, nil. C k OWII£aJtW

Based largely upon Good (19&1); ba..., map '" in Fig. 2.

Figu re 1 6- 3. Gener aliz @d dist r i b ution of the Dip t ero car paceae and Emp etrum.
Us ed wi t h p ermiss i o n.
 ~2 16

D,sT'lUIIfTJOI< OF n<~ AMUlCAN·AuntAI,ASIAN.MAl)~1I' FAWILY WIN'lO.Aa..U.

ModIfied from Smith (1!H3a. b); hue map aI in Fig. I.

MUQI GENlJ.ALlU:O DISTll.IIUT10N OF TIl( SUICOSMOPOUTAll AQUATIC SP~ CerlllQPhyllum

demeT$um
Modifio:d in PUI rrom Hulten (1968); base map aJ in Fig. 1.

Figure 16-4 . Di stribu ti on of the Hinteraceae and Cera tophyllum. Used with
permission .
16 353

Raphia, Sacoglottis , Symphoni a, and Vismia, are restricted to Ame r ica and
Africa . In analyzing the " Flora of illest Tr opi cal Africa" (Hutchinson and
Dalzie l, 1927-1936 . 1954-1968) , I listed 108 s pecies be lieved to be i ndigenous
just i n Ameri ca and Af r i ca . These, probably recent immigrants from one conti-
nent to the other , are a lmost all aq ua tic , semi-aquat i c , maritime, riparian ,
or rude ral plants.

The flo r isti c relationshi ps of Afri ca to Eurasia-Australasia and o f

Amer i ca to Eu ras ia-Austral asia are much stronger than those l inking Africa
a n d America . The v ast n umber of taxa on the two continents that have been
unable to span th e Atlantic far ou t weigh s the relatively few amphi-Atlantic
taxa. The greatly varying degree of divergence charac teri zing the American-
Af r ican disjuncts ref l ects roughly the varyin g l engths of time the vari ous
taxa have been separated on the t wo continen ts . Consequen tly, only long-
distance disp e r sal is adequat e t o exp l ain th e \"idely spaced events of i mmi gra-
tion across the Atlant.ic that must have continued througho ut the history of
the dispe rsal of seed plants in t he so uthern hemisphere . In v iew of a l l these
fa cts , nei ther continental di sp l acement via the currently popular tec t onic-
pla te movements nor trans-Atlant ic land - bridges are r e alist i c explanations fo r
most members of this categor y of major discontinuities.

X. NORTH AMERICAN- SOUTH ANERICAI.~

The taxa d iscontinuous hetHeen North and South America are best considered
under s everal subcat egories , de pending upon \"hether their areas are p r imar ily
tropical , temperat e (amphitropical), or b i polar .

1. Tropica l.

Possibly 3 , 000 seed-p l ant genera are endemic in trop i cal America (Good ,
1964) . Sure l y , a v ery large per centage of these , not a na l yzed for this study,
are conunon to North and South America. This is not surp r isin g considering
t he maSSively effective land - bridge furnished since Plio cene time by Central
America a nd the diagramatically placed \-les t I ndian is l ands lin kin g Florida to
nor thern Sou th America . I n subtropical Florida (Long and Lakela , 1971) a l one
th e r e are 150 neotropical genera that a r e di s junct be t ween the south e rn tip of
the peninsula and the Ant illes , Nexico , or South Ame r ica . Of 228 seed- plant
families indigenous in South America, all but 27 are represented also in Nexico ,
and 7 more h ave r eached at l east Panama in Ce ntral America . Twelve lar gely
tropical families a r e restricted to the two continents : Bixaceae , Brunel-
liaceae , Cannaceae , Cyclanthaceae , Cyril laceae. Kr ameri a ceae , Lenno -
aeeae, Marcgraviaceae, Ha r tyniaceae , Theophr astaceae , Tropaeolaceae (p r imarily
Andean) , and Zamiaceae . The North American Garryaceae and Sou t h American
Quiinaceae both r each the Gr eater Antilles and JI!iddle America; and 8 of the
12 American- African tropical families reach Hexico, at least , and most of them
rea c h Florida or farther no rth in North America . The Sarrac eniaeeae , pa r tly
tropical but largely temperate , are c lassical in their disjunction between the
Guyana Highlands (lleliamphora) , California-Or egon (Darlingtonia ), and eastern
North America ( Sar r a~e n i a).

2. Tempera te .

The amph i trop ica l American disjuncts have recently r eceiv ed considerabl e
att ention (Constan ce , Heckard , Chambers , Ornduff, and Raven , 1963 ; Thorne and
Lathrop , 1970 ) . At l eas t 65 primarily tempe ra te American gener a are wi dely
354 16

discontinuous acros s th e American tropics , in cluding Agose ris. Amsinckia ,

Bahia , Blennosperma. Camisson ia, Chor1zan the . Clarkia, Crvp t antha. Downingia.
Gaillardia. Galvezia. Gau ra . Gi l ia, Hap looappus . Lasth enia , Lepuropetalon ,
Lesguere lla , Nadia, Nonanthochloe, Orthoca rpus, Oxytheca . Phacf'-lia, Plec tri-
tis , and Sche donnardus . At least 90 more wide l y ranging temper ate genera
show simila r disjun ct ion s \.;rith i n the Net" Ho rld, as Adenoc Bulon, Androsac e.
Antennar ia . Calandrin ia , Descharnpsia. ~ hedra, Elatine, Fagonia, Fragar ia ,
Frankenia . Gle ditsia, J..epechinia . Lilaeopsis. Lotus . Lupinus . Henoclora, Hirnu-
Ius, NY050t15 , Nyo s ur us . Orobanche, Plagiobothrys . Sanicula, Taraxacum , and
Thlasp i . Raven (1963) lists 12 8 species , species-pairs , or species - groups
of seed plants t ha t have disj unct r anees in temperate No rt h and South America .
He treats in another g r oup t he temperate dis j uncts , many of them \.Joody , t hat
occur in the desert s of the tHO continents . Some of these taxa are espe cially
p rominent in the American South\.,rest : AtamisQuea emarginata Hiers, Errazu-
rizia, Fugonia , Gut ier r ezia , Helie t ta, 1I0ff manse ~ , Ko eberlinia soinosa
Zucc . , Larrea, Nenodora , Prosopid astrum, and Prosopis. Raven lists also
22 species or species-pairs of grasses and forbs that are likewise deser t
di sjun cts .

Raven (1963) concluded that the amphit rop ical disjunc t s probab l y mig rat ed
by long-distance dispersal in late Pliocene or Pleistocene t ime or even more
recently . The largely herbaceous temperate and bipolar di sjun cts mostly have
d is persed from North to South America; whe reas , the more \-lQody de sert dis-
juncts mostly evolved in South America or perhaps in part diverged from a com-
mon tropical ancestor. The considerations upon which he bases his logical
conclusions are t hat the unba lanced assemblage of disjuncts, mostly self-com-
patible and autogamous , a r e q ui te unrepresen tat ive of the distinctive floras
of the t\.,ro extratropical areas, \.,rhich have been distinct s ince the mid dle
Cretaceou s . Further , the disjunct s on ei ther side of the trop ics are closely
related, often conspe cific , and they occupy mostly open plant communit i e s, as
grasslands and vernal marshes ( Thorn e and La throp , 1970) , easily penet r ated
by migrant s .

An altern ative explanat ion with some merit is that the d iscontinuous taxa
may have migra ted by normal short-distance dispersal or by stepping-stone move-
ments alon g the more or less continuous Hes tern American mount ain system . Cer-
tainly th is exp lan ation is valid for t be dozens of genera like Ribes that are
nearly continuous in t h e American highlands from Alaska to Fueg~Some of
th e other genera with relatively cont i nuous montan e dis trib u tion across t he
American trop i c s are Alchemilla, Alnus , Berbe r is . Caltha , Carex. Cas tillej a,
Epilobium , Gentianella , Ge um, !Ivd~a, Hon tia , 'fh a li ct ru~leriana , and
Vi burnum . Unlike these, however , most of the temperate taxa discussed above
are unrep r esented on the Andean cordille ra acro ss the American tropics . Nost
are n ot montan e at all , and many are adap ted to the Hedite rrancan climat e of
the Pacific coastal areas . Thus , long-distance dispersal rather than con tinu-
ous migration is st r ongly indicated for them .

3. Bipolar .

Also discussed by Raven (196 3) '..Jere the bipolar disjuncts . He list ed 26

spec i es or species-pairs of seed plants \~ith bipolar distribution, all but two
of t hem circum-Arctic or circ umboreal as we l l . Nearly a dozen gen e ra conform
to th is pattern : Arme r ia , Catabro s a, F.mpet rum, Euphrasia, Hippuris, Honkenya,
Koeni&ia , Lit~re l la . folonolepis , Phipps ia, and Primula . Long-dis tan ce
16 355

dispersal fr om northern North America in Pleistocene or later times seems to

be indicated here , as Idth the temperate disjuncts . However , at least Euphra-
sia is an exception to this probable route . The South American spec i es of th i s
genus , according to Du Rietz (1960), are more closely re l ated to the southern
and Hestern Pacific spec i es than they are to the North Amer i can spec i es .

XI . SOUTH AMERICAN- AUSTRALASIAN

At least 48 genera and 7 families of seed plants are essential l y restric-

ted to temperate South America (some passing into the tropics along the Andes)
and Australasia , \~ith a fev follol"ing tropical highlands to southeastern
Asia or rarely reaching Hadagascar. An additional 28 more I"idely distributed
genera and 2 families are linked to Australasia by common species or c l ose l y
related species represented in the tHO regions . The largely Pacific genera ,
Coprosma , Haloragis, and Santa1um, reach the Juan Fernandez Islands but not
the South American mainland . These taxa are perhaps better analyzed by trea t -
ing them in th r ee minor subcategories .

1. South American-Aus tra1asian.

This is the largest subcategory, conta i ning those taxa with representa-
tives only in temperate South America (a fev pass i ng along the vestern Ameri-
can cordillera to Nexico) and the Australasian area , including Ne1,o.' Zealand
and its subantarctic islands , Tasmania , Australia , NeH Caledonia, and New
Gui nea (a fe", re a chi ng the mountains of Bor neo o r Tai ",an or even , i n the other
direction, the Po l ynesian and Hm...aiian mountains) . Groups that possess th i s
spectacular type of discontinuity are the Araucariaceae and Eucryphiaceae and
at leas t 36 genera : Gevuina , I.omatia, Oreocallis. and Ori tes , Amphibromu s,
Araucaria, Aristote1ia, Azorel1a, Celmisia, Co1obanthus, Discar i a . Donat i a ,
Eucryphia, Fuchsia, Gaimardia, Griselinia , Hebe , Hypsela, Jovellana, Laurel ia,
Libertia, Narsip pospe r mum, Huehlenbeckia , Nothofagus , Oreomyrrhis, Ourisia ,
Pernettya, Phyllachne , Pseudopanax , Rostkovia , Schizellema , Selliera , Tetra
chandra, Trichocline, Uncinia, and Vittadinia. Three American amphitropical
disjuncts, Calandrinia, Lilaeopsis . and Plagiobothrys , are a l so disjunct from
temperate South America to Australasia . Nost of the other 25 widely ranging
genera men tioned above reach only Aus tralasia , and hence also belong in th i s
subcategory .

2. South American-Australasian-Asian.

This subcategory is quite artificial, since it includes wide disjuncts

almost identical to the preceding except that they have species successfully
established on the mainland of southeastern Asia . Four prima r ily Aust r alasian
families have thus reached at l east mainland Asia to the 1:~est and temperate
South America to the east . They are the Cent r olep i daceae, Coriariaceae ,
Epacridaceae, and Stylidiaceae. Seven genera that like\~ise fit in this sub-
category are Coriaria , Dacrydium, Euphrasia (Hith also extensive holarctic
distribution). Gaultheria , Lagenifera , Leptocarpus , and Oreobolus. Coriar i a
i s some",hat anomalous her:e, for in addition to one species reaching Tahi t i
and north along the Andes and Middle American mountains to Nexico , it has
another in the Mediterranean region .

3. South Amer i can-Australasian-l'1adagascan .

Likelo!i se little different from the tuo preceding subcategories are those
356 16

that extend their wide discontinuities from South America to Au stralasia and
on to the Ma sca rene Islands or Madagascar , al so in some cases involving Male-
sia or southeastern Asia. The one family , IUnteraceae, with this type of
disjunct range has not been found on the Asiatic mainland hut members have
reached north to Nexico i n Ameri c a , north to Borneo and the Philippines in
Malesia , and south to the highlands of Madagascar . It is ... worthy of note he re
that this vesse lless family in most of its features is the most primitive liv-
ing family of t he Angiospermae. Five genera also with these enormo us gaps
across the Pacific and India n Oceans are Abrotanella and Astelia (to the
Mascarenes) , Nertera and \-1einmannia (to Madagascar ), and Dianella (on Nadagas-
car and also i n mainland East Afr i ca) .

For some of the plants of this t empera t e South American- Aus tral asian dis -
junct category, especially those ,dth fleshy , bird-d ispersed fruit that have
far-flung distributions on oceanic islands of the Paci fic and Indian Oceans ,
long- distance dispersal by birds i s the rather obvious explanation f or the
discont i nuities . Genera with fruit that mus t be especia lly attractive to
birds are Aristotelia , Astelia, Coprosma , Coriaria , Dacrydium , Dianella,
Fuchsia , Gaultheria , Gris elinia , Nerte ra , Pernettya, and Pseudopanax. Other
genera that app ear to require and be capable of long distance dispersal are
Abrotane l la, AzorelIa , Co l obanthus, Gaimar d i a, lIal or agis, Lasenifera , ~­
carpus , Oreobolus , Oreomyrrhis , Ourisia , Rostkovia , Schizellema, Se lliera ,
Tetrachondra, Vittad inia , Un cinia , and Weinmannia . There a r e several taxa ,
however , \"hich because of large, dry or heavy cones , fruits , or seeds are not
logically explained by bird-carriage or othe r forms of l ong-d istance dispersal.
Among these are Araucaria , Eucryph i a, Laur elia, Nothofagus , and the four gen-
era of Proteaceae .

~~ny author s (listed in Thorne, 1963) have s uggested that Antarctica,

much warmer, fo rested, and probably a more extensive archipelago i n Cretaceous
and Tertiary times , must have served forme rly as a "s,,,eepstakes " migration
route for many seed plants and an imals . Distances between Antarctica and
Tasmania- Australia, Anta rct ica and New Zealand, and Antarctica a nd South Amer-
ica may have been much less in the past . Biological evidence that Antarctica
eve r f urni shed a continuous land-bridge like Beringia , however , is unconvinc-
ing. Plate tectonics , now currently accep ted by many geo logist s , has mo st
recently been suggested (Raven and Axelrod , 1972) as the ultimate answer to
many of the prob l ems of Australasian- t emperate South Ame rican paleobiogeogra-
phy. According to this hypoth esis , about 100 million years before the present
(B . P .), Antarctica linked South America with New Zealand and with Australia ,
furni s hing an effective pathway for migration of temperate forest plants and
associated animals. Later the various tectonic plates moved apart t o their
present po sitions , causing conside rable tectonic disturbance i n the south-
western Pacific . This is a rather attractive hypo the sis which one hope s the
geologists and paleontolog ists can in time substantiate.

XlI . TEMPERATE SOUTH AMERICAN- ASIAN

This type of disjun ction in which the representatives of a taxon are sepa-
rated by the en tire width of the Pacific Ocean is exp ectedly quite rar e . The
outstanding example of this huge discontinuity is the f amily Lardizabal aceae,
'dth 2 genera , B09 ui l a and Lardizabala , in central Chile and S ill eastern Asia
from Japan and Korea to the Himalayas. A few other examples of this South
American-Asian disjunction were discu ssed by Stebbins (1940) . He suggested
that the l argely Andean mutisioid genera Hya l oseris and Proustia are possibly
16 357

closel y related to Catamixis of the northwestern IUrna laya a nd Nouelia of Yun-

nan , to liesperomannia of Ha\.,raii , and perhaps to St i fft ia of Braz il. He con-
siders to he even mor e closely related to the rnutisioid Leucomeris of south-
eastern Asia the genus Gochnatia of the Andes and Braz il north to Ne xico and
Texas. Hypochoeris of the Lactuceae has spe ciated heavi l y i n sout he rn South
America and has a secondary c enter in the Nediterranean Sea a r ea and a few
species across Europe to north eas t e rn As ia. There a r e no native spec i e s of
either Bypochoeris or of the La rd izahalaceae in North America . Stebb ins also
suggested that th e bignoniaceous perennial herb Argylia of Andean Peru to tem-
perate South America has its clo ses t relatives i n the equally h e rbaceous In-
carville a of the Sino-Himalayan region .

Of a ll the di sj unct categories disc ussed , this is the most d ifficul t to

explain . The species of Lard izabalaceae have baccate fruit, but t o explain
thi s dis continuity by direct bird-carriage between Chile and eas tern Asia
rather s trains cre dulity . The di strib ution of Coriaria may be instructive
here . Coriaria , ~"ith fruit a ps e udo-drupe o f cocci surro unded by pe rsisten t
fl eshy petals at maturity , is greatly favored by many birds and is almost
certainly dispersed by them . If the nor t hern South and Cen tral Ame rican ,
Pacific , and Hediterranean species were deleted from the map , i t wou l d show
a di st ribution pattern similar t o that of t he Lardizabalaceae. However, if
t he l atte r f amily dispe rsed l ike Coriaria across the southe rn Pacif i c , it
should ha ve left some traces along the Hay. One can as well a dduce the bipolar
di stribution of Empetrum . If the circumpolar, EuropE'an , and North American
parts of it s ran ge are eliminate d, a similar pattern again emer ges . Surely ,
however , as in Coriaria , Empetrum ' s di s junct distribution has recently been
a chieved by long- dis t a n c e dispersal ~dth birds as vect ors. The temperate
South American - Asian disj unction a ppe ars t o be , a t l east in par t. a relic t
pattern . I would e xpe c t that fossils of the Lardizabalaceae will ultimately
be identified from North American Cretaceous or Tertiary deposi t s .

XIII . CIRCUN-SQUTH TDfPERATE

Cons idering the pau city of l and in relation to ~"a ter in the s outh tem-
pe ra te zone , it i s under stand abl e tha t relatively few taxa are p r esent in ,
and largely res tricted to, the tempe rate port ions of t he three southe rn con-
tinents . Five families, the Cunoniaceae , Gunneraceae , Podocarpac eae, Pro-
teaceae , and Restionaceae , and two s ubfamilies , Esca llon ioideae of the Saxi-
fragaceae and Luzuriagoideae of the Liliaceae , are largely ci r c um-south tem-
perat e in their distribution . Yet a ll o f these have repr esenta t i ves in the
tropics , at least in the tropical highlands, and a ll reach north o f t he equa-
t or . The 9 genera which likewise have a primarily c ircum- south t empe rat e type
o f discon tin uous dis t rib ution are Acaena, Carpha , Cotu l a , Carpobrotu s , Gunnera,
Podocarpus , Pra tia, Hahlenbergia , and Tetragonia .

Some of th ese taxa are ~"ell eq uipped for bird o r sea-c urrent dispersal
over large distances, as Acaena, Carpobrotus, Gunnera, Podo carp us , Tetragonia,
and the luzur iagoi ds . f>lany members , however, of the Cunoniaceae , Proteaceae ,
and Esca llonioid eae are much less well adapted for l ong-distance di s persal
and wou ld seem to r equire more nearly comp l e t e land conne c tio n s for norma l
short- range or s tepp i ng-s tone dispersa l . Th eir present wide r anges may have
re s ulted from mi g r ation over a once t>'armer and more e xtensive Antarctic archi -
pelago b e tween South America and Australia . Thei r South African presence i s
more likely due to migr ation from Aust ralia through the Indonesian-Asian and
East Af rican highlan ds . The often called - upon breakup of a possible
358 16

Gondw analand , if it ever existed, must have taken place far too long ago to
have had much impact up on the pr esent distribution of see d plan ts .

XIV . CIRCtJH-ANTARCTIC

The land areas defined here as circum-Antarctic ·are t he Grahamla nd pen-

in sula of Antar ctica , the south ern tip o f South America (e spec ially Tie rr a del
Fuego), and the high latitude i s l ands surrounding the Antarc tic , s uch as the
Falklands. South Geor gia, Tristan da Cunha. Go ugh. Na rion . Crozet. Hea rd , Ker-
guel en , St . Paul, NeH Amsterd am , and Hacquarie Islands , mostly \,,1th10 or near
the nor thern l imit of drift ice . The suhantarc tic lands have a v ery s mall
flora. as might be expected from the strong cold winds , small and remo te land
a reas , and great amounts of permanent ice. Yet their meager fl ora of perhaps
100 species (Good , 1964) is a r ather charact e ristic, constant one, \"ith per-
haps 30 spe cies with wide dis tr ibutions in the circum-Antarctic region (Daw-
son , 1958) . Among these spec i es arc Acaena adscendcns Vahl , Azo r ella s elago
Hook . f . , Calli triche antar c t ica Engelm. ex lIegel , l-Iont ia fontana L . , ~­
phy llum e l a tinoides Gaud . , and Ranunculus bite r natus Sm . i n Rees . During
the Pleistocene ice age most o f tbese circum Ant arctic lands were covered
with ice , as were Hacquolrie , Ke r gue len , Heard and So uth Georgia Islands and
Grahamland . Con sequent l y , the circ umpolar f l o ra must be post-P l eistocene and
must have reached these remote is l ands by l ong-distance dispe rsal , presumably
via oceanic bird s (Falla, 1960; 1I0ldgate, 1960) mostly from so uthern Sou th
America and New Zea land .

XV . SUBCOSNOPOLITAN

Plant taxa that are not primarily tempe rate nor t r opi cal and that have
indigenous repr esen tatives on all the continen t s (includ i ng Aust r alia) can be
desc r ibed as sub cosmopolitan. Truly cosmopolit an taxa that have r ep r esenta-
tives o n all parts of the ea rth ' s su rfac e do not , of course , e x ist . The most
nearly cosmopo litan family of seed plants is the Poaceae, which has sp ecies
in all habitable par t s of a ll the con tinen t s and even in Grahaml and of Ant -
arc tica . Ninety families (among those listed by Thorne , 1968) and abou t 125
genera of seed plants are s ub cosmopo l itan in their distribut ion and in their
disjunction . It is certainly no coincidence tha t at least 72 of the subcos -
mopolitan genera are lar ge ly aquat ic or at least have species that inhab it
shallow wat er, ve t, open pl aces , mar itime si tuation s , or r iparian habita t s .
A few of these are Callitriche, Carex , Crassu la , Drosera , Elatine , Eleochari s ,
Juncus, Lemna, Nontia , Nyriophyllum , Najas , Nymphaea, Pot amogeton, Ranun culus ,
Ruppia , Triglochin , ~, Utricularia , Vallisneria, and l"olffi a. An addi-
tional 26 genera have species \"ith veed y tendencies, as Amaran thus , Cenchrus ,
Centaur ea , Cusc u ta , Euphorbia, Gnapha l ium , Lepidium, Oxalis , Senecio , Se t aria ,
and Urtica . Thus , about fou r- fifths of the genera have members that by habi-
tat are re adily dispersible . The rema in ing gener a are small- s eeded herbs or
woody plants with mostly s ucculent , b ird-dispers ed fr u i t.

Very few species can be des c ri bed as subcosmopo li tan , and these are almost
all of nece ssity aquatics , like Ceratoplwllum demersum L . Some othe r aquatics
that also have vast if sOT:lewhat sporad i c distributions on all o r nearly all the
continen ts are Anagal l is minimus (L . ) Kra use , Br asenia schrebe ri J . F. Gmel .,
Callit ri che ~ L ., Chenopodium glaucum L ., Cladium mar i scus L. {s.l . } ,
Cyperus flave scens L ., Juncus bufonius L . , !!.. effu sus L., Lemna gibba L .,
1.. min or L. , L . trisculca L. , Lud\"igia palustris (L.) Ell , Mont i a fon tana L. ,
16 359

}!yriophyll um spicatum L .• Najas marina L . , Phragmites australis ( Cav . ) Trio. ,

Potamogeton pectinat us L. , Ranunculus trichophyllus Chaix ., Ruppia cirrhosa
(Petag . ) Grande , Scirpus maritimus L .• Spirode la polyrhiza (L . ) SchIeid .,
Triglochin palustris L., and Zanichellia palustris L. Several of these are
mapped by Neusel , Jager. and IHnert (1965). There can be little doubt of the
efficacy of i.ater and shore birds in the di spe rsal of these species about the
world .

XVI . ANOI'lALDUS

The final catego r y of major types of discontin uous taxa is a \~astebasket

for those few genera that I am unable to fit comfortably into the preceding
c ategories . I n some instances , further exploration and discovery of the taxon ,
as is surely to be expected with the raff l esiaceo us para s ite Pilos tyl es , may
clarify its position, ,"hich, in this case , is already approaching circumt ropi -
cal . Nyr i ca also fal ls a little short of being sub cosmopol itan because of its
absenc e f r om Australasia. If the circumboreal di sj unct Gale belg i ca Dum .
(Nvrica gale L.) is removed from Nvrica . the genus is also unrepresented in
Europe and boreal North America. Th e present dis junc t range of l'lyrica is cer-
tain ly a relic of a once wor ld,dde dist ribution. The santalaceo us Thesium
falls somewhat short of being ci rcumtropi c a l hecau s e of its extreme l y weak
presence in southeastern Asia-Australasia and in south ern Brazil (map in Neusel ,
Jager , and l-1ine r t, 1965) . It might better be considered an African- Eurasian-
Aust ralasian disjunct with a smal l outlier in southern Brazil , as in the family
Pedaliaceae, in which only Rogeria r eaches the New Ivorld with a species in Bra-
zil (so a l so with the crassu1aceous Kalanchoe) . Likel"ise , the Ameri can-Afric an-
Nadagascan Urera of the Urticaceae transgresses its pattern by having a species
in the Hal.;raiian I slands . Perh aps th e most anomalous disjunction of all is
found in t he chiefly Indomalesian-American genus Thismia of the Burmanniaceae .
Among these f l eshy saprophytes the sma l l section Rodwaya has one species , I.
rodwayi F . v . Huell , i n Victoria , Tasmania , and New Zealand ; and another I .
ameri cana N. E . Pfeiff . , found a couple of times many years ago in a moist
prairie in Chicago, I ll inois .

STATISTICAL SUNNARY

Some authors, vide Good (1964) , divi de taxa geog raphically i nto three
majo r categories . dependin g upon ,.hethe r they are widely distribu t ed about
the world , are di scon t inuous, o r are endemic in a limited area. This divi-
si on is a rt ificial , since Iddely distributed taxa, because of the juxtaposi-
tion of continents and seas, are almost always discontinuous in their areas.
Thus , we a r e left prac t ically with on l y disj unct vs . endemic t axa . The l atte r
pres ent no great phytogeographic problems other than th e formerly much greater
a nd often discontinuous ranges of many of the relict endemics , or epibiotics .
Therefore , our interes t here is centered on the discontinuous taxa .

The larger the category of the taxon , the greater the likelihood that
its d ist r ibution wil l be more or les s wide-ranging and hence disjunct. Of
324 familie s that I curr ently recognize [a total somewhat larger t han in my
Synopsis (Thorne, 1968)] 254 , or 78.4 per cent , have intercontinental o r
equivalent discontinuitie s . If the 273 recognized additional sub f amil i es
are added to the family total , 462 of 597 families and subfamilies , about
the same percentage , 77 . 4 per cent , have major discontinuities in their
ran ges. Of the perhaps 12 , 500 angiosperm gener a usually recognized (Good,
1964) , a total of nearly 3 ,000 are counted in this st udy as having major
360 16

range disjunctions. Thus only 24 per cent of flowering p lant genera are int er-
continentally disjunct . Species have not been thoroughly analyzed for major
disjunctions, but a roug h estimat i on from this study \-Iould be about 2,000 with
wide natural range disjunctions. If 225,000 is accepted as a r easonable e sti-
mate of the named valid species of flowering p l ants (Good, 1964). fewe r than
1 per cent of angiospermous species are widely disjunct",

Of the many categories of major disjunctions, those encompassing the larg-

est number of genera are: African-Eurasian- (Pacific), 600; Asian-Pacific,
460; Pacific, 370 ; North American- South American, ca . 360 (surely a serious
underestimate since Niddle Ameri ca is disregarded her.e); Pantropical. 334;
No r. th Temperate . 310; Subcosmopo li tan, 1 25; Af rican-American. Ill; and Amphi-
Pacific Tropical , 85 . Those with the largest number of seed - plant families
are : Suhcosmopo litan, 90; Pan tropi cal , 59 ; North Tempera te, 20; African-
Eu r asian- (Pacific), 17; North American-South American, 12 ; African-American,
12 ; Tropical Amphi-l'acific, 11; South American - Australasian. 7; and Pac ific,
7.

GEOGRAPHIC EXERCISE

Select two species fo r geographi c study and determine the follo\dng dis -
tribut ion characteristic of each (consult local and regional manuals, Good
[1964] , and Blake and At\~ood [1942, pt. 1 and 1962 , pt . 2]).

DISTRIBUTION CHARACTER Ref. Species f!l Species H2

l. Biogeographic Region A- I
2. Biome A-II

3 . Flora Province A-III

4 . Community Type Chap . 15

5 . Habitat Type Chap. 15

Genus
6 . Range Pattern C or 0
Genus
7. Range Size B lIt
Genus Discont inuity
8 . Distribution Pattern C

9. Center of Distribution B-II

Origin of Establishment
10 . in Local Area (Genus) B-VI

11. Diaspore Type Chap. 15

12. Plant Dispersal Typ e H-VII

 ------- .. - - .

16 361

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362 16

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~~___ 1970 . The sea-g rasses of the 110 rld . Verhandelin gen !(on inklijke
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1972 . The sea -grass es of Brazil . Acta !3otanica Neerlandica 21 :
512 516 .
He r nandez-X ., E. , E. II . Cr um, W. B. Fox , and A. J . Sharp . 1951. A unique
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Hol dga te , N. 14 . 1960. Fina l dis cussion . 1 n: C. F . II . Pan tin (ed . ) . A Di s -
cussion on the Bi.ology of the Southern Cold Temperate Zone . Proceed in gs
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Institute of Science and Technology. Na nila .
Hu lten, E. 1937 . Outline of the !lis tory of Arctic and Borea l Biota During
the l1ua ternary Pe riod . Bokfor1ags Ak t eibolaget Thyle . Stockholm .
1958 . Th e a mphi-Atlanti c plants and their phytogeographic al con-
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279 maps .
1963 . Phytogeog ra phical connec tions of the Nor th Atlantic . In :
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16 363

Hulten , E. 1968 . Flora of Alaska and Neighboring Territories . Stanford Uni-

versity P ress . Stanford .
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2 vals . CrOHl1 Agents . London.
1954-1968 . Flo ra of \~est Tropical Africa. 2nd ed . R. 1-/ . J. Keay
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1.1 , Ii . L . 1952 . Floristic relationships 1etHeen eastern Asia and eastern
North Arnerica . Transactions American Philosophical Society, Nelv Series
42 , 371-429 .
Long , R. H., and O. Lakela . 1971. A Flora o f Tropical Florida . University
of Miami Press . Coral Gables .
Love , D. 1962 . Plan ts and Pleis tocene--problems 0 f the Pleis tocene and
Arctic . NcGill University Huseums P ublication 2 : 17-39 .
HcVaugh, R. 1952 . Suggested phylogeny of Prunus serotinC! and other I·,icle-
ranging phylads in North America . 13rittonia 7 : 317-346 .
Heusel , H. 1969 . Beziehungen in der Florendifferenzierung von Eurasien und
Nordamerika . Flora, Abt . B 158 : 537-56'1 .
--,-::-c~ ' E . Jager, and E . Hinert . 1965. Verg1eichende Chor010gie der Zentral-
europaischen Flora. Karten . Gustav Fisc her. Jena .
, and R. Schubert. 1971. Beitrage zur Pf1anzengeog raphi e des
Hesthimaljas . 1. Teil : Die Arealtypen. Flora 160 : 137-19/1 .
Hiranda , F ., and A. J . Sharp . 1950 . Character istics of the vegetation in
certain temperate regions of eastern l-!exico . Ecology 31 : 313-333 .
Ostenfeld, C. H. 1915. On the distribution of the sea-grasses . A prelimi-
nary communication. Proceedings Royal Society Vic.toria 27 : 179-191 .
1927<1 . t-leeregdiser 1. ~larine Hydrocharitaceae. In : E. Hannig
and 11. \.-l inkler (eds . ) . Die Pflanzenarealc . Vol. 1(3).
192 7b . Heeresgraser 2 . Narine Potarno getonaceae . In : E . Hannig
and I! . Hinkl er (eds.) . Die P flanzenareale . Vol . l ( 4) .
Polunin, P . 1960 . Introduction to Plant Geography . Longmans, Green and
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Raven , P . H. 1963 . Amphitropical relationships in the f loras of North and
South America . Quarterly Reviel·} of Biology 38 : 151-177 .
-,c:;-::::~ ' and D. r . Axelrod . 1972 . Plate t e ctonics and Australasian pa leo-
hiogeography . The complex biogeographic relations of the region reflect
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Schofield , \·1 . B. 1969 . Phytogeography of northHestern North America : bryo-
phytes and vascular plants . Nadrono 20 : 155-207 .
Setchell, \.J" . A. 1920 . Geographical distribution of the marine spermatophytes .
Bulletin of the Torrey Botanical Club /!7 : 563-579 .
1935 . Geographic elements of the marine flora of the North Pac i fic
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Agriculture . U. S. Government Printing Office . Hashington, D. C.
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efectuados in Nexico y Guatemala . Revista de la Sociedad Hexicana de
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1946b . Pinus strobus south of the United States . Journal of th e
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Shreve, F. , and 1. H. Higgins . 1964 . Vegetation and Flora o f the Sonoran
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Sleumer , H. 1955 . Proteaceae. Flora l'!alesiana 5 : lfI7-206 .
364 16

Smith, A. C. 1943a. The American species of Drimys . Journal of the Arnold

Arboretum 24 : 1-33.
1943b . Taxonomic notes on the Old l10rld species of Winteraceae.
Journal of the Arnold Arboretum 24: 119-164 .
Stebbins . G. L. , Jr . 1938 . The western American species of Paeonia .
Hadroi'io 4 : 252-260 . .... ..
1940 . Additional evidence for a holarctic dispersal of flm-lering
p l ants in the Hesozoic era . Proceedings Sixth Pacif ic Scientific Congress
4: 649-660.
----C---7. and A. Day. 1967 . Cytogenetic evidence for long continued stabil-
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(ed.). Pacific Plant Areas. Vol . 1. National Institute of Sc ience
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Honolu l u .
1965 . Floristic relationships of NeH Caledonia . University of
IOHa -S tudies Natural History 20(7) : 1-14 .
1968 . Synops i s of a putatively phylogenetic classificat ion of the
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1969 . Floristic relationships betHeen New Caledon i a a nd the Solo-
mon Islands . Philosophical Transactions Royal Society (London) . B. 255:
595-602 .
1973a . Floristic re l ationships hetl~een tropical Africa and tropi-
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Tropical Forest Ecosystems in Africa and South America : A Comparative
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The Quar terly Revie\o] of Biology 47(4) : 365-411.
__~~___ ' and E. H. Lathrop. 1970. Pilularia amer icana on the Santa Rosa
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St at e University . Bl acksburg, Virginia .
17 365

Chapter 17. FLORISTIC STUDIES

The plant taxonomist must have a holistic or ecosystematic approach to

have an understanding of individuals and populations in the field. This
approach involves (1) learning and observing the distinguishing and diagnos-
tic characteristics of taxa, (2) discovering the lill1iting facto r s in the
di stribution of species, (3) discerning the role of individuals in the eco-
system, (4) detecting evolutionary mechanisms at work, and (5) determining
the phene tic and phylogenetic relationships of diverse organisms. A field
student sho uld study the ecological characteristics--biotic, abiotic, spatial ,
temporal--of species and should become familiar ,,,ith the environmental rela-
tionships of the individuals within the population. The dynamic role of
individual organisms in the environmental processes--Climatogenics, Pedo-
genics, Biogenics , Phylogenetics--should he observed and determined \"henever
feasible . And a knm"ledge of "Hab ita ts" should be routinely applied in iden-
tifying geomorphic , geologic, and vegetational features in the field.

To help the student acquire the holistic and ecosystematic perspective

are inclusions in this chapter on habitats in North America north of Mexico,
a summary outline of ecological characteristics of species and populational
relationships, and a community sample study by means of community lists ,
mosaics or patterns, and pro f iles. The Elora or community spectral analysis
l~ill show some of the diversity Hithin the plant assemblage and give some
indication of orig i n and evolution of the flora or community .

Enough field trips should be taken during a studv for the student to
understand fundamen tal biotic, abiotic, spatial , and tempora l relationships;
to learn how to fi nd the maximum diversity in the reg ion; how to collect and
care fo r specimens; hOH to distinguish life forms and dispersal mechanisms;
how to classify or identify different cOnllnunities and habitats; hOH to recog-
nize families, genera, and dominant species in the field; and to be intro-
duced to f undamental population and geographic principles. If fou r trips are
taken during a course, habitats, patterns, and profiles could be emphasized
on one; life forms, dispersal, and sociability on a second; duration, phe-
nology, and taxon diversity on a third; and distribution, origin , and popu-
lation conce pts on a fourth (See Chaps . 15, 16) . Recognition of species
should be a basic part o f each trip and on each successive trip the emphasis
of the preceding trip should be reviewed.

Sound field work combines cerebration with vision. Good floristics

pract ice involves the making of presence li sts by community or habitat;
classifying the community or identifying the habitat; indicating the local-
ity and date of study; noting slope , exposure , and altitude; and determin-
ing the rock or soil type . In addit ion to collecting equipment , standard
accessories for fie ld Hark should be an altimeter , a Brunton compass or
clinometer and standard compass, topographic or political maps, geologic
or soil maps, and Hate~~roof writing equipment .

Successful field ....'ork involves adequate preparation hy the student on

Hell - presented assignments and objectives by the teacher prior to the actual
fie ld tri p. lIerbarium specimens arranged by conununity or family should be
available fo r study so that the student does not have to spend a major por-
t i on of hi s field time learning I"hich name tag goes with each plant .
366 17

Profiles of the communities \"1th10 the area to be visited should also be

availab l e early enough for study so that maximum effect i veness can be made of
the all too- limited f i eld time . A meaningful, organized field experience
should add to our basic knowledge of species biology; should be a conceptual
and observational chal lenge for a l l systematists; and should be fundamental
training for ecologi sts and geographers as t,fel1 as ta2'_o~omists .

Section A. HABITATS AND ECOLOGICAL CHARACTERISTICS

In describing habitat the geomorphic feature should be i ndicate d ; e . g.,

f lat, dome , lake and the mineral or rock composition should be g i ven where
possible; e.g., gneiss , alkali, syenite . For the later successional stages
the vegetation feature should he noted. The ecological cha racteristic and
populational relationships are included as an organized topical summary for
ecological review .

I. GEOMORPHIC OR GEOLOGIC FEATURES

Consult the annotated glossary at the end of this chapter . or any general
geology text glossary for definitions of geomorphic and geologic features
and Chapters 14 and 15 and any general ecology text for vegetation infor -
mation. An excellent topographic fo rm summary is in Lahee (1961) .

Bluffs and Cliffs Gravitational Features

Basalt Glacial Boulder Fields
Dolomite Fault Scarp Talus Slopes
Gneiss Sink Igneous 1nstrusions
Quartzite Stream Batholiths Sil ls
Sandstone Dikes Stocks
Depressions and Sinks Laccoliths
Gabbro Depressions Islands
Granite Depressions Barr ier and Estuarine
Lava Sinks Coral and Limestone
Lime Sinks Volcanic
Playas Lakes, Pools, Ponds
Dikes and Si lls Fluvial: oxbm,.rs , plunge basins
Diabase Syenite potholes
Pegmatite Quartz Glacial : cirque , kettle, moraine
Domes lakes
Gneiss Gran ite Longshore : bay-mouth bar ponds
Flats l'leteoritic : bays, pools
Alkali Hud Pluvial : dune ponds , playa lakes
Harl Peat Solution : lime sink lakes
Huck Sand Tectonic : fault or earthquake lakes
Glacial Depositional Features Volcanic : calde ra or f lm·, lakes
Drumlins Horaines Narine Depositional Features
Eskers Terminal Barrier Islands Spits
Kames Lateral Bars Continental Shelves
Outwash Plains Gro und ll<lrine Eros ional Features
Valley Tra i ns \.,rave cut cliffs
Gl acial Erosional Features Have-cut benches
Aretes Horns Hineral Deposits - Examples
Cirques Valleys Kaolin Olivi ne
Harl Soapstone
17 367

Honadnocks Sands
Granite Sandstone Bars Ridges
Hountains Dunes Scarps
Fault Volcanic Hills Strands
Fold Slopes
Plains Gentle Sheltered
Continental Shelf (old) Open Steep
Deltaic Lagoon, sound, St r eams
Flood estuary (old) Branches Fresh, brackish ,
Loess Outwash Creeks salt
Plateaus Rapids Black-{.,rater. mud-
Mesa Piedmont Rivers Hater
Peneplane Tidal
Ridges and Peaks Stream Depositional Features
Dolomite Quartzite Flood plains }Iud Flats
Gneiss Slate Deltaic plains Playas
Sandstone Outlvash plains Sand Bars
Rock Barrens Stream Erosional Features
Granite Serpentine Canyons Ravines
Limestone Shale Cliffs Valleys
Sandstone DraNS
Salt and Brackish Hater Valleys and Gorges
Bays River Embayments Fluvial Solution
Estuaries Sounds Glacial Volcanic
Lagoons Volcanic FlOl"s
Felsic or Rhyolitic Flm"
~mfic or Basaltic Flow

II . VEGETATIQI':Al. FEATURES

Aquatic herb : fresh, brackish , salt Glades

Balds : grass, heath :
~ !arshesfresh, brackish, salt
Bogs Prairie
Desert Succulents Savannah
Forests : needle Shrub or Sc rub : deciduous
Forests : deciduous , broad-leaved Shrub or Scrub : evergreen
Forests: evergreen, broad-leaved Tundra

III. ECOLOGICAL CIIARACTERISTICS AND POPULATIONAL RELATIONSHIPS (See Eco-

logical Evidence, Chapter 15, fo r Definitions)

A. Biotic B. Abiotic c. Spatial D. Temporal

A-I Genetic A-2 Nutritional A- 3 Symbiotic A-4 Reproductive

Genetic Autotrophic Commensal is tic Amphimictic

Phenetic Parasitic Hutualistic Apomictic
Phyletic Saprophytic Antagonistic
B-1 Climatic 13-2 Edaohic
.,
Cryothermic Euryphotic Acidophilous Psammophilous
Hicrothermic Nesophotic Basophilous Arg illophilou s
Mesothermic Stenophotic Neutrophilous Argillopsammo-
r.iegathermic Hyd roph ytic Calciphilous philous
Heliophilous Hygrophytic Halophilous Saprophilous
SciophiloliS Hesophytic Aerobic Eutrophic
Xero phyt ic Anae robic Mesotrophic
368 17

C-l Distributional C- 2 Numerical C-3 Areal C-4 Stratifica tional

Random Abundance Basal Area Canopy Herb
Clumped Random Density Cover Sub canopy
Unifo rm Frequency Size-Classes Shrub

D-1 Age Diversity D-2 Phenological D-3 Periodic _,_ D-4 Successional
(Life Cycle) (Annual Cyclic) (Daily Seasonal (Generation Cyclic)
Cyclic)
Birth- Germination Vernal Hydroperiodic Pioneer
Childhood-Seedling Aestival Photoperiodic Transitiona l
Youth -S apling Autumnal Thermoperiodic Climactic
Adult-Adult lIibernal
Old Age-Death

Section B. SA}-IPLE COHHUNITY LISTS, PATTERNS, PROFILES

In this section three approaches to a study of communities or vegetation

are presented: (1) community lists by province, (2) vegetation patterns
based on temperature and moisture (altitude and slope) , and (3) profiles
showing the distribution of communities in a local region \.,Iith the major
dominants listed . A sample species recognition list based on the flora of
several local communities is included as an illustration of species that
could be studied in fo u r afternoon field trips in the spring . A commun it y
report format is included as a guide to the \"riting of community analyses .

I. SUNHARY OF HAJOR PLANT COHHUNITIES IN EASTERN UNITED STATES

RY ~fAJOR PHYSIOGRAPHIC PROVINCE

Symbols : A, aqua tic ; B, bog; c, old field and disturbe d communities Hith 1,
2 etc . '" age; D, annual/ pioneer cormnunities; OF, deciduous forest; HF , ever-
green forest; PF , coniferous forest; G, grassland ; M, marsh; R, rock barrens;
S, shrub; OS , deciduous shrub or scrub; US , evergreen shrub or scrub; T ,
tundra; etc .

A. Communit i es of Coastal Plain : Atlantic and Gulf Coastal Plain, Keys ,

Nississippi Embayment
1. Dune - Grass (DG) 18 . Pocosin or Evergreen Shr ub Bog
2 . Sand Flat - Grass (SC) (5R)
3 . Dune Scrub (OS) 19 . Bay Forest or Bayhead (BHF)
4 . Naritimc Hammock or South- 20. Savannah & Prairie (TG)
eastern Broad-leaved Ever- 21. Pine Flatl.,TQods (LPF)
green Forest (HHF) 22 . Upland Coastal Pine Forests
5 . Tropical Har.1mock (Tllf') (UPF)
6 . Salt Hater-Aquatics (SA) 23. Upland Scrub (US)
7. Salt Flat - Annuals (SO) 24 . Turkey Oak-Hire Grass (TOOF-G)
8 . Salt Harsh (SH) 25. Blackjack Oak-Grass (BODF-G)
9. Salt Shrub (55) 26 . I ntermediate Oak-Hickory
10 . l>jangrove SHamp (HF) Forests (OHDF)
11. Brackish Uater (llA) 27 . Mes ic Oak-Hickory Fore sts (OHDP)
12 . Brackish Harsh (EM) 28 . Mesic Hard'.,IQod Forests (?·fl)F)
13 . Freshwater - Aquatics (FA) 29 . Inland Hammock Forests (IHF)
14. Freshl.,later Marsh (FH) 30 . Lime Sink Aquatics ( LA)
15 . Fresln,'<lter Shrub (PS) 31 . Cypress Domes, Strands, Forests
16. Swamp Forest (SOF) (CPY)
17 . Rag (B) 32 . Old F i eld Communities (Succession
(OFC)
Figure 1 7-1 369

VEe,:!"T to;, PATTERNS IN Tl!.!:;,.. SOUTI 'irJ: AI'I'AI.ACIi IAilS*

I
I
6500
SprucJ-Fir Forest (SPY)
I I He;l t l! !:~ I.d
I I (wi)
I 6000
~
~

I ~
"ecch-Iluckeye Fore t (IJDF) ~ 5500
I "
~

I
\~hitc
"" 5000
lleech F'I'rests
Red Oak
(ROTIlF)
Oak ""
~

10500 Ir_____-C__C7-C~'~I~e:,~i~'~TcY~P~'JL~s:el'"'ee~T~;>'
I
tlorther tl H'Hdwuous I I Table Nt.
(J;DF) I ~...., f Pine
(nlpF)
r l§- Tall fHeath
Henth
4000
I ... 1 I

,0'o"');' ~__________~--/'''----''i I'itch P i ne

(p}IPF)
3500

Chestnut Heath
3000 Oak Oak 3000
(COTIlF)
Co ve l!eath lieath
2500 Ilardlo'oous 2500
(CDF)
Scr ub Pine
2000 (S~lpF)
2:000

1500 1500
Coves F lnt s She lt ered Open Slopes Rid ges
~l es tc lIe c w s n Ill.' Peaks , Xeric
C.1nyons Dr.111S - Ravines Sl o pel!
se sw
Adapt ed f r om 'Ii hit taker (1956)
I . l;.astern Forest Sys t em C . Xerl:: Pine fo r ests
A. Nestc 1I0n-quercine fo rest s 9 . Virgini a pine fo rest
1. Cove and no rthern hardwood fo r est 10 . Pi t c h p ine -hea t h
2 . E,1ste r n hemlock forest 11. T.:lblc Mt . pine-heath
3 . Gray beech-buckeye fo r est D. Extreme exposu re type
S. Intermediate oak forests 12 . Grassy bald
4 . Mixed oak- pignut hickory forest 1[. lIoreal Forest System
5 . Chestn ut oak-tall heath forest A. Sub;l1r i ne furests
6. Chestnut oak-heath 13. Red spruce forest
7 . Rcti oak-ches tn u t fores t 14. Frase r fir forest
8 . I.'hlte oak-heath f o r es t D. i:xtreme exposure ty pe
15 . lIeath bald
*Thls vegetation rattern is based on the distribution of the communities in the
Great Smoky Hountains but is basically appllcable to the Dlue Ridge . Cumberland and
Alleg hany Pla t eau!> . CUr.lberl,1nd am] Alleghany rlounta i ns . :md the Ridge and Va lle y
Province. In the Pi ed':lOnt and Interior Low ]'Iateau with Altit udes 500' -- 1200 ' the
lowe I' elevat i on (150Q··:WOO) cOllUllunities a r e sti.ll present with th e mi xed oak-hickory
complex sp r eadi ng o ve; mos t of th e meso-xeri c center and the cove hardwoods and
heml ock limite d to the most mesic sites and the chest nut oak-heath and pine-heath
be ing limited t o t he most xeric sites . A hydromeslc .:l l iuviill woods co mmunity is
prese nt from 500 ' to ilpproxi.ma tel y 2000 ' in the major river s y!>t e ms .
370 17

B. Communi ties o f Piedmont : Pi edmont and Interior LOI" Platea us

1. Fresin.,rater Aquatics (FA) 6. Upland llardHood Forests (See
2. Freshl. . ater Harsh ( nl) Vegetation Patterns)
3. Shrub Harsh (FS) 7. Upland Pine Forests CPP}
4. Al l uvial Hoods or S~'amp 8. Flatrock Barrens (R)
Forest (SF) 9. Relict Nontane Forests
5. P r airie (TG) 10 . Old Field Communities
(Succession) (OFC)

C. Conununities of Nountains : Blue Ridge , Alleghany, Cumberland. Catskills,

Adirondacks , Green and \·:rhite. Cumberland Plateau , Alleghany Plateaus ,
Ri dge and Val l ey
1 . FreshHa t er Aquatics (FA) 8 . Hontane Pine Forests (HPF)
2 . Fresh'v'ater Harsh (171'1) 9, Hemlock Forests (HPF)
3. Shrub l'larsh CFS) 10 , Bogs (B)
4. Alluvial Hoods (SF) 11. Shrub Bogs (SB)
5 . No r thern Hardlwods (NDF) 12 . Shrub Balds (HS)
6. Cove Hardl"oods (CDF) 13. Grass Balds (AG)
(See Vegetation Patterns) 14 , Alpine Tundra (AT )
7 . Oak-Hickory Forests 15 . Old Field Communities
(See Vegetation Patterns) (Succession) (OFC)

II . CONHUNITY REPORT FORNAT

COHMUNITY
elevation

PHYS IOGRAPHIC PROVINCE

Geomorphic Feature(s}; County , State
Locality (distance and direction f rom a permanent landmark)

Sl ope : aspect __ , deg ree__ Canopy ht. Species DBH Age~_

Soils : ser i es te x ture , pH ts , pH ss
Al A,_ , B , C
Rock TYEe Geologic Formation

,~ *,~* *,~** *,', ,,< *,', ,', *," ,', ,,< ,', t: ,':1,1, 1,*,,< ,', ," ,',:,< ,,< :,< ,,<,', 1, *,,,:'< ,,< :',1,,', '" ,', *,,< ,', 1, ,',** ,', ,', ,', ,', ,', ,', ,', ,',1, ,'",< *,':,~* ,',,', :,<,'<1,,', *,', ,'"',,,:
CANOPY (Using percentage composition, list species i n order of descending
percentages . Using quarter point, list species by importance value ,
Using the releve ' , list species by descending cover values)
SUBCANOPY (For presence only , list species alphabetically)
SHRUBS (For presence only , list species alphabetically)
VINES (For presence only, list species alphabetically)
HERBS (List as Forbs , Graminoids, and Ferns . For presence only , list species
alphabetically in each group)

NOTES :
Indicate the follQloJing in this sequence : Sere type; succession stage; seed-
lings (list by species); transgressive size class es (li st b y s pecies) . Dis-
turbance (type and nature) . Adjacent moisture class communities. Adjacent
altitudinal con~unities .
\-Jhere and ,."hen relevant comment on factors that seeJ:\ to he controlling com-
munity type, e . g , : prevailing Hind, salt spray , tida l amplitude , Hater cur-
rent , type of substratum, rock or soil noncon f ormity . !'take other comments
that lJi11 add to an understanding of the community ,
 17 371
TIt . PROFILES
Coastal Plain : Sandhill Flora
Spout Sp rings , Harnett Co .• N . C.
ISOFC
~4~50;,~~~L--------2T~ODF_G
San dy Fiel d Coarse
(ISOFC) Sand
Turkey Oak-\-lire Gr ass
(rOOF- G) Clay
Pocosin (SB) laO
Turkey Oak-Upland Hillow Oak (TODF) f:

Blackjack Oak-Black Oak

~~-·----------_____ ~~________________~H~.i~C~kOry (BOHDF)
~-Iater Tab le - - -_ _ _ _ _ _.....:~~S;:.':8.....'8~H~F
Pocosin(SB) Clay
Upland Bay Forest ( BHF)

Sandy Field (lSOFC)

h-Striga lutea, l.JitchHeecl h-Krigia virgl.nJ.ca. OHarf Dandelion
h-Linaria canadensis. Toadflax h-Specularia perfoliata, Evening
h-Rumex hastatulus, Sorrel Looking-glass
h-Oenothera laciniata, Evening P . g- Festuca octoflora. Fescue

Turkey Oak-Wire Grass (roOF-G)

t-Quercus laevis , Turkey O. h Cnidoscolus stimulosus, Spurge
t-Quercus incana, Upland ~"i llow O. Nettle
t-Pinus palustris, Long- l eaf P . h-Trag ia urens, Nettle
t-Oiospyros virginiana , Persimmon h-Arenaria carol iniana , Sandwort
s-Gaylussa cia dumosa, O\~arf JI . h-Stipulicida setacca , l.,Tiry Sand\.JOrt
s-Vaccin ium tenellum, T.oHhush B. h-Pyxidanthera barhulata var . brevi-
g- Aristida stricta. Hire Grass folia , Pixie Ho ss
h-Lupinus diffusus, Lupine h-Aureolaria pectinata. YeIIO\~ Fox-
h- Baptisia cinerea, IUld Indigo glove

Blackjack Oak- Black Oak-Hickory (BOHDF)

t - Que rcus marilandica , Bl ackjack O. s -Ilex glabra, Gallberry
t-Quercus velutina, Black O. s-Vaccinium tenellum, LOHhush B.
t-Carya tomentosa . Nockernut II . s-Lyonia mariana . Sta gge rbush
t-Pinus palustris. Long-leaf P . s-Rhus toxicodendron , Poison Oak
t-Pinus taeda, Loblolly P . h-Tephrosia virginiuna , Goat ' s Rue
t-Diospyros virginian a • Persimmon g-Aris tida stricta, Hire Grass

Pocosin (Evergreen Shrub Bog) (SB)

t-Pinus serotina. Pond P . s-Rhus vern ix, P01 son Sumac
s-Cle thra alnifolia, \.,rhite Alder s-Cyrill a racemiflora . Titi
s - Gaylussacia frondosa, Iluckieberry s - Amelanchier obovalis, Shadbush
s -Lyonia ligustrina . Nale-berry s-So rbus arbutifolia . Chokeberry
s - Lyon ia mariana , Stagge rbush s-S tyrax americana, Storax
s-Rhododendron atlantic urn , Azalea v-Smilax la urifolia . Catbrie r
s-Vacciniurn atrococcum. Highbush B. h-Polygala llitea , Bachelor ' s-But ton
s-Vaccinium corymbosum , Highbush B. h-Sarracenia purpllrea, Pitcher Plant
s-Tlex coriacea, Gallberry h-Xyris flexllosa . Hard-head
s-llex glabra. Gallberry h-Eriocau lon decangulare . lIat-pin
 372 17

Piedmont : River Valley Flora

Deep River, 2 miles ...'est of Moncure , I.ee Co ., N. C .
pP'
SDf 340 '
IOfC Pineland
Alluvial (PPF)
Hoods (SDF) Alluvial Field (lOpe) Felsic Volc anic
290 ' Talus 1
River
Levee Old Oxbow Flow S ope
Sandy Loam Flood Plain Silt Loam Low l'lesic Hoods (CDF)

Low ~lesic

t-Betula nigra , River Birch

t-Salix nigra , Black \.jillol'"
t-Fraxinus pensylvanica , Green Ash
t-Carpinus caroliniana, Irolllwod
17 373

Piedmont : Granite Flatrock Flora

Nit chell' 5 Mill, 2 miles n . e. of Rolesville.

=,
Hake Co .• N. C.
o POHDF

~
Grimmia (D) D~; ~:preSSiO~ ~ A~" "':~""'''"
Annual Xerophytes
G R eepage
erna 1
Pool (GR-A) Piedmont
Durham Sandy Loam

(Gr-R) Pineland
(PPF) Pine-Oak-Hickory
(POIlDF)
Xeric Perennials (GR-D)

Annua l Xerophytes (GR- R)

h-Sedum smallii , Diamorpha h Crotonopsis elliptica , Rushfoil
h - Portulaca smallii, Rock Portulaca h-Hypericum gentianoides , Pine\veed
h-Arenaria groenlandica var . glahra, h - Dio dia teres , ButtOOl-leed
Sandlvort g- Cyperus granitop hilus

Vernal Hydrophvtes (G R- A)
h-L indernia monticola, False Pimpernel g Bulbostylis capillaris
g-Agrostis el liottiana , Rent Grass f-Isoetes melanopoda , Quillwort

Xeric Perennia l Herbs (GR-D)

h-Talinum teretifolium g Andropogon scoparius , Little Bluestem
h - Commelina erecta, Dayflmver g-Panicum laxiflorutn
h-Polygonaturn biflorum, Solomon's Seal f-S elaginella rupestris, Rock Clubmoss
h-Heuchera americana , Al um Root f-vioods ia ohtusa , \~ood Fern
h-Senecio smallii , Ragwort f-Cheilanthes lanosa , Hair-lip Fern
h-Yucca filamentosa, Beargrass f-Asplenium platyneuron, Ebony
h - Opuntia compressa , Prickly Pear Spleemvo rt

"P~i~e~d~m~o~n~t,--"P~i~n~e""lan d (P PF)
t-Juniperus v irginiana , Red Cedar v Smilax rotundifolia , Catbrier
t-Pinus virginiana, Scrub P . v-Lonicera sempervirens , Coral Honey-
t-Ulmus alata , Winged Elm suckle
s-Vaccinium arboreum , Sparkleberry h-Chimaphila maculata, Spotted
s-Rhus aroma tica , Fragran t Sumac I·lin tergreen
s-Callicarpa americana, French Hulberry h - Euphorbia corollata, Spurge

,P~i~n~e~-~O~a~k~-,"~i~c~k~o~rLy--,-F~o,r~e",st (POHDF)
t-Pinus taeda , Loblolly P . s-Chionanthus virgullcus , Fringe tree
t-Pinus echinata, Shortlead P . s-Euonymus americanus, Strawberry Bush
t-Quercus stellata , Post O. s-Viburnum rufidulum . Black Haw
t-Quercus velutina , Black o . v-Vitis rotundifolia , Huscadine
t-Quercus mari landica , Blackjack O. v-Parthenocissus quinquefolia. Vir-
t-Quercus falcata, Southern Red O. ginia Creeper
t-Quercus alba , hThite O. v-Smilax g l auca , Catbrier
t-Carya tomentosa, Hocl~ern u t Hickory h-Cypripedium acaule, Pink Ladyslipper
t-!lex opaca, American 1I011y h-Silene caroliniana, \..lild Pink
t - Diospyros virginiana , Persimmon h-Chimaphila maculata , Spotted Hinter-
t - Cornus f lorida, Flmvering Dogwood green
t - Oxydendrum arboreum , SouDvood h-Chrysogonum virginianum , Green and
s-Vaccinium stamineum, Deerberry Gold
 374 17

Piedmont : Ridge and Slope Flora

Occoneechee Hountain near Ena River , south of Hillsborough ,
Orange County , N. C.

__--~POI,
840 ' O~
o~ Ridge
C;
0' Xeric Pine Oak-Heath
North Slope 25' South Slope
(POhDF) 20 0
Chestnut Oak-lleath
( COtOF)

.~
~
cOf'" Cliff Pine Oak-Heath
-.J Scrub Pine-Heath (POhDF)
(SHPF)
Ena River Pastur e

Alluvial I-Ioods (SDF)

t - Betula nigra, River Birch s-Lindera benzoin, Spicebush
t - Carpinus caroliniana , I romwod s-Alnus serrulata , Tag Alder
t - Quer cus shumardii , Slvamp Red O. h-Erythronium americanum , Trout Lily
t - Fagus grand i folia , Beech h-Podophyllum peltatum , Nandrake
t - Liriodendron tulipifera, Poplar h-Dentaria heterophy l la , Tooth\vo r t
t-Acer saccharum spp . floridan um f-Polystichum acrostichoides , Christ -
Southern Sugar Haple mas F .
,S"c"r"u~b,--,Pc,in""e.::!.lI"e"a"tc'.'h. ( SHPF)
t - Pin u s virgin i ana , Sc r ub Pine s-Lyonia mariana , Staggerbush
t-Oxydendr um arboreum. Sounvood v-Smilax glauca , Catbrier
t - Acer rub ru m, Red Naple h-Ga lax aphylla , Gal ax
s-Kalmi a l ati f olia , Hountain Laure l h-Epigaea repens , Trailing Arbut u s
s-Rhododendron catawb i ense . Ros ebay h - Gaultheria procumbens , \~ in tergreen
s-Gaylussac i a baccata , Huckl e b e rry h-Hexastylis virginica, Hild Ginger
Chestnut Oak Heath (COtDF)
t-Quercus prinus, Rock Che stnut O. s Robinia hispida , Rristly locust
t - Nyssa sylvatica , Black Gum s-Viburnum acerifolium , Maple-leaved V.
t - Acer rubrum , Red Naple h-Chimaphil a maculata , Spotted Hinter-
t-Oxydendrum arboreum, SounlOod green
s-Styrax grandifol ia , Storax h-Galax aphylla , Galax
s- Vac cinium va c illans , Bluebe rry h-Hypoxi s hirsuta , YellO\·! Star-grass
s-Vaccin i uum stamin eum, Deerbe rry h-Hi eracium v en osum, Rattlesnake Heed
s-Lyonia ligustrina , Hale-berry h-Hexastylis arHolia , \-Jild Ginger
s - Rhododendron nudiflorum , \~ild Azalea g-Ca r ex nigromarginata
s - Fothergilla major, l-litch Alder f- Osmunda cirmamomea , Cinnamon F .
Xer i c Pine, Oak-Heath (POhDF)
t-Pinus virginiana , Scrub Pine t Cornus florida, FloHering DOgHood
t - Quercus marilandica. Blackjack O. s-Gaylussacia baccata , Huck l eberry
t-Que r cus stellata. Post O. s-Vacc i n i um vaci llans , Bl ueberry
t - Quercus velut i na , Bl ack O . s-Vaccinium sta mineum, Deerberry
t-Quercus pri nus . Rock Chestnut O. s-Kalmia latifolia , No unta in Laurel
t - Pr un u s serotina , Choke Cherry h - Chimaphila maculata , Spotted \-li nter-
t - Acer rubrum , Red Naple g r een
t - Oxydendrum arboreum , Sourwood h - Epigaea repen s , Trai l ing Arb utus
17 375

Coastal Plain : Barrier Island Flora

Dare Co . • N . C .

os
Dune
Scrub Hammock Forest
OG
(OS) (HHF )
Dune so
Grass (DG) J'J'
Sand Flat (SD) Salt Scrub (55) S-11 BA
~
Salt r-la r s h ( Si.?I);':J""''==''''~
Ocean (ED) Sound
Foredune 15 r Re ar Dune 65 '
At l antic Ocean Brackish Aquatics (llA)
Pamlico Sound

Beach Strand Succulents (BD)

h- Cakil e maritima , Sea Kale h Hydrocotyle bonariensis, Penny\~ort
h-Euphorbia polygonoides , Spurge h-Polygonum glaucum, !<notHeed
Dune Grassland (DG)
g-Uniola paniculata , Sea Oats h Erigeron pusillus , HorseHeed
g-Ammophila brevi ligula ta , Beach Crass h-Oenothera humi f usa, Evening Primrose
h-Hydrocotyle bonariens i s. PennYlw r t s-Iva imbricata , Iva
Sand Flat Herbs (SD)
g-Nuhlenberg i a fi lipes. Huhly h Gaillardia pulchella, Gaillardia
g-Chloris petraea , Finger Grass h-Gnaphalium obtus ifo lium , Rabbit
g- Cenchrus tribuloides. SandspuT tobacco
Dune Scrub (DS)
v - Parthenocissus quinquefolia , Virginia s-Horus rubra . Red Hulberry
Creeper s-Callicarpa americana, French Nulberry
v-Smilax auriculata , Catbrier s-Zanthoxylum clava-herculis, Tooth-
v-Vi tis aestivalis , Fox Grape ache Tree
v - Rhus radicans , Poison Ivy g-Uniola paniculata . Sea Oats
Haritime Hammock (l-filF)
t-Quercus virginiana , Live Oak sIlex vomitoria, Yaupon
t - Quercus laurifolia . Laurel Oak v-Gelsemium sempervirens , YellOl~
t - Ilex opaca , American Ho l ly Jessamine
t-Osmanthus americana , American Olive h-Hitchella repens, Partridge Berry
t - Cornus florida, FIO\~ering Dog\~ood h-Galium hispidulum, Bedstraw
Salt Shrub (SS)
s-Baccharis halimifolia , Goundse l s-Iva f r utescens , Narshelder
s-Nyrica cerifera , \-lax Hyrtle s-Borrich ia frutescens , Sea Ox-eye
Salt Harsh (SH)
h-Salicornia eur opaea , Glass~-1Ort g-Spartina patens
h-Suaeda linear i s, Suaeda g-Spartina alterniflora
h-Limonium car olinianum, Sea Lavender g-Distich l is spicata
h-Sesuvium po r tu l acastrum , Sea Purslane g-Juncus roeme rianus , Rush
Brackish Aquatic s (BA)
h- Ruppia ma r it i ma , Indgeon
-, Grass h-Zostera marina, Eel Grass
376 17

Blue Ridge : Montane Flora

Black Mountains , Yancey Co .• N. C.

SPF
6500'
Spruce Fir Peak
(SPF)
Ridge
>AS
SE Heath Bald
(lIS)
4500 ' NW

No rthern 1Vo Cove

Ha rdHoods J>
(NDF)
Carolina Gneiss 4000 '
Black Hountains

Heath Bald (HS)

s-Rhododendron catm.,rbiense, Purple 1. S Leiophyllum buxifolium, Sand Nyrtle
s-Rhododendron minus, Rosebay v-Smilax hispida, Catbrier
s-Kalmia latifolia , Nountain Laurel h-Galax aphylla, Galax
s - Leucothoe recurva , Fetterbush h-Krigia montana, Dandelion
s-Vacciniurn constablaei. Blueberry h-Hypericum graveolens, St . John's-wort
s - Gaylussacia baccata. Huckleberry h- Houstonia serpyllifolia, Bluet
s - Henziesia pilosa . Hinnie-bush h-Angel ica triquinata, Angelica
s-Viburnum cassinoides, Hitherod g-Scirpus cespitosus
s-Cornus alternifolia. Alternate- leaved g-Carex debilis
Dog\wod g-Carex brunnescens
s-Robinia hispida. Rose- acacia g-Danthonia spicata
Spruce-Fir Forest (SPF)
t-Abies fraseri, Fraser 's Fir s Sambucus pubens, Elderber ry
t-Picea rubens, Red Spruce s-Rhododendron cata\1biense , Purple L.
t-Betula lutea, Yellm! Birch s-Vaccinium erythrocarpon, Ht. Cran-
t-Betula papyrifera, Paper Birch berry
t-Sorbus americana , Hountain Ash h-Oxalis acetosel l a, Hood-shamrock
t - Prunus pensylvanica , Fire Cher ry h-Aster acuminatus, Aster
t-Acer spicatum, Nountain Maple h-Clintonia borealis, Clintonia
s-Ribes rotundifolium, Gooseberry h-Chelone lyoni, Turt l ehead
s-Viburnum alnifolium, Noosewood g-Carex intumescens
s-Nenziesia pilosa, Ninnie-bush g-Deschampsia flexuosa
Northern Hardwood Forest (NDF)
t-Acer saccharum, Sugar }Iaple v Aristo l ochia macrophylla, Dutch-
t-Aesculus octandra . Buckeye man ' s Pipe
t-Tilia americana , Basswood h-Eupatorium r ugosum , l.Jhite Snakeroot
t-Betula lutea , Yell0l1 Birch h-Claytonia caroliniana, Springbeauty
t-Acer pensylvanicum , Striped 11. h-Laportea canadensis, Stinging Nettle
t-Halesia ca ro l ina, Silverbell h-Vio l a canadensis , Violet
s-Hibes cynosbati , Gooseberry g-Ca r ex plantaginea
s-Leucothoe axillaris. Dog-hobble g-Carex pensylvanica
17 377

IV . SPECIES FOR RECOGNITION QUIZ

(Sample)

'~FERNS AND FERN ALLIES* Araceae

Lycopodium lucidulum, Shining Clubmoss Arisaema triphyllum . Jack-in-the-
(CDF) Pulpit (COF)
Selaginella arenicola , Sand Spikemoss
(TODFC) Juncaceae
Equisetum praealtum , Scouring Rush Luzula acuminata, Hood Rush (COF)
(OFC)
**,b~id:*>~**********,~**i,i,**1,*,~,'<****"/,*,~,~,\: Liliaceae
Adiantum pedatum . Maidenhair Fern (CDF) Erythronium americanum, Trout Lily
Athyrium asplenioides , Lady Fern (CDF) (CDF)
Camptosorus rhizophyllus, Halking Fern Nuscari racemosum, Grape Hyacinth (OFC)
(CDF) Polygonatum biflorum, Solomon's Seal
Cystopteris fragilis, Fragile Fern (CDF)
(CDF) Smilacina racemosa , False Solomon's
Osmunda cinnamomea , Cinnamon Fern (COY) Seal (COF)
Polypodium virginianum , Northern Poly- Smilax glauca , Cat-brier
pady (CDF) S. laurifolia , Bamboo (SB)
Polystichum acrostichoides. Christmas S. rotundifolia , Cat- brier (SB)
Fern (COP) Trillium grandiflorum , Trillium (COF)
Pteridium l atiusculum , Bracken Fern Uvularia perfoliata, Bellwort (CDF)
(UP) (TODF-G)
Arnaryllidaceae
''<GY}fNOSPER}IS* Zephyranthes atamasco, Atamasco Lily
Chamaecyparis thyoides. Hhite Cedar (PM)
(BlIF)
Juniperus virginiana, Red Cedar (PPF) Iridaceae
Pinus echinata , Short-leaf Pine (PPF) Iris cristata, Owarf Iris (POHOF)
P. palustri s , Long-leaf Pine (UPF)
P . pungens. Table Mountain Pine (T~WF) Orchidaceae
P. rigida, Pitch Pine (TIWF) Goodyera pubescens , Rattlesnake Plan-
P . serotina, Pond Pine (BHF) tain (HOOF)
P. strobus, t·1 hite Pine (PI1PF) Orchis spectabilis. Showy Orchis (CDF)
P. taeda, Loblolly Pine (PPF)
P . virginiana , Scrub Pine (PPF) ''<Oicots'"
Tsuga canadensis , Canadian Hemlock Salicaceae
(SPF) Salix sericea , Silky \'lillm" (FS)
T . caroliniana , Carolina Heml ock (SPP)
Myricaceae
"'NONOCOTS* Hyrica heterophylla , Hax Hyrtle (EHF)
Poaceae
Anthoxanthum odoratum , S\,reet Vernal Juglandaceae
Grass (OFC) Carya tomentosa, Nockernut Hickory
Alopecurus carolinianus, Foxtail (POHO¥)
Grass (OFC)
Aristida stricta , Hire Grass (TOOF-G) Betulaceae
Arundinaria gigantea, Cane (BHF) Alnus serru la ta . 'fag Alder (FS)
Bromus catharticus, Brame Grass (OHF) Betula nigra, River Birch (SDF)
Festuca sciurea, Fescue Grass (OFC) B. leuta, Cherry Birch (COF)
Poa annua, Annual Blue Grass (OFC) Carpinus caroliniana , Ironwood (SDF)
Cory lus americana, Hazelnut (SOF)
Cyperaceae
Carex nigromarginata (POHOF) Etc.
 378 17

Section C . FLORA AND VEGETATION--OBJECTIVES, AIDS, I\ND ANAl,YSIS

The objectives , the inclusions and aids, the spectra, and diversity types
are suggestions for comprehensive and basic organization, observation , analy-
sis, and synthesis in flora work. Flora and vegetation objectives and inclu-
sions are given to shm·] the similarities, differences.... and complimentary nature
of these two aspects of field study . I n add ition to learning ~Ihat ' s h'here when
or making a list of plant species within a given area at a given time, i.e., a
simple f lora, a field botany student should become acquainted \lIith plant habit
(Life Form) . g rowth patterns (Sociability), periodicity (Phenology), duration,
reproduction, spatial relat ions or pat terns of occurrence (Population and Com-
munity) , and distributional relations (Habitat and Geography) . Flora study
should add basic information about species biology to systematics .

I . OBJECTIVES OF FLORA AND VEGETATION STUDY

A. Flora (By Physiographic Region; Community; Soil , Rock, Habitat Type)

1. Determine and list species Hi thin study area by pattern, i.e . ,
community or habitat . (See Flora Summary )
2 . Analyze flora for life form, diaspore type, sociability , duration ,
and phenology. (See Spectra 1 , 2,3,4 , and 5)
3 . Analyze flora for indigenous and introduced species , disjuncts,
rel i cts , range ext e nsions, and new variations . (See Spectrum
8)
4 . Determine indica tor spe cies for soil and rock types and speCies
that are exclusive to restricted connnunities or habitats .
(See Spectrum 7)
5 . Indicate unique features Hithin a flora, e . g ., unusually large
specimens of a species, largest fruit , smallest flm']er, pecu-
liar pollinators, largest sym!)atric group , rarest taxa , endem-
ics, most variable popUlation , largest population, oldest pop-
ulation , largest tree , oldest plant, etc . ( See Flora Summary
and Spectrum 6)
6. Compare flora of study area Hith that of large r immediate area and
disjunct climatically similar areas as to speCies and larger
taxa similarities and/or differences . (See Flora Summary and
Spectrum 6)
7. Interpret flora as to origin, migration, and evolution . (See
Flora Sumillilry and Spect ra)
B. Vegetation (By Physiographic Region; Soil, Rock , Habitat Type)
1. Become familiar with flora of study area <lnd prepare a voucher
specimen of each species .
2 . Determine patterns (conuuunities) in \,]hich flora occurs .
3 . Determine distrihution of patterns (connnunities) .
4 . Analyze and map distribution of patterns (communities) in relation
to slope , exposure, elevation, etc .
5. Indicate unique pattern compositions and distributions .
6 . Compare community composition and distrib ution of communities with
those in similar climatic regions.
7. Interpret communities as to origil), e volution , and migration .

II . STANDARD INCLUSIONS IN FLORA AND VEGETATION STUDY (In addit ion to flora
l ists and field , herbarium, and labor ato ry data)

1. Location and geography of Study Area .

2. History of botanical exploration .
17 379

3. Survey of applicable physiography and topography .

4. Resume of pertinent pedologic and geologic data .
5. Summary o f meaningful climatological data.
6. Review o f previous pertinent work.
7. Description of present and past land use or abuse .
8. List of cited references .

III . RECONNAISSANCE AND PLANNING AIDS FOR FLORA AND VEGETATION STUDY

1. Topographic Naps (1) 6. Hydrologic Data & I-!ydographic

2. Geologi.c Naps (1) }!aps (1,4,5)
3. Soil Surveys (2) . 7. Climatological Data Summaries (3)
t•. Aerial Photographs (1,2,6) 8. Special Geological Reports (1 , 4)
5. Forestry & Vegetation Naps (2 ) 9. County Road Haps (6 )

Sources : (1) u. S. Geological Survey; (2) U . S . Department of Agriculture ;

(3) U . S . Heather Bureau; (4) State Geological Surveys or Departments of
Conservation & Development; (5) Corps of Engineers , U. S. Army; (6) State
Highway Department.

IV . FLORA SUHMARY

Species Lf Dia Soc Our Pbe Oi Conununities or Habitat

Note : Species should be listed according to a l-!anual or System, or systemat-

ical ly by family . See flora spectra classification and explanation fo r
meaning of symbols, and Section B for plant community symbols. Thi s flora
sununary arrangement can be easily progranuned for computer analysis in a
variety of '·JaYs.

V. DOCUMENTAT I ON

In flora and vegetation ,,'ork a s e t of specimens represen ting each

species studied or identified should be deposited in an herbarium as vouch-
ers for documentation. Standard label data should he included with each
specimen , and ecological and/or systemat ic data pertaining to each species
acquired during the study should be added to the voucher <. p,.~c i". eL . ,\ ;~trir
l.ihel ccr, taini n f, :: he ecologi.cal and/o r SYSt.€Tl:.:.ti.C 1n.i;orJl'.a t!.or'. pel l::'.npnt tn
the specimen should be added t o ti.(~ hotton .Jf ::!le :~l2 rbarl'. ,;n sh e er:. l'he
ecological and/or sY3 ,: emat :ic In:ormation ~h")t!id no t lJe i) ~dicat e d by symbols
or code but 'W!" Ltten cut so that future st\..ld-=nts \-jJ 1 have no trouble under-
,>ta!:dint~ the in ro rmdl-ivll on t_: e 1::ii;,~ J. .

Quantitative methods used i n 3'1aiv.;\.s JhOll~_,-l he dO<": .e l.enled \-,'itl: any

modifications of standarC: r:tocer:!u :es can, fl;l:!.:' ls.scr :.'. he--J . The types of
instruments used in acqu'isition or analysis of data should be ind i cated .
Literature used as a basis for any part of the \Vork; e . g. , physiographic
province nomenclature based on Fenneman, 1938, should be cited .

Documentation should be complete and comprehensive enough for the

reader of the report to be able to distinguish original observations ,
analyses, and syntheses [rom those previously published .
 380 17

VI. FLORISTI C OR COHHUNITY SPECTRAL ANALYS I S

(A sample form)
1. Life Form (Lf)
P-t P- s P v Ch G Th E S Pa Sa H
il of Species

% Total Flora I I I I I I I
2. Diaspore (Dia)
Spa Pte Pog Cyc Epz Syz Enz Eal Bar Hyd Ant Ate Pol
Ii of Species

% Total Flora

3. Sociability (Soc)
Soc I Soc 2 Soc 3 Soc 4 Soc 5
il of Species

% Total Flora

4. Duration (Our)
Evergreen
Ann Bien Per Decid Broad Needle
ii of Spec i es

% Total Flora

5. Phenolog:y (Phe) Leaf (L) - Flower (Fl) - Fruit (Fr)

V Ae Au H All Ds Ws
II of Species

% Total Fl ora

6. Taxon (Tax)
Psi Lyco Sphe Pter Pino Hagn Hagt Lilt
II of Species
Div./
% Total Flora Class

Mag Ham Car Dill Ros Ast Alis Arec Com Lil
If of Species
Sub-
% Total Flora Class

7. Communit:y (Examples)
DG SF DS SA SDF BHF TG LPF etc .
II of Species

8.
% Total Flora

Distribution (Di)
I I
Bar Cp Apr Tro Pra All Aut Dis ReI Spa Nat Dam
II of Species

% Total Flora
I IIIiIIIIIII I
Note : Hany other spectral analyses could be made, e.g.: those based on fidelity,
vitality, leaf size, lea f shape , corolla types, fruit types, poll inating vectors ,
etc.
17 381

Spectral Analysis Symbol Explanation

1. Life Form (L£)

P-t, Phane rophyte, tree; P-s, Phanerophyte, shrub; p-v. voody vine; eh,
Chamaephyte; G, Geophyte; Th. Therophyte; E, Epiphyte; S. Succulent; Pa .
Parasite; Sa, Saprophyte; H, Hemicryptophyte.

2. Diaspores (Dia)
Spa, Sporochore; Pte, Pterochore; Pog, Pogonochore; eye, Cyclochore; Epz. Epi-
zoochore; Syz, Synzoochore; Enz, Endozoochore; Bal, Ballochore; Bar, Baro-
chore; Hyd . Hydrochore; An t, Anthropochore; Ate. Atelechore; Po l, Polychore.

3. Sociability (Soc)
Soc I--Plants growing in one place, singly; So c 2--Plants grouped or tufted;
Soc 3- - Plants in small patches, or cushions; Soc 4--Plants in small colonies,
in extens ive patches, or forming carpets; Soc 5--Plants i n pure populations .

4. DUration (Dur)
Ann (l)--Annual; Bien (2)--Biennial; Per (3)- - Perennial; Decid (4)--Deciduous -
leaved; Evergreen Broad-leaved (5); Evergreen Needle-leaved (6). Note: Annu-
als could be classified as \~inter; spring - summer ; sununer-fall.

5. Phenol ogy (Phe)

Leaf (L)-- Time of Leafing; Flower (Fl) Time of Flowering; Fruit (Fr)--Tirne of
Fruiting . V(l)--Vernal or Spring; Ae(2)--Aestival or Summer; Au(3)--Autumnal
or Fall; H(4) --Hibernal or Hinter; All(5)--All Year; Ds(6)--Dry Season; Ws(7)--
Het Season .
Note: Phenology could be indicated by three numbers, the first, the l eafing
period ; the second, the time of f1m~ering; and the third, the time of f ruiting,
e . g . , 1-2-3 means leafing in spring, flowering in summer , fruiting in the fall .
Also instead of spring, summer, etc., phenology could be indicated by month,
e.g .• (1) Jan., (2) Feb., (12) Dec., etc.

6. Taxon (Tax)
See Classification System in Cronquist (1968)
Psi--Psi1otophyta; Lyco--Lycophyta; Sphe--Sphenophyta, Pter--Pterophyta; Pino--
Pinophyta; Magn--Magnoliophyta; Nagt--Magnoliatae; Lilt--Liliatae .

Nag- -Ma gno1iidae; Ham--Hamame1idae; Car--Caryophy11idae; Di11--Dilleniidae; Ros--

Rosidae; Ast - -Aster i dae; Alis - -Alismatidae; Arec--Arecidae ; Com--Comme1inidae;
Lil- - Liliidae.

7. Communities
See types of communities and symbol s in Section B.

8. Distribution (Di)
For definitions, see Geography in Chapter 16 .

Bor--Boreal; Cp--Coast"al Plain; App--Appalachians (which includes the Piedmont,

Blue Ridge, Ridge & Valley, Cumberland-Alleghany Mountains , Cumberland-Alle-
ghany Plateaus, and Interior Lm~ Plateaus); Tro - -Tropical: Pra--Prairie; All-
Allochthonous; Aut--Autochthonous; Dis--Disj unct; Rel--Re1ict; Spo--Sporad ic :
Nat--Naturalized, and Dom--Domesticated. Indigenous - native and Introduced-
brought in by man.
 382 17

VI. TYPES OF DIVERSITY IN FLORA OR CmfrlliNITY (Se e Flora Spectra Classifica -

tion and Origin and Establishment in Chapter 16)

1. Community (Habitat )-Area . Number of types of communities per unit

area.
2. Community-Species. Number of species per cnmmunity type .
3. Diaspore-Flora or Community. Number of types of diaspores Hithin
flo ra or community .
4. Diaspore-Species. Number of species per diaspore type within flora
or community .
5 . Distribution-Flora or Community . Number of prevalent ranges repre-
sented within flora or community--e . g . • species primarily prairie
or boreal in distribution, etc.
6 . Distribution-Species. Number of species per range type within flora
or community--e . g . , number of species primarily I/ith prairie dis-
tribution or boreal distribution, etc .
7. Duration-Flora or Community . Number of annuals, biennials, peren-
nials, or evergreen, deciduous species Hithin f lora or connnunity.
8. Duration-Species. Number of species per duration type within flora
or community .
9 . Life Form-Flora or Community . Number of types of life forms within
flora or community.
10 . Life Fo rm-Sp ecies . Number of species per life form type within
flora or community .
11. Phenology-Flora or Community . Number of phenological types l..rithin
f lora or community.
12. Phenology-Species . Number of species of each phenological type
I"ithin flora or community .
13. SOCiability-Flora or Community . Number of sociability types within
flora or community.
14. Sociability-S pecies . Number of species per sociability type I"ithin
flora or community.
15 . Spec ies-Area. Number of species per unit area.
16 . Species-Equitability. Number of ind ividuals per species within a
community or area.
17. Species-Number . Number of species per individual unit--e.g., num-
ber of species per number of stems or per number of separate plants.
18 . Species-Stratum. Number of species per stratum within a conununity
or flora .
19 . Taxon-Flora or Community. Number of divisions, classes, subclasses
etc . (ranks) represented in the flora or community .
20. Taxon-Higher Taxon. Number of taxa per next larger taxon Hithin a
flora or community- - e .g., number of genera Hithin a family, num-
ber families Hithin an order, etc. ; or number of higher taxa
within flora or community .
21 . Taxon-Species . Number of species l..rithin an order , subclass, et c.
I"ithin a flora or community -- e . g . , number of monocot s pecies
t..rithin flora or community .
COHBINATION DIVERS I TY. Any joining of two or more of the dive rsities listed
above--e . g., number of life form types per taxon or range type within the
flora or conununity ; number of species with a diaspore type and duration
type within the flora or community.
Note : Nany other types of diversity are present in a flora or community, e . g . ,
different fruits , pollinating agents, sexual conditions, etc . (See Phyto-
graphic Glossary) .
17 383

SELECTED TOPOGRAPHIC FORNS GLOSSARY

(Adapted from Lahee (1961) ; s ee Lahee [1961] or geomorp holo gy texts [e . g .,
Easterbrook, 1969] for a ddit ional for ms and information a bout all land or
topographic forms and geomorphic features .)

A. Basins. Topographic depr essions rimmed on all sides .

Barrier Basin . A dep ressio n formed by damming due to l ava flow , morain a l
debris , mud flow , landslide debris, or f luvial deposition .
810\,'-out . Usually an interdunal wind-blown depression .
Solution Basin. Sinkho le . Gabbro Depression . Depression forme d due to
chemical dissolution of underlying rock .

B. Beaches. Shoreline deposits of sand , pebbles or rock made by action o f

Haves and longshore curren ts .
Barrier Beach . Sand re e f bu i lt up offshore by deposition of churn ed
bottom san ds .
Bayhead Beach . Accumulated debri s frora ne i ghboring headlands de posited
a t the head of a bay .
Baymou th Ba r . Barrie r beach built across the en t ranc e to a bay .

C. Benc hes and Ter r aces . Relatively f l a t gently or horizon tally inc lin ed
s urfaces bounded by a s t eeper ascending slope on one side and a steeper
descend ing slope on the opposite side. Benches are usually cons ider ed
forms in solid rock and te rraces f o r ms in unconsolidated mate ria l s .
River Terrace . A gently sloping or horizontal depositional surface on a
flood plain .
Have- cut Bench . An eros i ona l flat rock floo r which te r minates land\mrd
at the foot of a seac li ff and seal..rard i n a s hore terrace.
Have-built Terrace . Gen tly ascending , beachlike form above high water
level .

D. Bluf f s , Cliff s , Scarps , Precipi ces , Escarpments . Vertical or very stee p

face of ro ck or par tially consolidated sedimen t s .
Fault Cliff or Sca rp . Cli ffs of varying height forme d by faultin g \..r H h
dislocation of bedrock structures.
Glacial Cl iff . S teep bluffs left aft er glacial scouring o r ice quarrying
a n d sapping .
Landslide Cli ff . Steep scars left by the fall of masses of rock or uncon-
solida ted materials .
Sea Clif f . Erosion blu ff fo r med by undercutting by sea waves .
Solution Cli ff . S t eep s l o pes l e ft by the sudd en collapse of the floor of
a l imesink .
Stream Cliff . Erosion bluff formed by lateral swinging and subsequent
undercuttin g by a s tream .
Volcanic Cliff. Abrupt edge s of lava flows . usually formed at the lava
front ; or steep slopes r es ultin g f rom a vol canic explo sion .
Heathering Cliff. Bluff resulting f rom normal weathering in horizontal or
inclined layers of rock of differential resistance .
-,
E. Hills , Ridges , Noun t ains , Peaks . Relief forms tha t proj ect upI..ra rd and are
sur r ounded on all s ides by lower land .
Block Hountains. Ridges due to fa ulting, u sually \.,rith one slop e gentle and
the other steep along the fault scarp .
Volcanic Cones. Peaks built from lava or volcanic fragment s or ejectaments .
Dunes . Ridges or hills b uil t by wind- blmm sand .
384 17

Drumlins . Oval-shaped hills formed from glacial drift under a moving

ice sheet .
Esker s. Na rrO\;f. elongate, \;finding ridges o f mixed sand and g ravel formed
in stream bed tunnels under a glacier .
Noraines. La teral, terminal, or recessional ridges or irregular deposi ts
laid dmm by valley glac iers or ice sheets . ....
Talus Ridges . Deposits of rock fragments at the foot of a rocky slope .
Monadnock. A residual hill or mountain on a peneplain .
Nesa . A f lat-topped mountain consisting of horizontal beds bounded by
steep erosional scarps .
Butte. An isolated hill or mountain forme d on a reduced or eroded mesa .
Domes. Rounded knobs formed by erosion or exfoliation of granitic rocks.
Hogback . Sharp ridge usually formed from erosion of steeply inclined
strata or beds.

F. Lakes, Pools, Ponds, Bays. \-later-filled depressions .

Fluvial Lakes or Ponds, Pools . l~ater-filled depressions due to stream
activity.
OxbOl". Hate r-f illed , cut-off meander on a mature flood plain.
Plunge Basin . Hat er-f illed depression formed at the base of a \>'ater-
fa ll.
Pothole. Sand-scoured, smooth-sided depressions forme d in hard rock
by a rapidly down-cutting stream .
Glacial Lakes. Hater-filled depressions formed by glacial activity.
Cirque Lake, Pond, Pool. A blocked, \"ater-filled amphitheater-like
valley or part thereof at the foo t of a mountain glacier .
Ke ttle . Hater-filled depression formed by melting ice left in glacial
drift .
Horainal Lake. j·later-filled depression behind a morainal dam that
blocks drainage.
Longshore Ponds or Bays. Bays or ponds formed by baymouth bars truncating
or blocking a stream or current .
Heteoritic Lakes, Pools , Ponds . hlater-filled depression s formed by met -
eoritic impact or shock\olBves .
Pluvial Ponds, Pools. Ra infi lled depressions.
Rock Barren Pools. Ephemeral, erosional t.ater-filled depressions in
granite and other hard rocks.
Dune Pools . Rainfilled depressions in blowouts or interdunal areas .
Playa Lakes. Ephemeral lakes in old lake bed areas.
Solu tion Lakes, Ponds, Poo ls . I·.later-filled depressions formed in rock
that has been chemically dissolved .
Tectonic Lakes, Ponds, Pools. Hater-filled depres sions due to b l ocking
by fault or earthquake movement of materials .
Volcanic Lakes. \!ater- f illed depressions caused by stream blockage by
lava fINIS or volcanic cone blO\;'Quts or eruptions producing a deep
depression .

G. Plains and Flats. Comparatively flat forms lvith l ow i nclinations usual ly

cover ing many acres or square miles .
Alluvial Fan or Cone. Gently sloping deposit fo rmed Hhere a stream l eaves
mountains \"ith an abruptly reduced load-carrying capacity .
Coastal Plain . Sea floor exposed due to relative upl ift of position in
respect to sea level .
De lta ic Plain . River deposit made by a stream in a large body of Iolater
when the stream loses its load-carrying pOHer .
17 385

Estuarine Flat, Hud Flat . Small exposed plain left at 10\.,T tide.
Estuarine or Lagoon Plain. Large flat le ft after evaporation or drainage
of the estuary or lagoon .
Flood Plain. Alluvial flats and terraces formed from sediments deposited
along a stream.
Lake-floor Plain . Flat left after evaporation or drainage o f a lake .
Outwash Plains. Glacio-fluvial d eposi ts in front of a cont inent al ice
sheet. Glacial delta plains Hould be built by deposition in a glacial
lake; frontal aprons are meltl"ater outHash deposits that ,,,QuId corres-
pond to alluvial plains; sand plains are glacial outwash composed mostly
of sand .
Peneplain. An erosional plain; a landscape of low r elie f formed by 100g-
continued erosion.
Playa . Plain underlain by clays alternating {"ith salt Hhich {"ere deposited
in broad, ephemeral intermittently water-filled basins .
Till Plain. Uniform deposits of glacial materials i n lands of low relief .
Volcanic Plain. Surfaces of broad sheets of lava and/or volcanic ejecta-
menta .

H. Valleys. Open topographic depressions with or {" ithout streams.

Glacial Valley. A rock-walled valley which is V-shaped in cross-section
and bears evidence o f glacial abrasion.
Fault Valleys. A valley formed from a fault in {·,hich blocks are tilted
so that a depression lies at the base of the fault; a valley formed
from rifts along multiple fractures; a valley formed hy the slipping
downward of a block between two fault scarps; a valley formed over a
block as two lateral blocks are uplifted.
Solution Valley . A valley formed over coalesced sink holes; a corrosion
valley over so l uble rock .
Stream Valley . Valley formed by the dov/U-cutting of a str eam usually with
a valley train or flood plain in the depression .
Gully, Gorge , Ravine, Canyon . V-shaped destruction valleys with compara-
tively steep sides.
Hanging Valley . A tributary glacial valley Hith a floor higher than that
of the trunk valley.

SELECTED ROCKY SUBSTRATES

(Adapted from Porter et al . [ 1973])

Basalt. Dark gray to black, fine··grained, extruded igneous rock.

Caliche. Calcareous hardpan .
Coralline Rock. Limestone derived from th e calcareous deposits of corals .
Dolomite. Harble or compact limestone rich in magnesium and calcium car-
bonates, often Hith a crystalline-granular structure.
Gneiss . Laminated or foliated metamorphic rock simi lar to granite in compo-
sition .
Granite. Fine- to coarse-textured int ruded igneous crystalline rock .
Hardpans . Relatively impermeable hardened strata beneath soils, cemented by
a variety of leacheclor deposited materials.
Igneous Rock. Hagmatic; originally molten, issuing from the interior of the
earth and solidified following intrusion or extrusion.
Lava . Igneous rocks extruded from volcanos .
Limestone. Sedimentary rock composed primarily of calcium carbonate.
Har!. A loose , crumbling calcareous sedimentary deposit laid down under still
water .
386 17

Metamorphic Rock. Sed imentary or igneous in origin , the basic structure

changed by heat and/or pressure follm.;ing deposition or intrusion .
Quartz. Crystalline silicon dioxide .
Quartzite . Netamorphosed san d ston e .
Rhyol it e . Acidic volcan ic rock t~ith a crystalline goundmass .
Sandstone . Sedimentary rock composed mainly of sand more or less firmly
cemented by other minerals . ~.
Sedimentary Rock . Composed of particulate matter deposited , accumulated, and
compac ted under Iva ter.
Serpentine . Consisting mainly of hydrated magnesium silicate and 101" in
calcium .
Shale . Finely laminated sedimentary rock composed of clay , mud, and/or silt,
the const i tuent minerals essential ly unchanged since d eposition , the lamina
parallel to the bedding plane .
Slate. Dense , fine-grained metamorphic rock formed by the compression of
shale , developing more or less parallel cleavage lamina in planes at any
angle to the bedding plane.
Soapstone . Soft, soapy-feeling rock composed mainly of talc, chlorite, and
often magnetite.
Talus (Scree). Rock fragments derived f rom the weathering of a cliff or
slope and accumulating at its base in a sloping pile .
Tuff (Tufa) . Porous and usually stratified rock composed of consolidated
volcanic ash .
Volcanic Rock. Igneous rock solidified after extrusion fr om a volcano or
similar vent .

GENERAL FLORISTIC LITERATURE

Braun , E . 1. 1950. Deciduous Forests of Ea stern North America . The Blakis-

ton Company. Philadelph ia .
Daubenmire , R. F . 1968. Plant Communities . Harper & ROi. . NeH York .
Lahee, F. H . 1961. Field Geology . 6th ed . NcGraw-lIi ll Book Company . Ne~·]
Yo rk .
Porter, D. N .• R. \.)" . Kiger , and J . E. Honahan . 1973 . A Guide for Contributors
to Flora North Amer.ica, Part II . An Outline and Glossary of Terms for Nor-
phological and Habitat Description (Provis iona l Edition) . Department of
Botany , Smithsonian Institution . Hashington , D. C.
Smith , R. 1. 1966 . Ecology and Field Biology . Harper & Ro\. . Ne\"l York .
\-lhittaker, R. H. 1956 . Vegetation o f the Great Smoky l-!ountains . Ecological
Monographs 26: 1 -80.
1972 . Evolution and measurement of species divers ity . Taxon 21 :
213-251.

flEW EXERCISES
(A limited number of suggestions)
I. Spectral Analyses--by class or ind ividual student .
1. List the species of a community, habitat, or area and make a spectral
analysis .
2 . l-!ake comparative spectral analyses of the floras ; e . g . , of a south
slope and a north slope; montane and piedmont selected communities ;
ridge and cove communities .
II . Diversity Studies--by class or individual student ; e . g ., determine leaf ,
root , stem types in a habitat; leaf or floral variation Hithin a popu-
lation; floHering or fruiting times for canopy and herb l aye r speci es
i n a climax community (phenology) .
III . Ecosystematics Problem (See Chapter 35) .
18 387

Chapter 18. COLLECTION AND FIELD PREPARATION OF SPECIHENS

Herbarium specimens are permanent records of a species (or population) as

it occurred at a given time and place . The future value and use of any speci-
men is largely dependent on the care \"rith which the collector selects, col-
lects and prepares his specimens . The follotdng directions and suggestions on
specimen preparation. field equipment and field records are given to assist
collectors in preparing high quality herbarium specimens accompanied by ade-
quate field notes .

1. COLLECTING OBJECTIVES A.l\JD PLANNING

The specimens to be collected depend on the objectives of the collector,

and the types of mat erial to be collected will determine the techniques used,
amount of material collected, and type of field data recorded . The following
suggestions can greatly aid the collector in getting maximum benefit from
valuable field time.

1. Outline objectives for a specific expedition.

2. Prepare a l ist of all equipment to be utilized on a field expedition .
3. Obtain localities from herbarium specimens if specific materials are
needed. It is best, when possible, to have several localities
in the event some areas have been disturbed . Check flm"rering and
fruiting dates on the specimens in the herbarium .
4. Check local t.;reather conditions and seasonal progress in the areas to he
visited.
5. Collect duplicate specimens (replicates) except in the case of very rare
or protected plants (check state la,.s and conservation lists) . Other
plants to be avoided include those noxious t~eeds or parasites and
their host plants under state or federal quarantine . Duplicates pro-
vide sufficient material for dissection or, i f needed, verification
by a specialist.
6. Prepare voucher specimens ( typical herbarium specimens) when collecting
materials for cytological, anatomical, and other studies .
7. Obtain collecting permits and other necessary permission in advance, i f
pOSSible, if a trip is planned to a specific area such as a national
park, foreign country, or other area. It is also advisable to obtain
permission for collecting on private lands t.;rhenever possible .
8. Collect and document as many different species as possible in floristic
studies . Do not overlook the inconspicuous (so-called belly plants)
or those plants t.;rhich are difficult to collect or identify . Collect
thoroughly in each habitat.

II. SUPPLIES AND EQUIPHENT

1. Field press. A press typically consists of t,.o hardwood frames, with

each fr ame made . as folloHs :

4 wood strips 3/4 x 1/4 x 18"

5 t~ood strips 3/4 x 1/4 x 12"

The five short strips should be equally spaced on the four longer
 388 18

strips vlhich are also equally spaced and nailed , or rivet ed securely
at the intersection of t he s tri ps . The complet e d frame should b e
12 x 18 inches . Repeat for the second frame .
Some coll ecto r s prefer the cheape r and of ten st r onger 12" x 18"
plywood press boards cut from 3/8" fir plywo od . A number of types of
pre ss es may be purchased from biolog ical s upply hou ses .
2. Driers (blotte rs). Exce llen t dr ier s may be ma de by cutt ing sheets 11 x
16 inches f rom l igh t weight builders deadening felt (unsaturated) o r
from heavy bl otting paper . Driers ar e also available f r om biological
s upply houses .
3. Newsprint. Cut paper 22 x 1 6 inches a nd fo l d to 1 1 x 16 inches . Hany use
newspapers as found on the newsstand but unused ne\"sprint may be pur-
chas ed i n ro l l s fro m l ocal print ers . Biological supply hou ses usually
offer precut pap ers .
4. Press st r aps (webbing st raps) . A pair of strong we b st r aps (pa r achute o r
c inch type) with elm" buckles are excellen t fo r field purposes . Sash
co rd or r ope is often al so us e d . The minimum length for press straps
is fo ur fee t .
5. Fie l d notebook . A pocke t-si ze book which will not dis integrate when wet
and pen cil or pe n \~i t h lJBter-proof i nk are necessa r y items . Some
prefe r to keep two books --one taken to the field and another \-,lhich
remains in a safe pl ace and into which f ie l d notes are copi ed . If
fie ld l abels are used, one i s placed in the pape r with the spec imen
and a copy kept by the col lec tor . Field no tebooks shoul d be permanent .
6. Stri ng t ags. H'aterproof s t ring t a gs are usef ul fo r l abel i ng pl an t s which
are not pressed immediately foll owing coll ection .
7. Digge r s and c lipper s . Both pruning shears or garden clippe rs and digging
tool s are necessary . A trowel (prefe r ably with a stee l shank) . geo l o-
gist pick , dandelion digger or heavy sheath knife are excellen t fo r
field use . A small trench shove l and pocke t knife a r e als o use ful.
8. Han d len s . For field obse rvation s and identificat ion s a small 5x or lOx
lens is des irable . The se a re genera lly available from bookstores and
biological supply houses .
9. Co l lecting bott l es . Glass or plast i c bottles with leak- proof screw caps
are oft e n desirab l e for collecting some materials. The siz e us ed
depends upon the mate ria l s to be col lected. Small via l- type bo ttles
a r e ideal fo r collec ting fl oral buds . f l owers for c l ear ing and o ther
mater ials to be pr ese rved i n liquid preservatives .
10 . Liquid preserva tive. The type of so lution use d will , of cour se, depend
on futur e use and type of mat erial . For gene ral anatomical purposes
and materials s uc h as lJood, leaves , flO\~ers and t he l i ke a mixt ure of
Fo rmalin -Acetic Acid- Al cohol (FAA) i s widely used and may be pre-
pa r e d in the follo\~i ng manne r: Ethyl alcohol (70 %) . ........ . .. 90 cc .
Forma lin (commerc ial s trength) . 5 cc .
Glacial Acetic ac id .... ... . . . . . 5 cc .
Cytologica l mate r ials are oft en fixe d in a 6 :3: 1 mixt ure of Ch l or o-
form , 95% e thyl a l cohol and gl acial acetic acid or i n Carnoy ' s Flu id
(3 : 1 ab sol ute e thyl alcohol and glaCial ace tic acid) . Carnoy ' s con-
tainin g 95% ethyl a lcohol is useful fo r clearing leaves ( see Chapter
7) • lvhen using the 6 : 3 : 1 mixture t he gl acial acetic ac id should not
be added until mater ials are r eady to be fixed .
After mat erials have been in f ixative fo r 24-48 hours the
fixative s hould be discarded and mate rial s stored in 70 percen t
18 389

alco hol. At Kew a solution of 50% alcohol , 5% Forma lin, 5% gly-

ce r o l and 40% water i s used fo r sto ring plant mate rials . Each
vial should contain a field l abel . Use pencil and sl ip s of bond
paper; the n umber s should cor r espond to those in the fie l d note-
book .
11 . Vascula a nd co llect ing bags. Plant mat eri als not pr essed i n the
fie ld immed iately may be stored in a metal containe r (vasculum)
in fold s of wet paper . Due to the cos t and bulkiness of this
container many now pr efer to use plastic bags ( turkey or fer -
tilizer) or rubbe r- lined canvas bags (military la undry bags) .
Lo cal conditions will determine in part the type of bag to be
used , but avoid expo sure to sun , particularly i f c l ear plastic
bags are used . Al l ma t erials should be carefully labeled to
avo id confus ion when materials a r e pressed .
12 . Waxed paper . A rol l of waxed paper avai l able in s upermarkets is use-
f ul in pressing man y plants , particu l arl y those that are vi sc i d
or which deliquesce. A singl e sheet pl aced ove r th e ma terial can
greatly facilitate the removal of materials f r om the pressing
paper.
13 . Manila coin envelopes . Small coin e nvelope s are excellent for col-
lectin g seeds, individual flm'J ers , leaves, polle n, pollinato r s
and other ma terials which cannot or should not be pressed . These
should always be carefully labelled with the co llection number .
14 . Trays , cans , ja r s fo r living materials . It i s of t en desirab le to col-
lec t living mat e r ia ls for experimen tal work i n the garden or
greenhouse. A variety of conta i ne rs may be used a nd wi l l depend
on the t ype of material to be coll e c ted and the l e ng th of t i me
ma t er i a l s must be stored.
15. Cardboard storage boxes . Herrill cases or other cordboard co ntain-
e rs which ca n be pur cha sed Elat make excellent storage for dry
mat e rials removed from the press in the field . It 1s advisa ble
to use a n insecticide o r r epe lle nt i f material s are to be stored
for an extended period of time .
16 . Insect i c i des and repellent s . Moth crys tals or f l akes (n aphthalene or
paradichlorobenzene) may be used as r epel l ents . Considerabl e quan-
t ities of paradichlorobenzene in a n ai r- t i ght container ma y be
used as an insec tic ide. An excellent t r eatment is to plac e PDB
in the fol ds of t he specimen paper , tie specimens into bund l es,
and sea l in a plas tic bag .
17 . Haps . Higiu.Jay , topographic, and geo logic maps are often very useful
in loca ting localities for particular spec ies . Deta iled county
road maps can be obtained , us ua lly at modest cos t , from state
highway departments .
18 . Camera . Photographs are of considerab le value when collect ing Hoody
or other materials l.Jhere en t ire plants are not pressed .
19 . Co l or Charts . Several colo r c har t s a r e available for de t e r minat i on
of flower colors in the field, e.g ., Horticultu r e Society Color
Char t, Nickerson Col or Fa n , Ho rt ic ultura l Co lor Char t, etc .
",
III . SELECTION OF MATERI AL

Vigorous , typica l specimens are to be selected . Avoid i nsect damaged

plan ts . Specimens should be representative of the populat i on but should
include the r a nge of variation of the plants , not tho se that best fit the
press . Roots, bulbs, and other underground parts should be caref ully
390 18

excavated and the dirt removed ,.ith care. In most cases flowering and/or
fruiting materials are necessary for identification purposes. ~!any collec-
tors prefer to add extra floHers and fr uits to their collec tions when pos-
sible to avoid dissection of the specimen proper. Plants too large for a
single sheet may be divided and pressed as a series of sheets (see discus-
sion heIO\,;) . In collect i ng large herbs, shrubs and trees, different types
of foliage , flowers and fruits should be collected from the same plant .
Collect sufficient material to fill an herbarium sheet and still leave
enough room for the label . Bark and wood samples are often desirable addi-
tions Hhcn coll ecting \-Ioody plants. Proper i dentification o f many plants
depends on several different characteristics--some roots , others seeds or
mature fruits , some flm"er color (Hhich should be noted in the fie ldbook) .
The following selected suggestions Hhich are largely adapted from
DeHolf (1968) , Fogg (1940) , Fosberg and Sachet (1965), and Smith (1971)
are given to assis t in the selection and collection of particular kinds of
plants .

Table 18-1 . Naterials for identif ication purposes.

ESSE NTIAL OR DESIRABLE

!>1ATERIALS FOR IDENTIFICATION SUGGESTIflNS FOR NOTE-
TA.'{ON PURPOSES TAKING Al~D PRESSING

Acanthaceae Flmlers and fruits important . rlm"ers often detach ea s-

ily after collect i ng .
Agrimonia Underground parts useful . Press immediate ly .
Alismataceae FIOl"ers and f ruits (essential). Note presence of perfect
and/or staminate and pis -
tillate flowers and posi-
tion of fruiting pedicels .
Allium Flowers, seeds , bulbs (Hith
coat).
Amaranthaceae Ripe fruits (essential in some) . Note Hhether plants are
monoeciou s or dioec i ous .
Colle ct both stamina te
and pistillate plants I"hen
possible .
Amelanchier FloHers , fruits and leaves Not e habit (erect or stolon-
from same plant necessary . iferous) .
Annonaceae FIO\"er s necessary (these may
open precociously and then
grow conSiderably before and
during anthesis) .
Apiaceae l'!ature fruits and basal leaves.
(Umbell i ferae)
Araceae Fruiting material alone of Peel epidermis 81"ay on one
little value ; flmlers , side to spee d ki lling and
inflorescences and under- drying or use alcohol or
ground parts of great value ; formaldehyde to kill and
also leaves (often d imorphic) . preserve in the field .
No te glaucescence , sap
colo r , vesture. For large
aroids see suggestions in
Fosberg and Sachet (1965) .
Araliaceae Flol"ers and fruits .
18 3~1

ESSENTIAL OR DESIRABLE
~~TERIALS FOR IDENTIFICATION SUCGESTIONS FOR NOTE-
TAXON PURPOSES TAKING AND PRESSING

Arccaceae Flowers and fruits, inflores- See Tomlinson (in Fos-

cence axis and bracts, leaves, berg and Sachet-,-1965)
petiole, leaf base , notes on for details on collectin~
stem . specimens .
Asclepiadaceae Flowers and/or fruits . Note flower colo r, number
of leaves and arrangement,
latex color , fr uiting
pedicels (erect or de-
flexed .
Asteraceae Ripe fruits and flowers , Note co lor of ray
(Compositae) bas al and median cauline and disk flowers; press
leaves and underground parts . some heads exposing upper
surface and othe rs Id t h
l ower surface exposed.
Sp l it heads and press .
Balanophoraceae Tuber sur face important. Plants sometimes dioe-
cious .
Ba l saminaceae Flowe r s a nd fruits desirable . Fl owers fragi l e and tend
to agglutinate on drying
under press ure. Liquid
preserved flowers desi r -
able . Note flower col or
and markings .
Bamboos Flot~ers and fruits rare but Record node numbe r (start -
desirable . Culm sheaths from ing at base of plan t) from
mid-culm nodes , portion of which materia l is taken.
c ulm and branch , rhizome . Collec t from 4th to 5th
node . (See s uggestions of
l>!cClure i n Fosberg and
Sachet for addit ional
handling) .
Begoniaceae Both staminate and pistillate Careful notes on colors,
f l owers and ripe fruits. pl ants fleshy and often
darken on dry ing .
Betul a Fruitin g catkins essentia l. Notes on bar k desir able .
Boraginaceae Flowers and ripe fruits . Sp lit and press some
flowe r s fla t .
Bra ssicaceae Flowe rs and fru i ts desirable ;
underground par t s and rosettes.
Burseraceae Fruit.
Cactaceae Flowe r s, stem with grooves, Photog r aphs des i rable.
t ubercles and spines . Stems may be cut. ho llowed
and pressed including both
transverse and longitudi-
.-. nal sections . Stems may
be dipped in bo iling water
or forma lin to ki ll t issue
and allow more rapid dry-
i ng . Handle ca ref ully
(preferably with gloves and
forceps) but do not remove
or cut all spines .
392 18

ESSENTIAL OR DESIRABLE
~~TERIALS FOR IDENTIFICAT ION SUGGESTIONS FOR NOTE-
TAXON PURPOSES TAKING AND PRESSING

Campanulaceae F!m..rers . Corolla shape important,

make carefu l notes and/or
drawings.
Cappar i daceae Flowers desirable (th ese op en
precociously and then grow
considerably before and dur-
ing anthesis . )
Caprifoliaceae Ripe fruits.
Carex !1ature perigynia, underground
parts.
Caryophyllaceae Flowers and f ru its . Note number of styles and
corolla color .
Casuarinaceae Stami n ate and pistillate Dioecious or monoecious .
flowers and ripe f rui t.
Celtis Nature fr uits. Shape and color of fruit
important.
Coccoloba Leaves from adventitious shoots
and from mature twigs; stami-
nate and pistillate inflores -
cences.
Commelinaceae F'lm.]ers, spathe , underground Flowers should be pressed
parts . between tissue and waxed
paper and/or preserved i n
liquid . Note corolla and
anther color . Spr ead
spathe and press flat .
Convolvulaceae Flowe rs and ripe fruits Spli t flowers and d~y f lat .
desirable . Note corolla color and
markings.
Cornaceae Na ture fruits . Note colo r of branchlets
and pith .
Crata egus Flowers and fruits from same Note anther colo r.
tree .
Cucurbitaceae Flowe r s and ripe fruits . Dio ec io us or monoecious.
Note corolla shape and
color and color of mature
fruits.
Cuscuta Flowers and fruits. Note nature of peta ls and
their appendages, shape of
fruit, and host plant .
Cyperaceae Ripe fruits and undergro und
parts.
Dilleniaceae Ripe fruits .
Dioscoreaceae Staminate and pisti ll ate in- Note direction of twi n i n g
f lorescences . axilla ry bulbils, of stems.
underground parts and base of
aer ial stem ; ma ture fruits i f
possible.
Dipterocarpaceae Ripe fruits .
Epac ridaceae Ripe fruits.
18 393

ESSENTIAL OR DESIRABLE
}~TERIALS FOR IDENTIFICATION SUGGESTIONS FOR NOTE-
TAXON PURPOSES TAKING AND PRESSING

Ericaceae Flowers and fruits . Note flower color and

\.,raxy "bloom" on fruit .
Euphorbiaceae Staminate and pistillate Note color of glands.
flo\.,rers, ripe fruits . (Sap may cause dermati-
tis. )
Fabaceae Flowers and fruits essential; Note flm.,rer color .
(Leguminosae) underground parts desirable.
Fagaceae Ripe fruits, mature leaves . Collect staminate and pis -
tillate flowers \.,rhen pos -
sible. Note shape and
size of plant .
Fraxinus Ripe fru its nearly essential, Nay be dioecious .
staminate and pistillate;
flowers desirable .
Calium Fruits often essent ial; Not e flower color and
underground parts . habit of plant (erect or
reclining) .
Gesneriaceae Flm...ers and fruits . Note colors, plants often
fleshy and darken upon
drying.
Hydrocharitaceae FloHers and fruits . Note position of f lm.,rers
(floating or submerged) .
Iridaceae Flowers and mature fruits, Press flowers immediately
underground parts. and use tissue and \.,raxed
paper .
Isoetes Fertile specimens. Note habitat and whether
plant was emersed or sub-
merged. Split some plants
vertically.
Jug landaceae Fruits, leaves. Note bark characters, size
and shape of tree.
Juncaceae Fruits. Note number of stamens,
shape of leaf (flat or
terete); expose ligule .
Lamiaceae Flm.,rers and mature nutlets, Split open some flowers
(Labiatae) base of stem and underground and press flat; note
parts. flower color and " markings.
Lemnaceae Flowers and fruits desirable. Note number of roots per
"frond. II Float specimens
out on paper; place in
packets .
Lepidium Flowers and fruits. Note presence or absence
of petals.
Liliaceae Fruits alone useless, flowers Note if leaves are flat
and u"nderground parts most or ro und. Hake longi-
desirable . tudinal section of bulbs;
leave bulb coats intact.
Loranthaceae Flm.,rers and fruit . Note f ruit color and host
plant .
Nalvaceae Flowers and mature fruits, Note flower color; split
underground parts helpful. open some flowers and press
flat .
394 18

ESSENTIAL OR DESIRABLE
HATERIALS FOR IDENTIFICATION SUGGESTIONS FOR NOTE -
TAXON PURPOSES TAKING AND PRESSING

Nelas t oma taeeae Flowers and f r uits . Pe t als fugacious .

Hen is per mace ae Staminate and pistillate l:Hoecious.
floHers ; fru i ts desi r able .
Moraceae Fruits desirable. Frequently dioecio us ;
note fr u it color .
Husaceae Axis of inflorescence and Photograph entire plan t
ripe fruit . I.;hen possible .
Nyristicac eae Ripe fruits desirable . Dioecious .
i1yrtaceae Fruits .
Najadace a e Fruits . Floa t specimens out on
white paper .
Nep en t haceae Pitchers of full gr own
cauline leaves essential .
Orch i daceae Flmolers and fruits ; fruits Note fl Ol."er color , mark -
alone almost useless . ings , and fragrance .
Preserve some flowers in
liqu i d .
Oroba n c haceae Ripe fruits desirable . Note host pl ant .
Pand anaceae Ripe fru i t , leaf tips . Note stem diameter ; photo-
graph or habit sketch very
desirab l e .
Picea l-la ture cones . Note co l or of foliage .
Pipe r aceae Fruits . Note colo r s (plan t s often
darken on drying) .
Poaceae !-Iature fruits, unde r ground Note anther color . Press
(Gramineae) parts and stolons . culms to shol." sheath and
ligule .
Polemon i aceae FloHers , fruits and under- Note flOl."er color and
ground paLts . split corollas open and
pre s s flat .
Polyga l aceae FloHers and f r uits and seeds; No te color , hahit , leaf
unde r ground parts. d i morp h ism, if no t press-
ing entire plant .
Po l ygon a ceae Frui t s a n d underground parts .
Pota mogeton Fruits and stipules. Press so stipules a re
clearly displayed .
Pt e rido p hy t a Fertile f r onds, rhizome or
rootstock , ster i le f r onds if
different from the fertile .
Tree fe r ns - petiol e , leaf
bases and leaf scars .
Ranunc ulaceae Fruits and unde r ground parts .
Ri be s FloHers and fruits . Note color of fruits .
Rosaceae Fruits ; floHering and sterile Note habit of plant.
shoots .
Rubiaceae Fru i ts very desirable .
Rubus Habit important , tl.;igs of Note if canes are arching
sterile and fruiting branches . or not , rooting at tip or
not .
18 395

ESSENTIAL OR DESIRABLE
}[ATERIALS FOR IDENTIFICATION SUGGESTIONS FOR NOTE-
TAXON PURPOSES TAK IN G AND PRESSING

Salix Nature staminate and pistil-

late catkins and l eaves from
the same p lant .
Sapindaceae Fruits desi r ab le .
Sapotaceae Fruits desirab le .
Scrophulariaceae Flowers . Sp lit fl m"e rs and press
flat ; make careful notes
on color and markings;
many must b e pressed imme-
dia tely; commonly darken
on drying ; note host .lhere
applicable .
Smilax Fruits and vegetative Note fruit color and
mat e rial. \vhether main axis is
smooth or spiny .
Solanaceae FlO\1ers and f ruits. Split open some flowers
a nd press flat .
Styracaceae Fruits desirable .
Symplocaceae Ripe fr uits .
Utricular ia Flowers , fruits. leaves . Float p l ant s out on white
paper , spread leaves . Ex-
cavate terrestrial species
carefully .
Vaccinium Flm.;ers . fr ui t . No t e fruit color and plant
habit .
Viburnum Ripe f ruit .
Zi ngib e raceae I n f loresc ence and underground Inflorescence b e st pre-
parts. served in liquid .

IV . PRESSING PLA..""IT SPEeD-fENS

A. Arranging and Pre par ing Specimens . After the s p ec imens have been dug
or cut they should b e pressed as soon as possible (s ee V f or
in f ormat ion on field storage of fresh specimens ). The care
given a specimen i n pressing will large l y determine i ts fut ure
value . Specimen s should be placed i n a single fo ld of ne\~s­
print or other suitable absorbent light\o.'eight paper. P lants
too l arge to fit the 11" x 16" f old of pap er may be bent into a
"V" , "N" or "N" f i gure . Bruis e the stem before bending and i t
\~ ill be less ap t to break . Specimens should not protrude from
the fold of paper . Protruding parts wi1l likely have to be
r emoved when specimens are mounted . A sp ecimen may be trinuned
to re duce bulk and expose certain characters advantageously if
sufficient material (e.g., leaf petioles, branch bas e s , etc . ) is
l eft so that the pattern o f branching , leaf arrangement, and
other feat ures are r eadily discernible. l.Jhen pressing large
plants , mate severa l sheets rather th an a single sheet with a
crumpled mass of material . Arrange specimens in such a wa y that
some upper and s ome lm,'er s u rfaces of the leaves are exposed.
Spread f l Olve rs or inflorescences to sh m~ as ma ny surfaces or
vie\~s as possible . Sec t ion some flowers longitudinally and
press flat to e xhibit the inner parts and thereby red u ce the
need f or dissect ion of the f ini shed specimens.
396 18

Excess ively bulky and fleshy parts such as stems and fruit s
may be split and both parts included. Fruits should be sectioned
\"hen possible in such a Hay that hoth transverse and longitudinal
sections are included. Succulent plants (e.g., cacti) may be
split and the fleshy inner parts removed. Some collectors prefer
dipping these plants in boiling \"ater. l£: lol or benzene before
pressing. Salt may be applied to cut surfaces to hasten drying.
Each sheet should bear a collector's number Hhich refers to notes
in the collector's field notebook. mleo preparing a s eries of
sheets for parts of one specimen, number each sheet with the col-
lection number and the numbe r of parts (e.g., #429. sheet 1 of 3);
duplicates should simply bear the collection number. Do not in-
clude more than one species in a single paper. Collect sufficient
material to make a full sheet but avoid crowding or overlapping of
specimens in the sp ecimen paper. See Smith ( 197 1) for an excellent
i llustrated account of specimen preparation.
B. Arranging the Press. Specimens (in the specimen paper) are placed
between two driers in the press. A ventilator is often inserted
before the next specimen paper and driers are added. For each
specimen pressed a drier unit (2 driers with ventilator betl"een
them) is added to the press. The usual sequence is ventilator,
drier, spec imen in specimen paper, drier, ventilator, drier, etc.
As the successive specimens are added and the press built, every
effort should be made to keep the press level for even distribu-
tion of pressure I"hen the press straps or weights are applied.
This will mean the use of alternate corners of the sheet fo r
bulky roots or other parts. Sheets or pads of plastic sponges
are very useful when placed over or around bulky specimens.

V. FIELD STORAGE OF SPECD-fENS

Although it is desirable to press collections inunediately, it may not

ah"ays be practical. Delicate materials should be pressed as soon as pos-
sible and other specimens properly stored. Vascula, plastic bags, or
rubb e rized bags can be used for storage if specimens are first wrapped in
moistened paper. Specimens may be kept in good shape without spoilage in
the containers if they are kept moist and are not packed too tightly.
Supersaturation I"ith water or drying out will spoil specimens. The speci-
men bags should be kept as cool as possible and a conscious effort should
be made to park a specimen-loaded vehicle in the shade on field t rips .
Upon return from the field, specimens in bags should be put into a
caldroom until the plants are pressed. Although immediate pressing is
again desirable, plants can be kept in this manner for several days.
Plastic gallon or half-gallon milk cartons or jugs make excellent
water containers for field us e . It is a good practice to wrap specimens
collected at one locality i n an uncut, numbered (with a wax pencil before
paper is soaked), double sheet of newspaper so that one end is open and
the other closed , bind Id th a rubber band, tag with a locality number
I.;hich i s entered i n the field notebook, moisten and place upright in a
bag, and then place the bag upright in the field vehicle. Moisten by
sprinkling I.;ater over the open top of specimen bags as needed. If it is
necessary to keep wet materials for long periods of time, as is neces -
sary for collectors in tropical regions, several preservatives may be
used: 1 . 2 parts commerical formaldehyde (40%):3 parts vater .
2 . 1 part fo rmaldehyde :2 parts 70% alcohol.
18 397

3 . 40-50% alcohol.
4. 1-2 % aqueous solution of Oxyquino1ine Sulfate.
Spec imens are dipped, spray ed or brushed with these solutions and encl osed
in airtight packages . For additiona l details consult Fosberg and Sachet
(1965), Lawrence (1951) , and Smith (1971).

VI . DRY ING FI ELD COLLECTIONS

Plants should be plac e d in the press and the press closed and
tight.ened. The faster the drying process the less difficulty ,~ith
mold, mildew and loss of color . Plants should be sweated in the fiel d
press f or 12- 24 hours and the press op ened . Any last arrange ment of the
specimens must be made at this time and the wet driers changed. For
exceptional results driers should be changed at least three times during
the first 48 hours. In many areas blotters may be dried in the sun (usu-
ally one hour is sufficient) . If specimens are to be dried without arti-
ficial heat, blotters should be changed daily until specimens are dry.
Automobile luggage carriers are excellent means of drying specimens pro-
vid ed ventilators with open ducts are used between the blotters .
If artificial heat is used , there should be maximum airflow through
the press . Use doublefaced corrugated cardboard or aluminum ventilators.
Never attempt to dry specimens in an oven. A metal or wooden box with
an open top which \.,till accommodate a press sideways (corrugations pointing
up) and equipped with an electric heater Hith a fan makes an excellent
drying chamber . A collapsible drying frame (either a wooden box or metal
frame with canvas skirt) may be used in the f ield and a camp stove or lan-
tern used as a heat sour ce . Electr i c heating coils or light bulbs may ,
also be us ed as heat sources but a fa n should be installed either i n or
above the chamber . Special drying cabinets are sold but most lack suffi-
cient ventilation fo r proper drying of specimens .

VII . DATA TO ACCOHPANY SPECIMENS

A. Field Notes. As mentioned earlier every collector should keep a

f i e ld book. This is not simply a road log. Each species col-
lected at a given place and time shou l d be given a collection
number . The best system to use is a chronological one begin-
ning ,>'ith number one and continuing from there. Avoid elaborate
numbering systems ,.,tith prefixes and cryptic notations or abbre-
viat i ons . Do not use the same number for any other collection .
All duplicates or replicates should bear the same collection or
collector's number. Although some abbreviations may be useful
and efficient in the field , these should be fully written out
,.,then permanent labels are made from the field notes. A speci-
men \~ithout field data (a t least locality and date) is of little
scientific value.
Data to be recorded in the field not ebook should include col-
lector's number (for reference) , exact locality , approximate alti-
tude , nature . 2f the habitat (t ype of soil, moisture conditions,
slope exposure, light conditions) , associated species, and other
pertinent information . With reference to the plant proper, record
those features "lhich Hill not be evident from the pressed speci-
men (height, branching, depth of root system, odor , etc . ) and
 398 18

those features \.Ih ich may he lost in drying ; e.g . , flower color.
Flm"er color may best be determined by using a color chart. The
more complete the field notes. the more complete the permanent
label can be and the greater the information content of the
specimen .
B. Permanent Label. The permanent label is the labe l affixed to the
mounting sheet \-lith a specimen . I nformation in c l uded in addi-
tion to the name of the plant and authority (e . g .• Clavtonia
caroliniana Nichaux) must corne from the collector's fie ld note-
book . Do not abbreviate or use symbols . Specimens are now used
throughout the ~lOrld and symbols and abbreviations are often dif-
fic ult to translate. Be specific in giving localities . If local
names are used, give some re ference to a city, major highHay, or
easily located reference point. Hinimum data for labels should
i nclude date, locality , county , state, col l ector and collection
number. Labels , unless done by offset pr i nting , should be type-
l,'ritten using a carbon r ibbon or written in longhand using india
or other permanent ink . Paper should be of high rag content--
pre ferably 100% . See sample labels in Chapter 31, page 774 .

VIII . IDENTIFICATION OF COLLECTIONS

Naterials should be identified using the appropriate manuals , floras ,

and monographs (see Chapter 30). If it is necessary to have materials
identified or verified by a specialist, one of the duplicates is sent
Hith a l abel to the specialist. Generally the specialist \"i11 keep the
specimen unless he has agreed to do othen.Jise. In the event that a dupli-
cate specimen is i dentified by someone else, the collector should enter
the plant name on the label follO\"ed by the name of the specialist and
• the date that the duplicate \-las determined : e.g . ,
Ip omopsis rubra (L . ) ~,rherry
duplicate determined by Verne Grant , 1974
Once materials have been labeled and identified, they are ready to be
mounted (see Chapter 31) .

COLLECTING LITERATURE

Davis, P. H., and V. I!. HeYHood . 1963 . P rincip l es of Angiosperm Taxonomy .

Van Nostrand Company, Inc . NeH York.
De\\lolf, G. P., Jr . 1968 . Notes on making an herbarium . Arnoldia 28: 69-111.
Fogg , J. M., Jr. 1940. Suggestions f or collectors . Rhodora 42: 145-157 .
Fosberg, F . R. , and 1'1. Sachet. 1965. Nanual for trop ical herbaria . Regnum
Vegetabi1e 39 . Utrecht.
Franks, J . \0,1. 1965. A Guide to Herbarium Practice. Handbook for }1useum
Curators , Part E, Section 3 . The Huseums Association. London .
Knudsen , J . I·i . 1972 . Collecting and Preserving Plants and Animals . Harper
and Row . New York .
LaHrence , G. H. 1'1. 1951. Taxonomy of Vascular Plants . The Hacmillan Company .
Ne\" York .
Pike , R. B. 1964. Plant pressing Hith plastic sponges. Rhodora 66: 172-176 .
Smith , C. E. , Jr . 1971. Preparing herbarium specimens of vascular plants .
Agriculture Information Bulletin No . 348. U. S . Government Printing Office .
hlashington , D. C.
 Chapter 19 . PALEOBOTANY AND SYSTE~~TICS*

Of fo u r basic ques t ions a systematic botanist can ask about a spe cies or
othe r taxon: what is i t? when did i t arise? \. . here did i t originate? and
how did it acquire its present distr ib ut ion? (see Smith, 1969), t he last three
can be an swered most convincing l y using paleobotanical data . Haterial pe rti-
nent to thes e ques tions and to the important role of paleobotany in plant
cla ssification is pres e nted belOl. . , but as these matters also relate to topics
covered else\. . here in this book , the r eade r is referred to Chapter 16 ( Geogra-
phy) and Chapt er 27 (St ructural Evolution and Phylogeny) for info rmation not
included here .

A comprehensive review of paleobotany \...-111 not be found in thi s chapter .

Ra ther, a s et of concepts and associa t ed terminology , a brief ou tl i ne of how
the plant fossil r ecord is used , a nd a series of biblio graphic r efe rences
potentia lly usef ul to bo t a n ists unfamili a r with the lit erat u re of paleobo tany
and geo l ogy are pr esented . Tertiary and Quaternary paleobotany are s tressed
because these subjects deal largely \"ith the angiospe rms , the plant gro up of
primary focus i n this book .

The oldest generally acc ep ted angiosperm fossi l s are f rom deposits
middle to l ate Ear l y Cretaceo us in age , a lthough some believe a n giosperms
or iginated l ong before this .

I . SOHE BAS IC TERHI NOLOGY AND CONCEPTS IN PALEOBOTANY

A. Plant f o ssi l s occur in sedimen tary rocks ma i nlY as :

1. Pet r ifications--thre e dimensional struct ural detail us ually He ll
pr eserved i n a ma trix of Silica , calci t e , dolomi te , or other
s ubstances .
2 . Compressions - - flat tened , often coalified plant s tructures with
variable amount of cellular organiza tion preserved; may be
removab l e i nt a ct f r om matrix us ing mineral acids .
3. Imp r essio ns - -essent i al l y no organic matter present, ex ternal
featu r es evi dent as an imprin t in the r ock; both comp ress i ons
and imp r essions leave two halves , a n obverse and a reverse .
4. Casts and molds--infi lled cavities , associat ed o r ganic mat te r
may be part l y or comp l ete l y decayed before in f illing , no
anatomical details preserved; provide dat<\ on external (and
sometimes i n te rnal), three- dimensional form .

Only slight l y compressed wood a nd plan t struct ures of other kinds

are frequent in li gni tes (brown coals) . Other gr ades of coal may con-
tain wel l - preserved b ut compressed plant str uctures .

Nonpe t r1f1ed , more or less carbonized p l ant r emains f r om Quater-

nary deposits a r .e-. often ca l led sub fossi l s .

Compounds characteristic of various metabo l ic pat hways in plant s

have been found in organic res idues ext racted f rom sedimentary rocks
of diver se ages , and as s uch they provide biochemi cal evidence of the

*Contributed by Norton G. Mi ller , The University of North Ca rolina at Chap e l

Hill .
400 19

nature and or~gl.n of life processes through time (see Eglinton &
Calvin , 1 967) .

B. Hega and microfossils. A distinction is made between megafossils and

microfossils, the former be ing seeds, fruits , leaves, wood, etc ., the latter
pollen, spores , and other small structures that req-!1.ire a compound microscope
for routine study . In North America. leaf compressions and imp re ssions are
among the most common Cretaceous and Tertiary plant megafossils . Numerous
leaf fl oras have been described from the western United States and from sites
on the Gul f and Atlantic coastal plains . Compressions of flm-lers and fru i ts
are infreq uent, although fru i ts (minus outer layers) may be abundant in lig-
nites (e . g ., the early Tertiary Brandon Lignite, Vermont) . Determination of
the spore and pollen content of rocks may provide data to support or refute
questionable identifications based on leaves .

C. Nomenc lature of fossil plants fol l ows the principles, rules, and rec-
ommendations of the International Code of Botanical Nomenclature (Sta f l eu et
a1. , 1972). A species based on fossil material may be placed in a genus ,..rith
extant species. An organ-genus is used t~hen a genus of fossil plants is
assignable to a family (either extan t or extinct ; but , if extant , not a mode rn
genus); a form-genus t~hen a genus is unassignable to a family . When propos-
ing new names , particul arly species in e x tant genera ,dth known fossil rec-
ords, both paleobotanists and neobotanists should check the standard catalogs
(e . g ., Gray Herbarium Index , Index Ke\~ensis, the catalogs of Andre\~s , Knowlton,
LaNotte , see oart B3 in General References) to avoid pub lishing illegitimate
names .
II. GEOLOGIC CONSIDERATIONS

A. Geologic time scale . That portion of earth history richest in fos -

sils comprises three Eras, the Paleozoic, Heso2oic , and Cenozoic (see Table 19-
1) . Each Era is divided into successivel y smaller units of time cal l ed Per-
iods , Epochs , and Ages . Rocks belonging to these time units comprise time-
rock units and are called Systems , Series, and Stages , res pective ly . One
speaks of Early, I-liddle, and Late Jurassic time, and of Lm..rer , Middle, and
Upper J urass ic rocks . The relative age of rocks is determined by their posi-
tion in a stratigraphic sequence . Achievement of an absolute time scale f or
geologic events depends largely on radiometric dating .

B. Stratigraphy . This is a geological discip line dealing with the distribu-

tion and age of stratified rocks and ~".ith episodes in earth history that can be
read from such rocks. A basic unit of stratigraphy is the formation. Forma-
tions have an eaSily recognized lithology and as such find wide usage in des-
criptive studies and mapping . Formational names are binomials and often in-
clude t he name of the locality at \..rhich they \<,ere first recognized. If a
formation consists of only one kind of rock, its lithology is part of the name ,
e . g ., Dakota Sandstone (Cretaceous , plant localities in Kansas, Neb raska , Ninne-
sota) . The te rm Formation is part of the name if the l ithology va r ies , e.g. ,
Fort Union Formation (Paleocene, Hidespread plant local ities, upper Great
Plains) .

C. Correlation. Determination of mutual time equivalents is called corre-

lation, and tHO uni ts occurring in d i ffe rent outcrops are correlative if they
are considered time equivalents of each other . Methods used to correlate
strata include tracing ind ividual beds, lithologic similarity , radiometr i c
dating (e . g . , lead-uranium-thorium, potassium-argon , carbon-l4 methods), and
19 401

Table 19-1. The Phanerozoic Time Scale

Age of Base in
Era Period Epoch Hi11ions of Years
Quaternary Holocene

________________________~p~1~e~i~'~t~o~c~e~n~e------------1c..~~-~2~~~~~~~
=~P~lji~O~C~e~n~e~==============:;
7
Cenozoic Hiocene 26
Tertiary Oligocene 37-38

------------~~--------~~~~---------665,------------
~Eo~c~e~n~e~--------------53-54------------
Paleocene

~~~~~__======~c~r~e~t;a~c~e~o~u~'==============================136--------
Hesozoic Jurassic
Trias 5 ic
225 _______ 190-19 5>-----------
________________

.--.---~p~e~rm~i~a~nc-----------------------------280~------------
Carbon-
________________
iferous-~Pce"n"n"'~YL1,v7"a"n~i"aen"-
Mississippian
Paleozoic "lY='=t=e=m='~~~~~==============================j345~
_ _______________
395~---------------
______ Devonian_____________________________430-440e_---------
~S~i~1~u~r~i~a~n

Ordovician
----~~~~~----------------------~c. 500'----------
Cambrian
Ages fr om : The Quarterly Journal of the Geological Societ~7gf London 1208 :
260- 262 (1964) •

comparing assemblages of contained fossils. Because of the universality of

progressive organic evolution, paleontologic data are commonly used to deter-
mine time equivalents on a wor1d-\.,ride basis .

D. Continental vs . marine deposits . Plant fossils, particularly Tertiary

leaf floras, are found predominantly in sediments deposited on continents or in
intermediate , near-shore marine environments (e.g ., estuaries). Only infre-
quently do plant megafossils occur in marine sediments , which may contain
spores, pollen, or other kinds of plant microfos sils and often abundant animal
remains . Many of the \.,rel l known Tertiary floras of the western United States
are from extinct fresh t.,tater lakes . Those I"hose sediments include diatomite
or volcanic ash often have remarkably I"ell-preserved plant remains, a l though
cellular details and cuticles may be absent. Glacial and nonglacial conti-
nental deposits of Pleistocene ag e are often rich in plant fossils and have
been extensively studied.

E. Depositional environments. This subject, o f importanc e to those re -

lating extinct f l oras and vegetation to past environments, draws heavily on
the methodology and principles of sedimentary petrology . Sediment origin
(including potential fossil origin), transport , deposition (both physical
and chemical aspects of this process), and post - depositional history may
relate to the nature of a fossil flora . For example, on both geologic and
biol ogic grounds, the Eocene Hilcox flora of the southeastern United States
is considered a strand flora . A consequence of this kind of depositional
environment is that plants of the uplands are poorly represented and a biased
sample of the total f1 or~ is present .

Further information on topics covered in A-E is found in Dunbar and Rodgers

(1957) , Kumme1 (1970) , and Pettijohn (1957).

F. Paleogeography. The geographical bearings of continents through time,

position and elevation of now eroded mountain chains, and distribution of
 402 19

epicontinental (epeiric) seas are kinds of paleogeograph ic al data use ful to

plant systematists concerned I,ith possihle geographic isolation, migration
routes, origin of floristic provinces, a nd related topics . Exp lanation s fo r
phytogeographic pattern s often incorporate paleogeographic al data .
1 . Land bridges . A general revi eu of the land bridge concept and its use
in floristic botany is provid ed by Van Steenis (1962 ), with special
refe r e nce to his extensive studies of tJ;:9pical pla nt s . Unfortun-
ately thi s report is marred by the author.'s re j e c tio n of continen-
tal dr if t. Land bridees fal l into tHO genera l c a t egories .
a. Isthmian links, e . g . , Central America [development of the Isthmus
of Panama near the end of the Pliocene ended the isolation o f
North and South America that had prevai led earlier in the Ceno-
zoic (see Hallam , 1972)], Beringia [s ee c.vidence summarized in
Hopkins ( 1967) for late Tert iary and Quaternary land connec-
t ions betHeen Siberia and Alaska) .
b . Insular stepping stones , e . g . , the various islands of the South
Paci fi c Ocean.
2. Contin ental dri ft . Postulation of nm" sunken l and connect ion s between
contin e nts to explain d istribution patterns in f lowering plants has
to some extent been r endered obsolete by the general acceptance of
the Theory of P l ate Tectonics Ivhich provides a mechanism for con-
tinental drift (see DeHey, 1972) . I!mvever certain authors
believe that angio sp erms evolved too recently fo r th e ir
present distr i hution to have been seriously affected by continen-
tal drift . Geologic data on the time and d irec tion of cont inental
movelllent s and hm,' pre sent con tinental masses originally fitted to-
gether are still accumulating , but some j"lorkers accept the exis-
tence of a s i ngle pre-drift super-continent, Panp,aea ( Diet z &
, Holden, 1970) , while other s tHO super-co n tinents , Gon dlvana (Sou th
America, Afri.ca , I ndi a, Antarctica , Australia ) and Laura sia (North
Amer i ca, Eurasia) . According to the Diet ::: and Holden model some
of the major events affect ing Nel-l \,forld paleogeography include :
a . Breakup o f Pangaea (beginnin g in Hidd1e Tria ssic time) .
b . Lauras ia split al"ay fro m GondHana (by late Triassic t i me).
c . North Atlantic basin \"c11 developed b ut North America and Eurasia
still attached via Greenland, initiat ion of South Atlantic by
rifting (by Late Jurass ic time) .
d. Hid e ning of South and North Atlant ic, rifting i nitiated bet\,'een
Greenland and Eurasia (by late Gretaceous time) .
e . Separation of North America , Greenland , and Eurasi a (during
Cenozoic time).

The breakup of GondHana has been used b y Jardine and HcKenzie (1972). Raven
and Axelrod (1972), and Schuster (1972) to explain dist ri bution patterns of
disjunct biota among land masses o f the ~;outhern Hemisphere .

The f ollm"ing refer ences contain use ful paleogeograph ic in f ormation for
various parts of the Hor1d : Harring ton (1962) , Khudo ley and Heye rhoff (1 972) ,
Schuchert (1955), Termier and Te rmi er (1960) .

III . smm HAYS FOSSIL PLANTS ARE USED IN BOTANICAL STUD IES
A. Mornho1og ica1 research. I nvestigations on both exte rnal and internal
structure of plant fossils are commonly unde rtaken . Apart from providing a
wealth of descriptive in fo rmation, s uch studies :
1- Establ ish and document evolutionary trends .
2. Permit recognition and circumscription of extinc t plants or plant
19 403

groups , e . g . , the Pteridospermae (seed ferns) by F . 11 . Oliver and

D. l! . Scott , Rhyn i a by R. Kidston and W. H. Land , the Progymno-
spermopsida (probable ancestral stock of many gymnospermous
groups) by C. B. Beck .
3. Enable interpretation of problematic st r uctures fo und in extant pl ants,
e.g . • clarification of conifer cone s t ructure (see Florin , 1955) .

Hany curren t concepts in plant morphology have been strongly influenced by

paleobotanical data .

B. Plant classification . Data obtainable through study of fossil plants

sho uld b e used extensively i n the formulation of a phylogenetic system of plant
classif ication, a major goal of systematic botany . Proposed phylogenies . how-
ever, ge nerally are based on o t her . usually less sat i sfactory criteria because
anteceden t s of IOOst extant taxa r ema in unkno ....rn or appear to be absent from the
fossil r ecord . This may be true even in r,roups that have a reasonably good
record . For example, though the main concern of t h i s book , the flm'Jering
p l ants , t;have the most extensive fOl'isil history of any group of comparab l e
rank . fossil angiosperms have probably had the l east i nfluence of a ny fossil
plants (except some of the IOHer plants) on classification of their living
representatives . The greater part of this record consists of vegetative
organs (leaves , wood) which are yet of little use and have in face been r e-
jected as a basis for eithe r fossil or living angiosperm classification "
(Chesters. 1964, p . 283) . Nevertheless . the fossil record provides othe r-
wise unobtainab le info rmation that will continue to influence classification.
pa r t i c ular l y the defini t ion and r efinement of higher taxonomic catego ries.
Some examples illustrating the influence of paleobo t anical research on classi-
fication f olloH (see Delevor yas , 1964. 1969).
1. Recognition of the Lycopsida and Pteropsida ( inc!. ferns and seed
plants) as opposed to the Pteridophyta and Spermatophyta of
earlier systems (by E . C. Jeffrey) .
2 . Separation of the Sphenopsida from the Lycopsida (by D. H. Scott ) .
3 . Re-evaluation of the relation ships of gynmospermous plants indicating
that as a formal concept the "Gymnospermae" is unnatural (see
Florin, 1955) .
4 . Par titioning the hete r ogeneous Psilophytales into three morphologi-
cally distinct groups of Devonian land plants, the Rhyniophytina ,
Zosterophyllophytina. and Trimerophytina (by II . P . Banks) .

c . Paleoecol ogy . Pl an t fossils are common ly used in paleoecologic al

resear ch , the ~oal of which is "to reconstruct an accurate picture of the
life and life conditions of t he past by inferences derived through a study
of modern biota that are comparable with fossil ones" (Cain , 1944 , p . 30) .
Acceptance of uniformitarianism, which includes the concept that "the present
is a key to the past, " is fundamental to the paleoecological method . As one
progresses hack in time , the more tentative paleoeco l ogical conclus ions be-
come . Reasons for this i n clude an increase Idth time i n the numbe r of extinct
taxa represented in the fossil record and the prob l em of dealing with races
uhich may have had differen t ecological tolerances tha n those c urre n tly l iving .

D. Floras and vege·tat1on . Studies of Tertiary and Quate rnary flo ras (a
taxonomic listing of species) and vegetation (combinations and abund ance of
species in a flora , many i n clude physiognomy also) have gone hand in hand,
but they are distinct con cep t s that involve a somewhat different research
approach and consequently yield different kinds of data.
404 19

1. Tertiary and Qua ternary floras (paleo floristics) . The identification

of spec i es (and other taxa) in widespread. isochronous deposits
al l ows maps to be pr e pared whi ch summarize d is tribution patterns
through time . Maps of this kind may be use ful in explaining
presen t ranges of taxa, pos tula ting center s of or igin and /o r dis -
persal , paths of migration , and r elated topics. In the No rthern
Hemisphere a Tertiary fossil r ecord has helped establish the
relict d is tributional stat us of numerou s genera including : Acer ,
Carpinus , Castanea, Cercidiphyllum, Fagus, Nyssa (Figure 19 -~
Platanus, Jug l ans . and many other woody angiosperms; and t he coni-
fe r gene r a (see Ferguson , 1967) Heta sequoia . Sequoia, Taxod ium,
and Tsuga to name a few . I den tifica t ions of bo t h po lle n and mega-
fossil s are us ed. Leopold and MacGinitie (1972) have presented
detailed synops es of th e pas t and present distribution of Platy-
carya, Pterocarya, and Engelhardtia (J u glandaceae) and Pachysan-
dra (Buxaceae) .

Figure 19-1 . Presen t and Tert ia r y distribution of Nyss a . Ranges of the

five extant species a re mapped . (From Hood, 1972 .)

Species accurately i dentified also provide a basis fo r the

recognit ion of flori stic boundaries and ev i dence that floristic
provin ces (and consequently vegetat ion types) have not been sta-
tionary through time . For example , southward l atitudina l d is-
pla cement of tundra and bor e al f loras dur i ng late phases of t he
Pleistocene glac i ation h as been documented by pollen and mega-
foss il identificat i ons in both Europe and North Ame ri ca . Deter-
mination of lineages (phy l ads ) that may have bearing on f loristic
history as \~ell as providing data on the history of s peciation
within gene r a depend on accurately identified species. The
occurrence of fossil material of an ex t ant species beyond its
present range has climatic impl ications , a topic t ha t will be
explored fur t her belO\L
2. Vegeta tion . Recogn it ion of a vege tation type or types from a fossil
flora in vol ves not only identifying species but also a quantita-
tive a ssessment of the ahun dance of these species i n a fossil
assemb lage . This allows dominance to be establishe~ and provides
a basis for recognizing communiti es more or les s eq uivalen t to
those i n exis tence today. Because plant mega fO SSi l s generally
can be transported for only sho rt distances and remain i denti-
fiable, they may be l ess useful i n deciphering aspects of
19 405

vegetational h i story than pollen and spores \.,Ihich often become widely
disper sed and buried in perfect condit i on and thus ser ve to document
both local and more distant communities .
Paleovegetation nups based on data from numerous individual
studies have been prepared for various times dur i ng the Tertiary and
Quaternary and for various regions. Two types of maps are generally
atte mpted : those showing the entire composition of the vegetation
(using pie-diagrams , hist o grams , etc . ) and those in Hh ich the fre-
Quency and d i stribution of species are presented as a series of maps ,
one map per species .
a. Tertiary vegetation . The Geoflora concept. Three major long-lived
un i ts of vegetation (Ceofloras) have traditionally been recognized
(Chaney , 1947 , 1959) in studies of plant fossils in North American
Tertiary deposits, which Hhile abundant in the Hestern part o f
this con tinent are rare north of the glacial boundary and to a
lesse r extent elsewhere in the East. The supposed migration of
these units through time and space has found Hide application in
explaining phytogeographic patterns in diverse plant groups .
1) Arcto-Tertiary Geof1ora . A temperate vegetation that occupied
high latit ude northern land masses around the tvorld early i n
Tertiary t i me . Its migration southHard in response to cooling
and drying , and in North America its extinction i n midconti -
nent regions (a probable result of decreased prec i pitat i on)
and restriction to the moister coasta l areas has been postu-
lated . During these times certain genera were e l imi nated from
all or various parts of North America but survived at p l aces
in Eurasia where they occur at present .
2) Ne otropical-Tertiary Geofl ora. A warm temperate to subtropical
vegetation occurring south of the Arcto-Te r tiary area i n mid -
and 1m" latitudes early in the Tertiary . In western North
America its migration southHard occurred during Oligocene and
Niocene times and by the Pliocene on l y a few members of this
geoflora persisted north of Nexico and in southern Florida .
It s present region of survival includes Nexico , Central
America , northern South America , and the Caribbean Islands .
3) Nadro-Tertiary Geoflora . Vegetation types included in this concept
are live oak-conifer t"oodland . chaparral , arid subtropic scrub,
p l ains (desert) grassland, and subdesert to desert (Axelrod ,
1958) . This geoflora appears to have originated from sub-
tropical to ,,,arm temperate groups in southHestern North Amer i ca
(where it now occurs) in response to an increase in area ex-
posed to dry climate .
The concept of an early Tertiar y (Eocene) , high latitude
Areta-Terti ary Geoflora has been rejecte d by 1-,10 1£e (1969 ,
1972, and references cited therein) who points out that tem-
per ate plant assemblages from this region, originally thought
to be from Eocene deposits , are actually Niocene and Pliocene
i n age . Eocene and Paleocene (and some Oligocene) floras in
Alaska (and elsewhere in the far north) a r e ma i nl y trop i c.al
and subt~-opica1 and are not temperate in character . The pos -
tulated stability and duration of geoflo r as has also been
questioned from a more theoretical standpoint , viz .• "It is
e xtremely improbable from a genetic and physiologi cal v i ew-
point that many lineages coul d have remained in association
 406 19

throughout the Tert iary; that is, that a given vegetational

type remained floristically unchanged ." (Wolfe, 1969)
It is of interest that many early Tertiary plant assem-
blages from western North America have an overal l pal eotropi-
cal affinity, while those from southeastern North America are
predominantly neotropical (see LeopoJd & HacGinitie, 1972) .
b. Quaternary vegetation. In North America paleobotanical studies of
Pleistocene and early Holocene vegetation have mainly involved
the use of pollen and spores for paleoecological purposes,
although seeds, seed-like fruits. and wood at times have pro-
vided important supplemental information. The common use of
radiocarbon dating (age determinations of organic material up to
30-40 , 000 years old , sometimes older) has permitted rather pre-
cise conclusions to be made about the development and composi-
tion of vegetation types recognized in this period of time. A
brief synopsis of some pertinent concepts as they relate to North
American studies follO\</s :
1) The Pleistocene is divided into four glacials (Nebraskan, Kansan,
Illinoian , IHsconsin) and four interglacial s (Aftonian , Yar-
mouth, Sangamon , the present). Glacials are divided into
stades (ice advanCing) and interstades (ice retreating) .
Both interglacial and interstadial deposits have yielded Sig-
nificant assemblages of fossil plants. The late phases of the
Hisconsin are often informally called fu ll-glacial (active
glaciation) and late-glacial (general ice retreat) . The Holo-
cene (post-glacial) began approximately 10 , 000 years ago
according to many workers .
2) An assemblage of pollen grains from one stratigraphic level is
'. called a pollen spectrum and series of spectra comprise pollen
diagrams . Numerical values for various taxa in a spectrum are
generally relative frequencies (percentages) calculated using
as the percentage base the total number of arboreal pollen
(AP) and nonarboreal pollen (NAP) at each level with that of
aquatics and spores of pteridophytes excluded. Prior to the
early 1960 ' s it was customary to exclude all NAP from the per-
centage base even if it was tabulated.
3) A few diagrams based on absolute pollen inf l ux (in grains/cm. 2 /yr.)
have appeared in recent years, and because s uch data are freed
of constraints imposed by percentage calculations , influx dia-
grams have allowed important new interpretations to be made .
4) Comparison of fossil pollen spectra with surface poll en assemblages
in an attempt to identify extant analogs of past vegetation
types is at present a frequently used methodology . Various
statistical techniques are employed in making correlations .
5) In North America more is knotm of extinct vegetation late in the
Quaternary than earlier, and Wright ' s revie\.,. (1971) is a
recent synopsis of this topic. Huch pertinent information
\.,.ill also be found in various INQUA volumes (see Wright &
Frey, 1965). The following points pertain to the sequence in
glaciated eastern and mid-continent North America (see Figure
19-2) .
a) Based on megafossils and pollen , late - glacial tundra (zone T
of New England zonation; zone designations vary from
region to region , pollen assemblage zones named and
treated in accordance with the Code of Stratigraphic
 19 W7

Nomenclature [see American Commission on Stratigraphic Nomencla-

ture , 1961] have come into current use) has been recognized in
Massachusetts . Michigan , Minnesota , and elsewhere .
b) Zone A forests rich in spruce (and I-.lith jack and/or red pine in
the East) replaced the tundra at sites where i t is recognized
or alternately is the oldest vegetation type at others .
c) The ab r upt end of the spruce zone according to pollen analysts
marks the beginning of the postglacial and zone B. The
species of trees replacing spruce differed between the upp e r
Micl,,,est and the East I.ith white pine and birch being important
in New England , white pine , birch . and oak in parts of Net-.!
Yor k State , Oak in Ohio. and birch and alder in Minnesota.
d) Later in postglacial time deciduous or deciduous-coniferous for-
ests of variable composition come into dominance. The first
of these , as r ecognized at Ne\" England sites , ~lith oak and
hemlock predominating (zon e C-l) is followed by oak and
hickory (zone C-2) and by oak and chestnut ( zone C-3) . Zone
C-2 has been traditionally interpreted as a ~larm/dry interval
between 4000 and 1500 years ago . The composition of zone C
forests shows considerable regional variat i on .

Vegeta t ion types recognized south of the glacial boundary in the East and
elsewhere in North America are summarized in the references cited above .

E. Climate. Because present vegetational formations and c limate have

been shown to be intimately r elated , climates of the past can be inferred
once former vegetat i on types and their analogs are determined . Paleoclimato-
logical research is multidisciplinary with i mportant data contributed by bot-
anists , geochemists , geologists, meteoro l ogists , oceanographers , and others .

J J~!i !! J ~~ ::1.
n.. ,. ,
~ li
,-"'Iii'"
n ~"' " ~ 1'is1 !
iI i
ii m H i !i liim ~diH
' '
: i
:: U H,
"
postglacial

late-glacial

Figure 19-2 . Lowest segment of relative frequency diagram from Allenberg Bog
(Cattaraugus County , N~w York) showing late- and early postglacial pollen
sequence and zonation. I t ems to the right of the sums column \"ere omitted
from the percentage base. Po llen from trees is graphed at left; percen-
tages from nontrees are bet ween the sums column and the cumulative AP-NAP
curve . (From N. G. Hiller. Net" Yo rk State Huseum and Science Service Bull e-
tin 420 , 1973.)
408 19

Morphological and anatomical studies of fossil plants can sometimes lead

to conclusions about past climates . For example , grm. . th rings and cycles of
leaf scars are often taken to indicate periods fa vorable for plant growth
alternating Hith ones not favorable.
1. Tertiary climates. Two methodologies involving fossil plants (pri-
marily leaf floras) are conunonly used to.. detennine Tertiary cli -
mates and climatic change (see Wolfe , 1971).
a . Floristic analysis. Estimating various temperature parameters
(incl . mean annual temperature and range of temperature)
based on the assumption that thermal requirements of extant
genera recognized in a fossil flora have not changed . This
involves linking the occurrence of a genus to some part of
an extant vegetational formation and this part to climate .
Accurate identif ications are essential.
b. Analysis of foliar physiognomy . Relating the percent of species
with ent ire leaf margins i n a fossil flora (ident ifications
need not be made) to comparable data assembled for extant
vegetation types and us ing the identified vegetation type to
infer climate. It is not possible to determine precise tem-
perature regimes by this method . Numbers of species with
drip tips and percent lianous taxa may provide supplemental
data . The following percentages of species Hith entire
leaves have been determined for the indicated extant mesic
vegetation types (l~olfe, 1969) :

76+%: Tropical rain forest

57-75% : Paratropical rain forest
40-5 6% , Subtropical rain fo rest
10-35%: Temperate forest

Tertiary temperature fluctuations infer red by this method in-

clude : "cooling during the Paleocene, maximum l.Jarmth in the
later Eocene, a strong decline in th e Oligocene, and Harming
in the l-liddle Miocene" (Wolfe, 1971).
2. Quaternary climates. Pollen analysts reconstruct Quaternary climates
by a several step process which generally involves determinat ion
of vegetation typ e from assemblages of fossil pollen, finding
extant analogs, and inferring past climatic parameters (usually
temperature and precipitation) from those de emed characteristic
of the reg i on occupied by the anal og. Individual species some-
times are us e d as indicators if present ranges can be causually
r e lated to climate and if former distributions are thoroughly doc-
umented for short spans of time. Although knowledge of Quaternary
climates in at least glaciated portions of North America is in a
state o f flux, with evidence that changes in climate in one region
Here not in phase with changes in another , there is general agree-
ment that an important change took place at about 10,000 years
ago (synchronous disappearance of spruce forest over a wide area)
and that the succeeding postglacial was marked by climatic amel-
ioration (at least during what has been termed the hypsitherma1) .
General climatic deterioration (cooler, wetter) during the past
1500 years is recognized by some .
U 4M

IV . SOME LABORATORY TECHNIQUES

A few methods used to study fossil plants , especially those of Tertiary

and Quaternary age, are summarized here. Paleobotanical techniques used pri-
marily in morpho l ogical and anatomical studies of Carboniferous plants are not
presented. Nany of these are reviewed in Andrews (1961) and Darrah (1960).
Various chapters in Kummel and Raup (1965) also contain helpful information.

A. Reference collections . A complete, thoroughly authenticated and

vouchered collection of extant plants or parts of plants obviously should be
used as a basis for identifying fossils suspected of belonging or being closely
related to extant species. Such a collection can take many forms . Pollen
anal ysts take pains to assemble extensive series of prepar ed pollen slides,
usually several for nearly every species in a flora. Paleobotanists working
with seeds or seed-like fruits have collections of these structures for com-
parative purposes. It is essential that a voucher from which pollen , seeds,
etc . were taken be placed in an herbarium so that identifications can be docu-
mented at a l ater date if necessary .

Both the number of proposed nel.. taxa and the accuracy of identification
of Tertiary l eaves have been criticized, and to remedy t he situation, a new
look at l eaf architecture in extant plants has recently been undertaken (see
Hickey , 1973). One outcome of this is the collection of cleared l eaves , pri-
marily of woody dicots , being assembled at the }!enlo Park, California , office
of the U. S . Geo l ogical Survey through the efforts of J . A. Wolfe . When com-
plete it is anticipated that the co l lection will contain l eaves from about
4500 genera and over 10 , 000 species . The collection is open to study by botan-
ists not associated I .. ith the Survey and is vouchered by specimens in herbaria .

B. Megafossil techniques.
1. Transfers . These are used with compressions and are particularly
useful if the embedded side of a leaf or other structure needs
to be revealed . The specimen is glued to a microscope slide
and the matrix surrounding the fossil i s dissolved in acid
(usually hydrofluoric) after the slide has been coated with
{.. ax. The specimen is then clear ed (i f necessary) and mounted
f or microscopic study .
2 . Cuticl e isolations. Several techniques are in use. That of
Dilcher (1960) follows :

-scrape cut i cle off rock i nto container o f 5% sodium

hypochlorite solution
-bleach (time variable), wash several times {.. ith water after
bleaching
- s tain (e . g ., 2% aqueous safran i n)
- a l cohol dehydration , xylene
-mount on slides .

Comparative cutisles from herbarium specimens may be prepared in the

same way.

A procedure for clearing leaves from extant species is found in

Chapter 7 (p. 186) .
410 19

C. Hic ro fossl1 techniq ues (pollen a n d s pores) . The object is to o b tain

a more or l ess pure res i due of microfossils from samples under investigation
and t he techniq ue employed varies depending on sediment l ithology. If cal-
cium carbonate is present, expos u re to h ydr och l oric acid is o n e of the f irst
steps; if the sample is s i l i ceous , hydrof l uoric acid is used. Heavy minerals
are removed by gravity separation using one of several liquids whose specific
gravity can be adjusted (e . g . , ZnC12 . bromoform) , Extraneous organic matter
i s removed b y sievin g follO\~ ing treatment in a solution of weak alkali (peat) .
Cellulose i s dissolved during acetolysis (see chapter 8) . Coals and we ll
indurated sedimentary rocks generally require special treatment . Palynologi-
cal techn i que s are outlined in Brown (1960), Faegri and Iversen ( 1964) , and
Kummel and Raup (1965) .

GENERAL GEOLOGICAL REFERENCES

The fo l lowi ng sources of paleobotanical and geological data have been

selected to guide botanists unfamiliar with t hese s ubjects to infor mation of
po t e n tial usef ulness in research . The emphasis is North American . Those
tracing the geological history of a genus or family will f i nd the catal ogs
l i sted of great value .

A. Geology .

1. Glossary .

Gary, H., R. HcAfee, Jr . , and C. L. Wolf (eds.) . 1972 . Glossar y of

geology . xiv + 805 pp. + A-I to A-52. American Geological Insti-
tute . Hashington , D. C.

2. General Bibliographie s .

Long , H. K. 1971 . A Bibliography of Earth Science Bib l iographies .

19 pp . American Geological Institute. tolashington, D. C. [List-
i ngs by state .]
t~ard , D. C. 1967 . Geologic Reference Sources . University of Colo-
rado Studies Earth Science 5 . x i i + 114 pp . [A bibliographical
guide to the literature of geol ogy , a general section , listings for
subdisciplines of geology and for political divisions of the world . I

3. Bibliographies- -North American.

Nickles , J . H. 1923. Geologic Literature on North America 1785- 1918 .

Pt . I (Bibliography) . United Sta tes Geological Survey Bulletin
74 6 ; Pt . 2 (Index ) . Ibid . 747 (1924). [Index organized by subject
under political regio~tate . etc . }.]
1931. Bibliography of North Amer i can Geology 1919-1928 .
Ibid. 823.
Thorn, E. H. 1944 . Bibliography of North American geology 1929-1939 ,
Pt . 1 (Bibliography) , Pt . 2 (Index) , Ibid. 937 .
King , R. R., et a l . 1957 . Bibliography of North American Geology
1940-1949, Pt . I (Bibliography ), Pt. 2 (Index) . Ibid. 1049 (in 2
vol s.) . [Further issues of the bibliography can be found in the
followin g U. S . Geological Survey Bulletins: 985 (for 1950) , 1025
(1951) , 1035 (1952-53), 1054 (1954), 1065 (1955), 1075 (1956) , 1095
(1957) , 1115 (1958) , 1145 (1959). [1195 (1950-59) J. 1196 (1960) ,
19 411

1197 (1961) , 1232 (1962), 1233 (1963), 1234 (1964), 1235 (1965),
1266 (1966) , 1267 (1967) , 1268 (1968) , and 1269 (1969) .J See
also "Bibliography and Index of Geology" in section 4 below .

4. Bib liographies --primarily other than North American.

Geological Socie t y of Ame ri ca . 1934-68 . Bibliography and Index of

Geol ogy Exclusive of North America . Vol . 1 (for 1933)--Vol. 32 .
{Geological literature by author (with an index organized by general
subjects and by topics under count ri es in all but last 2 volumes
which follow the format continued in the "Bibliography and I ndex
of Geology " (q . v . ); short abstracts included in all but the ear-
liest volumes. ]
1969-. Bibli og r aphy and I ndex of Geology . Vol. 33- .
[Continues t he "Bibliography and Index of Geo l ogy Exclusive of
North Ame r ica"; world - \-lide cove r age, issued monthly; literature
citations with des c ripto r abstrac t s arranged in 21 i nterest
fie l ds (incl . pa l eobotany), also subject and author indices . ]

5. I ndices to Geologic Names in North America .

Keroher , G. C. , and others . 1966. Lexicon of Geologic Names of the
United States for 1936-1960 . 3 vols. United States Geological
Survey Bulletin 1200 . ["A compilation of the geologic names
(i . e ., geologic formations and other units of stratigraphic
classification and nomenclature) of the United States, its pos -
sessions , t he Trust Te rritory of the Pacific Islands, and the
Panama Canal Zone ."]
l,'ilmarth, 1'1. G. 1938 . Lexicon of Geologic Names of the United
States (including Al aska) . 2 vals . Ibid. 896 .

B. Paleobotany .

1. Treatise
-
Boureau, E. (ed . ) . 1964- . Traite de paleobotanique. Hasson et
Ci e . Paris . [Vo l s. II--Bryophyta , Ps ilophyta, Lycophyta (1964).
IlI--Sphenophyta . Noeggerathiophyta (1964) , I V (Fasc . l) --Fi li.,..
cophyta (1970) have appeared; to be comple t ed in ni ne volumes . ]

2. Bibliography .
-
Boureau, E. (ed.). 1956- . World report on palaeobotany. I .
Regnum Vegetabile 7; II . Ibid . 11 (1958) ; III . Ibid . 19
(1960 ); IV . Jbid . 24 (196"'2')';V . Ibid . 35 (1964); vi. Ibid .
42 (1966); VII . Ibid . 57 (1968); VIII. Ibid . 78 (1971) ; IX .
Ibid . 89 (1973). [Alphabetical listing of publications by
author , the follmdng headings are used in a ll except Vo l s.
IV and V: General , Pre-Carboniferous , Carboniferous a nd
Permian , Hesozoic, and Tertiary and Quaternary . ]
412 19

3. Catalogs.

Andrews, H. N., Jr. 1970 . Index of Generic Names of Fossil Plants ,

1820-1965. United St a tes Geological Survey Bulle tin 1 300. 354 pp.
[Alphab etical catalog of generic names of fossil plants based on
th e U. S. Geological Survey's "Compend i utif ' I"ndex" (q. v.) and having
the same coverage; r eference s to original sources includ ed.]
Jongmans, H. J ., a nd S. J. Dijkstra . 1913- . Fossilium catalogus II.
Pl antae . Pt . 1 (Lycopod iales)- . Publisher varies . G. Feller ,
Neubra ndenburg ; W. Junk, Berl i n or's Gravenhage. [P t . 78 (Psi-
lophyta ) issued in 1970 ; ser ies of greatest usefulness to stu-
dents of l ycopods, sphenophy t es , fern s , se ed-fe rns , and cycads;
trea tment s of the fo llowing angiosperm fami lies have app eared:
Juglandaceae (19 15), Bet ulaceae (1916), Ulmaceae (1922), Sapinda-
ceae (1928), Anacardiaceae (1935), Cornaceae (1938), Vitaceae
1939), Rosaceae (194 2) , Symplocaceae (19 50), Cel astraceae (1955);
one pa rt treating dicot wood issued (19 31 ).]
Knowlton, F . H. 1898 . A Ca talo gue of the Cretaceous and Tertiary
Plants of North America . Unit e d States Geological Survey Bulletin
15 2 . 247 pp .
1919. Ca t alogue of Hesozo ic and Cenozoic Pl a nts of North
America. Ib id . 696 . 815 pp. [ Supplemen ted by: LaHotte, R. S.,
Supplemen t to "Ca talog ue of Hesozoic and Cenozoic Plan ts of North
America ," 1919-1937. Ibid. 924. 330 pp . (1944).]
LaNotte , R. S. 1952 . Cata logue of the Cenozoic Plants of Nor t h
America through 1950. Geo lo gical Soc i ety America Memoir 51. 381
pp. [Alphabetical index of fossil pl ants (pollen , spor es , and dia-
toms excluded) by genus and species f r om Paleocene through end of
Pleistocene, ages and complete bibl i ographic citations given .)
U. S. Geological Survey . Compendium Index of Paleobo tany . [An exten-
sive (ca . 150,000 entries), up -t o-da te fi l e to original sources by
genus and specie s (diatoms, pollen and spores, and many other types
of plant mic r ofossils exc l uded), world -~ i d e coverage , al l ages ; fo r
a discussion of t he usef ulness of this index , see R. H. Eyde,
Brittonia 24 : 111-116 (1972); no t published but available for con-
sult ation at the Paleobotany Library of t he U. S. Geologica l Sur-
vey , U. S. Nationa l Huseum, I~ashin gton, D. C. 20240; mail requests
for i nformation from the index are Q2! honor ed.]

C. Palynology.

1. Bibl i og r aphy .

Van Campo , M. , a nd C. Nillerand (eds . ) . Bib liogr aph ie sup p l ~ment 11

Pollen et Spor es . [Pr esently an annua l supplement to the journa l
Pollen et Spo r es , papers liste d and indexed, the first e ntry i n
Vol. 1: 92 (1959) , ent r y 12,986 i n Vol. 13 (1971) and continuing .]

2. Catalogs. Before using po llen and spore reports workers are urged to
read J . M. Schopf ' s analysis (see bel ow) of the s tate of sys tema tics
and nomenclature in pre-Quaternary palynology .

Po tonie, R. 1956 . Synopsis der Ga ttun gen der Sporae disp ersae. I.
Teil: Sporites . Beihefte Geologisches Jahrbuch 23 . 103 pp . .!l ill.;
I I . Teil : Spori t es (Nacht rage) , Saccites , Al etes , Praecolpates,
19 413

Polyplicates, Nonocolpates. Ibid . 31. 114 pp. 11~ , (1958);

III . Teil : Nachtrage Sporites, Fortsetzung Pollenites mit General-
register zu Teil I-III. Ibid . 39. 189 pp. 2~. (1960); IV. Teil :
Nachtrage zu allen Gruppen (Turmae). Ibid. 72. 244 pp. 12~.
(1966). [A valuable catalogue of spores found isolated in sedimen-
tary rocks. extensive nomenclatural information and original litera-
ture citations are included; genera arranged in an artificial supra-
generic classification that employs categories not sanctioned by the
I.C.B.N . ]
Potonie. R. 1962. Synopsis der Sporae in situ. Die Sporen der fos-
silen Fruktifikationen (Thallophyta bis Gymnospermophyta) im
natlirlichen System und im Vergleich mit den Sporae dispersae .
Beihefte Geologisches Jah rbuch 52. 204 pp . 19 ~.
Poton ie, R. 1967. Versuch der Einordnung der fossilen Sporae dis-
persae in das phylogenetische System der Pflanz enfamilie n.
Forschungsberichte Landes Nordrhein-Hestfalen 1761. 310 pp. [Part
I , Thallophyta to Gnetales, Part II, Angiospermae ; a synopsis of
records of spore and pollen types in extinct and extant plant
groups. ]
Schopf, J. M. 1969. Systematics and nomenclature in palynology, pp.
49-77 . I n: R. H. Tschudy and R. A. Scott (eds.). Aspects of
Palynology . vii + 510 pp. Hiley-Interscience. Ne\~ York. [See
especially pp . 49-51.]
Traverse, A., and H. Spackman (Current eds.) . 1957-. Catalog of
Fossil Spores and Po llen. Pennsylvania State University . Univer-
sity Park , Pennsylvania . Vol. 1. 182 sheets + index, looseleaf.
[Vol. 35 (1972) has appeared; republication of original descrip-
tions and , if available , illus trations of new pollen and spore
types fo llowing a standardized format; three indices have appeared
(every 10th volume), authorship , stratigraphic and geologic terms,
and taxonomic names are indexed.]

PALEOBOTANICAL LITERATURE CITED

American Commission on Stratigraphic Nomenclature . 1961. Code of strati-

graphic nomenclature. Bulletin of the Association of Petroleum Geolo-
gists 45: 645-665 .
Andrews, H. N. , Jr . 1961. Studies in Paleobo tany . xii + 487 pp. John
Hiley & Sons . New York. [Chapter 18.]
Axelrod , D. A. 1958. The evolution of the Hadro-Tertiary Geoflora. The
Botanical Revie\i 24 : 433-509.
Brown, C. A. 1960 . Palynological Techniques . vi + 188 pp. Pub!. by
author. Baton Rouge , Louis iana .
Cain , S. A. 1944 . Foundations of Plant Geography. x iv + 556 pp. Harper
/5. Brothers . New York.
Chaney, R. H. 1947 . Tertiary centers and migration routes. Ecological
Nonographs 17 : 139-14.,8.
1959. Composition and interpretation, pp. 1-134 . In: Miocene
Floras of the Columbia Plateau. Carnegie Institution of Hashington Pub-
lication 61 7. viii + 237 pp. 44~.
Chesters , K. 1. H. 1964 . Fossil plant taxonomy, pp. 239-297. In: H. B.
Turrill (ed.) . Vistas in Botany . Vol. 4. viii + 314 pp. Macmillan.
New York.
414 19

Darrah , W. C. 1960 . Principles of Paleobotany . vii + 295 pp. Ronald

Press . New York . [Chapter 3 .]
Delevoryas , T. 1964 . The role of palaeobotany in vascular plant classifica-
tion, pp . 29-36 . I n: V. H. Heywood and J . HeNeill (eds.). Phenetic and
Phy logenet i c Classification. xi + 164 pp. The Systemat ics As soc iation .
London. '"
1969. Paleobo tany, phylogeny, and a natural system of classifica-
tion. Taxon 18: 204-212 .
Dewey, J . F . 1972 . Plate tectonics . Sc ient ific American 226(5): 56-68.
Dietz, R. S. , and J. C. Holden. 1970. The breakup of Pangaea . Scienti fic
American 223(4): 30-41.
Dil cher. D. L. 1960 . Cuticular analysis of Eocene leaves of Oeo tea
obtusifolia. American Journal of Botany 50 : 1-8 .
Dunb ar, C. D., and J . Rodgers . 1957 . Principles of Stratigraphy . xii + 356
pp . John Wiley & Sons . New York.
Eg1inton. G.• and M. Calvin . 1967. Chemical fossils . Scientific Amer ican
216(1), 32-43 .
Faegri. K., and J. Ive r sen . 1964 . Textbook of Pollen Analysis (2nd Rev .
Ed .) . 237 pp. Hunksgaard . Copenhagen.
Ferguson, D. K. 1967 . On the phytogeography of Coniferales in European Ceno-
zoic. Palaeogeography. Palaeoclimatology . Palaeoecology 3: 73-110 .
Florin , R. 1955. The sys temati cs of the gymnosperms, pp . 323- 403 . In : A
Century of Progress in the Natural Sciences, 1853-1953. x + 807 pp . Cali-
fornia Academy of Sciences . San Francisco.
Hallam , A. 19 72 . Cont inental drift and the fossil record. Scient ific Ameri-
can 227(5) : 56-66.
Harrington, H. J . 1962 . Paleogeographic developmen t of South America. Bulle-
tin of the American Association of Petroleum Geologists 46 : 1773-1814.
Hickey , L. 1973. Classification of the archit ecture of dicotyledonous
leaves . American Journal of Botany 60: 17-33.
Hopkins , D. M. (ed . ). 1967. The Bering Land Bridge . xiii + 495 pp . Stan-
ford Univers i ty Press. Stanford , California .
Jardine , N.• and D. HcKen zie . 1972. Continen tal drift and t he dispersal and
evolution of organisms. Nature 235: 20-24 . [Paleogeography of Southern
Hemis phere continent s includin g reconstructions of their positions during
the Cenozoic. 1
Khudoley, K. M •• and A. A. Neyerhoff. 1972 . Paleogeography and geological
history of Greater Antilles . Geological Society of America Nemoir 1 29.
xv + 199 pp.
Kummel, B. 1970 . His tory of the Ear t h. An Introduct ion to Hi s torical Geol-
ogy (Ed. 2). xix + 707 pp . W. H. Freeman. San Francisco .
Kummel , B. , and D. Raup (eds . ) . 1965. Handbook of Paleontological Tech-
niques . xiii + 852 pp . ~,I. II . Freeman. San Francisco .
Leopold, E. B., and H. D. HacGinitie. 1972 . Development and affinities of
Tertiary flora s in the Rocky Nountains , pp . 147-200. In: A. Graham (ed.).
Floris tics and Pal eofloristics of Asia and Eastern North America . xii +
278 pp . Elsevier Publishing Company. Amsterdam.
Pettijohn , F . J. 1957. Sed imentar y Rocks (ed . 2). xvi + 718 pp . Harper &
Brothers . New Yo"c k.
Raven, P . H., and D. t . Axelrod . 1972 . Plate tectonics and Australasian
paleobiogeography . Science 176 : 1379-1 386.
Schuchert , C. 1955. Atlas of paleogeographic maps of North America . xi +
177 pp. John \.,Jiley & Sons . New York.
19 415

Schuster, R. M. 1972. Cont inen tal movemen t s , "Wallace ' s Line " a nd Indo-
malayan-Australasian dispersal of land plants: Some e clectic concepts.
The Botanical Review 38 : 3-86.
Smith , A. C. 1969. Systema tics and appreciation of reality. Taxon 18: 5- 13.
Stafleu, F. A. et al. (eds . ) . 1972 . I nt e r na t ional code of botanical nomen-
c l ature . 426 pp . Inte rnati onal Associ a tion for Plant Taxonomy . Utrech t.
Termier , H" and G. Termier . 1960. Atlas de paleogeographie . 99 pp. Masson
& Cie . Paris . [~.;rorld-wide cove r age but pre-drift cont i nental configura-
tions not given . ]
Wol fe , J . A. 1969 . Neogene floristic and vegetat i onal history of the Pacific
Northwest . MadrOnO 20 : 83-110 .
Wolfe. J . A. 1971. Tertiary climatic fluctuations and method s of analysis
of Tertiary floras. Palae ogeography , Pa l aeoc l imatol ogy , Pa l aeoecology
9, 27- 57.
1972 . An i nt erpretat ion of Al askan Tertiary floras, pp . 201-233 .
In: A. Graham (ed .). Floristics and Paleofloristics of Asia and Eastern
North America . xii + 278 pp. Elsevier Publi shing Company. Ams terdam .
Wood , C. E., J r . 1972 [1973]. Horphology and phytogeography: The classical
approach to the st udy of disjunctions . Anna l s of the Hissouri Bot anical
Garden 59 : 107-124. {O ther papers in this i ssue of the Annal s also t r ea t
dis junct ions. ]
Wright, H. E., Jr. 1971 . Late Quaternary vegetational history of Nor th
America, pp . 425 - 464 . In : K. K. Tur e kian (eds . ) . The Late Cenozoic
Gl acia l Ages. xii + 606 pp. Yale Uni versity Press . New Ilave n, Connecti-
cut.
Wright, H. E., Jr ., and D. G. Frey (eds . ) . 1965 . The Quat ernary of the
United St a t es. x + 922 pp . Princeton University Press . Prince ton , New
Jersey. [The Pr oceedings of the VII Congress of the In terna t ional Assoc ia-
tion for Quaternary Research , for which the ''''right and Frey vol ume was pre-
pared, have been published in six teen volumes. Nearly all of these contain
information pertinent to botanical studies, but of sp ec ial value is Vol . 7,
Quaternary Paleoecology , E. J . Cus hin g and H. E. Wright, Jr. (eds.) , Yale
University Pr ess . 19 67. ]
Van St eenis , C. G. G. J . 1962. The land- bridge theory in botany with parti-
cul ar reference to trop i cal plants . Biumea 11 : 235-372 .

...
416 Figure 19- 3
CON T I N ENTA L DRIFT

2
,
! The 200,000 , 000
year migration
and formation of
the continental
4 land masses from
3 the presumed
breaking of one
.
• 5 or two large
7 land masses,
Laurasia (1 & 2)
JI 6 and Gondwana

t
(3-7). into :
1 . North America
2. Eurasia
3. South America
4 . Africa
5. I ndia
2 6. Antarctica
7 . Australia

4
3 J

/ Iq , --

6
 Chapter 20 . DATA ACCUHULATION AND ANALYSIS

Knowledge i s acquaintance with facts; information is knowledge derived

from observation , read i ng or instruction and consists of either facts or data ,
or both . Data (datum , singular) can be either act ual facts ( " hard data ") or
assumptions (based on knOldedge) that are used , often toge ther, to make infer-
ences which may result in an hypothesis or , with more evidence , a theory. In
systematic botany , original obs ervat i on leads to the acc umulation of new facts
and thus increases o ur supply of data and our knOldedge about the plants actu-
ally observed. These observa t ion, or fac t s , are often extrapolated in various
ways (e . g .• statistic ally. philosophically) to make assumptions about evolu-
tionary relationships , and probable evolutionary pathways, that have a direct
bearing upon the taxonomic rank (Le . , classifica tion ) of a particular plant
or group of pl ants. Data accumulation thus involves bringing together a nd
interrelating all of the new and old facts and ideas about a given topic ; data
analysis (comparative , statistical , philosophical , or other) then indicates
what l ogical conclusions , and extrapolations, mieht be made from the total
in formation available that will hopef ully provide a better biological, and thus
systema t ic . understanding of the plan ts under st udy . Both data accumulation a nd
data anal ysis may i n volve many different approaches and can be carried on at
differen t leve l s of sophist i cation and degree of exactness . However , the mere
f act that a large amo unt of dat a has been accumulated and run through a highly
complex computer analysis program does not necessarily mean that th e taxonomic
value of thl:!; data has been increased significantly or tha t previously unsus-
pected taxonomic changes are in or der . Indeed, the keen observations and
"taxonomic intuition " of many of the earlier taxonomists ( see Chapter 2)
r esulted in conclusions tha t are mo re often than not uphel d by more intense
study and the accumulation of additional data made possible by ~odern tech-
nology .

Fo r much of the world ' s flora we have very scant data of any sort ! Less
than 5% of the higher plants have been stud ied biosystematically , and thousands
of spec i es are known only f rom relat i vely meager her barium materials which , at
best , can usually provide onl y incomplete morphological data, pe rhap s a bit of
ecological da t a (if the label is complete ! ) and, \-Jith proper preparation and
observa t ion might yield a sma ll amount of anatomical or biochemical data . Addi-
tional dat a . or " eVidenc e ," of all kinds (see Chapters 5-16) are thus badly
needed to h e lp fill in the many i nt eresting a nd biologica lly important evolu-
tionary detai l s necessar y to a more complete and more accurate underst andin g
of plan t r ela tion ship s at all taxonomic levels. ,',

*Note : As data become availab l e from one set of studies after another , and as
certain relation ships are indicated by one group of data while o t her possible
relation ships may be indi ca ted by another set of data, the value o f the t axo -
nomically neu tral "d eme" terminology (see Chapter I I, Ill ; also Evolution Glos-
sary) becomes apparent . If chromosome stud ies show feat ures of similarity, or
of relationship , bet\.feen the populations or cytological "rac es" investigated
these plants can be re i~·rred t o , and considered evolutionarily , as a " cytodeme . "
The plant s of similar e cologi cal requirements form , irrespective of some of
their other a ttr ibutes , an "ecodeme"; a group of interhreed ing individuals is
a II gamo deme, " and so on . The deme concept and termi nology thus frees the taxon-
omist from the more rigid "species concept " during (and sometimes after ! ) his
r esearch and thus makes for a more realis tic and preci se concept and t rea tment
of evolutionary pathways and relationships .
 418 20

No mat ter how carefu lly a research project is designed to collect spe-
cific data, there will be t i mes when unexpected and inte res ting questions
arise or when previously unexpected correlations seem likely and more, or
additional type s of, data are needed. TIle o f ten routine and mechanical accu-
mulation of raw data may represent up to 95 percent of the research time on
a part icular project and i ts impor tance cannot be oMerrated because without
"good" (or "valid") data there can be no valid conclusions , no interesting
synthesis of informat ion and no sound basis for any further research on the
g iven subject . The old saying that "good research is 95 percent perspira-
tion and 5 percent inspiration" still holds true . There fore it is of utmost
importance that every effo rt be made to insure that the data-gathering phase
of any research project be done as carefully and meticuously as possible to
insure the validity of the ult imate conclusions.

Section A. DATA ACCUHULATION

I. ORGANIZATION

The effective and efficient collection and use of data depend in l arge
part upon good organization of the research project, facility \.,rith the many
field and laboratory techn iq ues and procedures necessary for the collection,
preparation , and analysis of the plant mate rial , and a knm.,rledge of methods
of data presentation that will enable your results to be easily revie,.,red by
others . To this end a preliminary survey of the li terature and herbarium
material , coupled ( if possible) with f ield study , should precede any major
taxonomic research effort . Such a preliminary s urvey (,.;hich may ,.,rell take
a yea r!) should lead to :

1. A rathe r precise definition of the problem and some idea as to its

pract i cality in terms of the time and resources available .
2. A l ist of specific information needed and the reasons for its need .
Also , the production of a suitable data sheet for ease of record
keeping on an ind iv idual plant basis should have evolved from
the preparation of this list (see samp le data sheets at end of
chapter) .
3. A concept of particular me thods and approaches , either direct or
indirect, field or experimental, necessary to get the information
needed .
4. Definite figu r es as to the number of populat ions to be studied, the
number of samples needed, and the realist i cally optimum sample
size needed to ge t valid data.

Hany new items that may appear on the list mentioned i n #2 above (and the
appropriate methods concerning them) have been covered in previous chapters
pertinent to the various topics . Some of the additional descriptive as "'ell
as experimental approaches to data accumulation through current biosystematic
research are given in the fo11m.,ring sections of this chapter along "'ith brief
notes on the methods involved.

In the collection of biological data of any sort it is very important

that the material used for any comparisons or correlations be comparable and
that adequate documentation , in the form of herbarium specimens, as ",'ell as
photographs. dra"'ings. data sheets and other types of evidence, be pro-
vided. In the matt er of comparable material the state of the material (fresh,
dried , or preserved) is an important fa ctor : measurements of fresh flowers or
20 419

leaves, for examp l e , may not be comparable \-lith measu r e men ts from dried mate-
rial because of the shrinkage ; and color , scen t and even tex t ure are easily
changed by different treatments . Comparable materi a l also means c omparable i n
a deve l opmental sense--mature structures should be used Hhenever possible . A
mass sample of leave s , for example, might consist of one l eaf f rom a particu-
lar node on 100 p l ant s i n t h e popul ation - - not just 100 l eaves . Here again, it
i s i mportant to know how much wi t hin-plant variation exists before estab lish-
i ng a proposed samp l e size and sampUng method .

II . DATA ACCUNm.ATION

1 . t.J"here does one get data? Fo r the systematist t here are five possible
sources of informa t ion: the fie l d, t he herbarium , the library , the garden, and
t he laboratory . It is important t o r ealize t he value--and s hortcomings- - of
i nfonnation f r om each source . The library (see Chapter 30) may be t he source
of an idea for a resear ch problem as one peruses a long lis t o f synonyms in a
monograph or notes in an at las a peculiar distribution pattern for a particu-
lar plant or notes in a journal arti cle some unexp lained and striking s imilar-
ity in fo r m, chromosome number, chemical content, ecology o r some other aspect
of a particular species. Fur t her checking th r ough various bi ological abstracts
and indices may turn up more bits a n d pieces of related info r mation th a t can
e ither answer the que stion s rais ed or can add to the in t e rest of an emb r yon i c
res e arch project . If questions still remain , or if certain kinds of n ew ques -
tio ns arise , the next place to check is the herbar ium . liere one can qui ckly
get information on many aspects of general morphology of a given k i nd of plant
and , if th e label data are complet e , some informat ion on ecology , phenology ,
and dist ribu tion . Obv iously the more specimens. their deg ree of comple teness .
and the wide r the geographic a rea they represent . the more i nformation one can
ge t f rom a prel iminary look at the herbarium material.

Most of the early taxonomists were excellent naturalists and keen observ-
ers . Their detailed descriptions o ften l eave little room for improve ment a n d
their analytical mi nd s of t en put any anomalous situation in to r ea son ab le bio-
logical perspective. Today , with more basic knowledge, bet t er communication ,
and better t echnology , i t is poss ible to add a new dimenS i on, but r a r ely a new
con cept or explanation , to the many interesting systema t ic problems t hat con-
f ront us. Change does not necessarily mean improvement . To paraphras e Robert
Woodson , whose work on the Asc lepiadaceae s panned a profess ional l ifetime , no
systematist should either a dd to, or s ubtract f r om , the work o f another with -
out first pu tting in an equal amount of st udy on the organism or organisms
invo l ved ! This , then. usually means one must leave the library and the her-
ba rium and go into the fie ld to observe first hand and to ga t her specific
ma t erials for f urthe r labo ratory , garden or herbarium study . Or, i f the
problem first occurred in the course of field . herbarium or garden work , the
library , and then the laboratory . \'lOuld be the next phases of any preliminar y
investigat ions .

Unless prope rly organized well ahead of time fiel d work (see Cha pter 18)
can be eno r mo us l y \.,tastef ul ..o f time, energy and re sea r c h f unds . All t oo of t e n
seve ral hours or even days of trave l to reach a particular co l lecting sit e are
essentially was ted because al l of the necessa r y material and data were not
taken and a se cond trip must be mad e--pe rhaps a year later. when the plants
are again in bloom or fruit . Be prep are d ! Have a check- list of \.lhat mate rial
you need. or think you may need and . if the populati on si ze will pe rmit, ge t
plenty of material. IJith in r easonable limits , i t is much better to have
 420 20

material that you don't use on a research project than to need material and not
have it available !

Before the field trip begins go over the check-l ist and be sure that you
have all of the collecting equipment and materials necessary to do the job .
A fe\" tips :

a. Always label every plant , vial, or collection of any sort in the

fie l d as soon as collected and also enter the data in a-Yield
notebook. Decide ahead of time the codes and numbers you \-Jant
to use and i f individual plants or specimens are to be tagged
have the numbered tags prepared ahead of time and strung on a
\.,rire in such a way (in reverse order) that they can be taken off
in order from l--onward . This \.,rill prevent errors in numbering
plants as well as save valuable field time . Labels can also be
punched or coded with colored plastic tape if necessary to keep
material from different species separate . Do not trust any
"permanent" ink; use Ivax, or soft lead , pencilSfor making field
tags.
b. Living material carefully potted in the field and Ivatered with liquid
fertilizer (20-20-20, premixed and in gallon plastic jugs) sur-
vives much better than material carelessly dug and planted later,
perhaps after drying out a bit in a hot car; many plants, dug with
a small ball of earth around their roots , can be wrapped in news-
paper, labeled, and put in a waxed cardboard 1/2 gallon milk con-
tainer Ivhere they will "ride" for several days (if given a cup of
liquid fert i lizer) and can be potted up later with ease .
c. !lave colleeting vials and bottles already filled to the proper limit
(1/2-3/4 full) with the appropriate preservative o r fixative . Use
only good glass vials and jars with ~ caps . Heat will pop
out corks and plastic stoppers and the material Ivill be lost .
Some plastic vials melt when certain fixatives or other chemicals
are put in them. Use a soft lead (ltl or 2) pencil to write col-
lection data on small (1/2" x 2 " ) strips of bond paper to slip
into each jar or vial of material. I-lax pencil marks on glass , or
the plastic tops , is not permanent! A vial of precut label slips
will save time in the field .
d. Haterial to be used for any sort of comparison should be comparable
in developmental stage and/or position on the plant.
e. Although material for herbarium specimens can often be transported
for a day or tHO and then held another day or so in a cold room
Hhen prop e rly wrapped and wet down in the field, cytological
material , some chromatography samples , pollen samples and other
items for specialized study should be fixed , preserved, or other-
Hise prepared in the field at the time of collecting i f at all pos -
sible in order to keep from introducing another variab1e--deteri-
oration--into the study .

2. ~lat are the types of data? Data can be either quantitative (objec-
tive or measurable hy a uniform standard) or qualitative (subjective or the
result, in part , of a judgment). Quantitative data is more precise , easier to
Ivork with statistically, easier to duplicate or compare , and often easier to
understand or expla i n than qualitative data . HOivever , for some characters it
is sometimes much easier , and just as effective, to " score" a variable even if
it is somewhat suhjective rather than get an absolute or objective measurement
20 421

or count. Such a situation occurs when there is a great range of variability

(as in some leaf sizes or shapes or in certain degrees of trichome density)
and actual measurement ~~ould be either ve ry dif ficult or meaningless, or both.
Here"classes " of the character state are established, perhaps by making a few
actual measurements or counts, and s ub sequent samples are merely compared
visually with the repre sentative material, photograph or dra\"ing typical of
each class and then assigned a score or class value. l!m.Jever. the more objec-
tive the data the more objective the research !

3 . Samp l e size. The type of investigation will determine to some degr ee

the amount and kind of material needed . For a preliminary survey of variation
in a character, or a group of characters, a single plant (\.Jhich can often be
a herbarium specimen) from each of a dozen or so localities may be sufficient.
If the research is to be a more detailed study of variation in a particular
character over the geographic range of a species , sufficient samples should be
taken to give relatively uniform representation of populations from all parts
of the range of the species . Depending on the frequency of occurrence of the
plants within their range, the total size of the range, and the variability
found in the preliminary studies, anywhere f r om five to one hundred o r more
popul ations might be samp led, and each sample might consist of material from
one to five separate plants o r might consist of a single mass sample f rom each
population representing five, ten, or even 100 plants if the population size
and variation pattern so indicate. In general, the larger the sample the more
representative the sample and less extrapolation is necessary in final inter-
pretations of the resu lts . On the other hand a realistic time budget must also
be established for a given research project. Here the objec tives of the
r esearch , the degree of reliability desired . and the technology availab le for
use in the data gathe r ing as well as data analysis stages must be considered.
For example, i f a study involving chromosome numbers of plants from 100 popu-
lations is being undertaken, each collection ,dll require, perhaps. up to four
hours of laboratory time to get a chromosome count. If five such counts are
desired from each population the laboratory time required for the study in-
creases accordingly--to 2000 hours, or essentially a year ' s \.Jork at 40 hours
per week. And there is no machine to do da t a gathering . By contrast, a pol-
len size st udy invo lving measurement of 100 pollen grains from each of five
plants from each of 100 popula tions might involve only about 15 minutes time
per slide for a total of 125 hours of laboratory time to get and tabulate
measurements of 50 ,000 pollen grains. By use of ma ss samples , with the pollen
from the five plants of each population mixed on one slide . the laboratory
time can be reduced to about 25 hours . If a "Coulter Counter" is availab le (and
if i t is properly calibrated and its operation und erstood ) , a single mass sam-
p l e of 50.000 pollen grains can be measured in les s than t wo minutes ; o r the
100 mass samples coul d be run i n an hour or so. Here technology makes the
data gathering relatively fast and easy .

Often i t is possible to determine the realistic sample size needed from

preliminary observations or from the early data on population analysis. When
observations or test s show the pattern of variation i n a character to have
more or less s tabiliz ed . ~dditional measurements or observations are not essen-
tial even t hough they may be highly desirable . The same can be said for the
number of character s being investigated: i f preliminary studies and measure-
ments indicate that data from five characters or ten cha racters will support
the same concl usi ons as data from 30 or 40 characters it is not necess ary to
run a full analysis on the additional characters . 110wever , character co rre la-
tions , or lack of such cor relation s , cannot under any circumstances , be assumed
422 20

pas t the hypothesis stage of the r esearch but must be shown to exist , through
adequate preliminary obse rvat ion and documentation , before they have any value
in a syst ematic study .

Sample size will also be governed by whether or not the data being
gathe r ed are to be used for predictive purposes (where the final answer is
not known for certain) or descriptive purposes (in which previous obser vation
has s et the general descriptive limits) . In the f irst case l arge random sam-
pl es are called for wherea s in the second case smaller "r epresentative" sam-
ples are quite adequate and much more realistic to work with in eve r y r espect.
In th e selection of the representative sampl e consideration should be given
to proportionate representation of the different naturally occurring st at es
of the character under inve stigation; however , rare extreme forms might be
noted separately rather than included in the general data if such inclu sion
would distort the var i ation pa ttern i n an abnormal way. Because of such
occasio nal distortions i t is well to keep in mind that the statistical sig-
nificance of a given set of data is not always directly related to the bio-
logical significance of the data.

4 . Changes in variation ranges and pat t e rns . Ranges of variation and

var i a tion patterns us ual l y change as t he source of the samp l e material changes
from t he individual pl ant to the population and from a singl e population to a
se r ies of popUlations or the species as a whole . However , it is often valuable
to know something of the amount and kind of variation at each of the t h ree
l eve ls in establishing sample size and in evaluating the data on variation
within and between species and populations. (See the exercise on po l len size
variation, Section B, III A- 3 , further on . The same type of analys i s of
variation- -individual , population , species- - can, and should, be done for any
character being studied . ) If, for example , a s ingle plan t , or a s ingle popu-
lation shO\"s mos t of the range of variability i n a particular char acter , this
hi gh degree of uniformi t y , perhaps the r esul t of founder effect or a rather
strong selective force, \"ould make continued or extensive \-lOrk with this char-
acter seem uncalled for i n any study of variation .
III . DATA FRON GENERAL VARIATION STUDIES
All organisms vary to some extent and all variation occu r s within indivi-
dual genetic limits. If these genetic limits are very broad for a particular
cha ra cter , the primary phenotypic control of this character is most likely
environmental; if the gene tic limits are very narrow , primary phenotypic con-
tro l wi l l likely be gene tic . Since both evolu tionary and taxonomic studi es
are based on individual and population variation , i t is often importan t to
know t he source, and range , of t he variability o f a character in a population
or a group of populations . A number of methods exist to get t his in f ormation;
all involve careful planning , careful observa tion and adequate r ecord keeping
to amass sufficient data on comparable material to warrant later valid con-
clusions . In addition , appropriate experimental design is necessary to pro-
duce results which can help explain, or furthe r substantiate , the conclusion s .
Among s uch experimental app r oaches to morphologica l and developmental var ia-
tion are t he following:

A. ReCiproca l transplants. Move pl ant A from habitat A to the same hole in

habitat B from which plant B was taken ; transplan t plant B to habitat
A in the hole from which plant A was taken. If the transplants retain
their characteristics over several sea sons their diffe r ences are
genetic ; however , if transplant A now comes to resemble transp lant B.
20 423

and vice ve rsa, th e ir phenotypic variation is primarily t he re su lt of

their environments .

B. Uniform environment . This is often a more practical method to study var-

iation since plant s f rom a number of population s ar e all bro ught to-
ge the r i n an experimental garden at one location in an essentially
"un iform" envi ronment which mayor may not be optimum for one or more
of t he plants. Phenotypic changes, over several seasons , indi cate the
r ange of each genot ype (plant) and thus whe the r or no t the variation is
primarily environmental or genetic.

C. Growth chamber stud ie s . Where the equipment is avai lable t hese provide,
often in a rel a tively sho rt time, rather precise da ta on the tota l
range of phenotyp i c e xpress i on o f a se ries o f plan t samples unde r dif-
fe rent environments . Growth chamb e r s tudie s , when ap propriate l y de-
signed, can show clearl y specific cause and effect co rrel ation s between
di ffe rent combina tions of environmental fac tors (s uch a s temperat ure
and light intensity) and morphologica l and phYSio logical variation , the
l atte r easily evidenced by changed developmental re spons es . ~Hth vari-
ously programmed growth chamb ers it is also possib l e to s how the poten-
tial range of variability of a plant under environmental conditions that
do not currently ex is t in its natural hab i tat (e . g ., hi gher or lower
temperat ures, longer or sho rt er pho toperiods) and whi ch may be indica-
tive of prior adaptations to prior environment s .

D. Analysis of mass sampl es . It is often possible t o determin e the ac tual

and poten tial range of variation of a character \dthin a sp ecies , or a
serie s of population s , found i n more or l ess simi lar habit a t s, by an
analysi s of ma ss samples. For s tat is tical r e l evance , such samples
should invo l ve mea suremen ts or observations on 15 to 100 o r more pla nts
(depe ndin g on popula tion si ze) and from each of three or more popula-
tions . If the plants or samples are c ollec ted completely at random a
larger sample will be requi red to g i ve a picture of to tal variation
than would be the case if "representative " samples (of median and bo t h
extreme fo rms) were sel ected for measurement . Or, to put it another
way: a much larger sample is needed for " predic tive s t atis tics" t han
"desc riptive s tatistics . " It: might a l so be well to rememb e r that
be ca use of certain genetic and evo lu tionary facts (e . g . , linkage ;
polymorphism) and the ever pre sent possibi lity of samp le erro r, a
charac t er may show "statistical significance" out of proportion to i ts
"biologica l significance !"

A combination of both reciproca l transp lants and uniform envi r onment

studies (with the expe rimental plots at di f fer en t a l titudes) wer e used in
the classic stud ies of Achillea and Potentilla by Clau s en, Keck and Hies ey
( 194 8). In general , cloned material, wh i ch i nsur es gene tic uni formity , i s
the best for any studies of environmental eff e ct on phenotypic expression.
Fo r r ec iprocal t ransplants perennial s a re obviously better s uit ed than annu-
a l s; eithe r perennial s or annua l s can be used effec tively in uni fo r m ga rden
or growth chamber experiments , or in mass sample analy sis.

E. Data fr om breeding studies . Since variation patterns , and evolu tionary

rat es , are ofte n r e l a ted to the bre e ding system of a species of plants
(see Chapter 12 ), it is important t o know whether the plants of a popu-
lation norma lly reproduce by outcro ssing , se lf -pol l i nation , or apomixis .
424 20

This information can often be easily obtained by a series of observa-

tions and experiments as outlined in Chapter 12 , Section D.

F. Pollen data . Pollen size studies and pollen viability studie s (see Chap-
ter 8, Section C) can often make a valuable contr i bution to any sys-
tematic study involving actual or suspected hy.brid i zation or poly-
ploidy . Pollen data are general ly easy to gather with a minimum of
supplies . equipment , and training and provide good mate r ial for simple
statistical analys i s . The gathering and analysis of sets of pol l en
data also provides an interesting "term project" type of investigation
\.,rhich , if continued over a number of yea r s, HQuld be cumulative and
could provide a valuable block of biological information abou t certain
plants , or populations of plants, which might then be correlated with
environmental fluctuations or c hanges .

G. Chromosome data . Chromosome number, chromosome morphology and chromosome

behavior at meiosis are all important bits of biosystematic data often
necessary to an understand ing of int r a- and inter-specific evolution-
ary patlll"ays . See Chapter 10 on Cytological Evidence and Chapter 11
on Genetic Evidence for concepts and methods associated with the
va r ious aspects of chromosome data .

Section B. TREAnlENT OF VARIATION DATA

1. TilE USE OF SIMPLE STATISTICAL METHODS

Since variation is universal it is necessary for anyone ~Iorking with

biol ogical mate rial t o be able to deal effectively ~.,Iith the patterns of
variation found ,.,ithin and bet,oeen individuals , populations , and species of
organisms . This leads , in flor i stics for exampl e , to the ne c essary incl usion
of size ranges in species descri ptions (e . g ., leaves 7-18 cm long ) and , in
biosystematics , to the further inclusion of the mean, or average , and some
simple notation (the standard deviation) as to the extent of samp l e variation
around the mean (e.g . , range 7-18 cm , x
= 13.2 ± 2 . 3) . It should be noted
that the larger the standard deviation ( O' ) relat i ve to the mean the grea t er
the variability in a sample . Thus, if the mean leaf length for p l ants of
one popUlation is 13.2 cm and the standard devi ation is ± 2 . 3 cm, the plants
o f the popul ation are more uniform , as to leaf l ength , than the plants of a
s e cond popu l ation in which the leaf length range is again 7- 18 cm and the
mean 13 . 2 cm but the standard deviation is 3 . 4 . Likewise the figu r es
x'" 86 . 2 ± 9 . 3 indicates less variability (in ~"hatever is being measured)
than do the figures x = 12 . 8 ± 3.4 .

In addition to the mean there are t~w other " central tendencies " that
may be used on occasion . The se are the mode and the median . Although each
of these three terms designate a central tendency it is impo r tant to realize
that they do not f a l l at the same point of a distrib u tion graph unless the
distribution is per fe ctly synmletrical--a rare situation when dealing with
hiological T'lA.terial Hh ere distribution curves are usually skeV/ed , or asymmet i-
cal , as shmm in the example below .
 Class
mode'" 8 em Va lue Frequency fv
mean 6 . 9 em v f
median 6 em {Leaf {Tabulation (Frequency
l e ng th of number of times
12 0
in em) leaves of value)

•
11
~ 10
~ 9
8
1\ 1
2
3
each size)
1
1
2
1
2
6
~

~ 7 / 0 4 3 12
5 4 20
-
0 6 0
36
•
"',•
5
4 0
/ 6
7
6
10 70
3 / 8 12 96
" 2
1 0- /
/ 0
,
0
9
10
8
2 20
72

11 1 11
2 4 6 8 10 12 em 50 346
Graph of leaf length frequencies T;;~l-~~;h;;- -------------
sum of
(above) and the same data used to f leaves = n fv
determine the mean leaf length (right)

L fv or -x 346 or x ... 6 .9
x = 50
n

If the data on leaf length are carried through a few additional computations,
as shown on the following computation sheet, we will have additional valuable
information about the pattern of leaf l ength variation in the plant (or popu-
lation) under study : the standard error of the mean , the standard deviat i on,
and the coefficient of variability . These first two figures are normally
given along with the range and the mean and the four are often graphed fo r
ease of comparison with other , similar, sets of figures representing another
plant or another population . t-lhen so graphed the var ious items of data are
indicated as fo l low s:

Range 1-11 Cr.l

,--- d • + 2 .99-- ---,

,\ . e.~ . 0 . ~2 /
\ x = 6.;
~-
Figure 20- 1 . Graphic representation of standard statistical figures .
426 20

In some instances, as in the examp l es given belol>' for plants of several popu-
lations of three species and a hybrid of Physalis (see Hinton , Fred, 1970 ,
Brittonia 22 : 14-19 ) . the standard error of the mean is not graphed . It
tolOuld, ho\.;rever , be given in the tables of figures that support the graphic
presentation.

HET
Bt
SN HET
Bl
SN
-+
I
GR GR I
HYB Bt HYB Bt

Bt
sc
+
t
Bt
SC
VIR VIR
WE
+- WE
FR
+- FR

Ell
-+- Ell

LAN
WG
HO
+-
+- LAN
WG
HO I
RF
--+- RF I
HA
+-, a
HA

MICRONS -25
I
40
PER SQ. MM 0 , l2
, l6
, , ,
30 ,
35 , ,
45 50
,
UNDERSURFACE I

TRICHOME DENSITY - leaf GUARD CELL LENGTH

II . SYHBOLS AND DEFINITIONS

x '" mean cr '" standard devi ation

E sum of d = deviation of class from mean

v = class value V' = coeffi cient of variability

f = frequency S. E. = standard error

n = number (of ind ivid uals) P.E. probable error

Abscissa : the horizontal line of graph.

Average deviation : = Efd (not as good as 0) .

n
Class value or class center : average value for each class; if , f or example ,
a variable series of leaf-length figures are to be treated it might be
desirable, for convenience sake, to lump the leaves into size classes
(e . g ., 20-24 cm, 25-29 cm, 30-34 cm, etc . ) and then compute the mean by
use of the median class values (v) h'hich \.;rQuld be 22, 27 . 32 etc . f or the
above class ranges .
20 427

Coefficient of variability ("\I): us ed to compare amount of variability in

material measured in different units (e . g., length vs . we i ght ) . It is a
ratio of the standard deviation (0) to the mean ex) of the same population
r educed to a percentage . V = ~o_
x
Deviation of Class from l'!ean : (d) difference betl4een class value and mean.
i'jean: the o rdinary average , the sum o f the scores in a series divided by the
number of scores; x (bar x); each class value (v) is multiplied by class
frequency (0. the sum (1::) of these produc ts is divided by the number o f
individuals (n). Thus: x = l::fv
n
Hedian : the middle score in a series of scores after t hey have been arranged
in order froCi IOI-/est to highest. There are as many scores above it as
below it . It is not the middle of the range (except in rare cas es ,>,here
t he tHO coincide) .
Hade : is the scale value at which the frequency curve is highest.
Ordinate : the vertical line of graph.
Probable error (P . E .): same as standard error except a is multiplied by the
constant .67 45 and the mean of another random sample ,>,ould fall Hithin the
limit set by the P . E. in one out of tHO cases (50-50 chance).
Range : a single indicator of variability; the extreme scores or values in a
set of observations or, by definition, the spread betHeen the largest and
smallest scores.
Standard Deviation : (0 or sigma) i nd icates the amount of variation in a sam-
ple; e.g., t\o]Q samples o f ten leaves each, might have identical means, but
one sample would range in len g th f rom 40 to 50 em Hhile the second, more
variable, ranged from 20 to 70 cm and would have a larger stan dard devia-
tion. The small er the relative value of 0 the more closely the sample is
grouped around the mean. \>1ith normal frequency d is tribution approximately
2/3 of a given sample I"ill fall Hithin the range cove r ed by one rJ on each
side of the mean; occasionally it is desirable to include a greater per-
centage of a salllple in a given set of figures and then tlW standard devia-
tions are added on each side of the mean. I n this latter case , i f the curve
is skeHed, the doubled standard deviation may exceed the actual range found
in the sample. Reference back to Figure 20-1 ,viII clearly shDl" that an
increase in the standard deviation figure from 2.99 to
5 .9 8 would extend the s t andard deviation bar beyond the Ud
actual upper range limit of 11 cm. n
Standard error (s.e . ): the range Hithin Hhich the mean of another random sam-
pIe from the same population ",auld fall in tHO cases out of three .
a (The greater the s.e . or p.e. the l ess r eliable the data . )
s.e.x n
Variance or sum o f squares: as I'las the standard deviation, the variance is an
indication of dispersal around the mean. I t is , as one name implies, the
sum of the squared deviations , or differences, fr om the mean divided by n-l,
or Ud
S= n-l ' or for example , s'" 245 = 5 .0. Note that the standard devia-
49
tion is expressed in · the same units as the mean and range but that the
variance is not; the variance, hOl"ever , is the more useful figure in many
other mathematical treatments of the data .
428 20

III. THE DETERNI NATION OF VARI ATION IN A SINGLE PLANT AND A SINGLE POPULATION

Any character of a plant is the expression of the physiological processes

of the plant acting within the limits set by its genetic constitution . Thus.
while the various parts of a single plant have identical genetic constitutions,
the physiological processes from oue part of the plant to the next are not
usually identical, and di ffe r ences in size and development result . The follow-
i n g method is designed to shm. some of the variability within one plant and
between plants of one populat i on and to show how this variability may be shown
by a simple statistical treatment involving the mean , standard deviation ,
standard error of the mean and the coefficient of variabi l ity .

1. The material for the follm.,ing Hark should all be taken from the same
plant to illustrate the range of variability ~1ithin an individual.

(a) Using a minimum of 50 leaves construct a graph sho~o,Iin g the number of

leaves (ordinate) and the number of sinuses, or serrations , or
lateral veins , etc. per leaf (abscissa) .

(b) Using the same leaves construct two bar graphs showing the variation
in: (a) the length of the lamina
(b) the width of the lamina
using at least ten class intervals .

(c) Nmo,l make a slide (see Chapter 8. Sect i on C) of the pollen from a sin-
gle floHer (or inflorescence if more practical) of the plant you
are working \-lith and use an ocular micrometer to measur e as accu-
rately as possible the large st dimension of 100 pollen grains.
Record the f requency distribution of the measurements. This can
be done in "eye-piece units" for convenience , but be sur e to
calibrate the "eye- piece units" of the ocular micrometer ~o,Iith a
stage micrometer marked off in . 01 mm units (so that your measure-
ments will be in standard metric units a nd can be compared with
data gathered by others) and convert your final statistical calcu-
lations to these standard units.

(d) Use the data you have to determine the coefficient of variability for
each of the three leaf characters and the pollen size. lfuich of
the four characters vould be most satisfactory for use in a key?
1,.fhy?

2. Now rep ea t the exercise on a populat i on basis by using equal amounts of

material from a minimum of 5 plants from a singl e population; if
material from more than 5 plants is available it will , of course ,
give a more accurate "statistical descri ption" of the population
variation .
20 429

cmn>UTATION SHEET

Terminal Leaf Length

Naterial: Hass sample , Population 1 Hagnification: IX
4 July 1970 (cent imeters)

Class Class Frequency (f) Deviation

Range Value of class
from mean
V tabulation totals fV d d2 fd 2

1 I 1 1 -6 36 36

2 i 1 2 -5 25 50

3 II 2 6 -4 16 32

4 Ii i 3 12 -3 9 27

5 l ill 4 20 -2 4 16

6 h1-i-i 6 36 -1 1 6

7 !i1i-. lilI- 10 70 0* 0 0

S 11+1 1tt1 II 12 96 +1 1 12
1~ +2
9 ,o ' II! S 72 4 32

10 'I
, , 20 +3 9 IS

11 I 1 11 +4 16 16

Hean :
x- = rEV
n
= 346 =
50
6.9
r:." = 50 UV =
3~
*NOTE: For ease of calculation of
~d2 '" 245

d 2 and Ed (above) an as~umed ~

Standard deviat i on : of 7 \,'as u sed , rather than the
actual mean of 6.9, in o rde r to
Hork "/ith ,."hole numbers. The s.e.x
a=V Ud "V'4550 .~=' . 99
n
so obtained is adequate for prelimi-
nary use. \~hen dealing with deci-
mals , calculations are generally
Standard e rror of mean : carried to one more place than
occurs in the individual base
S .e. x • 2 .9 9 measur ements .
-7-= 0.42

Coeffic ient of vari ability :

299 "" 4) . )
6.9
 ~
w
o

Impatiens, _____________________________ Population No . ___________________________

Date.________________________________ Locality_______________________________

Plant number

Spur length mm

Sac length nun

Angle of spur/sac

Corolla spot-ting

0E:ening \~idth nun

No . Seeds/capsule

No . Fruits/plant

Leaf length

Leaf width

Serrations/em

Internode diam. nun

Node diam . nun

Longest internode em
N
o
x pollen diem . .u
20 431

Data Sheet, Solanum Project

Population I! Species______________________________

Plant No . Da te Collected,_______________________

Herbarium If Co unty and 5tate'_ _____________________

Notes:

Habit Flower color

Heigh t Flol'Ter s/lnfl .

Leaf color Corolla diam.

Infl. type Lobe l eng th

Peduncle leng th _ _______________ Sepal orien t .

Leaf s hape Fruit color

W/L ratio Fruit diam .

,
Basal If . anglec-_____________ Seeds / fruit

Nargins Seed size

Pubescence Seed shape

Tr ichome densi t y________________ Concretions

Sepal length Chromosome II

S tyle type Pollen diam .

Style l e ngth Guard cell size _____________________

Anther length St em XS

S tem pubesc .
432 20

EXPERUIENTAL AND ANALYTICAL LITERATURE

I. General References

Bel l , C. R. 1967. Plant Variation and Classification. Wadsworth Pub-

lishing Company. Belmont, California .
Briggs, D. • and S . N. I,Taiters . 1969 . Plant Variation and Evolution .
HcGraH-Hill. Ne\~ York.
Clausen, J . , D. D. Keck, and I·i. Hiesey . 1948 . Experimental stud i es on
the nature of species: III. Environmental responses of climatic
races of Achillea . Carnegie Instit ution of j.,lashington Publicat i on
(1581.
Cochran, H. G. 1953. Sampling Techniques . John Hiley & Sons , I nc . New
York .
Mode, E . B. 1951. Elements of Statistics . P rentice Hall, Inc . Engle-
Hood Cliffs. New Jersey .
Postlethv.'ai t. S . N. , and N. J. Enochs. 1967 . Tachyplan ts suited to
instruction and research . Plant Science Bul l etin 13(2) : 1-5.
Sakal. R. R., and F . J . Rohlf . 1969 . Biometry . H. H. Freeman and Com-
pany . San Francisco .
Solbrig. O. T . 1970. Principles and Nethods of Plant Biosystematics .
The Hacmillan Company. New York .

Also see the references at the end of Chapter 12 on Reproductive Biology and
Systematics , Chapter 21 on Data Presentation and Documentation, Chapter 22
on Analysis o f Character Variation in SystenBtics . Chapter 23 on Numerical
Aids to Taxonomy, and Chapter 26 on Var.iation and Evolutionary Relation-
ship s .
21 433

Chapter 21. DATA PRESENTATION AND DOCUHENTATION

Although detail ed description is the usual method of presentation of much

taxonomic informat i on, nothing can take the place of accurate drawings and
clear photographs i n the presentation of morphological data, or pre cise charts
and graphs in illustrating clearly the trends shown by large amounts of tabu-
lar or statistical data. A page of line drawings costs no more to print than
a page o f printed material . Photographs , and of course color photographs ,
cost considerably more to reproduce than line drawings . Photographs should
generally not be on the same plate as line drawings . Regardless of type--
photograph or drawing-- graphic material that is not adequately labeled loses
much of its value and impact . Hhere specific detail is involved the actual
voucher or herbarium specimen associated with the figure should be given in
the legend. All data should be referab le to a pertinent standard herbarium
voucher, or other acceptab l e form of documentation (also see Chapters 18 & 22) .

I. DRAHINGS

1. Line Dral"ings. The most common form of botanical illustration is by

means of line drawings. Although such drawings may appear decep-
tively simple, good drawings r equire considerable skill . There
is more art than mechanical ability in just drawing an ink line,
or stippling a drawing . Top notch dra"IVings by a professional
artist are an essential investment in scientific accuracy .
,
Detailed draldngs ·of unfamiliar objects require more labels
than less detailed drawings of standard objects (compare Figures
21-1 and 21-2). Appropriate scales or other indications of magni-
fication or reduction are essential. Also, each figu re, plate ,
chart, graph, or table should carry a legend unless all pertinent
information needed to understand the material is included as part
of the figu r e , plate, chart, graph or table. The legends for
Figures 21-1 and 21-2 (from unpublished thesis of Drapalik, D. J.
1969, used with permission) read as follows :

Figure 21-1. Typ i cal seeds, seedlings , leaves, and fruits of the
four groups of Hatelea knO\ffi from the s outheastern USA. Each
group is represented by a vertical column . The "Odontostephana
group" is represented by!:!. baldlryniana , and each other group is
represented by its one and only species .

Figure 21- 2. Floral Structure of Matelea baldwyniana.

A. Top view shm"ing the relative positions of the gynostegium ,
corona, corolla lobes , and calyx lobes.
B. Side viel' of the gynostegium and corona I"ith tHo-fifths of
the upper corona removed .
c. Radial . ~ection of the corona, gynostegium, and carpels .
D. Abaxial view of a pollinium apparatus. The cellular
patterns of the pol linium walls are not portrayed . The
outline I.as drawn with a camera lucida .

Drawings may be effectively combined with maps to show character

variation over a geographic range. Such a p i ctorialized distribution
434 21
Fi g ure 21-1

. ,-
~I

~I

,,
,

'\\ \

\\
M. baldwyniona M. alabomensis M. pubif loro M. gonocorpo
 435

Figure 21-2

calyx lo be

ani her center

carano lobe

A sinus between carano oPP'"' ages

',pc, sSlon
. in sfIgmo apex
corpu5culum

corona appendage
corono lobe
B
corpuscu lum

in stigmatic
te rslaminol hsri
I (opening to a
camber)
lmm

stigmo
corpuscutum

stigma
t ube growth)
undersurf oce ( polh of pollen

st yle

c
carpel wall

corpusculum

corpusculum furrow
0.2 mm
oem
arca of pollen tube growth from the
pollinium

pollinium
D
436 21

Figure 21-3

o C. verticillata 2x

• 0
0
0
0

00
o 0

° 0
..®®

®
4x

•• 0 00 00
0

•• • •
•• • 00
00 o 0 0

• •
• • • • 00 • 00 0
00

• •• . .. •• ~ .
•
• •
• .'
•
' .•
•
...
•••®
• •
: ®
• C. major 2x

•
• •

"C . delphinifolia 8x

SC ~L E ,,. ""L ~! '?"

,
'C . .... < QUOC .:;. "0"0''''''

.. / "
.'

COREOPSIS DISTRIBUTION
21 437

Figu re 21-4. Xe roxed "drawing " of

a pressed plant of El cocha r is obtusa .
438 21

map for different species and chromosome races of Coreopsis are

shown in Figure 21-3 (from unpublished work of Anna M. Mueller,
used with permission). Documentation, always essential to any
taxonomic study, would he in the form of an extant (and available)
herbarium collection to correspond with each dot on the map. The
collection data for this group of specimens could be presented in
the form of a table (in an appendix, i f necessary) and should
include for each species the following minimal information in
columns, as shown:

Collector Colle ction Number Date County State Herbarium

2. Drawings from xerox copies of plant material. For those plant struc-
tures that are more or less laminar. or that can be f lattened
without undue loss of character, an extremely accurate outline
can be obtained merely by having the material xeroxed (Figure
21-4). The xerox copy can then be traced to get an outline draw-
ing, or it can be inked in to get a silhouette. This method is
especially valuable for getting comparative drawings of seedlings
and of irregular or highly divided leaves, petals, inflorescences
or other plant parts. The natural size drawings can be photo-
graphically reduced as necessary for publ i cation.

3. Camera Lucida drawings. Camera Lucida drawings, or drawings made with

any other projection device attached to a standard light micro-
scope, are both accurate and easy to produce with little or no
artistic talent. Such drawings are commonly used to illustrate
either meiotic or mitotic chromosomes (Figure 21-5 A and B).
lfuere chromosome morphology is important the somatic chromosomes
may be redrawn to scale and with the centromeres on a line, in a
stylized form known as an idiogram (Figure 21-5 C).

A c

Figure 21-5. Chromosomes of Erigeron simplex: A, meiotic; B, mitotic; C,

ideogram. [ From unpublished data of Stephen Spongberg. Used with permission ]

I I. PRESENTATION OF CHROMATOGRAPHIC DATA

Chromatographic information (see Chapt e r 13) is often presented as shown
in Figures 21-6 and 7; when the number of di f ferent i nterspecific chromato-
graphic spots is the main point of the i l lustration the spots common to all
species are not shown (Figure 21-7). I f the chromatographic data involve
 2l 439

0 ) 0 )

1
11

1 Q)CD G) 20 Ib~ ®

Oe
(0
0)
@@ ct8 @~
7

0
06
0) G G e
Physalis heteroph~ll a
8°& Physa lis lanceoJata

Figure 21-6. Drawing of chromatographs of 2 species of Physa lis showing all

of the spots obtained on each chromatograph. Equivalent numbers indicate
equivalent spots .

.j- .-j .-
.-- .--
. ~

0 0 j 1 <m>- 0
c

0
0 0 0
0
0 0 0
hybrid
•
~ v;rginiana
•
•, -P
@.

lanceolalll
"
- heterophylla

Figure 21-7. Chromatographs, showing variation in leaf extract s of three

species and one hybrid of PhysaliS , c learly indicate that Physalis l anceolata
is not, as once postulated, a hybrid between f. virginiana and f . heterophy lla.
Note that in this presentation of chromatographic data only those spo ts t hat
are unique to the plant s of a species are shown - the many pigment spo ts the
related plants may have . in common a re omitted from the chromatograph for the
sake of clarity. Compare wi t h the " total spot" drawings s hown in Figure 21-6.
[From Hinton, ''' . F. 1970 . The taxonomic Significance of PhysaliS lanceolata
(Solanaceae) in the Carolina Sandhills . Brittonia 22: 14-19 . Used with
permission . ]
 440 21

CD CD

CD

Figure 21-8. Pa ired Affinity Indices of Arctic and Alpine Erigeron. Dots on
each arm", 50% mark, outer e nd of ea ch arm '" 100% af f inity. Ea ch arm, a s
l ett e re d in graph 1, represents o ne taxon, as follot... s :

A. Erigeron uniflorus spp . eriocephal us G. E. melanocephalus

B. B.. simple x spp. simplex H. E. humi lis
C. f. simplex spp. petiolaris I. E. aureus
D. K. simplex spp . f lettii J. E. hYl2erboreus
E. E. grandiflorus. northern alpine race K. E.
F. E. grandiflorus . s outhern alpine race L. E. -----
lanatus
vagus
Graph l. Erigeron uniflorus ssp. eriocephalus
2. E. simplex ssp . simplex
J. E. simplex s sp. pet io laris
4. E. simplex ssp. flettii

[From unpublished da ta of Stephen Spongber g . Use<' Ivith permiss ion.]

21 441

a large number of spots and i f t he use of the informa t ion is to illustrate the
degree of chromatographic similarity or difference between species . a "paired
affinity index" is computed for each pair of species involved in the study (see
formula. below) and the resulting percentages are then plotted on the appro-
priate arms of a polygonal graph (see Figure 21-8),

Paired Aff ioity Index (PA) ., !s"p~o~tss!L!i~nc-"c~o~mmo~"n,-:-f::oeEr.,.!'s~p,e~c"i~e~s,;;,A,-,a.n""d--"B

total spots for A + B

Since , obvious ly, a species will show a 100% affinity with itself, the "peak"
of the polygon will shift from species to species (arm to arm) around the
graph; it is the general relationships shown on the other arms of the graph
that are of interest. Through a summation of the PA i ndices of one taxon with
each of th e others to which it is being compared a numerical expression of
chromatographic similarity (called a Gro up Af fi nity Value) can be obtained. as
shown by Ellison , et al. (1962 ) .

III. POLYGONAL GRAPHS AND HI STOGRAMS

Polygonal graphs, with up to 16 arms, each representing a character , are

a l so frequent l y used to compare graphic ally the variation patterns within or
among a se ries of populations or species . The graphs (Figure 21-10) can be com-
posed of many lines vith each line r e presenting an individua l plant, or a single
line which repres ents t he mean values for a population or a species . l-Then the
polygons so formed by the da t a for a population are placed in the proper po si-
tion on an outline map the pattern of geographic variation within a speci es is
quickly evident. A "master graph," with each arm clear ly labeled as to what
character it represent s and the uni t s of measurement clearly marked should be
given with each set of po l ygonal graphs presented.

Histograms and r egular line graphs (Figure 21- 9) are commonly used to
illustra te frequency distribution .
•

1\
12

"
/
1O

.
9

•- •-
8

D D 7
E
o
E
0 • •
2 2 5
4
/
•
3 /
2
. _/
/
• •
'- .
1213 [4
Leaf length in em . I 2 3 4 5 678910
Leaf length in em . "
Figure 21-9 . Frequency distribution of the data given in Table 20- 1 by
histogram (left) and line graph (right) .
442 21

.."," ..",,".
/

....., $C""

...".".".
" ".".
...",".".,."
."""
~'.!..!':'El
/

~ACC I N I UM Vi\CILLANS V VAC IL LANS . TENELLUM VACCIN IUM TEN(l LU M

Figure 21-10. Polygonal graphs of eight leaf characters (see center graph)
from plan ts of t,,,,o species of Vaccinium (Ericaceae) and plants of a presumed
hybrid b etween them. [From unpublished data of Sabina Nueller. Used with
permission.]

IV . HYBRID INDEX

A series of measure ments , scores, or size classes can be used to con-

struct a numerical Hybrid Index (see Table 21-1) in which the basic charac-
teristics of a group of individual plants are sco r ed on a 1-2-3 basis (or a
1-3-5 basis if a more accurate scoring of intermediates will resul t) , and
then summed for each plant . If the n on-linked cha racteristics are involved
in t he index the plants of one taxon will have a total score of 1 0 (if they
are typical fo r the taxon) , plants of a second taxon woul d have a score of
SO (if typical) and intermediate plants ,VQuld have intermediate scores . I f
a histogram is made of the Hybrid Index scores for a population the distribu-
t ion pattern will readily illustrate the composition of the population in
terms of the different variants (Figure 21-11) .

V. SCATTER DI AGRANS

If the states of a second character are used on the ordinate instead of

freq uency , the graph shoHs the relationship between the tHO characters for
the individual plants (or populations) plotted . This type of graph is called
a s catter diag ram. If , th ro ugh a pictorial code , each dot on the scatter
diagram is made to represent additional characters, as Has done by Edgar
Ande rso n (1949) we have a "Pictorialized Scatter Diagr am" Hhich presents a
considerable amount o f correlated in fo r mation (about either individual plants
in a population or individual population of a species) very effectively and
quickly (Figure 21 -1 2 ). Similar pictor ialized or coded spots can be used
with outline maps (re fer to Figure 21-3) to show the geographic d i stribution
of plan t s with particul ar sets of characteristics .
21 443

Index Index Index

Character Value 1 Value 3 Value 5
Leaf width: length ratio 0.8-1.0 0.4-0 . 7 0 .1- 0.3
Leaf pubescence glabrous puberulent tomentose
Stomate density mean per 0 . 1 mm 2 153 256 )11
Plant height 1-3 dm 4-7 dm 8- 11 dm
Flower color red pink Hhite
Sepal margin entire serrate fimbriate
Pollen size (mean) 32-36 IJ 37- 41 ]..I 42-48 )l
Style length 2-4 mm 2-8 mm 9-12 nun
Stigma shape globose ellipsoid cylindrical
Seed per capsule 1-9 10-19 20- 29

Total Index Value 10 )0 50

Taxon R Taxon S Taxon T

Table 21- 1 . Hybrid Index [From Bell. C. R. 1967 . Plant Vari ation and
Classification. t.Jads\.,rorth Publishing Company. Used with permis -
sion . ]

20 R S T

,
'"
r-
z
«
-'
a.
"-
0 IND EX VA LUES
a:
w '0
'"
:!'
::>
z "
'0
5

INDEX VA L UES

Figure 21- 11 . Hybrid Index Histogram. Two types of distribution patterns of

Hybrid Index values for 50 plants from one popul ation that would show
two entities and their presumed hybrid (top) , and var i a t ion in one entity
(bottom) . A similar graph could also represent the Hybrid Index averages
of samples from 50 diff~rent populations; the top figure would then
probably best be interpreted as representing three distinct taxa , but
the bottom figure would again represent the va r iation in a single t axon
over its geographic range with Rand T only representing the extremes in
a normal variation curve . [From Bel l, C. R. 1967. Plant Variation and
Classification. Wadsworth Publishing Company . Used with permission . ]
 444 21

mm
" o U. cornuta (cho'mogamou . How", )

• •
o U. iuncea (c!c;stog3mou. Ho .... )
"

,"
""
~
Z
t -
'h-+Jt
w
."
,
~
~
•
+t , ~+
w
•••
,
It' r-
,
,

• - ---0-

P7' lip leo\I,h

• • -.-" .
/ ;I.-...-1le<!g1n p;.1i Ieng.h ..,,!'It, 1''''1Oth
1 - 5""
° 01 _ 10. ""
0
01-1.4 eo
0
0.1 - 0.7 ""

<> 5 - )0.""

,,- ,,. 6 -3.5 ""

0 1.1 _ 1.0. ....
0
1,5 - 34 ....

0- - 0
"0.8 - 15'"

1.1 3.5-59 eu 1.6- 2 ~ ""

L---t---t---t---t----t--,r---rj---t!---t;---i16.--,\I\,--,I~,---,k\---il~---il~,mm
SPUR LENGTH

Figure 21-12. A scatter diagram of Utricularia characters. [from Kon do, K.

1972. A comparison of variabil ity i n Utricularia cornuta and Utricularia
j uncea. American Journal of Botany 59 :23 37. Used with permission .]
21 445

Utricularia Spur Length, (in mm) . (eH " chasmogamous flower,

eL cleistogamous flower) .

Popu Number
Species lation Heasured X S.D. S . E.X C.V. Range

U. cornuta U-ll 50 8. 4 0.91 0.13 10 . 9 5 . 8- 10 . 7

u-12 50 8. 9 0.78 0.11 8. 9 6.9-11.4

U- IS 50 11.3 1. 05 0 . 15 9.3 8.6-13 . 9

U-17 50 12 . 9 0 . 89 0 . 13 6. 9 10 .1-1 4.6

U- 18 50 11.0 0.85 0 . 12 7.7 8 . 6-12 . 5

U-19 50 10 . 2 0 . 79 0 . 11 7. 8 7.6- 12 . 1

U-20 50 11.6 0.63 0 . 09 5 .4 10.4-12.9

U-21 50 10 . 9 0 . 88 0.12 8.0 9 . 0-1 2.5

U-22 50 11.4 1.14 0 . 16 10 . 1 6.5- 13 . 2

juncea U--2(eH) 50 5.6 0 .9 3 0 . 10 16.6 4.0- 8 . 3

11· 0.64 0.09 40.0 0 . 5-4 . 1
(eL) 50 1.6
U--6(CH) 50 5. 3 0.89 0.12 16.8 2. 9-7.4
(eL) 50 1.8 0.64 0 . 05 20 . 0 0.7-4. 3
U-13(eH) 50 4. 6 1.00 0.14 21.9 3 . 1-7.8
(eL) 50 1.6 0.57 0 . 07 36 . 4 0 .5-4 . 1
U-14(eH) 50 6.6 0.78 0.11 12 . 0 0.5-8.4
(eL) 50 1.4 0 .71 0.07 50.0 0.7-4 . 1
U-23(eH) 50 4. 8 1.00 0 . 14 20 . 8 3. 0- 7. 8
(eL) 50 1.4 0 .64 0 .09 45 . 7 0 . 5-4.3

u- II Table 21-2 . Comparison of single

2 U-12 versus double spacing of tabular
~ u _ 15 material (above) and a graphic
C
~
o
V'I7

V-16 -
-
-i--
......., - summary of the data (left) . [ From
Kondo , K. 1972. A comparison of
o u·!9
variability in Utricu1aria cornuta
::l u- 20
u_ 21
-+-
~-i--
and Utricularia juncea. American
,- " Journal of Botany 59: 23-37. Used
mm 10 I, .. with permission . ]
v-z _ _ _ _
00~ _ o ..
,
,

- -,- -- ~
_+ _ __ ct

_~ _ C><

U-!'3 ____

_+ __ ,
~ '"

Cl

SPUR LENGTH
 446 21

VI . TABULAR MATERIAL. VI . TABULAR MATERIAL

Although tabular material is often summarized graphically in some form,

the actual tables of specific data should accompany the charts and graphs
derived from them whenever possible. perhaps as an appendix to the body of
the research paper. In the tables themselves more details of the analysis of
a char acter can be given (e.g., the coeffic ient of variability, number of
measurement, standard error of the mean) than are generally shown on graphs.
Also , the actual figures are easier to compare in detail than a r e the graphs
(compare Table 21-2 and the accompanying graph). Tables, as figures , graphs
and all other inclusions, should be fully labeled and captioned to be self-
explanatory. Also, the entries (species, populations, etc . ) in a series of
tables and graphs should always be in the same order to facilitate compari-
son .

VII . CROSSING DIAGRAMS AND PHENOGRAMS

The graphic illustration of conc epts of relationship can take many forms
from the phylogenetic " cac tus" chart of Bessey (see page 602) dealing with
the orders of angiosperms to the simple lines and branches , de t ermined by
computer, of a "corre lation phenogram" for plants within a s i ngle genus as
shown in Figure 21-13. In between these seemingly objective diagrams of
monophyletic relationships are the many admittedly speculative (to greater
or lesser degrees) evolutionary diagrams that seek to express the often
rather involved reticulate evolution found in a " species complex" or a
"polyploid complex" as shown in Figure 21- 14.
L
Crossing diagrams. which show t he relative degree of gene exchange or
hybridization possible between the members of a homogamodeme (a ser ies of
more or less i nt erfertile pl a n ts belonging to several species , varieties,
races or populations) are often used to summarize the res ults of breeding
experiments. In these diagrams the t hi ckness of the line connecting two
parent sources indicates the relative degree of interfertility between them
(Figure 21-15). The direction of the c ro ss or crosses is indicated by an
arrow on the line from the pollen par ent to the female paren t.

There is an almost infinite variety of graphic form used by individual

taxonomists to express plant relationships as indicated by the data collected
during a specific research program. Since it is not possible he r e to treat
in detail th e many combinations of forms and types of drawings, charts ,
graphs, and photographic techniques. reference should be mad e to several
curr ent volumes of the periodicals listed at the end of Chapter 22 for further
ideas on data presentation . As to t he actual preparation of the material .
there are numerous references on the methodology of both line drawing and
photography which should be consulted for the details of preparation . A
few such references are given in the following Literature List .
21 447

CORRELAT ION OTU

- 0 .3 250 -0 . 0750 0.1750 0.4250 0 . 6750 0 . 9250

1
2

5
I
8
'-
9

14

- 12
I
I 13

10

16

6
.- I
I 18

11

I 17

15

Figure 21-13 . A cor relation phenogram of 18 sets (OrU's) of plants of

Haplo papp us based o n 31 c ha r acte r s. [F r om Jackson, R. C. and T. J . Crovel lo .
1971. Brittonia 23:54-70 . Used with permission.]
448 21

E. • UNIFLORUS 2x
E. HULTENII E. MUIRII

E. GRANDIFLORUS
3x =•

m
(Jl

;;:
SIERRAN
ERIGERON .R '?
"m
r
X
U>
U>
~

(Jl
;;:
"
r
m
X

ERIGERON ?
'"
~

Figure 21 - 14 . Int errela t:i.onships Hithin the Er i geron uniflorus-simplex complex ,

[From unpublished data o f Stephen Spongberg , used with per mi ssion . 1
21 449

_ _, Mono l oke
Yosemite Jet, creek

Mather, meadow Soli Lake City

Mether, spring Centerville

5834

Bear Lake
MI. Oso
Big Collonwood Conyon
Yosemite Jet., marsh

lee Vining Canyon

A.eraqe Seed Set per Capsule"
Pacific Grove

Pescadero

Chew's Ridge 15 -28

MI. Diablo 33 - 50

59 • 100

Figure 21-15. Summary of seed set in various crosses between plants of

different populations of Mimulus guttatus. [From Vickery, Robert J .• Jr.
1959. Evolution 13: 300-310. Used wi th permission.]
450 21

DATA PRESENTATI ON AND DOCU1ffiNTATION LITERATURE

Anderson , E. 1949 . Introgressive Hybridization . John Wiley and Sons, Inc.
Net" York .
Blaker , A. A. 1965 . Photography fo r Scientific Publication. Freeman Books .
San Francisco.
Blun t , W. J . l~ . 1950 . The Art of Botanical Illustration . Collins . London.
Bracegirdle , B. 1970 . Photography for Books and Reports . David and Charl es,
Pubs. Newton Abbot , England .
Ellison . lif , L .• et a1. 1962. Methods of presentation of crude biochemical
data for systematic purposes , with particular reference to the genus Bahia
(Compositae) . American Journal of Botany 49: 599-604.
Engel, C. E. Ced.) . 1968. Photogr aphy for the Scientist . Academi c Press.
New York .
Hodgkiss. A. G. 1970 . Haps for Books and Thesis. Universe Books , Inc. New
York.
Knudsen, J. H. 1966. Biologica l Techniques : Collecting, Preserving, and
Illustrating Plants and Animal s. Harper and Row. New Yo r k.
Lloyd , B. 1935 . Handbook of Botanical Diagrams . University of London Press .
London.
Papp , C. S. 1968 . Scientific I l lustration, Theory and Practice . 1•.Jilliam C.
Brown Company . Dubuque, I owa .
Poole, L., and G. Poole . 1965. Scientists Uho Hork IHth Cameras. Dodd ,
Mead and Company, I nc. New Yor k.
Rose, G. G. (ed.). 1963. Cinemicrography in Cell Biology . Academic Pre ss.
New York .
Zweifel , F. hI. 1961. A Handbook of Biological Illustrations. The University
of Chicago Press. Chi cago.

The Consumer Harkets Divi sion of Eastman Kodak Company, Rochester , New York
14650 , publishes excellent inexpensive handbooks which cover every aspect
of photography from basic information on films, lighting , developing and
enlarging to highly technical information on spec i a l ized photographic
me t hods.
22 451

Chapter 22 . ANALYSIS OF CHARACTER VARIATION IN SYSTEHATICS*

Knowingly or not , all sys t ematists a r e engaged in the analysis of cha r ac-
ter variation. The purpose of this paper is to outline app r oaches to t he study
o f character va r iation in systematics . Perhaps this chapter wi l l be of greatest
value if it motivates us to get out of our usual "thought -r uts." Some people
have not re- evaluated their idea s , purposes , or methods of analysis sinc e their
graduate days . For example , many people st il l use Anderson ' s pictorialized
scatter diagram as the only approach to the study of character variation due
t o i ntrogression . As creatures of habi t, we tend to th ink along the path of
l east resistance . So , if one explanation is given for a set of phenomena , it
is easiest f rom that point on not to keep searching for nove l explanation s .
The same concept is true, to some degree , in methodology: while I have tried
to includ e a ll major areas of syst ematics in this review , I am aware that my
own background and interests influence the emphasis given to each area and
method. Hopefully , a gl impse at the many methods of study ing character varia-
tion u sed i n systematics today wi ll have a stimulating effect and permit the
development o f new pa th s of t ho ught and research . One cannot help being im-
pressed with the potential ben efits to both the descript i ve and explanatory
phases of systematics ~"hen its workers consider the use of the contents and
methods of each o ther ' s fields as well as those developed in ecology, other
areas of biology and areas outsid e of biology, e . g ., psychology , business)
and engineering .

I find it interesting t h at with the proposed Flora Nor t h America Program

and a growin g numb e r of projects to " computeriz e herbar ia" (see Crovel lo,
1972 for references) , botanists have forged ahead of zoologists in the use of
computers fo r information retrieval. Unfortunately , plant systematis ts ar e
far behind in the use of computers to analyze character va riation . Per hap s
this book will help to alleviate the situation .

NECESSARY PRINITIVE NOTIONS

To minimize misunderstanding , certain essential concepts , or primitive

not ions , must be agreed upon before any comparison of approaches can be made.
An object is an individual o rgan ism or a collection of organisms grouped by
some common property. Sokal & Sneath (1963) designated the object an opera-
tional taxonomic unit, i. e ., t he unit to be classified. A system is a collec-
t ion of objects . their properties, the interactions among them, and all
"environmental" forces acting upon them . A character is a property of an
object or of a system that cannot logical l y be subdivided further. This com-
pares with Sokal & Sneath's (196 3) operational definition of character . A
character state i s the part icular expression of a c haracter of a giv e n object
or of a system . For example , in a taxonomic study of wil lows , each taxospecies
may arbitrarily be designated as an object . Each has the character stamen num-
be r. The charac ter state fo r object (species) A may be one stamen; fo r object
( species) B i t may be five stamens, etc., depending on the i ndivid ua l s obse r ved
in each taxosp e cies.

The choice of objects , of systems , of characters , and of character st ates

is arbit r ary , and the choices vary with the investigator and with the purpose

*Contributed by Theodore J . Crovello , University of No tre Dame .

214 Figurp. 8- 1

POLLEN MORPHOLOGY

..----:-:1 pole

Polar view Equatorial view EQuotorial view

T r icolpa te Distal polor view Tr ica lporote Zanoc olpate
Monocolpate

Polar vIew Eq uo toria l view

Triporate Pant aporat e Zonopora te Dyad

baCUlum ge mmo

Tetrahedral li near Rhombo id al Tetragonal

tetrad tetra d tetrad te trad

Erdt mon Foeg ri

E~tose ~in e Tectum

Sexlne Eklexine

Colume llo
Exine

",," , Fool 'o~ef

Nexine

fnline
{ Nc .. nc 2

Nn lne 3
)'",.. ".,
Not recognIZed
I nline

St r uctu re of Pol l en Wall

(Modi/ied, o ller T5Ch ... d~ ond Scolt)
8 215

Verrucate. Pollen with \~art-like processes ahlays broader than high and
always higher than llim. The process is a verruca (pl . verrucae); e . g . ,
Berberis.

H. Stru cture and Stratification of {.,)'all. Concepts regarding the stratifica-

tion of th e pollen wa l l have been and stil l are in a sta te of confu-
sion. For a brief review of thi s subject the reader is referred to
the editorial, Review of Paleobotany and Palynology 10(1) : 4-37 . 1970.
The following terminology is only that in common usage.
Columella (pl. columellae) . Pillar- like elements. supporting a layer or
cro~led by a single element. Columellae may be well deve loped, moder-
ately \"ell developed , or reduced in size .
Exine . The part of a pollen wall outside the intine; is acetolysis
resistant .
Intectate. Po llen grain without a tectum or tectal areas .
Intine . The cell \"all proper, inner laye r of a pollen wa ll (removed
during acetolysis) .
Nexine . Inner united homogeneous layer of the exine .
Sexine . Outer layer of the exine .
Tectum . Outermost closed layer of the sexine . The pollen grain is
tectate .
NOTE : The exine is divided by some workers on chemical differences into
the endexine. an inner homogeneous layer, and an outer ektexine . In
order to determine exine structure the student i s encouraged to l earn
th e t echniq ue of L-O analysis (see Erdtman , 1952, 1969) .

I. Nuclear State. Pollen is shed in either the binucleate or trinucleate

stage. The phylogenetic significance of this character has been dis-
cussed by BrC\"baker (1967) .

Section B. SCANNING ELECTRON HICROSCOPE POLLEN STUDIES

Hith the scanning E . lI . the intricate surface features of pollen grains

can be studied in detail at magnification s up to 20 , OOOX and features that
cannot even be seen \"ith the light microscope can be accurately measured and
compared. The great depth of field possible (see Fig . 8- 2) and the relative
ease of preparation (many specimens can be observed in the living condit ion
without either fixation or coating) make thi s method of pollen study of spec i al
value and interes t to the taxonomist with access to a scanning electron mic r o-
scope . NOTE : The S . E . H. has also been used to advantage in studying the
minute surface features of such things as seeds , leaves , trichomes and apical
meristems. [See Falk , et a1., 1971; also Chuang , T., and L . Jleckard . 1972 .
Seed coat morphology in Cordylanthus (Scrophulariaceae) and its taxonomic
significance. American Journa l of Botany 59 : 258-265 ; and Whiffen , T., and
A. S . Tomb . 1972. The systematic significance of seed morphology in the
neotropical capsular-fruited Melastomataceae. American Journal cf Bo tany 59 :
411- 422. ]

Sect ion. C.. POLLEN SIZE AND VIABILITY STUDIES

In addition to systematic information that can often be gained from the

intricate detail of the exine or outer pollen wall as beautifully revealed by
the scanning electron microscope , the role of the po llen grain in the repro-
ductive process makes it a valuable tool in fertility studies and the relatively
determinate nature of pollen size (a characteristic shared with guard ce lls)
216 8

a b

c d

Figure 8-2 . Scann ing electron ph otomicrographs of Vernonia (Asteraceae) pollen

grains : (a) equatorial view of entire pollen grain of V. cinerea, ca . 30~ in
diameter ; (b) surface viel, of same grain at greater mag~ificat i on to 8hm" c r ests
and lacunae, each lacunae is ca . 10]1 in diameter; (c) equatorial viet" of e nt ire
pol l en grain o f Y.. . triflosculosa, ca. 32\l in diamete r; (d) surface detail o f
crests , spines and microapertures of poll en grain of V. l indheimeri , the spines
are ca . SJJ long . [From Jo nes, S . B .• Jr . 1970 . Used Hith pe rmiSSion . ]
offers some degree of as si stance in pollination biology stud ies and in the
possible detection of polyp l Oidy . For these latter t\o/O types of obse r vation
only standard liAht microscopes are necessary. Expanded pollen, meas ured
across the l argest dimension (usually th e diamet er) at the highest practical
magnification will give the most accurate in for mation in regard to si ze studies ,
and the following survey method provides good preliminary informa t ion on pollen
size varia tion \dthin one flot.;er , for one pla nt and for one o r more populations.
Th e sets of data can b e easily analyzed fo r comparison by the simple statisti -
cal me thods shm.,rn in Chapter 20 . section B; fo r pollen sli de pr epa ration see
methods 1-4 in the section below .

Hake 5 pollen s l ides using pollen from five flot"ers of ~ plant; this ,,,il l
indicate something of the range of flot"er to flower pollen size variati on
in one plant .

}take 5 pollen slides using pollen from one or more flo\;rers from each of five
different plan t s from one population (or geographic area); this will
indicate the degree of plant to plant pollen size va riation in one popu-
lation.

}take 5 po lle n slides using pollen from one or more flowers from each of
five differe nt plants from a second population (or geographic area) ;
this \"ill give a preliminary estimate of the pollen size range of the
species in the geographic area sampled .

Th e ~ method of slide preparation must be used for all fifteen slides .

If the pollen is mounted in one of t h e gl ycer ine jelly preparations ,
the cells may continue to s~lell for several days ; thus all measurements
should be done after the pollen size has stabilized and at about the
same time after the slide was prepared .

Heasure 100 pollen grains on each slide and compute the size range, mean
pollen size, the standard deviation , and standard error of the mean
for th e pollen on each slide (see Chapter 20 , section B) . Also compute
the same statis t ics for each of the three samples (of f i ve slides each)
and th e total sample of 15 s l ides . If two or more taxa are involved
similar sets of s l ides and calculations should be made for comparative
purposes .

Section D. }tETHODS OF POLLEN SLIDE PREPARATION

1. Methyl green in glycerine jelly (semi-permanent; gr ains expan ded) .

Place small amoun t of po l len in center of s lide , add a drop of alcohol
and allow to partly evaporate . A second , third , or fourth drop may be
added if neces sary. The alcohol spreads out and l eaves a ring of dis-
solved oils from the poll en . Hipe oily ring of f with cotton moistened
in alcohol . Before specimen dr ie s completely place a drop of hot
methyl-gr een glycerine jelly on the slide and stir the pollen into it
with a clean n eed le. Keep slide warm on warming plate, or by passing
over flame. Cover -with a No . 0 coverslip and heat briefly. The
methyl-green stains only the exine . If contrast is desired the ce ll
contents may be stained on the s l ide before starting the alcoho l with
a weak aqueous eosine solution ; blot off excess stain . Pollen color
is most brillian t after about 3 days ; good up to a year , then fades ,
but th e pollen can be restained .
218 8

Brandt's Glycerol Jel l y

Soak 40 gm . sheet of granulated gelat in 2-3 hrs . in cool Ivater .
Drain of f excess Hater.
Nelt hydrated gelatin in (,'ater bath.
Add 60 ml. of glycerol and 1-2 grams phenol crystals; mix thoroughly.
Filter through glass ("001 .
Bottle in 50 mI . (or 25 mI . ) containers . '-.'

To prepare stain (for 50 mI . portion) add 2 . 5 mI . sa turated solution of

methyl green in 50% ethanol, and also 2 ml. of a sat urated solution of
Phloxine (or rose Hengal) in 50% ethanol to 50 mI . of melted glycerine
j el ly . The jelly-dye mixture keeps only 4-6 months so should be made
up fresh; also. reheating causes the jelly to deteriorate, so use of
only small quantities of the jelly Ivill prevent eXCesS Haste.

To prepare slides:
a . tap pollen onto slide
b . wash off oils and resins Hith a drop of 70% ethanol
c . remove ring of sludge I"ith a cotton swab moistened in 70% ethanol
d . place a drop of melted jelly on the pollen on the slide
e . stir gently to get even distribution of the pollen (the slide may
be kept I"arm to facilitate mounting. but excess heat Idll
rupture many pollen grains ) .
f . add 110 or 1/ 1 cover slip (a fter slide is removed from heat . i f used)
g . allaH slide to cool.

2. Gentian violet in aniline oil (permanent; grains not expanded) .

Place pollen on slide and add 2 or 3 drops of anili ne oil just tinted
purple Ivith gentian violet. Heat gently over small flame until grains
are deeply stain ed . AllaH slide to cool to room temperature, dral'" off
excess oil {>lith filter paper; Hash by repeated ly adding and blotting
off xylol until all! oil and unabsorb e d dye is removed . Haunt in clarite .

3. Aceto - carmine in Glycerine Je l ly (for pol len fertility counts) .

Aceto- carmine is frequently used to d ifferentiate betlveen good and bad
pollen but \'Ihen quantitative estimates of pollen fertility are required
the low viscosity of an aqueous solution is a dis advantage. Empty
pollen grains being lighter t en d to move to\vards the edges of the cover
slip when it is Im"ered . so that a random distribution of good and bad
pollen is not obtained . Counts, therefore made near the center of the
cover slip are biased in favor of good pollen. This error can be
avoided by using a viscous mounting med ium . The aceto-carmine glycerol
jelly described here has the advantage of being liquid in bulk at room
temperature. setting to a jelly when used on a slide and giving a per-
manent preparation \"hich can be scored at any time that is convenient.

The medium is prepa red in tlW stages as follows:

a. Stock jelly . Add 0.25 gm. of finely powdered carmine to 50 m!. of
mel ted Brand t 's glycerol jelly . Nix thoroughly so as to obtain
an even suspension of carmine particles throughout the jelly
when cooled. This concentrated jelly is convenient for storage;
small quantities can be used to ma ke the final medium, as
required .
b . Aceto-carmine jelly. Add 40 mI. of 45% acetic acid to 20 mi. of
melt ed stock jelly . Boil gently until the solution is clear
and all the carmine has dissolved ; this can be checked by
8 219

examin in g a drop of the medium under the microsco pe .Finally ,

add 4 ml . of a saturated so lution of ferric acetate in 45 %
acetic acid ; this cau seR the med ium to acq uire a d e ep port Hi ne
color .

Macerate an und ehisced anther in a dro p of th e medium on a slide, remove

debris a nd sti r the dro p with a needle to distribute the pollen grains
even ly . Th e dro p of medium u sed shou l d be such that no excess ,..d l l exud e
round the edges of the cover slip ,.,rhen the latter is lowered into posi-
tion . The prepar ation can b e used after a few hours by which time stain-
ing and di f ferentiat ion have occurred and the mount is permanent . [Adapt ed
from G. E . Harks, Stain Technology 29 : 277 , 1954 . 1

4. Cotto n Blue in Lactophenol (for pollen fertility count s) .

For tempo rary o r semi -permanent pollen s lides Cot ton Blue (Anilin Blue)
in lactophenol is convenient . Th e fertile pollen g ra ins stain a dark
blue and ster i l e g ra ins generall y do not s t ain or stain faintly. The
lactophenol is prepa red f rom: 20 ml. melted phenol (u se a water bath),
40 ml. glycerin, 20 ml. l actic aC id , 20 ml. distilled wat e r . To the 100
mI . of lactophenol add 1-5 mI. of 1% aqueous solution of Cotton Bl ue ,
depending on the int e ns it y of colo r desired . Prepa r e slide s as in 03 .
5. Double Staining of Pol len Grains .
The medium consists of 50 ml. of glycero l jelly t o which 2 . 5 ml. of
me thyl green and 2 ml. of phlOXine , both in 50% alcohol ic solut ions .
have been add ed . Prior to the application of the jelly-dye mix tur e
the pollen is washed \dth 70% ethanol to remove adher ing oi l s an d
r esins . The staining reaction is diffe r ential and pe r mits the rapid
c l assific ation of pol l en grains . The " functional " pollen expands and
stains \1ith both dyes "'lhereas the abo rted gr ai ns remain sh runken and
take only the methyl green wall stai n . The same reaction functions
with orchid seed . [Adapted from A. 0i1czar zak , A rapid method fo r
mounting pollen grains . with special regard to ste rility studies .
Stain Technology 27 : 249-251 . ]

6. Hoyer' s Haunt ing Nedium (for pollen and spores) .

A general mounting medium that gives expanded . unstained po l len grains
[o r s tudy \-J ith the light microscope .

Distilled \Jater SO gm. Chloral hydrate 200 gm.

Arabic gu m lump 30 gm . Glyce rine 20 gm .

The Arabic gum should be soaked in the distilled water a bout 24 hours .
Add c hlo ral hydrate and let the sol u tion stand unt il a l l the materia l
dissolves ( this may r equi r e several days) b e fo r e the glycerine is ad ded
and the so lution is ready for use .

7. Acetolysis Technique .
The acetolysis method for the prepara tion of pol l e n g r ains from dried
o r prese r ved p l ants- is the most ,.,tide l y used palynological techniq ue.
Nearly all of the ,"orld ' s pollen reference col l ec t ions have been pre-
pared in thi s manne r . As \o,'i1l be no ted. t his acid hydrolysis process
e mploys very caustic chemicals which leave on l y the resistant oute r
exine wall layer of the pollen grain intact . The pol len of a few
angiosperms does not s urvive this tr eatme nt and must be prepared in
other \1ays . Follol>'ing acetolysi s , grains are frequently brownis h in
220 8

color . Some Horkers have found it desirable to sta i n the prepared

pollen lightly (e.g . • basic fuchsin). Although slight variations
exist in the acetolysis technique. the basic procedure as outlined
beloH should give satisfactory results .
CAUTION: THIS PROCEDURE SHOULD BE DONE UNDER A HOOD . DO NOT BREATHE FillffiS

a.
-..
OR GET ACETOLYSIS MIXTURE ON SKIN OR CLOTHES .
Prepare acetylation reagent-9 parts acetic anhydride:l part concentrated
sulfuric acid. Add acid slowly to the anhydride . Prepare just enough
for one day ' s use; i t Hill not keep longer .
b. Remove perianth parts of the flower and pOt~der dried stamens through a
fine mesh screen into a funnel placed in a centrifuge tube . Het or
fresh pollen must be dehydrated in glacial acetic acid first . Note :
1 to 6 samples may be run simultaneously in separate tubes .
c. Pour 5-10 cc of the acetylation mixture over the polleniferous contents
of each centrifuge tube . Hash pollen off the sides of the funnels
with the above mixture also.
d. Place centrifuge tubes in a water bath under a hood . Equip each tube \-lith
a glass stirring rod and stir contents at regular intervals. Bring
water bath to a boil and cook for 10 minutes . Remove stirring rods
and let bath cool .
e. Centrifuge tubes at about 1500+ rpms for 1 - 2 minutes , then decant. All
large debris should be decanted. REHEl-illER: IF ACETOLYSIS MIXTURE
IS ADDED TO l·lATER A VIOLENT REACTION IHLL OCCUR . DECANT SLOHLY.
f. Hash pollen in tubes Hith glacial acetic acid, centrifuge and decant.
g. Add distilled water to the tubes, shake thoroughly , centrifuge and decant.
h. Add a fet.! drops of equal parts glycerine and distilled water to the tubes.
Allow to stand for ca . 10 minutes, centrifuge and decant .
i. Mount pollen grains . Use a bacteriological loop to remove pollen from
centrifuge tubes . The question of the most satisfactory mounting
medium has not been resolved. The most widely used med i um is glycerine
jelly, although some workers use lactic acid or other media. Cover
slips may be sealed using paraffin or left unsealed .
j . REMEl-IEER: AS IN ALL TAXONOHIC RESEARCH , IT IS VERY DESIRABLE TO HAVE A
VOUCHER SPECIHEN FROH HIn CH POLLEN hTAS REMOVED PLACED IN AN HERBARIUH .

PALYNOLOGICAL LITERATURE

I. Periodicals
Grana- -An international journal of Palynology . Uppsala , SHeden . 1954+ .
(until vol. 10(1), 1970, this journal Has published as Grana Palynolo-
gica, Stockholm, SHeden) .
Japanese Journal of Palynology. Shizuoka, Japan. 1967+. [In Japanese ]
Journal of Palynology . LucknOl.;r, I ndia. 1965+.
Pollen et Spores. Paris . 1959+. (Bibliographic supplement published
regularly) .
Review of Palaeobotany and Palynology . Amsterdam, The Netherlands. 1965+.
II . General References
BreHbaker , J . L . 1967 . The distribution and phylogenetic significance of
binucleate and trinucleate pollen grains . American Journal of Botany
54 , 1069-1083.
Cranwell , 1. H. 1953 . NeH Zealand Pollen Studies . The Monocotyledons .
Harvard University Press . Cambridge .
Erdtman, G. 1943 . An Introduction to Pollen Analysis. Chronical Botanica
Company. Haltham. Hassachusetts .
 Erdtman , G. 1952 . Po llen Horphology and Plant Taxonomy - Angiosperms . (An
Introduction to Palynology , vol. I) . Almqvist and Hiksel l. Stockholm .
1963 . Palynology . In : Vistas in Botany , IV . I,' . B. Tu rrill (ed . ).
HacBillan and Company . NeH York.
196 9 . Handbook of Palynology . Horphology-Taxonomy-Eco10gy. An
Introduction to the Study of Pollen Grains and Spores . Hunksgaard . Copen-
hagen .
Faegri, K., and J . Iversen . 1964. Textbook of Pollen Analysis . Ed . 2 .
lIafner Publishinp, Company . NeIll York .
Felix, J . C. 1961 . An Introduction to Palynology . In : H . N . Andre",'s,
Studies in Paleobotany . John l-liley and Sons. New York .
Germeraad , J. II ., and J . Nuller . 1970. A comput er-based numerical coding
system for the description of pollen grains and spores . Review of Paleo-
hotany and Palynology 10: 175-202.
Hyde, H . A., and K . F . Adams . 1958 . An Atlas o f Airborne Pollen Grains .
Hadlillan Company . London.
Ikuse , H . 1956. Pollen Grains of Japan . HirokaH3 Publishing Company. Tokyo .
Kapp, R. O. 1969. How to Knm.,T Pollen and Spores. Uilliam C. Brmm Company .
Dubuque, IOI"a .
Kremp , G . O . h' . 1965 . Horphologic Encyclopedia of Palyno logy . Universi ty
of Arizona Press . Tucson .
Reitsma , T . 1970 . Suggestions tOl"ards unification of descriptive terminology
of angiosperm pollen grains . RevieH of Palaeobotany and Palynology 10 : 39-
60 .
Richard, P . 1970 . Atlas pollinique des arbres et de quelques arbustes
i ndi genes du Quebec . I-IV . Naturaliste Canadien 97 : 1-34 ; 97-161 ; 241-
306 .
Tsch udy, R. 11 . 1969 . The Plant Kingdom and Its Palynological Representation .
In : R. H. Tschudy and R. A. Scott (eds . ) . Aspects of Palynology . John
Ihley and Sons. NeH Yo rk.
j.]odehouse, R. P . 1935 (reprinted in 1960) . Pollen Grains : Their Structure ,
Identification, and Signi ficance in Science and Hedicine . HcGra\oJ-Hill
Company . Ne\" York .
Horld Pollen Flora. 1970- . G. Erdtman (ed . ) . Hafner Publ ish ing Company.
Ne,'" York . (to be published in parts. Part I appeared in 1970) .

III . References Illustrating Taxonomic Application of Pollen Norphology

Dahl, A. O. 1952. The comparative morphology of the I cacinaceae, VI. The

pollen . Journal of the Arnold Arboretum 33 : 252-295 .
Fergus on, I. K., and D. A. Hebb . 1970 . Pollen morphology in the genus Saxi-
frilga and its taxonomic significance . Botanical Journal of the Linnean
Socie ty 63 : 295-311 .
Graham, A., and S. A. Graham. 1967. Pollen morphology and taxonomy of
Cuphea (Lythraceae). Revie\" Paleobotany and Palynology 3 : 155-162.
Jones , S. B. 1970 . Scanning elect ron microscopy of pollen as an aid to the
systematics of Vernon ia (Compositae) . Bulletin, Torrey Botanical Club 97 :
325-335 .
Lewis , H. H. 1965 . Pollen morphology and evolution in Iledyotis subgenus
Edrisia (Rubiaceae)· . "- American Journal of Botany 52: 257 264 .
Skvar1a, J . J . , and B. L . Turner . 1966 . Systema tic implications from elec-
tron microscopic studies of Compositae poll en - a reviel". Annals Hissouri
Botanical Garden 53 : 220-256 .
 222 8

\~alker ,J . hI , 1971. Pollen morphology, phytogeography, and phylogeny of

the Annonaceae . Contributions from the Gray Herbarium 202 . 131 p .

IV . References on Technique

Brown, C. A. 1960. Palynolog ical Techniques. Pr-ivate l y published by

the author, Baton Rouge , Louisiana .
Erdtman, G. 1960. The acetolysis method. A revised description . Svensk
Botanisk Tidskrift 54(4): 561-564 .
Falk, R. H. , E. M. Gifford , Jr ., and E. G. Cutter . 19 71. The effect of
various fixation schedules on the scanning electron microscopic image
of Tropaeolum majus. American Journal of Botany 58 : 676 -6 80.

PALYNOLOGICAL EXERCISE

From slides prepared by the student or provided by the instructor obtain

pol len data for one or more species .

Family Family

Species I Spec ies II

1. Pollen units . .... .... ....... .. .•••• •

2. Symmetry ......... .... ........ •••••••

3. Polarity ....................... ... . .

4. Size (Polar axis and equatoria l

axis} ............................ .

5. Aperture Type ... .. .. • • ••..•. • • .... ..

6. \,lall Structure .....• • • . . . ... ........

7. Wall Sculpture ..... • ... .. .. .... .....

8. Nuclear State ... . ........ ....... ... .

Prepare an accurate drawing showing the above features.

For an exercise dealing \-lith pollen size variation see the pollen exercise
used as an example for data accumulation (Chapter 20 , Section F).
9 223

Chapter 9 . EHBRYOLOGICAL EVIDENCE

If broadly defined. the term embryology inc ludes the successive stages
of megasporogenesis and mic rosporogenesis (meiot ic development of the func-
tional megaspore and microspore), megagametogenesis and microgametogenesis
(development of the female and male gametophyte) , and the post-f ertilization
stages of embryogeny (development stages from the fertilized egg to the mature
embryo and its surrounding seed coats) . Information derived from embryology
has proven to be of systematic value ; hOHever. obtaining these data requires
considerable effort. Not only do flo,,,ers have to be collected at critical
periods during their development, but they must be skillfully fixed , sec-
tioned and stained . Bjornstad (1970) states that an embryological study can
be helpful to the taxonomist in three Hays : (1) systematic questions may be
decided ",hen two or more positions have been proposed on the basis of other
characters. (2) net-! research may be promoted on a taxon \"hen embryological
data make the present systematic position doubtful or improbable , and (3)
confirmation of the present systematic pOSition of a taxon by emb ryo logical
information . The following t\velve embryologica l characters are considered
by HaheshlVari (1950 , 1964) to be taxonomically significan t .

A. Anther .
1 . Number and arrangement of loculi .
2 . Structure and thickening of the endothecium ( " fibrous" or not) .
3 . Nature of tapetum .
a. Glandular ( secretory) tapetum . Tapetum composed of highly
vacuolated cells Hhich remain stationary .
b . Amoeboid (plasmodial) tapetum . Tapetum in I"hich the cells , sub-
sequent to \vall breakdo\ffi, move I"ithin the loculus.
c. Huclear cond it ion (uni-, bi- , multinucleate , polyploid).
4. Nature of periplasmodium.
5 . Type of l"al1 fo rmat ion (Basic , Dicotyledonous, Monocotyledonous,
Reduced) .

<
endothecium
secondary
parietal cell

<
middle layer
Basic primary
parietal cell
middle layer
secondary
parietal cell
tapetum

Dicotyledonous
.< <
. '..

pn.mary
parietal cell
secondary
parietal cell

secondary
endothecium

middle layer

parietal cell ---------- tapetum

7 167

Chapter 7. ANATOHICAL EVIDENCE

This chapter deals with anatomy as a systematic too l. As stated by

C. R. Hetcalfe (1 968) anatomy of the vegetative organs of flowering plants
can be taxonomic ally useful in the following Hays: (1) the i dentifica tion
of fragmenta r y ma ter ial, (2) the preliminary identif i cation of herbarium
spec imens , and (3) as an aid tOHard establishing the interrelationships of
taxa at and above the species l evel .

Anatomical characters have been employed for systematic purposes well

ove r a hundred years. The most significant guiding principles of systematic
ana t omical study resulting from this long history have been eloquently dis-
cussed by 1. I,' . Bailey (1951 , 1953, 1957), and include: (1) anatomical data
tends to be most useful at the level of the genus and at higher taxonomic
categor ies, (2) anatomical characters are no more or less reliable than
characters from other parts of the plant, (3) similarities in structural
specialization do not necessarily imply close relationship but may be the
result of parallel and convergent evolution , (4) anatomica l data have proven
most reliable in statements of negation of close relationship rather than
positive assertions of relationship, (5) the assumption that a single sample
of an organ or tissue provides reliable data for the anatomical characteri-
zation of a spe cies or genus is invalid, (6) it is essentia l that more
become knm-m regarding the ranges of variability of anatomical characters
within the same individual and different individuals of the same speCies ,
and (7) only when anatomical information is coupled Idth evidences from
other parts of the plant will a natural classif i cation be attained. It is
important, fu rthermore, to emphasize that anatomical information is fre-
quently taxonomical l y useful I,d thout having obvious evolutionary or phy lo -
genetic interpretation . Info rmation on floral anatomy and evolutionary
morphology may be found in Chapter 27 . Unless otherwise stated, terms are
defined in accordance I~ith K. Esau (1961) .

Section A. SECONDARY XYLE~I

This sect i on Hill be concerned with the anatomy of twod . Included are
characters of significant systematic value . I n no other vegetative tissu es
of the plant are the trends of evolution as t.;rell understood (see Chapter 27) .
As a result , wood anatomy has assumed considerable importance in inte rpreta-
tions of vascular plant evolu tion (see Chalk , 1937, 1944; Tippo , 1946).
Transverse, radia l and tangen t ial sections of \.fQod shou l d be prepared and
studied to understand the organ ization of the tissues . Permanent macera-
tions also should be made to obtain statistical data on cell l ength a nd
other features where data on individual elements of the secondar y xylem are
desired .

The follow ing list of diagnostic characteristics for descriptions of

dicotyledonous t.;roods has .,peen t aken with modif i cations from Tippo (1941).
Inc l uded among this list are features of proven taxonomic and phylogenetic
value. Terms are mostly defined in accordance Hith the ~!ul til ingual Glos-
sary of Terms Used in Hood Anatomy (Committe e on Nomenclature, International
Association of Wood Anatomists , 1964). For the most part, categories of
measurement fol low the suggestions of the Committee on Standardization of
Terms of Cell Size (1937, 1939). An abbrevia t ed data Iwrk sheet is provided
168 7

Figure 7-1. Cell types of the secondary xylem, a-d, Iwod fibers . a,
librHorm wood f ib e r. Note the simple pits and pointed ends of th is most
advanced of \~ood fiber types. b-d, fiber -tracheids . Note the progressive
loss of the pit border from d to b. e-f, tracheids . The cell figured in
e has circular-bordered pits whereas the element pictured in f has scalari-
form pits. g-k, vessel elements . g . the cell pictured here represents a
primitive vessel element type h'ith many-barred scalariform perforation
plates that are fully bordered (inset) and scalariform intervascular pitting.
h, this cell <;Ilso has scalariform perfora tion plates but the number of bars
is reduced in number and they are bordered only at the ends (ins et ). The
intervascu la r pitting is composed of circular bo rdered pits arranged in an
opposite manner. i, this vessel element represents an intermediate stage
in evolution I"ith a scalariform perforation plat e at one end and a simple
perforation plate at the other e nd . I ntervascular p i tting is still essen-
tially opposite . j, the cell pictured here is f urther advanced as evidenced
by the exclusively simple perforation plates Hhich are tending to become
t ransverse in orientation. The intervascular pitting is best d escribed as
transitional betHeen opposite and alte rnate . k, this vessel element is the
most advanced type illustrated Hith simp le, transverse perforation plates
and the alternate intervascular pitting . Also note its short length and
increase in IJidth . I-n, crystal types . 1, druse crystal. ro, prismatic
crystal. n, raphides . 0, septate libriform lJOod fiber . p, vasicentric
tracheid . Note the irregular shape of this element.
 •
•
(o!
, 0

, %• ~,
,
• e
• ,
,
,
,
, •,
I'•
,
~

•
,
, .'s'

e
, ®
0
• • ~

b
,
c ~-
d

o
170 7

to record observations. For more complete wood descr i ptions a form like the
one used by Zamuco (1965) is r ec ommend ed .

1. GROWTH LAYERS rANNUAL RI NG S] (PRESENT OR ABSENT ).

A layer o f Iwod or bark produced apparently during one grow ing

period; f requent ly, espec ially in Iwods of the temf!erate zones , d i visi-
hIe into early and late wood or hark . '" -

II. TRACHEIDS AND FIBERS (IMPERFORATE TRACHEARY ELENENTS).

A. ~ (see Figure 7-1).

1. Tracheid . An imperforate Hood cell l"ith bordered pits to conge-
n e ric e l emen ts.
2. Vasicentric tracheid . A short, irregularly-formed tracheid' in
the immediate proximity of a vessel and not fo rming part of
a definite axial rm... .
3. Vascular trach eid . An imperfo rat e cell resemb ling in f orm and
position a small vessel member .
4. Fi ber. A general term of convenience in wood anatomy for any
long , narrm" cell of \wod or bast other than vessel s and
parenchyma .
NOTE : Often further qualified as wood fibers or bast fibers;
the former including both the tracheids of gymnosperms and
libriform I"ood fibers and f iber-tracheids of I"oody angio-
sperms . Al so used loosely for Iwod elements in general.
5 . Fiber-trache i d. A fiber - like tracheid ; commonly thick-walled
I"ith a small lumen, pointed ends , and bordered pit pairs
having lenticular to slit-like apertures . This term is
applicable to the late Iwod tracheids of gymnosperms as
I"ell as to the fiber-like tracheids o f Iwody angiosperms.
6. Septate fiber-tracheid. A fiber-trache id I"ith thin transverse
walls across the lumen .
7 . Libriform Iwod f iber. An elongated, commonly thick-HaIled cell
with simple pits; usually distinctly longer than the cam-
bial init ial as inferred from the length of vessel members
and parenchyma strands.
8. Septat e wood fiber . A fiber with thin transverse walls across
the lumen .
9 . Gelatinous fibe r . A fi ber having a more or less unlignified
inner wall with a gelatinous appearance .
B. Hall Thickness, Sculpture and Cell Length
1. Hall thickness .
a. Very thin - lumen much greater than thickness of walls.
b . Thin - lumen greater than thickness of wall s .
c. Thick - lumen less than thickness of walls.
d . Very thick - lumen almost completely c l osed .
2. Size of pits - minute, small, large .
3 . Length (measure 100 cells from macerations) . Record range, most
frequent range, and mean . [llID = micrometer]
a. Very short ............... up to lOOOVm
b . Short .................... 1000 - l SOOvm
c. Long ......... .. .......... 1500 - 2000vm
d . Very long .......... .. .... over 2000vm
4 . Spiral Th icken ings (Record presence or absence) . Helical ridges
on the inner face of , and part of, the secondary wall .
7 171

Ill. VESSELS ( PERFORATE TRACHEARY ELEl-IENTS) .

A vessel is an axia l series of cel ls (vesse l member s or elemen ts )

that have coa l esced to form a n articulated tube- l ike str uct ure of inde-
ter minate length ; t he pi t s t o congene r ic elements a r e bo r dered . Amon g
dico t yle dons , the p r imitively vesselless condition is enco un tered in
the Trochodendraceae , Tet r acentraceae , Hinteraceae , Nonimiaceae and
Chloran t haceae .
A. Fregu ency and Dist rib ut ion ( see Figure 7- 2).
1. Number per sq . mm. ( as seen in transverse section) . Coun t from
several field s. Record range, most frequent range , and
mean .
a . Very few ................... ... . up to 2
h. Few .............. . .... . • • •• . ... 2- 5
c . Hoderately few . ... ... . .••• •. ... 5- 10
d . l-loderately n umerou s .. . ..•••• . .. 10- 20
e . Numerous . . . ...... ... .... . ..... . 20-40
f . Very numerous . .. .. . . ...... .... over 40
2. Pore (vessel) dist r ibut ion .
a . Solitary pores . A pore complete l y surrounded by ot he r ele-
ments .
b . Pore chain . A se r ies or line o f adjacent so l itary pores .
c . Pore mul t i ple . A group of two or mor e pores crOl"ded to-
ge t he r and fla ttened along the lines of contact so as
to appear a s s ubdivisions of a s i ngl e pore . The mos t
common t ype is a r adial po r e multiple, in whic h t he
po r es are i n r adial files l1i t h f l atten ed tangent i a l
walls be t ween them . Another type is a pore cluste r,
in which the grouping is irregula r.
3 . Diffuse- po r ous wood. Wood ill \~hich t he po r es are of fairly uni-
form or on l y gradually changing siz e and distribution through-
out a grOl"th ring .
4. Semi-ring porous wood . ~~ood in which the early wood is ma r ked
by a zone of ( a ) occasional l arge vessels , or (b) numerous
smal l vessel s .
5 . Ring- porous wood . ~olood in I"hich the pores of the early wood
are distinc tly larger than those of t he la t e wood and f o r m
a well - def ined zone or ring .
B. Vessel Elemen t Shape , Hall Thickness and Size .
1. Angular or ci r cular ( as seen in tran s ver se section) .
2. Thin or thick walle d .
3. Diameters (as me a s ur ed in transverse s ection) . Heas u re 1 00
cells . Reco rd r a nge , most frequen t range , and mean .
a . Extremely sma l l ... .. ... ... ....... up t o 25vm
b . Ve r y sma l l ......... . ............. 25- 50vm
c . Noderate l y smal l . . . . . . . . . .... .... 50- 100vm
d . Nedium - sized . ....... .. . . . . . . ..... 100-200um
e . Hoderate l y large . . .. . . .. . . .. . .... 200-300lJm
f . Very l arge .. . ........ . .... .. ..... 300- 400um
g . Extremely large ....... .. ......... ove r 400).l1n
4 . Length of vessel elements (total length - tip to tip) . Heas ur e
cells fro m mace r ations . Record range , most freque n t range,
and mean .
a . Ext r emel y sho rt . . ......... less t han l75vm
h . Ver y sho r t ........••••. . . . ..... 1 75 - 250urn
172 7

Figure 7-2 . Transverse sections of dicotyledonous Iwods. All X 117 .

A. Trochodendron aralioides (Trochodendraceae), a primitively vesselless
angiosperm showing the secondary xylem composed of tracheids (T) . Distinc-
tion can be noted between tracheids of the late \Vood (Ul) and early wood
(EI,l). B, Annes l ea crassipes (Theaceae). This wood shows solitary . very
angular vessel elements (V) and mostly diffuse axial parenchyma . The
imperforate tracheary elements are fiher - tracheids. C. Dillenia pentagyna
(Dilleniaceae) . Although thi s wood is still of a comparatively primitive
nature the vessel elements are tending to become rounded in outline and
the axial parenchyma is both para tracheal scanty and apotracheal diffuse
to diffuse- in-aggregates . The imperforate tracheary elements are fiber -
tracheids (FT). D, Scytopetalum tieghemii (Scytopetalaceae) . I n this wood
the vessel elements are distributed as solitary pores and radial pore mul-
tiples. Imperforate tracheary elements are thick-tmlled libriform fibers.
E, Bombax ellipticum (Bombacaceae). In this section note the rounded vessel
elements and the diffuse-in-aggregates parenchyma. F, Acer saccharum (Acera-
ceae). In this comparatively advanced wood the vessel elements occasionally
occur in pore clusters . The pores are circul ar in outline and the imper-
forate tracheary elements are libriform fibers . Rays are homocellular.
7 173

uniseriole ro\' diffuse parenchyme

(opot rach eol)

, [;0 \; . t\.
')'rf ~
: ~,

,
\< ~
•
J, r,'),~ B
,~
, (

, ~:\ X . ,, ,f;
,
I~ f-' , c
,

, ,

c 'JT ,\,- ,
1
, ~~ ;r
seanly parenchyma
{porOlro cheo l) FT

di ffu se-in- aggregates porenchyma

 174 7

c . Hoderately short . . .. . .... . .. .. .. . .. ... 250- 3501.lm

d . Medium-sized . . . . . . . . . . . . . . . . . . • • • • . . . . 350- 800\.lm
e . Noderate1y long .. ..... .. . . ..• ..... . . . . 800-l!OO\.lm
f . Very long ..... ... .... .. .... . ... . .... . . llOO-1900 \.lm
g. Extremely long .. .. .. ... . . ..... . . ..... . over 1900\.lm
C. Per foration plates. A ter m of convenience for the area of the wall
( o riginal l y imperforate) involved in the "'co'alescence of two mem-
bers of a vessel .
1 . Types (see Figure 7-1) .
a . Perforation plate scalariform . A plate with multiple perfora-
tions elongated and parallel. The remnants of the pla t e
between the openings are called bars . NOTE : Record num-
ber of bars (range , most frequent range, and mean) , and
t"hether bars are completely bordered, bordered to middle ,
bordered at ends, or non-bordered .
b . Pe r foration plate simple. A single and usually large and
more or less rounded opening in the perforation pl ate .
c . Perforation pl ates simple and scalariform.
d . Perforation plate reticulate . A plate t ... ith multiple perfora-
tions having a net-like appearance (as in certain Big-
noniaceae) .
2 . End walls . Record t... hether oblique (express as a ng l e) or trans-
verse .
D. Pitting . A recess i n the secondary ",all of a cel l, together "'ith its
externa l closing membrane ; open internally to the lumen .
1. Intervascular pitting . A term used (a) in a t..ride sense for pi t -
ting between tracheary elements , and (b) in a narro",e r sense
i n wood anatomy for pitting bet",een vessel members . Record
spa r se or crot.,rded, shape of pits ( circular , squar e , penta-
gonal, etc.) , and size of pits - Ninute ( l ess than 41.1m) ,
Small (4-7lJm) , Nedium (7-l0)Jrn ) , Lar ge (lO-lS)Jm). and Very
Large ( over l5)Jm) . (See Figure 7-1) .
a . Scalariform . Pitting in which elongated or l i near pits are
ar r anged in a ladder-like series .
b . Opposite . Hult i ser i ate pitting in ",hi ch the pits are in hori-
zontal pairs or i n short horizon t al rot...s . NOTE : when
pits are crot...ded the outlines of the bor ders tend to
become rectangular in surface v i et..r .
c . Alternate . Nultiser i ate pitting in which the p i ts are in
diagonal rmvs . NOTE : tvhen the pits are crm...ded the
o u t l ines of the borders tend to become hexagonal in
surface v i e", .
2 . Ray-Vessel pitting . Pitting bet\...een a ray cel l and a vessel mem-
ber . Record s i ze , shape , whether scalariform , oppos i te ,
al t ernate , or irregular in distrib u tion. and whether bordered
or simple .
E. Other Features .
1 . Tylosis . An outgrowth from an adjac ent ray or axial parenchyma
cell through a pit cavity in a vessel wall , par t ially or
co mpletely b l ocking the vesse l lumen . Ty l oses may be few
or many crowded together ; thi n - or thick- walled ; pitted o r
unpitted ; t..ri th o r without starch , crystals , resins , gums ,
etc .

IV . RAYS .

A r i bbon - like aggregate of cells extending radially in the xylem a n d phloem.

7 175

A. ~ (see Figure 7- 3) .
1. Heterocellular ray . Ray tissue in Hhich the i ndividual rays are
composed of both procumbent and square or upright ce l ls .
2 . Homocellular ray. Ray tissue in Hhich the i ndividual rays are
composed Hholly of pr ocumbent cells ; or all square or upright
cells .
3 . Aggregate ray . A group of small , narr m", xylem rays appea r ing to
the unaided eye or a low magnification as a single la r ge ray .
B. Abundance . Number per sq . mm . (as seen in tangential section) .
1. Very f eH ................. . ... up to 2
2 . Fe\" . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 2- 4
3. Hoderately n umerous ............. 4- 7
4 . Numerous ..... . .... . ............. 7-10
5. Very numerous ................ over 10
C. Si ze.
1. l'l'idth (number of cells \"ide) .
a . Uniseriate ray . A ray one cell wide as seen in tang ential
section .
b . rtultiser iate ray . A ray t\"O or more cells \"ide as seen in
tangential section . Record range and most frequent range.
2. Height (number of cells high) . Reco r d range and most frequent
range .
D. Pitting (bet\"e en r ay cells and other parenchyma cells) . Record s ize
and number - feH , many , clustered .
E. Specialized Cells and Contents .
1. Lignified cell \"alls or nonlignified cell Halls .
2 . Intercellular canals . A tubular inte r cellular space of inde -
termina t e length, genera l ly serving as a repository for
r esin, gum , etc . , secreted by the epithelium .
a . Lysigenous canal . Originat i ng by dissolution of cell s .
b . Schizogen ous canal. Origi nating by separation of cell Halls
a l ong the middle lamella .
3. Latex t ubes . A l at i c i fer enclosed by a r ay .
4 . Oi l cel l. A spec i alized cell of the ray or axial parenchyma con-
taining o il, typicall y rounded in outline. NOTE : limited to
\"oody dicotyl edons and similar to a mucilage ce ll, except for
contents .
5. Crystal lifer ous cell . A cell containing one or more crys t als .
6 . Sheath cell . One of a series of up r ight ce l ls on the margins o f,
and tending to form a sheath around , the proc umbent cells of
a multiseriate ray as seen in tangential sect i on .
7. Til e cell . A s pecialized type of apparent ly empty upright ray
cell of ap p roximately the same he i ght as the procumbent ray
cells and occu r ring in indeterm i nate horizontal series us u-
a l ly interspersed among the procumbent cells . NOTE : common
in ce rta i n of the Tiliales and Hal va l es .
8 . Scle r otic ray cell . Ray cell Hith thick , often l ignified secon-
dary Hall s .

V. AXIAL PARENCHYNA .

Par enchyma cells derived from fusifo r m cambial i nit i als .

A. Di stribut i on (as seen i n transverse section) . (See Fi gure 7-2) .

176 7

Figure 7-3. Tangentia l se ct ions of dicotyledonous weods . All X 117.

A, Eupomatia laurina (Eupomatiaceae). The wood of Eupomat ia is comparatively
pr i mitive in structure . The multiser iate rays are decid edly heteroc e llular
and composed of nearly all upright cells . Sheath cells (S) are of freq uent
occurrence . B, l-Ia gnolia grandiflora ( Hagn oliaceae) . Th i s wood shows both
uni seria te and biseriate h eterocellula r rays . Th e unise r iate wing on the
biseria te rays is frequently reduced t o a s ingle ce ll. C, llex apaea (Aqui-
foliaceae) . This \-load also illus tra t es the occurrence of both un iser iate and
multiseriate r ays . The un .iseriat e rays are composed almost exclusively o f
uprigh t cells whereas the mai n body of the mult iseriat e ray consists o f pro -
cumbent ce lls with only the cells of the uniseriate wi ng havin g an uprigh t
orientat ion . D, Quercus a lba (Fagaceae) . This species of oak illu s trates
the very Idde nature of some mult iseria te rays. The rays are nearly homo-
cellular . E , Gonvstvlus I-Jarburgianus (Thymelaeac eae). The rays of this
species are predominantly uniseriate a nd almo st homocellular . F , Populus
de l toides (Sali caceae) . The rays of t his plant are exclus ively un ise riate
and homocellular . Two adjoining vesse l elements (V) are also shown .
7 177

biserial e

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••,•.",.. •. 1'l ::..:-..
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: ' ; I ,

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· ....• ...1 ,; Ir.
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multi serio Ie
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 178 7

1. Apot ra cheal parenchyma. Axial parenchyma typical l y independent of

the pores or vessels .
a. Termina l parenchyma . Apotracheal parenchyma cells occurring
either singly or forming a more or less con tinuous layer
of variable width at the c lo se of a season ' 5 g r o\.,rth .
h. Diffuse parenchyma . Single apotrache-a"l '- parenchyma st r ands or
ce ll s d i stributed irregul a r ly among fibers .
c. Diffuse - in-aggregates . Apotracheal parenchyma cells that tend
to be grouped in short tangen tial lin es from ray to ray .
d. Banded apot ra cheal pa r enchyma. Ax i a l parenchyma forming con-
centric lines or bands . NOTE : termed Apo tracheal Banded,
if typically independent of the vessels; Paratrachea l
Banded , if associated \"ith the vessels .
2. Paratracheal parenchyma . Axial parenchyma assoc iated with the
vessels or vascular tracheids .
a . Scanty paren chyma . I ncomplete sheaths or occasional pa ren-
chyma cell s around the vessels .
b . Vas i centric parenchyma . Paratracheal pa r enchyma form i ng a
complet e sheath around a vessel , of variable width and
circular or slightly oval in cross sectio n.
c. Aliform par e n chyma. Para t racheal parenchyma with Hing-like
l ateral exten sions , as seen in cross sect i on .
d. Conf luent parenchyma . Coa l esced a l iform parenchyma fo r mi ng
irregular tangent ial or diagonal bands as seen in cross
section .
B. Pitting (between ax ial parenchyma cells) . Record s ize and number--
few, many , or clustered.
C. Special ized types.
1. Fusiform parenchyma cell . An axial parenchyma cel l, de r ived from
a f us iform initial without subd ivision.
2 . Septate parenchyma cell . An axial o r r adial parenchyma cel l ~.ith
one or more thin transverse walls across its lumen .
NOTE : in these elements the protoplast divides after the for-
mation of the secondary cell wall .
3 . Crystallif erous axial parenchyma cell.
4 . Chamber ed crystalli ferous cell . A crystalliferous cell that is
divided into compartments by septa .
S. Sc lerot i c pare nchyma ce ll.

VI . OTHER FEATURES .
A. Storied . A te rm applied to the axial cells and rays in wood when these
are arranged in horizontal series on tangential surfaces. NOTE:
the term is applied to particular tissues, e.g . • " storied paren-
chyma" or used in a gener al sense, as in "woods with stori ed
structure . "
B. Crystals (present or absent) . (Se e Figure 7-1) .
1. Acic ula r. A slender , needle-shaped crysta l. NOTE: not to be con-
fused with a styl oid, \~hich is a columnar crystal.
2 . Crystal Sand . A gr anular mass of very fine crys t als .
3 . Druse . A globular cluster of crystals , sometime s Hith an o rganic
core , eithe r attached to the wall by a peg or l ying fre e in
the cell .
 7 179

4. Raphide (pl . raphides). A needle - shaped (acicular) c rystal occur-

ring typically as one of a c lo sely packed, sheath-like bundle .
5 . Styloid . An elongated c ryst al , typically abou t four times as long
as broa d, ,.,rith pointed or square ends .
6 . Prismatic. A solitary crystal either a rhombohedron or octahe-
dron .
e. Intercellular canals. Tubular intercellular space of indetermina t e
length .
D. Included phloem. Phlo em strands or layers inc l uded in the secondary
xylem of certain dicotyledonous \'-'Oods .
E. Vestured pits. A bordered pit with the pit cavity \.,rholly or par-
tially lined with pr oject i ons f rom the tertiary wall.
F. Fibriform vesse l e l ement . A vessel element of relatively small
diameter bearing a r esemblance to a fiber-tracheid .
G. Disjunc t ive tracheid . A tracheid partly disjoined laterally from
another during differentiation; contact is main tained by mea ns
of tubular processes .
II . Disjunctive parenchyma . Axial or rad i al parenchyma cells part ial ly
diSjoined during the process of diffe r ent iat ion ; contact is
maintained by means of tubular processes .

SECONDARY XYLEH LITERATURE

I. Periodicals .

Tropical \~oods . Ne\" Haven , Conn . Yale Universi t y School of Fo r estry .

(This journal began publication in 1925 and terminated with No . 113
in 1960) .

II. General Refer ences .

Bailey, I . H. 1957 . The pot enti alit ies and limi tations of \'-'Ood atlatomy
in the phylogeny and classification of angiosperms. Journal of the
Arnold Arbo r etum (Harvard University) 38: 243- 254 .
Chalk , L. 1937 . The phylogenetic value of certain ana t omical features
of dicotyledonous \~oods. Annals of Botany, N. S . , 1 : 409-428.
1944. On the taxonomic value of the anatomical st r ucture of
the vegetative organs of the dicotyledons. 2 . The t axonomic value
of Hood anatomy. Proceedings Linnean Society of London 155 : 214-218 .
Committee on Standa rdi zat ion of Terms of Cell Size , International Asso -
ciation of Hood Anatomist s , 1937. Sta ndard terms of l ength of vessel
members and \.,rood fibers . Tropical Hoods 51 : 21-
1939. Standard terms of size f or vessel diameter and ray
<.-ddth . Tropical Hoods 59: 51-52 .
Tippo , o . 1941. A list of diagnostic characterist ics for descriptions
of di cotyledonous woods . Tr-a nsactions Illino i s Academy of Scienc e
34 : 105-106 .
1~46. The role of wood anatomy in phyl ogeny . Ame r ican Hid -
land Natur alist 36 : 362 - 372.
Zamuco , t . T . 1965: ' .... Bark and wood anatomy of fou r Philippine bast-
f iber producing trees. The Phi lippine Journa l of Forestry 21 : 127-
146 .
 180 7
7

TECHNIQUES EHPLOYED IN THE STUDY OF HOOD ANATOl-fY

I. MACERATIONS.

Maceration is a chemical method of dissociating elements by dissolv-

ing out the intercellular cement substance, thus allm.,ring the investiga-
tor a clear three - dimensional picture of the cell - types comprising the
tissues .
1. In macerating dry woods, spl it the material into slivers no thicker
than a toothpick, then boil in Hater until all pieces are thor-
oughly saturated.
2. Transfer the slivers to a shell vial and cover them lvith Jeffrey 's
macerating fluid (equa l volumes of 10% aqueous nitr ic acid and
10% aqueous chromic acid) . Stopper the vial and place in a 60 Q
paraffin oven for 24-48 hours . The time varies with the hard -
ness and thickness of the material, so check regu l ar l y at the
end of the first day; Hhen the maceration proce ss has progressed
sufficiently, the Hood elements \"ill separate readily \"hen
teased apart with needles.
3 . Hash the material thoroughly in running water to remove all traces
of the acids , then dehydrate rapidly by decanting and adding
increas ingly stronger alcohols; e . g. , 50, 75 , 95, and 100% .
4. Staining is effec ted by transferring the Hood slivers into a 1%
solution of light green in 75 parts clove oil and 25 parts
absolute alcohol . If a red stain is preferred , stain 4-6
hours in a 1% solution of safran in in 70% alcohol before com-
pleting the dehydration series.
5 . After about ten minutes the material can be removed from the stain
and the exc ess stain can be removed by blotting with a filter
paper . A sliver of \"ood is then placed on a clean g1.;ISS slide
in a drop of xyl ol and mounting medium , and the elements can
be teased apart and carefully spread around with dissecting
needles . Avoid too much material on one slide .
6 . Add cover slip to slide and place on a slide warmer to dry .

II . USE OF THE SLIDING HICROTOl'lE .

1. Soft Hoods can often be sectioned successfully on a sliding micro-

tome without embedding, follmving their saturation in boiling
water . Hard woods may also be cut unembedded either by the use
of a jet of super- heated steam directed upon the wood clamped
in the microtome , or by preliminary softening of the \"ood in
hydrofluoric acid . Some woods can be sect i oned successfully
only after being embedded in celloidin.
Trim the wood carefully into blocks measuring approximately
1 x I x 2 em . , and prepa r e them by boiling in water and pumping
in cold water unti l the wood is thoroughly saturated and sinks .
Clamp the knife firmly in the knife ho l der of the sliding
microtome and adjust the three clamping screl"S to give the
desired cutting and clearance angles . (The best cutting angle
for the knife edge, \vith reference to the line of travel, usually
ranges from 30 to 35°. A greater angle is necessary when cutting
harder objects . ) Fasten the block of wood in the split- ball
clamp , and adjust the universal jOint until the desired plane of
sectioning is parallel to the plane of travel of the knife . A
 thickness of 15 to 20~ is satisfactory for most \.,roody subjects .
Cut cross , radial, and tangential sections and place them in
95% alcohol in a Syracuse \.,ratch glass. In the event that sec-
tions tend to curl after removal from the knife , press the
sections , Hith the concave side down, on a slide covered \.,Tith
a film of glycerine- alcohol or use a detergent solution . Then
place a dry slide over the sections, \.,Teight with lead blocks and
cover with \.,Tater in a Petri dish or finger bowl. The water I.;ill
render the sections f lexible and permit them to f latten , so that
they can be properly stained and mounted.

2. Staining .

Run sections down to water rapidly, using a section lifter or

camel ! s hair brush to make the transfers beth1een solut io ns in
\.,ratch glasses . Hordant 10-20 minutes in a 4% solution of ferric
ammonium sulphate ("iron alum" ), follm.,red by at least three \.,rashe s
in tap water . Care should be taken that neither of the first two
\.,rashes remain very long on the sections or fer ric hydroxide will
f orm in the tissues and spoil their staining quality . The las t
tap water wash should be allowed to stand on the tissues 9-10
minutes, follm..red by a distilled water rinse to remove the salts
dissolved in the tap water . Transfer sections to a watch glass
containing 3- 4 drops of O. 5% aqueous Heidenhain's he matoxylin fo r
10-20 minutes, or until cell \.,Talls appear grayish-black when
observed in Hater under the micro scope . \~ash out excess stain
in three changes of distilled water. (If sections are over-
stained , destain in 2% iron alum, \.,Tashing this out carefully \.;rith
tap and distilled \.,Tater as before) . Trans fer sections to a \.,Tatch
glass containing distilled \.,Tater plus 4-5 drops of a 1% solution
of safranin in 50 or 70% alcohol, and leave in t he stain over-
night. Use three changes of distilled \.,Tater, then dehydrate in
50, 70, 95, and absolute alcoho l , allm.,ring about 5 minutes in
each . Transfer sections to equal proportions of absolute alco-
hol and xylol , and complete the clearing in t\.,TO changes of xylol,
ca . 5 minutes in each. Hount transverse, radial , and tangential
0
sections all on one slide in balsam . Dry slides on a 50 I"arm-
ing plate for 24 hours, then place lead I"eights on the cover
slips to force out the excess dammar and to f urther flatten the
sections while drying .
 182

Section B. LEAF ANATO~~

Carlquist (1961) has stated that the leaf is "perhaps anatomically the
most varied organ of angiospe rms, . . . " . It is not surprising, therefore , that
this organ possesses many anatomical features of potential taxonomic signifi-
cance . Although features of the l eaf arc often diagnostic at the genus and
spec ies level , these charac ter s cannot at our present sta t e of knowledge be
readily interpreted evolutiona ril y or accounted for by environmental condi-
tions . If fr esh o r prese rved specimens are available , caref ully prepared
freehand sections often revea l considerable a natomical detai l; however, par -
affin- embedd ed and sectioned leave s are most desirable . In many instances,
epidermal characters can be studied by making simp le epidermal peels . Among
the characters which have proven to be of systema tic value are the following:

I. CUTICULAR CHARACTERS .

Record distribution and orientation of papillae , striae , and rods .

II. EPIDERNIS.

The outer layer of ce ll s primary in origin. If it is multise riate

(multiple epidermis), only the outermost laye r differentiates epidermal
characteristics .
A. Uniseriate or multiseriate . Record number of cells in latter .
B. Thicknes s of cell walls. Record thin-walled , thick-walled or unevenly
thickened.
C. Sclerification of epidermis . Record presence or absence.
D. Size , shape and con tents of cells.
1. Isodiametric, pol ygonal or transverse l y extended in surface views.
Antic linal walls wavy or straight .
2 . Ce l ls in fi l es .
3 . Differentiation into long a nd short cells (Gramineae).
4 . Distribution of cork cells (Cramineae) .
5 . Epidermis papillose. Record occurrence on Im"er, upper, or both
surfaces ~ and size and shape of papillae .
6. Epidermis mucilaginous. Record occurrence and nature of mucilage.
7 . Distribution and nature of bulliform and colorless cells (Gramineae).
A bulliform cell is an enlarged epidermal cell occurring in
longitudinal rows of similar cell s in leaves of grasses .

III. STOHATA.

A pore in the ep idermis .

A. Subsidiary cells . Epidermal cells associated with a stoma and at

least morphologically distinguishable from the epidermal cells
composing the ground mass of the tissue. Also called accessory
cells .
1 . Classification of angiosperm stoma ta based on appearance of mature
stomata . Definitions after Van Co t them (1970). (See Figure 7-4).
a . Anomocytic (irregular-celled, ranunculaceous type). Stoma
surrounded by a limited number of cel l s that are indis-
tinguishable in size, shape, or form from those of the
rema inder of epidermis .
7 183

b. Anisocytic (unequal - celled, cruciferous type). Stoma sur-

rounded by three ce lls of which one is dis tinctly small e r
than the other ttw .
c. Paracytic (parallel-celled, rubiaceous type) . Stoma accom-
panied on either side by one or more subsidiary cells
parallel to the long axis of the pore and guard cells.
d. Diacytic (cross-celled, caryophyllaceous type). Stoma en-
closed by a pair of subsidiary cells whose common wall
is at right angles to the guard cells .
e . Tetracytic. Four subsidiary cells are present, two lat eral
and two terminal. Characteristic of many monocotyledons .
f. Actinocy tic . Stoma surrounded by a circle of radially elon-
gate subsidiary c e lls .
g . Cyclocytic. Stoma surrounded by four or more subsidiary
cells which form a ring around each stomata .
h. Hexacytic . Stoma accompanied by six subsidiary cells con-
sisting of two lateral pairs parallel to the long axis
of the pore and tHO polar (terminal) cells ; the second
lat eral pair as long as the stomatal complex . (Th is
type could be described as a modification of the tetra-
cytic type with an add itional pair of lateral subsidiary
cells. )
2 . Classification of angiosperm stomata ba sed on ontogeny.
a . Hesogenous . Stoma in ~"hich the subsidiary cells are pro-
duced from the same mer istemoid (initial) as the guard
cell initials .
b . Perigenous . Stoma in Hhich the subsidiary cells do not have
a common origin with the guard cells , but are formed by
cells lying around the meristemoid that divides to form
the guard cells.
c. Hesoperigenous . Stoma in which at least one of the subsidi-
ary cells has a common origin t.Jith the guard cells , but
the others do not .
d . Helicocytic . A type of mesogenous stoma in Hhich a helix of
four or more subsidiary cells surround the guard cells .
e . Allelocytic . A type of mesogenous stoma in which an alter-
nating complex of three or more C-shaped subsidiary cel l s
o f graded sizes occurs . This pattern may be further
divided into:
1 . parallelocytic. Guard cells develop parallel to the
subsidiary cells .
2 . diallelocytic . Guard cells develop at right angles to
subsidiary cells .
3. Classification of gymnosperm stomata based on ontogeny .
a. Haplocheilic. Stoma in t"hich the subsidiary cells are not
derived from the same meristemoid as the guard cel l s .
b . Syndetocheilic . Stoma in which the subs idiary cells are
de r ived from the same meristemoid as the guard cells .
B. Guard ce lls. In stoma . Two cells which by changes in turgescence
open or clos~t he stomatal opening. Variable features include
cell shape (symmetrical, slightly asymmetrical , or markedly
asymmetrical) and size .
C. Distribution. Stomata may be confined to the upper leaf surface
(ep i stomatic), lower leaf surface (hypostomatic), or occur on
both upper and 10l"er surfaces (amphistomatic) .
184 7

IV . HYPODERl>lIS .

Gene r al te r m for layer, or layers , of cel ls beneath the epidermis whic.h

are morphologicall y distinct from under l ying cor ti cal layer s. Record pres-
cence or absen ce , location , number of layers, and sclerotic condit i on .
..... , ..
V. SCLERENCIIYHA .

A. Sclere i d (5) . A sc l erenchyma cell , varied in form, b ut typically not

much elongated , and having thic k lignified secondary ",'a11s ""ith
many pits.
1. Types based on form (see Figure 7-4) .
a . As trosclere i d . A branched (ramified) t ype of sclereid .
b . Brach ysclereid . A short, ro ughly isodiametric sc l ereid .
resembling a pa renchyma cell in shape . A " stone cell. "
c . Filiform sclereid . A much e l ongated, slender sc l ereid resem-
bling a fiber.
d . l>lacrosclereid . A somewhat elongated sclereid with unevenly
distr ibut ed secondary \"all s .
e . Osteosclerei d. Scler eid hav ing the shape of a bone, with a
columna r middle pa r t and enlargements at bo th ends .
f. Tri chosclereid. A type of b ran ched sclereid with thin hai r -
like branches extending into intercellular. spaces .
2. Types based on distribution .
a . Diff use sclereid . Sc l ereid dispersed i n the leaf mesophyl1.
b. Termi nal sclereid . Sclereid confined to the ends of small
veins .
B. Fiber(s) . An elongated tapering sclerenchyma cell with a more or
less thick secondary wall , \~ith o r witho ut lignin .
1. Girders . A scle r enchyma t ous connection between a vascul a r bun -
dle and the epidermis .
2 . Incomp l e t e (partial) girder s . A sclerenchymatous extension from
the b undle sheath \"hich does not reach the epidermis .
3 . Strand . Sclerenchyma f r ee from the vascul ar bundle sheath and
I~hich may be subepidermal.

VI . NESOPHYLL .

Photosynthetic parenchyma tissue of a leaf located be t \"een epidermal

layers .

A. Tvpes.
1. Bifacial. Palisade parenchyma on one side of the blade and
spongy parenchyma on tbe othe r side .
2 . Iso l at er a l. Palisade parenchyma on both sides of the bla de .
B. Cons truction .
1 . Relative differentiation into palisade and spon gy l ayers and
number of layers in each .
2 . Distribution a nd shape of chlorenchyma .
3 . Shape of mesophyll ce lls. Record isod1ametr ic , s t ellate , thick-
wa lled, e t c .
4. Presence or abs ence of air-lacunae . Hesophy l l " loose " or " compact."
C. Presence , abs ence and distribution of secreto ry ca nals or cavitie s .
D. Form and distribution of c rystals (see Sect ion A. SECONDARY XYLEH) .
Figure 7- 4 185

STOMATAL TYPES

Quord
ce ll

Poro cyl ic

Anomoe ylic Anisocy l ic

Oio ey li e Aet inoe ytic Tetrocy ti c Cyclocy'ic

SCL E REID TYPES

Aslros elereid 8roc hy sclereid Filifor m $clereid

D= < =3
~ '~',
Mac roscler eid Osleoscle re id Trichosclereid
 186 7

VII . SILICA BODIES .

Record shape and distr i bution in all tissues . Characteristic of

mono c otyledons .

VIII. VENATION .

A. Petiole . (see Section C) .

B. Lamina . (see Section D) . Record major venation pattern, minor vena-
tion patte rn , including islet size and veinlet termination number,
and nature of vein sheathing (parenchymatous , sclerenchymatous ,
or double sheaths of one or both of these types , and size and
shape o f sheathing cells) .

IX . TRICHOl>lES . (See Section E) .

X. OTHER CHARACTERS .

A. Presence , absence and size of hydathod e s. A hydathode is a water

pore or water gland, a structure that releases water.
B. Presence or absence of nectaries . A nectary is a multicellular
glandular structure secreting a sugary liquid . Occurs on
flowers (floral nectary) or on vegetat ive plant parts ( e xtra-
floral nectary) .
C. Ecological modification such as presence and structure of stomatal
crypts .

LEAF ANA'roHY LITERATURE

Bokhari , N . H., and B. L . Burtt. 1970. Studies in the Gesneriaceae of the

Old Horld XXXII : Foliar sc1ereids in Cyrtandra . Notes f rom the Royal
Botanic Garden , Edinburgh 30 : 11 - 21 .
Payne , H . H . 1970 . Helicocytic and allelocytic stomata : unrecognized
patterns in the Dicoty l edonae . American Journal of Botany 57 : 140-147 .
Van Cotthem, toJ . R. 1970 . A classification of stomatal types. Botanical
Journal of the Linnean Society 63 : 235-246.
1971. Vergle ichende morphologi~ch e Studien uber Stomata and
e i ne neue K1assifikation iherer Typen . Berichte del' Deutschen Botanischen
Gesellschaft 84 : 141 - 168 .

TECHNIQUES ENP LOYED IN THE STUDY OF LEAF AND FLORAl, ANATOl'fY

1. CLEARING.

In addition to the standard microtome sectioning procedures , the tech-

nique of "clearing" leaves and fl oral parts in order to study vasculation
patterns is extremely useful and widely employed . This techniqu e is appli-
cable to fresh, preserved, or dried (herbarium) specimens . See Payne , l~ .
1969 . A Quick Nethod f or Clearing Leaves . \~ard ' s Bulletin 8(61) : 4-5 .
The following schedule should g ive good results.

1. Plant parts should be small enough to be placed in a petri dish (in

the case of leaves some cutting may be required) . Dried or liquid-
preserved materials need no special prior treatment; however , fresh
leaves should be boiled in alcohol or placed in Carnoys (95%) for
a while, to remove chlorophyll before the clearing procedure .
 7 187

2. Flood tissue with 5% NaOI1 (or KOH) and place in an oven at about 37 0 C.
fo r one to many days depending upon the nature of the ti ssue .
3. After tissue ha s become transparent , or nearly so , \va sh briefly in wate r .
NOTE : some plan t s contain various dark pigments that can be remove d
at thi s pO int by flooding tiss ue with Stockwe l l ' s Bleach . St ockwell IS
is composed 0 f::
Hater ...... . ........... .... .. . 90 cc .
Potassium bichromate ......... . 1 g .
Glacial acetic acid ...... . ... . 10 cc .
Chromi c acid ................. . 1 g .
This solut i on s houl d r ema in on the t i ss ue fo r one to severa l hours at
r oom tempe r ature until all pigments are r emoved . Pour off bleach and
~lash thoroughly .
4. Some find it desirable at this point to use a conce ntrated chloral hydrate
solution which will continue the clearing process if the tissue is not
complete l y transpar ent . In many cases t his is not necessary .
5. Tissue should nOl." begin to be dehydrated by pouring on 30% and 50% ethyl
alcohol for 5 minutes each .
6. Staining is accomplished by using 1% Safra nin in 50% alcoho l. Generally
only a fe\." minut es in the stain is sufficie nt with gentle agitation .
7. Continue dehydration using 70% and 95% alcohols . The alcohols will de-
sta i n (Le . , r emove the safranin) so care should be taken not to leave
t hem on the ti ssue too long . Remembe r, hOl."ever , the t i ssue must be
completely dehydra t ed .
8. Pour on absolute alcohol for several minutes . Very little destaining
wil l occur in the highe r alcohols.
9. Pour on a solu tion of 1 /2 ahs olu te alcohol and 1/2 xylene for several
minutes .
10 . Clearing is the final step af ter dehydration is complete using xyl ene .
If tissue is not completely dehydr ated it will t urn a mi l ky \."hite;
if this happens, go ba ck to absolute alcohol.
11 . 1I0unt on slide usin g a synthetic res i n or ba l sam . For some flora l parts
it is some times desirabl e not to mount tissue on a s lide but store in
a v i al so all aspects and viet."s of the vasculation pa ttern can be
studied.

II. EPIDERHAL PEELS .

Although various techniques have been pub lished for making epidermal
impressions (see previou sly cited paper by Payne , 1970 , fo r a method
using c ellulose acetate), epidermal cell patterns can f r equently be
studied b y si mply taking a fresh or preserved leaf and with the aid of
a sharp r azor b l ade peeling off the adaxial and a baxial layer , mounting
in water , and observing under the mic r oscope . Since this preparation is
a t empo rary one it is desirable to prepal"e a sketch of the epidermis using
a camera lucida .

TECHNIQUE LITERATURE

Arnott, H. J . 1959 . Leaf clearings . Turtox News 37 : 192 - 194 .

Lersten , N. R. 1967 . An annotated bibliography of bo t anical clearing m{~c hods .
Iowa Sta te Journal of Science 41 : 481- 486.
Payne . Ioj". w. 1969 . A quick method for clearing leaves. 11ard 1 s Bullet J.I1
8(61), 4- 5 .
 7
188 )

LEAF ANATOMY EXERCISE

Species I Species II i,
f
Cutic ular Characters tl
o
t'
Uniseriate or
c·
Multiseriate
c
u
d
Thickness of walls
c
Epidermis
i
Sclerifi cation +-

Size, shape, content

of cells

Trichome Type

Stomatal Type (prepare drawing)

1
Hypodermis + -
(number of layers if +)

Sc.lerenchyma
(type and distribution)

Sifaeial or isolateral 1:

Number of palisade layers

Mesophyll

Shape of ce ll s

Crystals (type and

distribution)

~~jor Venation Type

Hiner Venation Pattern

Type of Vein Sheathing

Other Features of In terest

 7 189

Section C. PETIOLE ANATOHY

The taxonomi c usefulness of pe tiole vasc ula rization patterns has become
i nc r eas in g l y apparent in recent years . The most comp l ete system of c lassi-
fication of majo r venation pattern s i n t h e petiole of wood y dicotyled ons is
that of Howard (1962). This c l assification is presented below. I n st ud i es
of this nature , it is important to examin e sect ions throughout the length of
the petiole to unde rstand completely the changes wh i ch occ ur in the vascular
configuration . It will be no t ed , accordingly, that t he system below considers
c hanges wh ich occur i n the vascular ti ssue from the time it leaves t he stem
until it become s relativ ely stable in the leaf . NOTE : This c l assif ication
does not account fo r all of the variation patterns that may exist , but does
cove r the more common maj or patterns . Also, terminology with respect to pet -
iole vasculation patter ns has not been standardiz ed (see Figure 7-5) .

t. NODE 3 : 3 ( SEE SECTION F- I), BUNDLES FREE .

A. Three b undles th roughout petiole; e . g . , Pedilanthus .

B. Lateral traces divide .
1 . Petiole with five free traces; e . g ., Pittosporum.
2 . Petiol e '''ith many traces in " U" -shaped configura tion; e . g . , Niconia.
3. Free traces in a ring with medullary bundle s ; e . g ., Aesculus .
4. Free traces in a ring without med ullary bundles ; e . g . , Sambucus .
C. Nedian trace divides and the division products ass ume a dorsal posi
t i eD; e.g. , Hibiscus .
II . NODE 3 : 3 , BUNDLES FUSE TO FORN AN ARC .

A. Bundles f use and form flat arc; e . g . , Lo nice ra.

".
C.
Bundle s
Bundles
fuse
fuse
a nd form a flat a r c with dorsal free traces; e . g . , Cornu s .
in !l U" shaped arc; e . g . , Betula .
D. Bundles f u se and invag i nate a t e ad s ; e . g . , Congea .

III . NODE 3 : 3, BUNDLES FUSE TO FORN A SIP IIONOSTELE .

A. Siphonostele Simpl e .
1 . Formed by simp l e fusion of t races ; e . g . , Co t i nus.
2 . Formed s ubsequent to division of the median, the b r a nc hes of
which fo r m dorsal bundles ; e . g . , Acer .
B. Siphonostele with accessory bundles .
1. One la r ge accesso r y bundl e si tuated dorsa l ly; e . g . , Hamame l is .
2 . Small multiple accessory bundles dorsal ; e . g . , Carya .
3 . Accessory bundles in med ulla r y position ; e . g ., Bauhinia.

IV . NODE 3:3 , BUNDLES FUSE TO Foru-1 HaRE COHPLEX PATTERNS .

A. By i nvagina t ion fo rming one or many med ul l ary bundles or plates;

e . g . , Que r c u s or Ti lia
B. Siphonostele inv:aginat i ng forming included or dor sa l accesso r y bun-
dle; e . g ., Fagus.
C. Siphonos t e l e fo rmed , then late r al invagina t ions give rise to dorsal ,
smalle r siphonostele or plat e over "u" shaped arc ; e . g ., Ca r pinus .
D. Polystelic typ es .
1 . Axillary bud included , pet iole not compressed; e . g ., Platanus
type .
188 7

LEAF ANATOMY EXERCISE

Species I Species II

Cuticular Charact e rs

Uniseriate or
Multiseriate

Thickness of walls
Epidermis

Sclerification +-

Size , shape. content

of cells

Trichome Type

Stomatal Type (p r epare drawing)

Hypodermis + -
(number of layers if +)

Sclerenchyma
(type and distribut ion)

Bifacia l or isolateral

Number of palisade layer s

Mesophyll

Shape of cells

Crystals (type and

distribution)

Major Vena tion Type

Hinor Venation Pattern

Type of Vein Sheathing

Other Feat ures of Interes t

 7 189

Section C. PETIOLE ANATO~N

The taxonomic usefulness of petiole vascularization patterns has become

increasingly apparent i n recent years . The most complete system of classi-
fication of major venation patterns in the petiole of l,roody dicotyledons is
that of Howard (1962) . This classification is presented below. In studies
of this nature, it is important to examine sections throughout the length of
the petiole to understand completely the changes which occur in t he vascular
configuration . It will be noted, accordingly, that the system be l ow considers
changes which occur in the vascular tissue from the time it leaves the stem
until it becomes relatively stable in the leaf . NOTE : This classification
does not account for all of the va r iation patterns that may exist, but does
cover the more common major pa t terns. Also , terminology with respect to pet-
iole vasculation patterns has not been standardized (see Figure 7- 5).

1. NODE 3 : 3 (SEE SECTION F-I) , BUNDLES FREE .

A. Three bundles throughout pe tiole; e. g., Pedilanthus .

B. Lateral traces divide .
1 . Petiole with five free traces; e . g . , Pittosporum .
2 . Petiole Ivith many traces in "U"-shaped configuration ; e . g. , Hiconia .
3. Free traces in a ring with medullary bundles ; e.g. , Aesculus .
4. Free traces in a ring without medullary bundles ; e . g ., Sambucus.
C. Hedian trace divides and the d i vision products assume a dorsa l posi
tion j e.g ., Hibiscus .

II . NODE 3 : 3, BUNDLES FUSE TO FOR}[ AN ARC .

A. Bundles fuse and form flat arc; e . g . , Lonicera .

B. Bundles fuse and form a flat arc with dorsal free traces ; e . g . , Cornus .
C. Bundles fuse in "U" - shaped arc; e . g . , Betu l a .
D. Bund l es fuse and invaginate at ends; e.g. , Congea.

III. NODE 3 : 3 , BUNDLES FUSE TO FORM A SIPHONOSTELE .

A. Siphonostele simple .
1. Formed by simple fusion of traces; e . g . , Cotinus .
2 . Formed subsequent to division of the median, the branches of
which form dorsal bundles ; e.g . , Acer .
B. Si phonostele with accessory bundles .
1 . One large accessory bundle situated dorsal l y; e . g ., Hamamelis.
2 . Small multiple accessory bundles dorsal; e . g . , Carya .
3 . Accessory bundles in medullary position; e . g . , Bauhinia .

IV . NODE 3 : 3, BUNDLES FUSE TO FOR}1 HORE COHPLEX PATTERNS .

A. By invagination forming one or many medul l ary bundles or plates;

e . g., Quercus or Tilia
B. Siphonostele inv.aginating forming included or dorsal accessory bun-
dle ; e.g., Fagus.
C. Siphonostele formed , then lateral invaginations give rise to dorsal,
smaller siphonoste l e or plate over "u" shaped arc ; e.g ., Carpinus .
D. Polystel i c types.
1. Axillary bud included , petiole not compressed; e . g., Platanus
type.
102 6

Tepal. A perianth member or seg- Hhor! . A cyclic or acyclic group

ment; term used for perianth of sepals, or petals, or sta-
parts undifferentiated into mens, or carpels.
distinctive sepals and petals .

B. Flm.,rer Types _v"- (Figure 6-6 , p . 101)

(Classi f ication bas e d on evolutionary flOt.Jer pollinator relationships--from
Leppik [1957])
Actinomorphic . Flmlers \vith radial hemispheric form ; petals or tepals
symmetry and parts arranged at colored; parts numerous; e . g .,
one level; \-lith definite numbers Nymphaea, Hagnolia .
of parts and size; e.g . , Anemone , Pleomorphic . Ac tinomorphic wi th num-
Caltha . bers of parts r educed; e.g ., Tri-
Amorphic or Paleomorphic . FloHers pogandra .
Hithout symmetry; usually \·lith an Stereomorphic . Flowers 3-dimens ional
indefinite number of stamens and I>'ith basically radial symmetry;
carpels, and usuall y subtended by parts many or reduced, and usually
bracts or d i scolored upper l e aves; regular ; e . g ., Narcissus , Aguilegia.
e . g .• Salix discolor , Echinops Zygomorphic . Flowers with bilateral
ritro (mos tly fossil forms) . symmetry; parts usually reduced in
l:Iaplomorphic. Flowers ,,,ith parts number and irregular; e . g ., Cypri -
spirally arranged at a simple pedium, Salvia.
l evel in a semispheric or

VIII. PERIANTH AND I-lYPANTHIUN

A. Perianth Parts (Figure 6-5 , p . 100)

Anterior Lobes . The l obes al>'ay from Hood . A cover- shaped perianth part,
axis, t01·,rard the sub tending bract; usually Hith a turned d01m margin .
abaxial lobes. Horn. A curved, pointed and hol101'"
Anterior Ridges, Lines , Grooves. protuberance from the perianth.
The lines, grooves , ridges in or Hypogynium. Perianth-like structure
on the dorsal side, abaxial , of bony scales sub tending the
within the pE'rianth . ovary, as in Seleria and other
Base . Bottom or lower portion . members of the Cyperaceae .
Beard . A tuft , line or zone of Keel. The two united petals of a
trichomes. papilionaceous flO1"er; any strue-
Bristle . A stiff, strong trichome , ture ridged like the hot tom of a
as in the perianth of some members boat .
of the Cyperaceae . Ligule . The strap-shaped portion of
Callosity . A thickened, raised area, a ray or ligulate corolla.
,,,hieh is usually hard ; a callus. Lip or Labellum. Either of tHO
Car ina . Keel. variously shaped parts into which
Cla'v. The long , narrow petiole- a corolla or calyx is divided,
like base of a sepal or petal . usually into an upper and lower
Corona . A crown; any outgrol>'th lip, as in the Lamiaceae and
between the stamens and corolla Orehidaceae .
which may be petaline or stami- Limb . Expanded portion of corolla
nal in origin . or calyx above the tube, throat
Dorsal Side . Back or abaxial side, or claw .
or the lower side of a perianth Lobe . Any, usually rounded, segment
part . or part of the perianth .
Faucal Area . The throat area . Lodicule. Abortive perianth part in
Fringe . The modified marg i n of a the Poaceae ; hyaline scales at ba s e
petal , sepal , tepal or lip. of ovary in the Poaceae .
6 103

Palate. The raised area in the Spur . A tubula r or pointed pr ojec -

throat of a sympetalous corolla. tion from the perianth.
Pa ppus. Bristly or scaly calyx in Standard. Banner . or Vexillum . The
the Asteraceae. upper. usually \.,ide petal in a
Petal . A coro l la member or segmen t; papilionaceous corolla.
a unit of the corolla. Tepal. A member or segment of per-
Posterior Lobe . The lobe next to ianth in which the parts are not
axis, away from the subtending differentiated into distinct sepals
bract; adaxial l obe . and petals .
Posterior Ridges , Lines , Grooves . Throat . An open , expanded tube in
The lines, gr ooves , ridges in the perianth .
or on the ventral side , adaxial, Tube. The cylindrical part of the
\"ithin the perianth. per ianth .
Pouch or Sac. A bag-shaped struc- Ven t ral Side . Top side or upper
ture . side of a perianth part .
Scal e . Small , scarious to co r ia- Hing. Lateral pe t als , as in the
ceous flatte n ed bodies wit h i n Fabaceae; a f l attened extensio n,
the perianth , as in the Cypera- appendage or pr ojection from a
ceae and Asteraceae . perianth part .
Sepal . A calyx member or segment;
a unit of the cal yx .

B. Hypanthium Parts
Base . Bottom or l ower portio n of Neck. Narrm.,ted portion of hypanthium,
the hypanthium . between the base and a f l a r ed l imb .
Limb . Free , f l a r ed portion of T~be or Casing . Cylindrical part of
the hypanthi um. the hypanth i um .

C. Perianth Types
(Classification based primarily on fusion of parts)
Ach lamyde.ous . Without perianth . Di c hlamyde.ous . With perianth c omposed
Apetalous . No petals or coroll a . of distinct calyx and coro lla.
Apopetalous or Choripetalous . Homochlamydeous . With perianth com-
tnth separ ate petal s . posed of similar parts , each part
Aposepal ous or Chorisepalous . a tepal .
With separate sepals. $ympetalous. Wi t h fused petal s .
Asepalous . No sepals or calyx . Synsepalous . With fused sepals .
Chlamydeous . j·lit h perian th o

D. Pertanth and Hypanthium St r uctural Types (Figure 6-6 , p . 101)

Actinomorph ic. \o/'ith radially ar Coronate. Tubul ar or fla r ing perlanth
ranged perian t h parts; ray-l i ke or stamin al outgr m"th; petalo i d
f i gure . appendage .
Bilabiate . Two -l ipped, with two Cru ciate. Four separate petals in
unequa l div i s i ons . cross form .
Calcarate . Spurred . Cucul la te . Hooded .
Calceolate . Slipper-shaped , as in Galeate . Helmet - shaped , as one sepal
the coro lla of Cypripedium . i n Aconitum .
Campanulate. Bell shaped; l11th Gibbous. Inflated on one side near
flaring tube about as broad as the base .
long and a flaring limb. Globose . Round .
Carinate . Kee l ed . I n fundibular. Funnel-shaped.
Cor niculate . Horned . Ligulate or Ray . Strap-shaped .
104 6

Papilionaceous. '.lith large pos - Saccate . Pouch-like.

terior petal (banner or stan - Salverform . Trumpet-shaped . Hith
dard) tlW lateral petals (1),1ng5) slender tube and limb nearly at
and usually two connate lOHer right angles to tube .
petals (keel) ; as in the Fabaceae . Subgl obose. Almost round or spheri-
Personate . THo - lipped with the upper
arched and the lower protruding
cal. -.
Tubular. Cylindrica l.
into corolla throat . Unguiculate. Clawed .
Rotate. ~'1heel - shaped , ~1ith short Ur ceolate. Urn-shaped .
tube and wide limb at right Ventricose . Inflated on one side
angles to tube . near the middle.

IX . ANDROECIUM

A. Androecial Parts
Stamen . Hale sporophyll within the Stamina! Disc . A fleshy, elevated
flower; f l oral organ that bears cushion formed from coalesced
pollen in angiosperms . staminodia or nectaries .
Staminodium . Steri l e stamen, may be
modified as a nectary or petaloid
structure.

B. Androecial Types (Figure 6-7 , p . 105)

(Classification based primarily on fusion of parts)
Apostemonous . With separate sta- Monadelphous . Pith one group of
mens . stamens connate by their fila -
Diadelphous. l-lith two groups of ments .
stamens connate by their fila- Petalostemonous . With filaments
ments. fused to corolla. anthers free .
Gynandrial or Gynostemial. With Polydelphous . l"Tith several groups
fused stamens and carpels of stamens connate by their fila-
(stigma and style) as in ments .
the Orchidaceae. Syngenesious . Hith fused anthers .

C. Stamen Parts
Anther. Pollen-bearing portion Filament . Stamen stalk .
of stamen .

o. Stamen Structural Types (Figure 6-7 , p . IDS)

(Classification based pr imarily on structure of fi l ament and anther)
Note : In this classification intermediate types of stamens do occur ; shapes ,
apices , and bases of anthers sho uld be described separately and indepen -
dently of stamen type.
Appendicular . Typical stamen with a Petaloid . Petal - like stamen with-
variously-shaped or modified, pro- out distinct anther and filament
truding connective, as in Viola . but with marginal microsporangia,
Laminar . Leaf-like stamen without a as in Magnolia nit ida.
distinct anther and filament but Filantherous or Typical . Stamen
with embedded or superficial ivith distinct an t her and filament
mi crosporangia , as in Degeneria. ~.,ith or without thecal appendages .
Petalantherou$ . With a terminal as in Rhexia or Vacciniurn .
ant her and distinctly petaloid
fi l ament, as in Saxifraga .
Figure 6-7 105

ANDROEcrUM

STAMEN POSI T IO N

Aolisepolous Cryptootherous Diplostemonous Epipetolous Ph one rantherc us

(e . s~'l e d)

STAMEN AR RAN GEMENT ANQROECIAL TYPES

DidYnOrnOuS FOsc'lcled Tetrodynomous Di odelphous Monodelphous Syngenesious

STAM EN STRUCTURAL T Y PES THECA L ARRANGEMENT

Xl. .1

!~~
nin
/Y

Fi 10 nlheraus Tr ansve rse

Appendicular Lam inar Pela laid (Typical) Oivergent Oblique Poralle l Or Explan ale

ANTHER TYPES and A TTA CHM ENT

E.lrcrse Introrse Poricidal Trans verse Bosifited Dcrsifi.ed

106 Fi g u re 6-8

GY N O E C IUM

GY NO E CIAL T Y PES

w@ M~ 1~ l® 1~w A~
Apoca r po~s Semica rpou5 Syncarpous Synova"ous Syns tylov arLOus Unic a, pello us

C h O l a,o~_ CAR P EL and OVULE

'. \ "o,jpodol
po lo. nu(: l ei - { , I~
,".~
\ . te l l
owo. l l u <
ou Te r __ _
; n t"~um"" \ \
i!-U
\ j , I I .mbrjo.ac
0. 9
r ap he

'n n ~ r
i n t~Qum.nl - ' - '/ " /" . - ccru nCle

m ic rop yl e - - - - - - - - -

CARPE L T YP E S

n,,,,m.
~
" , . mo

""m.
4
, ~ ' l y le

~
."'m. -<- s t yle

~.d;cel - .. ,p.
(~p e ~iC" 1 - pod ic . ' _
_ " 'p e
Hd;cel

Ast ylocorp ell ous Asty loCa r pepod ic Slylocorpellous Sty lacor p epO d l c

OV ULE TY P ES

~ ~,r .".,
Amph i tropous
bo dy
..:- fu n ic ulu"
A.no tro pous
~ """ "",m~"
~
Compylotro po us
Hem l<¥\o tropous
or Homitropous Or t ho trap ou s

S TIGMA T Y PES

9 V (jO
Ca pit a te Cl ava te Crestea Decu rr ent

Discoi d Fi mbriale Lin€ote Lo be d Terete

STYL E TY P E S

Condu pli co\ e

@
Eccen t"c

HeterQSTyl ous Homos tylous Pet o io ,d Stylo podic Tcrcte Tu ber cu lole Umbrncu\ote
6 107

E. Anther Parts (Figure 6-7, p . 105)

Connective . F11amen t extension '-"=='--'P"o""l'l:':en Sac . Hale sporangium .
between thecae. Theca . One half of an t her con-
Locul e . Compa rtment of an anther. taining tt-IO pollen sacs or male
Polle n Grain. Young ma l e gameto- sporangia .
phyte .
F. Anther Types (Figure 6-7. p . lOS)
(Classification based on dehiscence)
Longitudinal. Dehiscing along long Pericidal. Dehiscing through a
axis of theca . pore at apex of theca .
Ex trorse . Dehiscing longitudi- Transverse . Dehiscing at right
nally outward . angles to long axis of theca .
In trorse. Dehiscin g longitudi- Valvular . Dehiscing through a pore
nally inward. covered by a flar of tissue.
Latrorse . Dehiscing l ongitudi-
nally and laterally.

G. Anther Attachment (Figure 6-7 , p . 105)

Basifixed . Anther attached at its Subbasifixe d. Anther attached near
base to apex of filament . its base to apex of filament .
Dorsifixed. Anther attached dor- Versatile . Dorsifixed but anther
sally and medially to apex of seeming ly swinging free on the
filamen t . filamen t .

H. Pol l en
(See pal ynology in Chapter 8)
Dyads . Grains occurring in clusters Polyad . Grains occurring in groups
of two . of more than four .
Filiform . Thr ead-like . Tetrads. Grains occurring in groups
Nonad . Grains occurring singly . of four.
Pollinia . Grains occurring in uni-
form coherent masses .

X. GYNOECIUN

A. Gynoecial Parts (Figure 6-8, p . 106)

Carpel. Female sporophyll wi t hin Placenta. Ovule-bearing region of
f l ower; f lora l organ that bears ovary wall .
ovules in angiosperms . Stigma . Pollen-receptive portion of
Carpopodiurn. Short , thick , pistil- pistil.
late stalk. Stipe . Pistillate stalk .
Locule . Ovary cavity . Style . Attenuated, non - ovule-bearing
Ovary . Ovule-bearing part of pistil . portion of pistil between stigma
and ovary .
B. Gynoecial Types (Figure 6-8. p. 106)
(Classification based on fusion)
Apocarpous . l-lith carpels separate . styles and stigmas separate .
Semica rpous . With ovaries of adja- Synstylovarious . With ovaries and
cent carpels partly fused , stig- styles of a djacent carpels com-
mas and styles separate . pletely fused, stigmas separate .
Syncarpous . l.,Tith stigmas , styles , Unicarpellous or Stylodious . Wit h
and ovaries completely f us ed . solitary, free carpel in gyno e-
Synovarious. With ovaries of adja- cium.
cent carpels completely fused ,
108 6

C. Carpel Parts (Figure 6-8, p. 106)

Funiculus . Stalk by which ovule is Placenta . Ovule- bearing region of
attached to placenta. ovary wall.
Lacule . Ovary cavity. Stigma. Pollen receptive portion of
Ovary. Ovule-bearing part of carpel carpel .
in simple ovary . Stipe, Podogyne, Carpopodium . Basal
Ovule. Embryonic seed consisting of stalk .
integument(s) and nucellus . Style . Attenuated portion of carpel
between stigma and ovary.

D. Carpel Types (Figure 6-8, p. 106)

(Classification based on presence or absence of style and stipe)
Astylocarpellous. Hithout a style Stylocarpepodic. With a style and
and a stipe . stipe.
Astylocarpepodic . Hithout a style, Stylocarpellous. With a style and
with a stipe . without a stipe, the normal carpel.

E. Stigma Types (Figure 6-8, p . 106)

(Classification based on shape)
Capitate. Head - like . Discoid. Disc-like.
Clavate. Club -shaped. Fimbriate. Fringed.
Crested or Cristate. Hith a termi- Lineate. In lines, stigmatic sur-
naI ridge or tuft. face linear .
Decurrent. Elongate, extending Lobed . Divided into lobes.
downward . Plumose. Feather-like .
Diffuse . Spread over a wide surface . Terete. Cylindrical and elongate.

F. Style Types (Figure 6-8, p. 106)

(Classification based primarily on shape)
Astylous. Style absent. Homostylous. Wi t h styles of same
Conduplicate. Folded with a 10ngi- sizes or lengths and shapes .
tudinal groove . Involute. Hi th margins infolded
Cristate. Crested . longitudinally , ~"ith groove
Eccentric . Off-center style. present.
Fimbriate. Fringed. Petaloid. Petal- like .
Flabellate . Fan-shaped. Stylopodic. Hith a stylopodium or
Geniculate. Bent abruptly . discoid base, as in the Apiaceae .
Gynobasic. Attached at base of Terete. Cylindrical and elongate.
ovary in central depression. Tuberculate . With hard , swollen, per-
Heterostylous. With styles of dif- sistent base or tubercle .
ferent sizes or lengths or shapes Umbraculate. Umbrella-shaped, as in
within a species. Sarracenia .

G. Ovule Parts (Figure 6-8, p . 106)

Chalaza . End of ovule opposite Nucellus. Female sporangium within
micropyle. ovule; megasporangium in seed
Embryo Sac. Female gametophyte. plants .
Integuments. Outer covering of Raphe . Longitudinal ridge on outer
ovule; embryonic seed coat. integument.
Micropyle. Hole through integument (5) .
6 109

H. Ovule Typ~s (Figure 6-8 , p . 106)

(Classification based on orientation of ovule body in relation to the funicu -
Ius and micropyle)
Amphitropous. With body bent or Hemianatropous or Hemitropous. Hith
curved on both sides so that the body half-inverted so tha t fun i -
micropyle is near the medially culus i s attached near middle with
attached funiculus. micropyle terminal and at right
Anatropous. Hith body completely angles .
inverted so that funiculus is Orthotropous or Atropous . Hith
attached basally near adjoinin g straight body s o that funicular
micropyle area . attachment is at one end and
Campylotropous. With body bent or micropyle at other .
curved on one side so that micro-
pyle is near medially attached
f uniculus .

XI. FRUITS

A. Fru i t Parts
Carpophore . Floral axis extension Placenta. Region of attachment of
between adjacent carpels , as in seeds on inner fru i t wall.
the Apiaceae . Replum. Persistent septum after
Ectocarp or Exocarp . Outermost dehiscence of f ruits, as in
layer of pericarp . the Brassicaceae .
Endocarp . Innermost diffe r entiated Retinaculum , Jaculator or Echma .
layer of pericarp. A persistent indurated , hook-
Funiculus . Seed stalk . like fun i culus in the fruits of
Hericarp . A portion of f ruit that Acanthaceae.
seemingly matured as a separate Rostel lum or Beak. Persistent styl ar
fruit . base on f ruit.
Hesocarp . Hiddle layer of pericarp . Seed . A matured ovule .
Pericarp . Fruit \o,Ial1. Septum or Dissepiment . Partition.

B. Fruit Structural Types

(C lassification based pr i marily on orLgLn , texture, and dehiscence ; types
grouped as simpl e , aggregate , multiple, accessory.)

1 . Simple Fruits
(Fruit derived from the ovary of a solitary pistil in a single flower)

a. Dr y I ndehiscent Fruit Types (Figure 6 - 9, p . 112)

(Fruits that do not split open at matur ity)
Achene . A one-seeded, dry , inde- Capsule, Indehiscent . Dry f r uit
his cent fruit with seed attached derived from a two- o r more
to fruit wall at one point only, loculed ovary, as in Peplis .
derived from a one- loculed super- Caryopsis or Grain . A one seeded
ior ovary . dry, indehiscent fruit with the
Balausta . Hany-seeded , many- seed coat adnate to the fruit
loculetl indehiscent "ftuit with wall , derived from a one-Ioeul ed
a tough , leathery pericarp , as superior ovary.
in Puniea. Cypsela . An achene derived from a
Calybium . A hard one-Ioculed dry one- Ioculed , inferior ovar y .
fruit derived from an inferior Nut . A one-seeded , dry , indehiscent
ovary , as in Quercus . fruit with a hard pericarp , usual l y
derived f rom a one - l ocu l ed ovary .
110 6

Nutlet . A small nut . Utricle. A small , bladde ry or in-

Samara . A t..dnged, dry fruit . flated , one-seeded. dry fruit .

h. Dry Dehiscent Fruit Typ es (F igure 6-9 , p . 112)

(Fruits that split open at matur ity)
Caps ule. Dry, dehiscent fruit de- Loment . A ~ legume that separates
r ived from a compound ovary of transversely between seed sec-
2 or mor e carpels . tions.
Diplotegium . A pyxis derived from Si l ie le . A dry, dehiscent fruit
an inferior ovary. derived from two or more carpels
Fol lic l e . A dry, dehiseent fruit that dehisce along two sutures
derived from one carpel that and which has a persisten t par-
splits along one suture . tition after dehiscence and is as
Legume . A u sually dry , dehiscen t broad as, or broader , than l ong .
fruit derived from one carpel Silique . A silicle type fruit that
that splits along two sutures. is longer than broad .

c. Capsule Types (Figure 6- 9 , p. 112)

(Classificat i on based on type of dehiscence)
Acrocidal Capsule. One that dehis- Loculicidal Capsule. One that dehis-
ces through terminal slits, or ces longitudinally into the cavity
fissures , as in Staphyl ea. of the locule, as in Epilobium .
Anomalicidal or Rupturing Capsule. Ope rculate Capsule. One that dehis-
One that dehisces irregularly, ces through pores, each of which
as in Ammannia. is c overed by a flap , cap , or
Basicidal Capsule. One that de- lid , as in Papaver .
hisces through basal slits or Poricid al Capsule. One that dehis-
fissures, as in some species ces through pores , as in Trio-
of Ari stolochia . danis.
Circumsc i ssle Capsule or Pyxis . Septicida l Capsule. One that dehis -
One tha t dehisces c ircumfer- ces l ongitudinally through the
entia lly, as in Plantago. sep ta, as in Pen stemon.
Denticidal Capsule . One that de- Valvular or Septifraga l Cap sule.
hisces apically , leaving a One with v alves breaking away
ring of teeth, as in Cerastium . from the septa, as in Ipomoea .
Indehiscent Capsule . One t hat does
not dehis ce at maturity , as in
Pepli s.

d . Schizoca rp ic Frui t Tvnes (Figure 6-9 , p . 112)

(Fruits der ived from a simple , two- or more locul ar compound ovary in which
the locules separate at fr ui t maturity simula ting fruits derived from the
ovar ies of simple pistils)
Schizocarpic Achenes. Separating Schizocarpic Carcerules . Separating
achenes which are one-seeded , carcerules which are dry , few-
dry , indehiscent fruits with seeded , indehiscent locules , as
seed attached to fruit wall at in Althaea .
one point only , derived from a Schizo carpic Follicles . Separa ting
superior ovary , as in Sidalcea . follicles which are dry , dehis-
Schizocarpic Berries. Separat ing cent fruits der ived from one
ber r ies which have a fles h y caroel, splittin~ a long one suture ,
perica rp, as i n Phytolacca . as in Apocynaceae .
6 III

Sch izocarpic Hericarps . Cremocarp . indehiscent 4- parted fr uits with

or Carpopodium. Separa ting a hard peri carp around a gynobasic
mer icarps which are dry, seed- sty le, as in the Bor aginaceae and
like fruits derived f rom an i n- Lamiaceae .
ferior ovary , as in the Ap i aceae . Schizocarp ic Samaras . Separa ting
Schi'zoca rpi c Nutlets . Cenobium. samaras {"hieh are winged , dry
Separating nutlets wh ich are dry, fr uit s, as in Acer .

e . Fleshy Fruit Type s (Figure 6-9, p . 112)

Amphisarca . A b e rry -like succulent Hesperidium . A th ick- s kinned septate
fruit with a c ru staceo us or berry with the bulk of the fruit
woody rind , as in Lagenaria. de rived from glandula r hairs, as
Berry. Fleshy fruit, with succulent in Citrus .
pericarp, as in Vitis . Pepo . A berry \·lith a l eathery non-
Drupe . A f leshy fr uit with a stony septate rind derived from an in-
endocarp. as i n Prunus. ferio r ovary , as in Cucurhita.
Drupele t. A small drupe, as in Pyrene . Fleshy fruit with each seed
Rubus . surrounded by a bony endocarp, as
in I l ex .

2 . Aggregate Fruit Types (Conocarpium) (Figure 6- 10 , p . 113)

(A group of separate fruits developed from carpels of Otle flower)
Achenecetum . An aggregation of Follicetum. An aggregation of
achenes , as in Ranunculus . foll i cles, as in Cal tha .
Baccacetum or Etaerio . An aggre- Samaracetum. An aggregation of
ga tion of be rries , as in Actaea . samaras, as in Li riodendron .
Drupecetum . An aggregation o f
drupelets , as in Rubus .

3 . Hul tiple Fruit Types

(Fruits on a common a xi s that are usually coalesced and der ived f rom the
ovaries of several f lowers)
Bibacca . A fused double be rry, as Sycon ium . A syncarp wi t h the
in Lonicera . achenes borne on the inside
50rosis . Fruits on a common axis of a hol l owed-out r ecept acle
that are usually coalesced and or peduncle, as in Ficus .
derived fro m the ovaries of
several flowe r s , as i n Horus.

4. Acce ssory Fruit Types (Figure 6-10, p . 113)

(Fruits derived from simp le or compound ovaries and some non-ovarian tissues ,
as the hypanthium ; c lassification ar r anged alphabetically; types of acces-
sory structures given in parentheses be l ow .)
Bur (Invo l ucre) . Cypsela enclosed in Gl ans (Involucre) . Nut s ub tended by
dry involucre , as in Xanth i um . a cupulat e , dry involucre, as in
Coenocar pium (Various Str uctures) . Quer cus .
Hul tiple fruit derived f r om ova- Hip or Cynarr hodion (Receptacle and
ries, f l oral pa r ts , a nd recep- Hypanthium) . An aggregation of
tacles of many coal esc~d f lower s , achenes s urrounded by an urceol ate
as in Ananas. receptacle and hypanthium, as i n
Diclesium (Calyx) . Achene or nut Rosa.
surro unded by a persistent calyx , Pome (Receptacle and Hyp anthium). A
as in Hirahali s . berry-like fruit, a dna te to a
112 Figure 6- 9

FRUIT TYPES

D R Y IN DEHISCENT FRun T Y PE S

"" (,

Colyb.",m Cap sule

(); <~-
:-::
.~,

,
~~

Ny tie l Som (lro Ul ri cl e

DRY DEHISCENT FRUIT TYPES

f olloel\! -
L omen l $,I;tle $ihqu"

CAPSULE TYPES

Po .ie ld ol Sep l ,ei dol Valvular

Locul,e, (l OI Op e,c ulaTe

SCHIZQCARPIC FRUIT TYPES

5th., Faillele$ SC IUl. Nut1e l ~ 5C hi. So m{l ' o~

SChi zogerr t 5th"_ M~,icarp

FLESHY FRU I T TYPES

V ,·"
G eo"

OJj~
Pyrene
Be" ~ P, upe
Figure 6- 10 113

FRUIT T Y PES. Cont'd .• SEED PARTS and EMBRYO TYPES

AGGREGATE FRUIT TYPES

~. .

Achenecetum
~~
~rro
Baccacetum
, ~
Drupecetum Fo tl ice tum
~ So mor acetum

ACCESSORY FRUIT TYPES

B",

Pseudocorp Pseudodrupe Syncon.um Tr~ma Winged Nut

~YPOCOlyl~ep i COlyt

nypocOlyl
i\~e ,

~c.
rn;"·", ,u
,od i cl e ~~~ colylodon
h.lum--.!,\

. eedCOOl

Lima Bean Corn CaS lo r Beon

EMBRYO TYPES

Rudimentary B rood Capitate L ateral Peripherol Lin ear

BI'ISAL PERIPHERAL AX IAL

Spotvlote Bent Fa Ided Investing Dwarf Micro

f - - -- - - - -- FOLIATE - - - - - - - - - - ; > - - -- - M IN IATURE ----<
114 6

fleshy receptacle, with cartilagi- Syconium (Receptacle , possibly pedun-

nous endocarp, as in Halus . cle). Hultiple fruit surrounded
Pseudocarp (Receptacle). An aggrega- by a ho llow , compound , fleshy
tion of achenQS embedded in a r eceptacle, as in Ficus .
fleshy receptacle , as in Fragaria. Tryma (Involucre) . Two-four l oculed
Pseuclodrupe (Involucre). Two four nut surr ounded by a dehiscent
loculed n u t surr ounded by a involucre at mat urity, as in most
fleshy involucre , as in Juglans . species of Carya .
\Hnged Nut (Bract) . Nut enclosed in
a winglike bract, as in Carpinus .

XII. SEEDS AND SEEDLINGS

A. Seed Parts (Figure 6-10 , p. 113)

Aril. Outgro\"th of funiculus , Hilum . Funicula r scar on seed
raphe , or integuments; or coat .
fleshy integuments or seed Nicropyle . Hole through seed
coat , a sa rco testa . coat .
Chalaza . Funicular end of seed Perisperrn .Food reserve in seed
body. derived from diploid nucellus
Embryo . Young sporophyte consist - or integuments .
ing of epicotyl, hypoco t yl, Raphe. Ridge on seed coat formed
radicle , and one or more coty- from adnate funiculus.
l edons . Seed Coat. Outer protective cover-
Endo sperm . Food reserve tissue in ing of seed .
seed derived from fertilized
polar nuclei; or food reserve
derived f rom megame t ophyte in
gymnosperms .
B. Seed Types
(Classification based on type of nourishing tissue)
Cotylespermous . l-li th food r eserve in Hypocotylespermous or Nacropodial.
cotyledon , derived from zygote . Hith food reserve s tored in hypo-
Endospermous o r Albuminous . l-lith cotyl, derived from zygote .
food r eserve 1.n endosperm or Perispermous . With food reserve in
albumen , derived from fe r til- perisperm, derived from diploid
ized polar nuclei. nucellus or integuments.

C. Embryo Parts (Fi gure 6-10 , p . 113)

Coleoptile . Pro tective sheath that gives rise to shoot system.
around epicotyl in grasses . lIypocotyl. Embryonic stem in seed .
Coleorhiza . Protective sheath located belmY' cotyledons .
around radicle in grasses . Plumule. Embryonic leaves in seed
Co t yledon . Embryonic l eaf or derived from epicotyl .
leaves in seed . Radicle . Ba sal end of embryo axis
Epico t yl . Ap i cal e nd of embryo axis t hat gives r ise to roo t system .

D. Embryo Types (Figure 6-10 , p . 113)

(Classification based primarily on embryo shape . size, and position--adapted
from Hartin (1 946 1)
Bent . Foliate embryo with expanded Broad. Basal, globula r or lenticular
and usually thick cotyledons i n embryo in cop io us endospe r m.
an axile position bent upon the Capitate . l-Iore or l ess basal head-
hypocotyl in a jacknife position . like or turbinate embryo in copi-
ous endosperm .
6 115

Dwarf , Axial embryo variable in "'licro .Axial embryo in minute seeds ,

size relative to seed , small to less fuan 0 . 2 mm. long; minute
nearly total size of seed: seeds and undifferentiated to almost
0 . 2-2 tmIl . long . total size of seed .
Folded. Foliate embryo with cotyle- Peripheral. Peripheral embryo
dons usually thin and extens ively large and elongate , arcuate.
expanded and folded in various annular , spiro l obal, or
ways . straight; cotyledons narrow
Investing. Axial embryo usually erect or expanded; perisperm centra l
with thick cotyledons ove rlapping or lateral.
and encasing the somewhat d\"arfed Rudimentary . Basal, small non-
hypocotyl; endosperm wanting or periphera l embryo in small to
limited. large seed ; relatively undiffer-
Lateral . Basal or base- la te ral entiated; endosperm copious .
embryo, discoid or l enticular. Spatulate . Fo l iate, erect emb r yo
usual l y surrounded by copious I"ith variable cotyledons, th i n
endosperm . to thick and s l ight l y expanded
Linear . Axi al embryo several times to broad .
longer than broad, straight,
curved or coiled ; cotyledons not
expanded; endosperm present or
absen t.

E. Aril Structural Types and Selected Seed Surface Features

1. Aril Structural Types

Ar illate . General term for an out- Fibrous . With stringy or cord-like
gr owth from the funiculus, seed seed coat , as mace in Hyristica .
coat or chalaza; or a fleshy seed Funicular . {.,rith a persistent elon -
coat . gate funiculus attached to seed
Caruncula te . \.,rith an excrescent coat , as in Hagnolia.
outgrowth from integuments near Sarcous . \-lith the seed coat fleshy .
the hilum, as in Euphorbia . Strophio late. \.,rith elongate a ril or
stroph i ol e in the hilum reg i on .

2. Special Seed Surface Features

(See Surface io Section B of this chapter)
Alate. \noged. Crested . I·li th elevated ridge or
Circuma l ate . IHnged circumferen- ridges, rap hal.
tially. Umbonate . With a distinct projec-
Comose . l.,rith a tuft of tric homes . tion usually from the side .
Coronate . l-lith a crown . Verrucose. Harty .

F. Seedling Parts
(Specialized parts only adapted from Duke (19691)
Cataphyll. Rudimentary sca le l eaf Eophy ll. Ter m applied to first few
produced by seed l ing , us ually i n leaves with green, expanded l amina
cryptocotylar species . developed by seedlings; transi-
Coll et . External demarca~ion be- tional type leaves developed before
tween hypocotyl and root . formatio n of adult leaves.
?-Ietaphyll. Adult leaf .
116 6

G. Seedling Types
(Classification based on position of cotyledons in germination)
Cryptocotylar or Hypogeous . With the Phanerocotylar or Ep i geous. Hith
cotyledons remaining inside the the cotyledons emergent from
seed; seed usually remaining be- seed, usually appearing above
low grou nd. ground·,

Section B. GENERAL CHARACTERS AND CHARACTER STATES

I . POSITION AND ARRANGill-1ENT

A. Location or Environmental Position

(Classification based on position of organs or parts in their surrounding
environment)

1. General
Aerial or Epigeous. Above the Floating. Upon the surface of the
ground or water ; i n the air . water .
Emergent. Hith part (s) of plant Submersed . Beneath the surface of
aerial and partes) submersed; the water.
rl.s~ng out of the water above Subterranean or Hypogeous . Below the
the surface . surface of t he ground .
Epipetric . Upon rock. Surficial or Epigeous . Upon or spread
Epiphytic. Upon another plant . over the surface of the ground.

2 . Special
(Selected location terms . See habitat prefixes i n Chapte r 15 and word stems
for plant organs and parts in Chapter 4 for mean ings of other location
terms . )
Aerocaulous. Hith aeria l stems. Flotophyllous . \.J'ith floating leaves.
Aerophyllous . Hith aerial leaves . Geoflorous. Hith subterranean
Amphicarpous. Hith f ruits in two flowers .
environments ; e.g ., aerial and Petrorhizous. With roots on rock.
subterranean . Submersicaulous. With submersed
Emersifolious. Hith emergent leaves . stems.
Epirhizous. With roots upon another Surcarpous . With fruits on surface
plant. of ground .

B. Position
(Class ification based on location of parts or _organs with respect to other
dissimilar parts, or or gans)

1. General
Apical or Terminal . At the top. Discontinuous . Basal and lateral,
tip, or end of a st ructure . basal and terminal , or lateral
Basal or Radical. At the bottom and terminal ; not continuous .
or base of a structure. Lateral or Axill ary. On the side
Continuous . Basal, lateral , and of a structure or at the nodes
terminal. of the axis.

2. Special
(Classification based on positional terms usually appl icable t o individual
par t s)
6 117

B. Androecial Position
(See Perianth and Stamen Position , h . and k.)

b.Branch Position
Acrocaulous . Hith terminal branches. Subbasicaulous. With b ranches at or
Basicaulous. Hi th basal branches. near base of main stem.
Caulous . With branches more or less Zonocaulous. Hith branches inter-
evenly spaced along trunk. mittently spaced along main
Suhacrocaulous. With branches at or stem .
near tip of main stem .

c. Cotyledon Position (Figure 6-12, p. 121)

Accumbent or Pleurorhizal. Recl i- Incumbent or Notorhizal. Reclinate
nate with cotyledon edges against with sides of cotyledons against
hypocotyl. hypocotyl.

d. Flower , Fruit, Inflorescence , Infructescence Position

Acrocaulous. At the tip of the Geoflorous. Geocarpous. Flowers or
stem. fruits below ground, as in
Amphiflorous, Amphicarpous. Flowers Amphicarpurn.
or fruits above and below ground, Infrafoliar. On the stem below the
as in Amphicarpum. leaves, as in the Arecaceae .
Axillary. In axil of leaf. Interfoliar. On the stem between the
Basicaulous. Near base of stem. leaves, as in the Arecaceae .
Cauline or Caulous. On old t"oody Leaf-opposed. On stem opposite the
stem. base of the leaf, as in Alchemilla.
Epiphyllous. From a phylloclad or Suprafoliar. On the stem above the
peculiar bract , as in Tilia . leaves, as in the Arecaceae.
Terminal. At or near tip of branch.

e . Leaf Position
Acroramous. Leaves terminal, near Cauline or Ramous . Leaves more or less
apex of branch . evenly distributed on stem or branch .
Aphyllopodic . Without blade-bearing Phyllopodic. \\fith blade-bearing l eaves
l eaves at base of plant . at base of plant .
Basiramous . Leaves on lower part of Radical . Leaves basal, near ground ,
branch . usually from caudex or rootstock.

f . Ovary Position
Inferior. Other fl oral organs hypanthium adnate to lower half
attached above ovary \"ith hy- of ovary.
panthium adnate to ovary. Superior . Other floral organs
Half- inferior. Other floral organs attached below ovary.
attached around ovary ....,ith

g . Ovule Position (Figure 6-12 , p. 121)

(Based on posit ion of ovule in locule and orientation of the micropyle and
raphe--adapted from Bjornstad [1970J)
Epitropous, dorsal. Ovul'e pendulous Epitropous, ventral . Ovule pendu-
or hanging, micropyle above, lous or hanging, micropyle above,
raphe dorsal (away from ventral raphe ventral ( toward ventral
bundle) . bundle.
 118 6

Heterotropous. Ovule position not Pleurotropous , dorsal. Ovule hori-

fixed in ovary . zontal , micropyle toward ventral
Hypotropous, dorsal. Ovule erect, bund l e, raphe above.
micropyle below, raphe dorsal Pleurotropous , ventral. Ovule hori-
(a~"ay from ventral bundle) . zontal, micropyle toward ventral
Hypotropous , ventral. Ovule erect , bundle:; raphe below.
micropyle below, raphe ventral
(toward ventral bundle) .

h . Perianth and Androecium Position (Figure 6-11, p . 120)

(Classification based on insertion of Floral Parts--Corolla, Calyx, and
Anclroecium--the androperianth)
Epigyny. The condition in which the attached to the floral or hypan-
sepals, petals , stamens are at- thium cup above the ovary with
tached to the floral tube above the lower part of the hypanthium
the ovary with the ovary adnate completely adnate to the ovary .
to the tube or hypanthium. Hypanepigyny . The condition i n which
Epihyperigyny. The condition in the sepals, petals , stamens are
which the sepals, petals. stamens attached to the elongate floral
are attached to the floral tube tube or hypanthium above the in-
or hypanthium surrounding the ferior ovary , as in
ovary ; a combination perigyny Oenothera .
and partly inferior ovary. Hypogyny. The condition in which the
Epihypogyny . The condition in which sepals, petals, stamens are attached
the sepals. petals, stamens are below the ovary .
attached about half- way from the Pedgyny . The condition in which the
base of the ovary to the partly sepals, petals, stamens are attached
adnate hypanthium tube; half- to the flora l tube or hypanthium
inferior insertion of parts . surrounding the ovary with the
Epiperigyny. The condition in \"hich tube or hypanthium f ree from the
the sepals, petals , stamens are ovary .

i. Placenta Position (Placentation) (Figure 6-11, p. 120)

Axile . l.Jith the placentae along the Harginal or Ventra l. With the pla-
central axis in a compound ovary centa along the rna rgin of the
with septa. simple ovary .
Basal. l.Jith the placenta at the base Parietal. Hith the placentae on the
of the ovary. wallar i ntruding partitions of a
Free-central. With the placenta unilocular compound ovary.
along the central axis in a com- Pendulous, Apical. or Suspended .
pound ovary wi thout septa. With the placenta at the top of
Laminate . With the placenta over the the ovary.
inner surface of the ovary wall .

j . Radicle Posit i on
Antitropous . With radicle pOinting Syntropous . I-lith radicle pointing
away from hilum. toward hilum.

k . Stamen Position (Figure 6-7, p. 105)

Allagostemonous . Having stamens Cryptantherous. With stamens in-
attached to petal and torus eluded.
alternately . Diplostemonous. Hith stamens in two
Antipetalous. Opposite the petals. whorls, outer opposite the sepals ,
Antisepalous. Opposite the sepals. inner opposite petals.
6 119

Epipetalous . With stamens attached Obdip lo stemonous. Inth stamens in

to or insert ed upon petals or two whorls , outer opposite pet a ls,
corol l a . inner opposite the sepals .
Episepal ous . Hith stamens attached Phaneran t he rous. ""ith st amens ex-
or i nserted upon sepals or ca l yx . serted .

1. St y l e Position
Gynobasic . At the base of an invagi- Subapical . At one side near apex of
nated ovary. ovary .
Lateral. At th e side of an ovar y . Terminal or Apical . At the apex of
t he ovary .

C. Arr angement
(C l assific a tion based on locat ion of pa rts in relation to each other)

1. General (Figure 6- 16 . p . 136)

Alternate . One l ea f or other struc- Symmetry o f arrangement b roken ,
ture pe r node. wi th uneven lengths of inte rnodes .
Clustered, Conglomerate , Agglomer- Loose, Di stant , or Scat t ered . Par ts
ate, Cr owded . Aggregate . Parts \.,Iidely separat ed from one another ,
dense , us ually irregularly over- us ually irr egularly .
lapp i ng each other. Opposite . Two leaves o r othe r
Continuous. Symmetry of arrange- structures per node , on oppo-
ment even, not b roken. site sides of stem o r central
De cussate . Opposite leaves at right axis .
a ngle t o pre ceding pair . Polys tichous . Leaves or othe r st r uc-
Distichous . Leaves 2-ranked, in tures i n many r ows .
one plane . Ros ulat e . Leaves in a rose tt e .
Equitant . Leave s 2- r anked with Secund or Unilateral . Flowers or
overlapping bases , us ually ot her structures on one side of
sharply fo l ded along midrib . axis .
Fasciculate . Leaves or other st ru c- Tetrastichous . Leaves or other
tures in a cluster from a com- struct ure s in four rows .
mon point . Tris t ichou s . Leaves or other struc-
Geminate or Binate . Paired ; in t u res in three r ows .
pa i r s . l~ o r led , Radiate , or Ve rt icillate .
Imbrica t e . Leaves or other struc- Three o r more leaves o r other
tures over l apping. s tructures per node .
Interrupted or Discontinuous .

2 . Special
(Class ification based on arrangement with s pecia l terms applicable to indivi-
dual pl ant part s )

a . St amen Arrangement (Figure 6-7, p . 105)

( See General Arrangement f o r additional terms)
Didymous. With stamens i n two Tet r a dynamous. Hith stamen s i n two
equal pai rs . g roups , usually fou r lon g and
Didynamous . With stamens ....i n two two sho r t .
unequal pair s . Tridynamous . Hith stamens in two
e qual groups of thr ee .
120 Fig ure 6-11

POSITrON and POSTURE

PERlA NTH and ANDROECIAL POSITION

Epihyperigyny Epi hypol;Jyny E pi per igyny Hyp onepi gyny Hypogy ny

PLACENTAL POSITION

Ax i te B 0 5 01 fr ee - ce ntrol Lorn ioot e Marginal Pariet a l Pend u lou s

TRANSVERSE POSTURE

Cern uou s Fl e ~u ou s Ge niculole Incurv cd R eCu rved Squarrose Und ulate

LONG I TUDINAL POSTURE

Condup l icO le Gen icu late Ind u pl ic ot e , ,\Vo lute

Re volute Roll ed Sin uo te Straigh t To rtu ous Va lua te

Figure 6-12 121

EMBRYONIC POSITION
AESTIVATION

Alternate Con volute IndupliCOle Volvole

PTYXIS (Cotyledons)

~
..,----" ...,'
- "" ..
- .-" . ' ,:}
.

e., ....• ".'

Conduplicote Contortupl ;cote D;plecolobol Sp;rolobol

PTYXIS

Circinate CondupliCOle Convolute Corrugate Cu rvote Inc linote

Involute Plicate Reclino te Replicate Revolute Supervolute

_k, COTYLE DON POSITION

l!A"~:?W
r'" rodic le
c = cotyledon

I ncumbent

OVULE POSITION

~~ . .~ \~
Epitropus Epitropus Hypotropous
?'"-
Hypotropous
if ~ .~
Pleu rotrapous PleurOlrapouS
dorsal ventral dorsa I ventral dorsal ve nl ral
I 122 6

b. Thecal Arrangement (Fi gure 6- 7. p . lOS)

(The thecae i n this c l assification can be conjunctive or disj unctive.)
Divergent. Thecae or anther cells Parallel. Thecae Dr anther cells
divaricate or separated from one along side of the connective or
another at an acute angle to the longitudinal to each other .
connective or filament . Transverse.... or Explanate. Thecae or
Oblique. Thecae or anthe r cell s anther cells with maximum diver-
lower on one side of connective gence of about 90° from the con -
than the other. nective or filament.

D. Orientation
(Classification based on arrangement of parts in relation to vertical angle
of divergence from a central axis or point)

1. General
Acroscopic. Facing apica l ly . I n flexed . Bent abruptly inward or
Agglomerate, Con glomerate, Crowded , upward .
or Aggrega te . Dense structures Patent. Spreading .
with varied angles of divergence. Pendulous . Hanging loosely or
Antrorse . Bent or directed up\~ard . freely .
Assurgent. Direct ed upward or for - Reclinate . Ben t down upon the axis,
ward. no angle of diverge nce.
Basiscopic. Facing basally . Reflexed. Bent or turn e d downward .
Connivent . Convergent apically with- Retrorse . Bent or directed dovn-
out fusion . vard .
Conto rted. Twisted around a central Salient , Porrect, or Projected .
axi s; twisted . Pointed outwar d, us ually said of
Dec l inate. Directed or curved down- teeth.
ward . Sinist ror se . Rising helically from
Deflex ed . Bent abruptly downward . left to right , a characteristic
Dextrorse. Rising helical l y from of twining stems .
right to left , a characteristic Twining . Twisted around a central
of twining stems. ax is .

2 . Special
(Classification based on stated degrees of divergence)
Appressed or adpressed . Pressed Horizontal l y . Spread ing outHard
c losely to axis upward \vith at 90° from vertical axis or
angle of divergence lsD or l ess . plane.
Ascending . Directed upward with an Inc lined . Ascending at 46-75° angle
angle of divergence of 16-4 5° . of divergence .
Depressed . Pressed closely to axis Rec lined. Descending at 106-135°
downward with angle of divergence angle of divergence .
of 166- 180° . Resupinate . I nve r ted or twisted
Descending . Direct ed downward with an 180° , as in pedicels in the
angle of divergence of 1 36-165° . Orchidaceae .
Divergent, Patent , or Divaricate . Hare
o r less horizontal l y spr eading with
angle of divergenc e of 15 ° or less
up or down f r om the horizontal.

E. Transverse Posture (Figure 6-11 , p. 120)

(Classification based on position of ends of single s tructure in relation to
its center or transverse axis )
6 123

Applanate or Plane . Flat, without Lorate. Hith elon gate vertical waves
vertical curves or bend s . in the margins or s ides at right
Arcuate . Curved like a crescent , angles to the l ongitudina l axis .
can be dm"m"ard or upwa rd. Rec urved . Curved outward or down-
Ce rnuous. Drooping . wa rd .
Flexuous . Hith a ser ies of long Squarrose . Us ually s harply curved
or open vertical c urves at down\"ard or outward in the apical
right angles to the cent ral region , as the bracts of some
axis . spec ies of Aster .
Geniculate. Abrup t ly bent verti- Undulate . Wit~e ri es of vert i cal
cally , usually near the base . curves at right angles to the
Inc urved . Curved inward or upward. cent ral axis .

F. Longitudinal Posture (Fi a ure 6- 11, p . 120)

(Classification based on posit i on of the sides of a singl e st ruc tur e in rel a-
tion to its cent ral axis)
Condup licate . Longitudinall y fold e d Re volute . ~mrg i ns or outer portion
upward or downward a long the cen - o f sides rolled outward or dmmward
tral axis so that ventral and/or over lower or dor sa l surface .
dorsal sides face each other . Rolled . Sides enrolled , usually
Geniculate . Abruptly bent horizon- loosely , over upper or lower sur-
tally , usually in ser ies . faces .
Indupl icate . Having margins bent Sinuate . Long horizontal c urves in
inward and touching margi n of the body of the structure parallel
each adjacent structure. to the central axis .
Involute . Hargins or outer portion St r a i ght . t..'i t hou t a curve, ben d, or
of sides rolled i nwa rd over upper a ngle .
or ventral surface . Tor tuous . Irregularly twisted.
Plicate . With a ser ies of longi- Valvate . Sides enrolled , adaxia l ly
tudinal folds; plaited . or abaxia lly so that margins touch .

G. General Struc tural Position

(Pertains to regional locations on a structure)

1. General
Abaxial. Away from the axis ; the of an outer face o f organ ; lower
l ower surface of t he leaf; dor- s ide of leaf; abaxial .
sal. Narginal . Per taining to the border
Adaxial . Next to the axis ; facing or edge .
the stem; ventral . Nedial. Upon or along the longi-
Ap ica l. At or near the tip . tudinal axis .
Basal . At or near the bottom . Peripheral. On the outer surface or
Central . In the middle or middle edge .
plane of a structure . Proximal. Near the point of origin
Circumferential . At or nea r the or attachment .
Circumference ; s urrounding a Subbasal. Near the base.
rounded struc tur e . Subterminal . Near the apex .
Distal. Away from the point o f Ventral. Pertaining to the surface
origin or attach ment-~ A .... nea r es t the axis; inne r face of
Dorsal. Pertaining t o the surface an organ ; the upper s urface of
most distant from the axis ; back the lea E; adaxial.
 124 6

2. Special
(Selected terms for location on a str ucture . The meanings of many addi tional
terms can be determined f rom the pos itional prefixes and word stems of
plant organs and parts in Chapter 4 . )
Ac r ocaulous . At t i p of s tem . Laterosper mous . On the side of the
Bas ipetiolar. At the base of the seed . ...~
petiole . Pericar pous . Around the fruit .
Cent roramo us . At the cen t e r of t he Suprarhizous . On top of t he r oot .
b ranch . Ven tristipular. On vent r al side of
Dorsila minar. On dorsa l s ide of s ti p ul e .
blade .

H. Embryonic Position
(Position of immature organs or parts)

1. Aestivation or Pref loration (Figure 6-12 , p . 121)

(Classification based on position of emb r yonic perianth parts . Cal yx and
corolla may have d iffe r en t aestiva tion types . )
Al te rna t e . Havin g struct ure in two I mbricate. Hav i n g margin s overlapping .
rows or series so that t he inner I nduplicat e. Having margins bent i n-
str ucture has its mar gins over - ward and touch ing margin of adja-
lapped by a marg i n from each cent structur e .
adjacent outer structure . Quin cuncial . Having five st ructures ,
Co chleate . Having one hollow or two of Hhich are exte r i or , two
helmet-s haped struct ure which interior , and a fifth \·dth one
encloses or covers the others . ma rgin covering i nterior struc-
Contorted . Havin g several struc- ture and other margin covered by
, tures in a who rl or close that o f one of t h e exterior
s piral with on e ma r gin cov e r- s tructures.
ing the margin of an adjacent Val va te. Having margins of adjacent
st ruc ture . st ruc ture s t ouch ing a t edges only .
Convolute. Havin g one leaf o r Vexillate . Having one struc ture lar ger
perianth part rolled in another , than others which is fol ded over
usua lly twisted apically . smalle r enc l osed s tructures .

2. Cotyledon Ptyxis (Figu re 6-12, p . 121)

(Clas sifica t ion based on pO Sition of co tyledon s in seed)
Co nduplicate . Cotyledons fo lded Dipl eco lobal. With inc umben t
lengthwi se alon g mid r ib with one cotyl edons fold ed two or more
coty l edon covering othe r and inner times.
co t y l ed on covering hypoco t yl. Spir oloba l. Hith i n cumbent coty-
Con t o rtupl ica t e . I-lith weird l y fo lded ledons fo l ded on ce .
corru ga t e cotyledons .

3 . Ptyx is (Figure 6-12 , p . 121 )

(Classif i cation based on rollin g or folding of ind ividual e mbryonic leaves
and arrangement of embryonic leaves within a struct ure ; vernation accord-
i ng to most au thors ) . Adapted f r om Davis and Heywood (1963) .
Cir c inate. With lamina rolled from Convolute . \-1ith one l amina enrolled
apex to base wi th apex i n cente r in another l amina .
of coil . . Co rrugate . Uith lamina i rre gularly
Con duplicate . I-lith lami na f o ld ed folde d in a ll direc tion s, wrinkled.
once adaxia lly along midr ib o r Curvative or a r cua t e . Wit h lamina
mi dvein . fo lded transversely i nt o an arc .
6 125

Impl icate . t·lith both lamina margins Reclinate . Hith lamina folded or
folded sharply i nward . curved bach~ards fro m near its base
Inclinate . Hith lamina folded or so that embryonic blade is parallel
curved transversely near the to its petiole , hypocotyl , or stem.
apex . Replicate. Ivith lamina folded once
Involute. Hith lamina margins en- abaxially along midrib or midvein .
rolled adaxially. Revolute. lnth lamina margins en-
Planate or Plain . Hith lamina flat, rolled abaxially.
without folds or rolls . Supervolute . \.;rith lamina with one edge
Plicate . toJith many longitud i nal tightly en rolled and with the other
folds in lamina . loosely enrolled covering the first ,
loosely convolute .

II . NUMBER AND SIZE

A. Number
(Perta i ns to sel ected terms dealing tvith numbers. See number prefixes and
word stems in Chapter 4 for meanings of additional terms . )
Ancipital. Two-edged . Monadelphous . Hith one group of sta-
Bicarpellate . Two-carpelled . mens connate by their filaments .
Bidentate . Two-too thed . Honocarpellate. One- carpelled .
Biflorous . ~w-flowered . Honocephalous . One- headed, as in com-
Bifo l iate . Ttw-Ieaved . posites.
Bilabiate . 'fuo- lipped . Nonochasium. Cymose inflorescence in
Bilocular . Ttvo- I ocular . which each central axis bears one
Bina te . Ttdnned . lateral axis.
Biseriate . Ttvo - rowed ; in two Monochlamydeous . Hith perianth com-
series . posed of similar parts, each being
Bisexual. Both sexes in same a tepal; with one whorl of acces-
f lower (monoclinous , perfect). sory parts .
Diadelphous . Hith two groups of Monocotyledonous . Hith one cotyle-
stamens connate by their fila- don .
ments . Nonoecious . With staminate and car-
Dianclrous. Hith ttva stamens per pellate flowers on same plant.
flower . Nonophyl lous. One-leaved .
Dichasium. Cymose inflorescence Nulticellul ar. }!any- celled .
in which each axis produces a Nulticipital. Hith many axes or
pair of lateral axes . stems from one rootstock or
Dich l amydeous . Hith two perianth caudex .
parts, a distinct calyx and Multicostal . Many- r ibbed .
corolla . Hultilocular. Hany-locular .
Diclinous. Having the stamens and Hultiseriate . Nany-rowed; i n many
ca r pels in separate flmvers, series .
imperf ect , either monoecious or Nultistriate. Many- lined .
dioecious . Pentagonal . Five-angled .
Dicotyledonous . Hith two cotyle- Pentandrous . With five stamens .
dons . Polyad . Po llen grains in cluste r s
Dioecious . With staminate and car- of more than four.
pellate flowers on s.~R~rate Polyandrous. Hany- stamen ed.
plants . Polycarpel late . Nany- carpellate.
Dipterous . Two-winged. Polycephalous . Many- headed, as in
Dyad . Pollen grains occurring in composites .
clusters of two. Polydelphous. With several groups
Monad . Pollen gra ins occurring of stamens connate by their fila-
singly . ments .
 126 6

Tetrad . Pollen gra i ns in cl us t ers Trifo l iat e . Three-leaved .

of four . Trifoliolate . \.Jith th ree leaflets .
Tetragonal. Four-angled . Tr1gonou5 . Thr ee-angled .
Tet r ahedral. Having the form of a Triquet r ous. Three - angled with the
tetrah ed r on . sid e s usually concave .
Tetral ocular. Fou r-loc ular . Unilocular . One- locu lar .
Tetrandrous . With four stamens . Uni seriate . One- rowed; in one seri es .
Tricarpel late . Three - ca rpellate . Uni sexual. l.Jith only one sex in each
Tr if10rou5 . Thr ee-flowered . flower.

B. Size
(Selected terms r elated to s i ze and f r equent l y shape . See size prefixes and
' . . ord stems of organs and parts in Chap ter 4 for meanings of add itional
tenns . )
Amp lia te o Enlar ged ; dilated. Hete rostichus . With unequal rows .
Angustate. Narrow . Homanclrous . \~ith stamens of same
Anisocarpous. Inth unequal car- s ize and shape .
pel s . Homocarpous. \\'ith carpels of same
Anisocot y l ous . Hith unequal coty- size and shape .
l edons . Hypophyllous . I~ith sma l l leaves , as
Aniso l ateral. \o!it h unequa l sides . bracts, sca l es , cataphyl l s .
Anisopeta l ous. \.J'ith un equal pe t a l s . I nequilate ral. Hith unequal sides .
Anisophyllous . Hith unequal l eaves . Isocotylous . Hith cotyledons of same
Anisostylous . Hith unequal styles . si z e and s h ape .
Depauperate. Sma ll and us ually Isodyn amou s . \~ith e qually develop ed
poorly developed. structures .
Dila ted . Hidened; expanded . Isopetal ous . IUth petals of same s ize
Dwarf. Very s ma ll . and shape .
Gi gantic. Very large . I sophyllous . Hi th l eaves of same s ize
Hete r androus . With s tamens of and shape .
different sizes a nd/or shapes . I sosepalous . l-lith sepals o f same size
He t eroblasty . Inth juvenile foliage and shape .
distinctly differen t from adult I s ostichous . With equal rows.
foliage in size or shape. Latiflorous. With broad- f l owers.
Heterocarpous . With carpels of Leptophyllous . IUt h leaves to 25 s q .
different s iz es and/or shapes . rom . in size .
Hete r oc l ado u s. IUth stems of dif - Major . Greater i n s ize .
f erent sizes and/o r shapes . Hinor . Smaller i n size .
Heteropetalous . Hi th pe tals of Hinute . Very sma l l.
dif f e r e n t s i zes and/or shapes . Nanophyllou s . Hi th l eaves to 225 sq .
Hete r ophyllous . l-li th leaves of tml. in size .
different s izes a n d/ or shapes . Pla t ycan thous . With flat and usua l ly
Heter osepalous . With s epa l s of large spines .
different sizes and/or shapes. Reduced . Decreased in size .
Robus t . Large .

C. ~
(Pertains to number of whorls of flora l pa r ts , l eaves , or stems)
Acar pous . No carpels or carpellate Arhizous . Hithout r oots , no whorls
who r l ; no pistil. of root s .
Achlamydeous . t~ithout perianth. Asepal ous. No sepals or calyx .
Ape t a l ous . No petals or corolla . Astemonous or Anandrous . No stamens
Aphyllous. Without leaves. no or androecium.
whorls of leaves . Chlamydeo us. With p erianth .
6 127

Complete . I~ith fo ur types of fl ora l Nonocyclic . One-whor led .

parts . Oligot axy . Reduction in numbe r of
Dichlamydeo us . Hith perianth com- Hhorls .
posed of distinct calyx and Pentacyclic . Five -whorled .
corolla . Pleio t axy. Increase in number of
Dicyc lic . Two- whorled . '''horl s .
Homochlamydeous . With pe r ian th com- Polycyc lic . NanY- I"horled .
posed of s i milar parts , each part Tetracyclic . Four - whor l ed .
a tepal . Tr icyclic. Three-uhor led .
Incomp lete . One or more types of
floral parts absent.

D. f>lerosity
(Pe rtains to n umber of parts wit hin ,,,horls of f loral parts , l eaves , o r stems)
Dimerous. l.fue r ! with two members . Pleiomerous . Hit h increase in number
Heteromerou s or Ani somero us . Hi th of members wit hin whorl .
di ffere n t numb er of me mbers in Polymer ous . hlhorl with many members .
different whorl s . Pseud omonomerous. ~~ orl seemingly
I somerous . With same number of with one member which is a f usion
member s in different whorls . product of two or mo r e parts.
Honomerous. Whorl with cne member . Te trame r ous . l.fhor l with fo ur mem-
Oligomero u s . With red uc tion i n ber s .
number of members within whorl. Trimerou s . \<1I"\O rl with three members .
Pentame r ous . II1horl with five member s .

E. Fusio n
(Pertains to f usion of members wi thin and between whor l s of floral part s)

1 . Gene ral
(Based on Porter et a1. , [1973))
Adherent . hlit h unlike p.:lrts of or- Connat e . With like parts o r organs
gans jOined , but only s uperfi- int egral l y fus ed t o one ano ther
cia lly and without actua l histo- with histol ogical continuity.
l ogical cont inuity . Conti guo us . Touchin g but not adnate,
Adnate . lUth unl ike parts or conn ate, adhe rent , o r coherent.
organs in teg rally f used to one Di stinct . With l i ke par ts or organs
another with histol ogical con- unjoin ed and separate from one
tinui ty . another .
Coale sced . Hit h like or unlike Fasciat ed . Unna turally and o ft e n
par ts or organs incomplete l y monstro usly connat e o r adnat e,
separated ; par tial ly fused in the coal esced parts of t en unnat-
a more or l ess irregular fashio n . u rally pr oliferated in s i ze a nd /
Cohe r en t . With like parts or or- or numbe r; e . g ., inf l orescen ce of
gans joined , but only s up er fi - Celosia .
cially and without actual his- Free. Unlike parts or organs unjoined
tological continuity . and separate f rom one another .

2 . Spec i al
(Se 1.es ted terms pertaining to fusion)
Anthocarpous. Having a body of Apopetal ous or Choripeta l o us . With
combined floral a nd fru it parts , separate petals .
as in multiple fru its . Aposepalous or Cho r isepalous . i·lith
Apocarpous. Wi th separate carpel s . separate se pals .
Apostemonous . \\lith separate s tamens.
 128 6

Column, Gynostemium or Gynandrium . Petalostemonous . Wi th fil aments

\hth fused stamens and carpels fused to co rolla , anthers free.
(stigma and style) as in Orchis. Polydelphous . t.Jith several groups of
Conjugate. Fused pair s , as the stamens connate by their f ilamen t s .
fruits of Lonicera . Sympetalo us. J.1ith fused petals.
Diadelphous . With two groups of sta - Syncarpous . l.]ith f used carpels.
mens connate by thei r f ilaments . Syncotyly . Cotyledons coalesced ,
Hypanthium . Fused flo ral parts fo rm- forming a ...funnel or trumpet .
ing an envelope around the ovary. Synsepalous . With f used sepals .
Honadelphous. t..lith one group of sta- Syngenesio us . With fused anthers .
mens connate b y their filaments .
3. Hypan thium Adnation
(Based on fus ion with ovary)
Absent . No hypanthium present . Free-adnate . Hypant hium fused with
Adnate . Hypanthium completely fused ovary and having a free limb
to ovary . ar ound or above ovary .
Free . Hypanthium s urrounding but Partly adnate. Hypanthium adnate to
completely free f r om the ovary. part of the ova r y and with no free
limb or tube.
F . Divis i on
Bifid. Cut or divided into two Palmatifid . Cut palmately .
lobes or parts . Palmat i sect . Sectioned or divided
Bi f urcate . Div ided into two forks palmately i nto distinct segments .
or branches . Parted. Cut 1/2-3/4 of di stance to
Cleft . Cut 1/4-1/2 of distance of middle of structure .
middl e of structur e . Pectinate . Having closely parallel
Dichotomous . Divided into t\.JO toothlike projectionsj comblike.
equal parts. Pedate , Bipalmate. Palmately cleft
Dimidi at e . Divided into unequal halves. or divided with l ateral lobes
Dissected . Irregularly cut into cleft or divi ded .
numerous segments . Pinnatif id. Cut pinnately.
Divided . Cut 3/4 to a lmost e nt ire Pinnatisect . Sectioned or divided
distance to middle o f structure . pinnate l y into distinct segments .
Furcate . Forked. Quadrif i d . Cut or divided into f our
I ncised . Hargins sharply and deeply lobes or parts .
cut , usual ly jaggedly . Septate . Divided by internal parti-
Lacerat e . I rregularly cut , appear- tions into locules or cel ls.
ing torn . Se r rated . Cut in to sa\.Jlike tee th.
Laciniate . Cut int o closely parallel Trifid . Cut or d ivided into three
ribbonlike or straplike p roj ec t ions . lobes o r parts.
Lobed . Round-toothed , cut 1/8-1/4 of Tr ifu rcate . Divided into three f orks
distance to middle of structure. or bran ches ; three- fo rked .
III. SHAPES

A. Shapes- Plane and Solid (Figure 6- 13, p . 131)

1. Symmet ric Figur es . Based on terminology in Taxon (Vol. 11) a nd Stearn (1966) .
A. Elliptic. Hith widest axis at midpoint of structure and with margins
symmetrically curved .
Plane L/t-! Sol i d L/D
a . Narrowly e l liptic more than a. Narrowly ellipsoid more than
6 : 1- 3 : 1 6 : 1- 3 :1
b. Elliptic 2 :1-3 :2 b . Ellipsoid 2 :1- 3 : 2
c . Hi dely e ll i ptic 6 : 5 c . Broadly ellip so id 6:5
d . Circul ar 1 : 1 d . Sphe r oid 1 : 1
e . Oblate 5 : 6 e . Oblo id 5 : 6
f . Transvers ely elliptic 2 : 3-1 :2 f . Transvers ely ell i psoid 2 : 3-1:2
g. Narrowly transversely elliptic g. Lenticular 1 :3-1:6 or more
1 : 3-1: 6 or more
130 6

h. Trul l ate 2 : 1-3 : 2 b. Trulloid 2 :1 -3 : 2

c. Widely trullate 6 : 5 c. Broadly trulloid 6 : 5
d. Very widely trullate 1:1 d. Very broadly trulloid 1 : 1
e. Hidely depressed trullate e. Broadly depressed trulloid 5 : 6
5,6 f. Transversely depressed trulloid
f. Transversely depressed 2: 3 -~_: 2
trullate 2 : 3-1 : 2
G. Obtrullate . Inversely trul1ate .
Plane L/11 Solid LID
a. Narrowly obt rullate more a. Narrowly obtrulloid more than
than 6 : 1-3 : 1 6 : 1-3 : 1
h. Obtrullate 3 : 2-2:1 b. Ohtrulloid 3 : 2-2 : 1
c. Widely obtrullate 6 : 5 c. Broadly obtrulloid 6:5
d. Very \"idely ohtrullate 1 : 1 d. Very broadly obtrulloid 1 : 1
e. Indely depressed obtrullate e. Broadly depress ed obtrul1oid
5:6 5:6
f . Transversely depressed f . Transversely depressed obtrul-
obtrullate 2 : 3-1:2 laid 2:3-1:2
H. Triangular . \~ith th r ee sides and three a ngles.
P1aneLN Solid L/D
a . Linear-tr iangular mor e a . Subulate more than 12:1
than 12 : 1 b . Narrm..rly pyramidal 6 : 1-3 : 1
b . Narrm..r1y triangular 6 : 1-3:1 c . Triangular 2:1-3:2
c . Triangular 2:1-3 : 2 d . Broadly deltoid 6 : 5
d. Hide1y deltate 6: 5 e . Deltoid 1 :1
e . De1tate 1 : 1 f. Shallowly deltoid 5 :6
f. Shallm..r1y deltate 5 : 6 g . Shallowly pyramidal 2:3-1 : 2
g . Shallowly triangular 2:3- h . Very shallowly pyramidal 1: 3-
1 :2 1 : 6 or more
h. Very shallowly t riangular
1 : 3-1 : 6 or more
I . Obtriangula r. Inversely triangular .
Plane L/FJ Solid L/O
a . Linear-obtriangu1ar or a . Linear-obpyramidal or narrowly
narrowly cuneate more cuneiform more than 12 : 1
than 12 :1 b. Cuneiform 6 : 1-3:1
b. Cuneate 6:1-3: 1 c . Obpyramidal or broadly cuneiform
c . Obtriangular 2 : 1-3 :2 or 2:1-3 : 2
widely cuneate d . Broadly obdeltoid 6:5
d . Hide1y obdeltate 6 : 5 e . Obde1 toid 1 : 1
e . Obde1tate 1:1 f. Shallowly obdeltoid 5 : 6
f. Shal l owl y obdeltate 5 : 6 g . Shallowly obpyramidal 2 : 3-1 : 2
g . Shallowly obtriangu1ar 2:3- h. Very shallowly obpyramida1 1:3-
1 :2 1:6 or more
h . Very shallowly obtriangular
1 : 3-1 : 6 or more
2. Special Plane Figures--Out1ine
a . Acicular . Needlelike, round d . Oimidiate. Inequilateral with
or grooved in cross section . one - half \..rho1ly or nearly
b . Auricu1iform . Usually obovate \..ranting.
with two small rounded , e . Falcate. Scimitar-shaped .
basal lobes. f . Filiform . Threadlike , usually
c. Cordiform. Heart - shaped . flexuou s.
132 Fi gure 6-14

SHAPES

..,
,:-.

:/ ' :i
'f

BOh,lliform Capitate Clavate Cochlea!, Conicol Cotyl,form

Croleri/arm Cymbiform Discoid Dolobri form Folca te Flobe11i lorm

·
o O
\
."' ,;
. "

. , , _. _", ''~'-

Fusiform Len liCulor Lingulate Nap iform NOdifarm Obconic

Pa te llifo rm Pis ifor m Pyriform Rec tan gular Sp irol Ste llo te

..
" ~'

T ere le Hol/ - terete Turbinale Tu rgid UmbonOIE

6 133

g. Hastiform . Triangular I"ith 1. Peltifo~m. Rounded \"ith petiole

two flaring basal lobes. attached to center of blade or
h. Lunate . Crescent-shaped, apparentl y to laminar tissue .
with acute ends. m. Rectangular. Box-shaped , longer
i . Lyrate . Lyre - shaped ; p i nna- than Hide .
t i Ei d Hith large terminal n . Reniform . Kidney-shaped , I"ith
lobe and smaller lower shallow sinus and widely
lobes . rounded margins .
j . Obcordiform . Inversely cordi- o. Runcinate. Oblanceo l ate with
form . lacerate to parted margins .
k . Panduriforrn . Fiddle- shaped ; p . Sagit tiform. Triangular-ovate with
ohovate with sinus or in- two straight or slightly incurved
dentat i on on each s i de near basal lobes .
base and with tl"ro small q . Spathulate or Spatulate . Oblong
basal lobes . or obovate apically with a long
attenuat e base .
3. Special Solid Fi gures (See caly x, corolla, and perianth types f or special
3-dimensional shapes, Sec t . A, VIII) (Figure 6-14, p . 132)
a . Acerose . Needle- shaped ; x . Half-terete . Flat on one side ,
sharp . terete on other ; semicircular
b . Annular . Ring- like . in cross section .
c . Arcuate . Ben t like the arc y. Hippocrepiform. Horseshoe-
of a circle . shaped.
d . Botuliform. Sausage-shaped . z . Lenticular. Biconvex, usually
e . Cap i llate . Hair-shaped . elongate and fl attish .
f . Capitate . Head-like . aa . Lingulate . Tongue-shaped , p l ano-
g . Clavate. Club-shaped . convex in cross section .
h . Cochleate. Snail-shaped . bb . Neniscoidal. Thin and concave-
i . Compressed or Complanate . convex .
Flattened . cc . Napiform. Turnip-shaped.
j . Conical . Having figure of dd . Navicular . Boat-shaped .
true cone . ee. Nodiform or Nodulose . Knotty or
k . Coroniform . Crmm- shaped . knobby , as the roots of most
1 . Cotyliform. Cup - shaped . of the Fabaceae .
m. Crateriform . Shallow cup- ff . Obconic . Inversely conical .
shaped as the involucre of gg . Pate l liform. Knee-shaped ; d i sk-
some species of Que r cus . shaped .
n . Cruciform or Cruciate . hh . Pisiform . Pea-shaped .
Cross-shaped. ii . Pyriform . Pear-shaped .
o . Cy l indric . Long-tubular . jj . Rectangular. Boxlike, longer
p . Cymbi form . Boat - shaped . than wide .
q . Discoid . Orbicular \"ith kk . Spiral . Twi sted like a corkscrew .
convex faces . 11. Stel l ate . Star- shaped .
r. Dolabriform . Axe-shaped . rnrn. Strombiform. Elongate 5nail-
s . Excent r ic . One-sided ; off- shaped .
center . nn . Te rete. Cylindrical.
t . Falcate o r Seculate . Sickle- 00 . Torose . Cylindrical with con-
shaped . tractions at inter vals .
u . Fi stulose . Hollow , as a culm pp . Turb i nate . Top-shaped ; obcon i c.
without pith .- ~- qq . Turgid . Tumid or swollen .
v . Fl abe ll iform. Fan-shaped . rr . Umbilicate . Depressed in the
w. Fusifo r m. Spindle - shaped ; center .
b r oadest in middle and 5S . Umbonate. Round with a projec -
tapering to each end . tion in center .
134 6

tt . Umbraculiform . Umbre lla-shaped . uu . Ve r miform . Horm-shaped.

B. Apices and Bases (Figure 6-15 , p . 135)

(Leaves , petals, sepals , scales , bracts or other flatte ned struc tures)
1. Apices and Bases t"Hh Sinuses .
a . Retuse . Lobe rounded; sinus e . Reniform.. _ Lobe rounded ; sinus
depth to 1/16 distance to depth variable; oute r margin con-
midpoint of blade ; margins vex or stra i ght , inne r concave .
convex . f . Auriculate . Lobe rounded ; sinus
b . Emarginate . Lobe rounded; depth variable; outer margin con-
sinus depth 1/16-1/8 dis- cave, inner convex or straight.
tance to midpoint of blade ; g. Lobate . Lobe rounded; sinus depth
margins straight or convex . variable ; ou ter and inner mar -
c . Cordate (Ap ex obcordate) , gins concave .
Lobe rounded; sinus depth h . Sagittate . Lobe pointed and
1/8-1/4 distance to mid- oriented downward or imvard in
point of blade ; margins relation to petiole o r midrib;
convex and/or straight. sinus depth variab l e; mar gins
d. Cleft . Lobe rounded; sinus variable.
depth 1/4-1/2 distance to 1. Hastate . Lobe point ed and ori-
midpoint of blade ; mar- ented outward or divergent in
gins convex and/or straight. relation to petiole or midrib;
sin us depth variable; margins
variable .
2. Apices and Bases \~ithout Sinuses.
a. Truncate . Cut straight e . Acuminate (Base narrowly cuneate) .
across ; ending abruptly Hargins straight to convex
almost at right angles to forming a terminal angle of
midrib or midve in. less than 45° .
h . Rounded . Hargins and apex f . Caudate (Base attenuate) . Acumi-
forming a smooth arc . nate with concave margins .
c . Obtuse . Margins st raight g. Hastate . Narg ins variable ; lobe
to convex, forming a pointed; oriented outward or
terminal angle more than divergently in relation to
90° . petiole or midrib .
d. Acut e (Base cuneate ) . t-lar- h. Cuspidate . Acute but co riaceous
gins straight to convex and stiff .
formi n g a termina l angle i. Spinose or Pungent . Acuminate
45° _90° . but coriaceous and stiff .
3. Apices with Hidrib. Hidvein or Vein Extension .
a . Apicula te. Nore than 3 : 1 d . Hucronate . Less. than 3:1 lhv,
l/w , usually slightly straight and stiff.
curled and flex uous . e . Nucronulate. 1 : 1 l/w or broader
b . Aristate. Nore than 3:1 than l ong; s traight.
l/w, usually prolonged , £. l-Iuticous . Without a vein exten-
straigh t and stiff . sion, awn or hair .
c . Cirrhous . More than 10 :1 g. Piliferous. More than 20 : 1 lh.,
l/w , coiled and flexuous . hair-like, flexuous .
4. Specialized Bases a nd Leaf Attachments .
a . Amplexicaul . Completely c. Connate or Connate - per foliate .
clasping the stem . Having bases of opposite leaves
b . Clasping . Partly surrounding fused around th e stern .
the stem . d . Decurrent. Extending along stem
downward from leaf base .
Figure 6-15 135

LEAF APICES , ATTACHMENTS AND BASES

Acum ina te
) (~'<A ,~,~) ,Wi,
Acute Apiculote Ac ista!e Couda Te Ci rrhose

~}~ ,~\I ~~\ ~@

Clef! Cusp idate Ema rginate Mucronate Mucronulo te Qbco,dole

~ ~ ~.~Obtuse Re tus e Ro unded Sp; nose

(([ IIi1
Tru ncate

Ample~ic au le Attenuate Auricu late Conno Te per/ohore Co rd ate Cu neate

~ l ff l~\ . ltY1~
'f
Oecurrent
.~
Hoslele LiQulale Oblique
..~~w
Ocreote Pellole

.~ I v~~ ~ ~ II
Perlaliale Petiolate Renif or m Sessil e
136 Figure 6-16

MARGI N S

Cle f t Crenate Crenu lale Crlsp ole

!'h
'{,il \J. f(~\
'I' ~
,!
\

•
}

Dent icu l ate

" '
w
Dlv ided Do u ble -s errafe Entir e
~

Erose
,
'I

Lobe d

;<1\
Polmo lif. d
tPoned

'I~- (;~ ,
! I I
i !
\~ i
~
I
1
P i nnoti fid
'"
Rev olute Se rraTe Se rru late S inuole Undulate

ARRANGEMEN T

,
AlTe rn aTe Dec usso te Ois l ictlO U5 Wh Orled Eq uilonl Im b ricote Fascicled

D i c hOTomous Palma te ly netled ? "lnol e ly nelte d P en ni- p aro lle l Ret iculate Parallel
6 137

e. Ligulat e . Hav i n g a tongue- i. Peltate . Usually having petiole

like outgrowth at base attached near the center on the
of blade o r t op of shea t h . unde r side of blade .
f. Obl i que . Having an asymmet- j. Perfo liate . Having base com-
rical base . pletely s ur rounding s tem.
g. Oc r e at e . Having a stipular k. Pe tiolat e . Wi th a pe tiole .
tube surro unding stem 1. Sess ile . Wit hout a petio le.
above insertion of petiole m. Shea thing . Having tubular struc-
Dr blade. ture enclosing stem below appar-
h. Ocreolat e . Diminutive of ent insertion of blade or petiole .
ocrea te ; us ually applied n. Surcurrent . Extending a long stem
to bract bases . upward from leaf base .

C. Ha rg i ns (Figure 6- 16 , p . 136)
(Leaves , petals , sepa l s , bracts, scales o r ot he r fla tt ened str uc tur e)
Note : Fo r precison in ma r gin description the type (as desc r ibed below), t he
--symme try of the individua l tooth, the ma r g in s of the i nd ividual ;;ooth, the
apex of the individual tooth, the type of sinu s (round ed or angled) , the
number o f t eet h per unit of ~rgin measuremen t , the spacing (regular or
irregular ) of the teeth , the natur e of teeth (simple or compo und i n two
or more size groups) s hould be ind icated or described .
1 . Margin Type s j . Denticulate . Diminutive of den-
a . Aculeat e . Prickly . tate , cut to 1/16 distance to
b . Bi crenat e o r Doubly-crenate . midrib o r mid vein.
\"ith smaller rounded teeth k . Divided. Indentations or inci -
on l arger rounded teeth . s i ons cut 3/4-a l most compl etely
c . Biserrate o r Doubly-serrate. to midrib or midve i n .
Wit h sharp l y cut t ee th on 1 . En tire . Without indentations or
the margins of larger incisions on margins ; s mooth .
sharply cut teeth . m. Eros e . I rreg ularl y , shallowl y
d. Ciliate . I.;ith trichomes pro- toothed and/or lobed margins;
trud ing from margins . appearing gn~wed .
e . Cleft . Indentations or in ci- n . Filamentose or Filiferous . \.Jith
sions c ut 1/4-1/2 distance coa rse marginal fibers or threads .
t o mi d rib o r midvein. o . Fimbr iat e . Harg in s fringed .
£. Crenate . Shallowly ascend- p . Fimbriola t e. Ni nu tely fimb riate.
ing round- toothed , or teeth q . Incised. ~~ r gins sharpl y and
obt use ; teeth c ut l ess than d eepl y cut, usually jagged l y .
1 /8 way to midrib o r mid- r. Involute . Margins r olled inward.
vein. s . Lacerate . Mar gins i rregularly
g . Crenulat e . Dimin utive of cut , appear ing torn .
crenate , t ee th c ut to 1/16 t. Laciniate. Margins cu t into
distance to midrib or mid- ribbon-l ike segme n ts .
vein . u. Lobed . Large , round-tooth ed, cut
h . Crispate . Cu rled ; margins 1 /8-1 /4 distance to mjdvein.
divided a nd twis ted in mo r e v . Pa l ma t i f id . Cut pa l ma tely.
than one plane . w. Par ted . Inden tations or inci s i o n s
i. Denta t e . Margins with cu t 1 /2-3 /4 dis tance to midrib.
roun de d or sharp, coa r se x . Pinna tifid. Cut pinnatel y.
teet h that point outwards y . Repand . Sinuate with indentions
at right angles to midrib le ss than 1/16 distance to mid-
or midvein , cut 1/16 to rib or midvein.
1/8 distance to midrib or z . Re trorsely Crenate. Rounded
midvein . teeth direct ed toward base.
138 6

aa . Retrorse l y Serra te. Sharp or ee . Sinuate. Hargins sha llOldy

pointed teeth direc ted toward and smoothly i nd e nted , ~Javy
base . in a horizontal plane , ,,,rith-
bb . Revolute . l'lorgins roll e d uncle r . out distinctive teeth or
ce . Serrate . Saw- toothed; t eeth lobes, indented 1/16-1/8
sharp and ascending . but cut d istan ce to midrib or mid-
1/16-1/8 distance to mi drib vein.
or mid vein . ff . Undulate . Hargins shal lml'ly
dd . Se rrulate . Diminutive of and smoothly i ndented, wavy
serrate, but cut to 1/16 dis - i n a vertical plane .
t ance to midrib or midvein .

IV . SURFACE-VENATION-TEXTURE

A. Con fig ura tion of Sur face

(Cla ssific ation based on patterns of configuration)
Aciculate . Finely ma r ked as with Plicate or Plaited . Fluted, longi-
pin pr i cks. fine lines usually tudina lly fo l ded.
randomly arranged . Punctate . Covered Hith minute im-
Ala te . I"inged . pressions o r depressions .
Alveo l ate . Honey-combed. Pustulate . IHth scattered bl ister-
Areo l ate . Divided into many angu- like swellings .
lar or squarish spaces . Reticulate . Netted .
Bullate . Puckered or blist ered. Ribbed . Hith longitudinal nerves .
Canaliculate . Longitudinally Ringed . With old bud scale scar rings.
grooved, usually in relat ion Rugose . Covered with coarse reticu-
to p etioles or midribs. late lines .
Cancel late or Clathrate . Latticed . Ruminate . Coarsely wrinkled , appear-
Corruga t e . Ridged . ing as chewed .
Costate. Coarse l y ribbed . Scarred . Ihth old l ea f base , stipular
Fenest r ate. I\lith windowlike holes and/or b r anch sca r regions .
thr ough t he l eaves or other struc- Smooth or Plane. l.athout configura-
tures . tion .
Flexuous . Coarsely undulate \.,Iith Striate . \-lith longitudin al lines .
fold s at r ight angles to long Sulcat e. Hith lon gitudinal grooves.
ax is . Tort uous. Having the surfa ce vari-
Foveo lat e . Pitted . ously t\.,Iisted.

B. Venation
(Class ification based on ve i n pattern. See Chap t er 7 . sec tion D for detailed
s tructure and page 194 for illustrations . Based on Hickey [1973])

1- Genera l
Acrod romous. Hith two or more pri- Actinodromous . Hith three or more
ma r y or strongly developed secon- primary veins diverging radially
dary vein s diverging a t or above from a single point at or above
the base of the blade and runn ing the base of the blade a nd r unning
in convergen t arches toward the toward the margin , reaching it or
apex over some or all of the not .
blade length . the arches not
bas ally c urved .
6 139

Brochidodromous . Inth a single pri- diminishing distally within the mar-

mary vein. the secondary veins gin and interconnected by a series
not terminating at the margin but of cross-veins without forming con-
joined together in a series of spicuous margina l loops.
prominent upward arches or mar- Hyphodromous . With a single primary
ginal loops on each sid e of the vein and all other v e nation absent,
primary vein. rudimentary , or concealed within a
Campylodromous. I,hth several pri- coriaceous or fleshy blade .
mary veins or their branches Palinactinodromous . Actinodromous , the
diverging at or close to a single primary veins \"ith one or more sub-
point and running in strongly sidiary radiations above the pri-
developed , basally recurved mary one.
arches Hhich converge toward the Parallelodromous . With two or more
apex, reaching it or not. primary veins originating beside
Cladodromous . Hith a single pri- one another at the blade base and
mary vein, the secondary veins running more or less parallel to
not terminating at the margin the apex where they converge .
and freely ramified tOHard it . Reticulodromous . With a single pri-
Craspedodromous, Mixed . Hith a mary vein, the s econdary veins not
single primary vein, some of the terminating at the margin and los-
secondary veins terminating at ing their i dent ities near the mar-
the margin and an approximately gin by repeated branching, yield-
equal number otherwise . ing a dense reticulum.
Craspedodroffious, Simple . With a Semicraspedodromous . \-lith a single
single primary vein , all of the primary vein, the secondary veins
secondary veins and their branches branching just \"ithin the margin ,
terminating at the margin. one branch from each terminating
Eucamptodromous. I·lith a single pri- at the margin and the other form-
mary vein , the secondary veins ing a marginal loop and joining
curved upward and gradually the superadjacent secondary vein.

2. Special (Figure 6-16 , p . 136)

(Traditional classification)
Dichotomous. With veins branching Palmately or Digitately Netted .
or fork ing in pairs equally . With three or more primary veins
Netted or Reticula te. Inth veins arising from a common point .
fo rming a network . Parallel. \-lith veins extending from
Pinnately Netted . \-lith secon- base to apex, essentially parallel .
dary veins arising from mid- Penni-parallel. With v e ins exte nding
rib or midvein . from midrib to margins , essentially
parallel.

C . Surface (Fi~ure 6-17 , p . 141)

(Classification based usually on epidermal outgrowths or excrescences , but not
trichomes)
Asperous . Having a rough su r face. Glaucescent. Sparingly or slightly
Bristly . Beset \"ith bristles. glaucous .
Echinate . Covered with spines . Glaucous . Covered Hith a bloom or
Farinaceous . Nealy . "' ~ ..... smooth , t~axy coating .
Glabrate or Glabrescent . Becoming Glutinous . Having a shiny, sticky
glabrous. surface .
Glabrous. Smooth; devoid of Granular . Finely mealy , covered
trichomes . Hith small granules.
Glandu l ar . Covered with minute, Greasy or Unct uous. Slick , Oily,
blackish to translucent glands . slippery to touch .
140 6

Huc ilaginous. Gummy or gelatinous . Resinous . Having a yellowish,

Huricate . Covered with short, hard sticky, exudate .
protuberances . Roridulate or Oe...,y . Covered \-lith
Huriculate . Minutely muricate . \.,raxy platelets , appearing det.,ry .
Opaque . Hith a dull surface . Scaberulent . Approaching scabrous.
Prickly . Hi th prickles. Scabridulous. Minutely scabrous.
Pruinose, Frosted, or Sebiferous. Scabrous. Having a harsh surface.
\\li th a heavy wax coat. Shining, Nitid, or Laevigate . Lus -
Pubescent . Covered Hith dense or trous , polished.
scattered trichomes. Spiculate. hlith crystals in or on
Pulverulent. Covered tdth fine, the surface.
pO\.,rdE'ry Nax granules. Squamose . Having coarse scales .
Ramen taceous. Having many thin Subglabrate. Almost glahrous.
scales, as on the epidermis of Tuberculate or Verrucose. l<lith a
some ferns . Harty surface.
Viscid . Sticky or glutinous.

D. Vestiture or Indument (Figure 6 - 17 , p . 141)

(Classification based on type of hairy cover, see Chapter 7, section E for
individual types of trichomes and page 21)0 for" ill ustrations.)
Aculeate . Prickly. Inermous. Unarmed . Without prickles
Arachnoid. CobHebby . or spines .
Barbed . l<lith short, rigid reflexed Lanate . Covered with long, inter-
bristles or processes. t,dne d trichomes, cottony .
Barbellate . Minutely barbed. Lanuginose . Cottony, similar to 1an-
Bearded or Barbate . Hith long tri- ate but trichomes shorter.
chomes usually in a tuft, line Lepidote or Squamu1ose . Covered \~ith
or zone . minute scales .
Bristly . Covered with stiff, strong Paleaceous. Hith small membranous
trichomes . scales, chaffy.
Canes cent or Incanous . Covered with Pannose or Fe l ted . IJ ith matted,
dense, fine grayish-white trichomes. feltlike layer of trichomes .
Ciliate. With conspicuous marginal Papillose: Covered with minute
trichomes. tubercles.
Ciliolate. Hith tiny or small mar- Pilose. Hi th soft , shaggy trichomes.
ginal trichomes . Puberulent. Minutely pubescent.
Comose . Hith a tuft of trichomes, Pubescent. Usually with straight,
usually apical. slender trichomes .
Dm-my. Covered ,~ ith short, ueak, Scurfy or Lentiginous . Hith exfoli-
soft trichomes . ating scaly incrustations.
Floccose. Covered with dense, ap- Sericeous. Hith long, Silky trichomes,
pressed trichomes in patches or usually appressed.
tufts . Setose , Setaceous . Having setae or
Glabrate. Hithout trichomes . bristlelike trichomes .
Glandular . Covered with secretory Strigillose . Diminutive of strigose .
or excretory trichomes. Strigose. Covered with sharp, coarse,
Glochidiate. Hith barbed trichomes, bent trichomes usually with a bul-
glochids, usually in tufts. bous base.
Hirsute. Cover ed ,~ith long, rather Tomentose. Covered with dense, inter-
stiff trichomes . woven trichomes .
Hirsutullous or Hirtellous. Ninutely Urent or Stinging. Hith erect , usually
hirsute . long trichomes that produce irrita-
Hispid. Covered ,-lith very long, tion t~hen touched .
stiff trichomes . Velutinous . Covered with dense ,
Hispidulous . Approaching hispid, straight , long and soft trichomes;
minutely hispid . pile-like .
 "; .

Figure 6-17 141

BRANCH I NG PATTERNS

Rame ntoceous Res inous

VESTITURE

Barbed

Ci liole Comosc F l accose Gl oCh ,dl0l!;

' f".

c=::J,/\~\
i" ,t.,. 'j
. JI, \ .
r·,,. J
6·:,'·~?- "~·:':J3
Ponnos e Pop il lo se

>c,i/"
~' '!I'Jt:
~L Urent
~)

Velut inous Vi llous

 Figure 6- 18
142

INFLORESCENCE SEX
FLOWER SEX

AQomOQynOU$ Androhef~ Hermophr o- Gyneh e,- He r mC phrO- Agamohe r - Hermcphro -

mophr odi liC d o ndrau $ mop hro d i !iC Q'g1n QUS mooh rodinc d o go mOU$

FERNS

Dimorp~ IC Pinnoe
Dimorph ic Fronds
Fr on d

LOWER VASCULAR PLANTS

S e lo(l' nello
Ly copodium E(j u isetum

REPRODUCTIVE STR U CTURES

$o rU$ Sp o ran gLum I ndus ium

6 143

V11105U10U5 . l-!inutely villous. Villous . Covered ,-lith long , so f t,

crooked trichomes .

E. Texture
(Classification based on consistency of l eaves , scales , petals , fr uits and
oth er part s)
Baccate . Juicy and very s uccu lent . Hyaline. Thin and translucent or
Carnase or Sarcous . Fleshy . transparent .
Cartilaginous . Hard and tough but I ncrassat e. Th i ckened .
flexible . Indurate . Hardened .
Ceraceous . h1axy . Ligneous . 1,.1oody ,
Chartaceous . Papery, opaque and Nembranous , Thin and s emi-translu-
thin . cent; membrane-like ,
Coriaceous . Thick and leathery . Osseous . Bony .
Corneous . Horny . Pannos e . IHth a fel ty te x tur e .
Crustaceous . Hard, thin, and Pellucid . Clear, transparent .
brittle . Sc ar i ous . Thin and dry, appearing
Diaphanous . Translucen t . shriveled .
Fibrous . Having loose, \.,roody Sclerous . Hard .
fibers . Spongy . Cellular; sponge-like .
Flaccid . Lax and \,'eak . Suherous. Corky.
Gelatinous . Jellylike; soft and Suffrutescent . T...'oody basally ,
quivery . herbaceous apically .
Herbaceous . Soft and succulent . Hoody . Hard and lign ified .

v. SEX

A. Flm,'er Sex (Figure 6-18, p . 142)

(Classification based on sexual condition of the individual floh'er)
I mperfect or Unisexual. IHth sta- Per fec t or Bisexual. ',hth bot h sta-
mens or carpels absent in the mens and carpels or pistils in
fl m"e r . the f lmier .
Neuter or Agamous. Hithout sta- Pistillate , Carpella te, or Female .
mens and carpels in flQl,!er; or Ihth pistils or carpels only in
sex organs abortive. the flQl.;er .
Staminate or ~la le . I'lith stamens anI,:
in the floHer .

B. Inflorescence Sex (Figure 6-18, p . 142)

(Classification based on sexual condition of ind ividual flOHers in different
parts of inflorescence; arrangement of flOI,'e rs by sexual condition in an
inflorescence)
Note : If plant sex is staminate or pistillate or hermaphroditic or ne u ter,
then inflorescence sex h'ould be r espectively the same . The term in ( )
beloH is an example of t he sexual condition of one inflorescence type , the
head, as in composites .
Androgynecandrous . Inf lorescence Agamandrous . Inflo rescence 1.Ji th
with staminate flOHers above and neut er floh'ers in side or abovE'_
belm.; pistillate, as ~ in the spikes and staminate outside or
of some species of Carex . (agamandrocephalous )
Androgynous . I nflorescence I"ith Agamogynous . I n floresc ence I.; i th
staminate flm.;ers inside or neuter flOl,'ers inside or above
above and pistillate outside or and pistillate outs i de or bela,,! .
belm,' . (androgynecephalous) (agamogynecephalous)
144 6

Agamohermaphrodi tic . Inflorescence Gynehe rmaphrodi tic . Inf lorescence

\·;i t h neuter floHers in side or Hith pistillate flOt"ers inside
above and hermap hrodi tic out side or or above and hermaphroditic out-
belm". (agarnohermaphrodicephalous) side or below . (gynehermaphro-
Andragamous. Inflorescence with stam- dicephalous)
inate flowers inside o r above and Hermaph r ooa·gamous . Inflorescence
neuter EIOl.'e rs outside or beloH . with hermaphrod it ic floHe r s in-
Candragamocephalous) side or above and neuter outside
Androhermaphrodi ti c . Infl or escence or belOt,'. (her maphrodagamocepha-
"'ith staminate flowers in side or lous)
a~oveand hermaphroditic outside or !lermaphr odandro us . Inflorescence
b eIOl" (androherrnaphrodicephalous) I~ith hermaph r oditic flol,/e r s in-
Gynagamous . InflotE:scence I.;ith pis- side or above and stamina te
tillate floh'e r s inside or above and outside or below . (hermaphrodan-
neuter flol·.'ers outside or below . drocephalous)
(gynagarnocephalous) llermaphrodigynous . In florescence
Gynecandrous . Inflorescence with pis - Idth hermaph roditic flo\-lers in-
tillate flm.le rs inside or above and side or above and pistilla te out-
staminat e ou t sid e or belm,' . as i n si de or helol';. (h ermaphrodigyne-
spikes of some species of Carex . cephalou s)

C. Plant Se x
(Classification based on sexual condition of all flowers on the pl ant(s) within
the species)
Androdioe cio us . Some plants \lith He rmaphrodi tic or Honoclinous . Plant
staminate flmle r s and some Hith I-.'ith all floHe r s perfect .
perfect flOt"ers . ria Ie or Stamina te. Plant Hi t h stami-
Andromonoecious . Plant Hith stami- nate flowe r s only .
nate and perfect flOi-:ers . Nonoecious . Plan t t·Ji th all flm-/ers
Dicl i nous . Plant Idth impe r fect i mperf ect , but st aminate and
flm-.'ers; st ar.1ens and ca rp els in pist ill ate flolJe r s on same plant .
separate flOt~ers. Polygarnous . Pl an t \-lith pe rfect and
Dioec ious. Plan t l11th al l flOI,'ers ir.1p erfec t flol-l ers .
imperfect , bu t staminate and Polygamo- dioecious . Plants dioecious ,
pistillate on sC!pdrate plants . but t-li th some perfect flowers on
Female , Carpellate , or Pist i llate . stami n ate o r pistillate plants or
Plant I-lith pistillate flOl-lers ho th.
on l y . Pol ygamo-monoecious . Plant monoe-
Gynodioecious . Some plants Hith per- cious , but \-lith some perfect
f ect ilm.:ers and SOMe 141th pis- flOl4e r s .
ti llate . Trioecious . Pl ants stamin ate, pis -
Gynomono ecious . Plant "'ith pistil- tillate or perfect .
lat e and pe rfect flOl-.'e rs.

D. Gene ral Sexual Terms

Agamo us or ~leu t er . I.'it hou t s ex ; Hete rosexua l . Inf lo rescences or
se xual organs abortive . fl m-le rs l,tithin t he plant t-lith
Heterocephalous , Heterocymo us, Hete r o- diff erent sexua l condi tions.
spicous. Heads , cymes , spikes with Homocepha lo us. Homocymous . Homo-
flOt-fers of different se xual condi - spicous . Heads , cymes , spikes
tions . Note : other inflorescence I.. ith Elm"ers sexually uniform .
loJOrd stems could be used for app ro- Homose xua l . Inf lores cences or
priate inflorescence type . flOt4ers sexually uniform .
6 145

VI. POLLINATION AND FERTILIZATION

A. Pollination Types
(Classifica t ion based on agent by which pollen i s t r ansferred from t he pollen
sac or anthe r to the s tigma or ovule)
Anemophi l y . Pollinated by wind. Myiophily . Pollinated by diptera.
Anthropophily . Pollinated by man. Hyrmecoph ily . Pollinated by ants .
Cantharophily. Pollinated by Necrocoleopterophily. Po llinated
beetles. by carrion bee t l es .
Chei ropterophily. Pollina ted by Ornithophi ly. Pollinated by birds .
bats . Phalaenophily . Pollinated by
Entomophily . Pollinated by insects . moths .
Hydrophily . Poll ina t ed by water . Psycho phily . Po llinated by butter-
Hymenopte rophily . Pollinated by flies .
bees . Sapromyiophily . Pollinated by
Mal acophily. Po llinated by s nail s carrion or dung fl ies .
or slugs . Sphingophily . Po llinated by hawk
Me littophily. Pollinated by bees. moths and noc t urna l l e pidop-
Mic romelittophily. Pollinated by tera .
smal l bees .

B. Pollination Pathways
Chasmantheric Poll ination . Pollen Cleis t antheric Po llin ation . Po llen
transferr ed f r om a normally de- not transferred from a normally
hisced anther by a pollinating dehisced anthe r by a pollinating
agent with pollen gra i n germin- agen t ; pollen gr ain germinates
ation on the s ti gma and subse- t·lithin the anther wit h s ubsequent
quent grOtY'th of t he pollen tube gr owth of the pollen tube through
through stigma, styl e and the the anthe r wall and ovary wall
ovule into the embryo sac . into the ovule and embryo sac .

C. Fertilization Types
(Refers usually to sources of male game tes in pollen parent)
Allogamy or Xenogamy . Cross - Autogamy . Self- fertilization in a
f e rtilization in pl a nt s . s ingle f l owe r.
Alla utogamy . Cross- a nd se lf- Geitonogamy . Fert ilization of one
ferti lizat ion in same plant, flmY'er by anothe r on the same
as i n Viol a . plant.

D. Fertiliza tion Pathways

(Classification based on place of entrance of pollen tub e i nto ovule )
Cha lazogamy. Polle n tube entrance Porogamy. Pollen tube e nt rance
through chalaza . through the mic ropyl e .
Pleurogamy. Pollen tube ent r ance
through side of ovule .

VII. TEHPORAL PHENOHENA

-; ~.... A. Per iodicit y

Aestival. Appearing in s umme r. Anno t i nal or Year ly . Appearing
A1anthous or Semperflorous . With yearly .
flowers appearin g throughout Autumnal. Appeari ng in a utumn.
the year. Biferous . Appearing twice yearly .
146 6

Biflorous. FIO\"ering in autumn as Hatutinal. hlith flowers opening in

well as in spring . the morning .
Bimestrial . Oc cur ring every two Nocturnal. Ope ning during n i ght .
months . Seasonally. Occurring during a
Diurnal. Opening during the day . seasonal cycle , or each season .
Equinoctial. Havin g flowers Hhieh Serotinal. .... ·Opening late; appearing
expand and close regularly at in late summer .
particular hours of the day . Vernal . Appearing i n spring.
Hibernal or Hiernal. Appearing in Vespertine . Hith flowers opening
the winter . eve ning or night ; appearing or
expanding in the evening .

B. Naturation
An thesis. Time of flOl-/ering ; opening Precocious . Developing unusually
of flo\"er \"ith parts availabl e for early.
pollination. Proanthesis . Flower i ng befo r e no r -
BIas tocarpous . Germination of seeds mal period, as spring Elm,ers in
,... hile wi thin the pericarp , as in the fall .
Rh izophora. Protandrous . With s tamens o r anthers
Coetaneous . Flmveri ng as the leaves devel oping befo r e carpels or
expand; synantherous. stigma .
Dichogamous . Hith maturation of sta- Protan t herous . Nith leaves appearing
mens o r anther and carpels or before flowers .
st i gma at differe nt times. Protogynous . Hith ca r pels or stigma
Homogamous . Hi th maturation of sta- maturing before stamens or an-
mens o r anther and carpels or ther s .
s ti gma at same time . Synantherous . \.,Iith leaves and
Hysteranthous. With leaves appearing flOHers appearing a t same time.
after flowers.

c. Duxation
Annual . Living one year or l es s . Fugacious . Ephemeral , usual l y
Biduous. Lasting two days . applied to plant parts .
Biennial. Living two years , usually Narcescent . Usually ephemeral with
f lowering se cond year . persistent remains; withering
Bimestrial. Lasting t ....'o months. persistent.
Caducous. Dropping off ver y early, NonocarpiC, Hapaxanthic . Perennial
usually applied to floral parts . or annual , flowering and fruiting
Cau locarp ic . Plants having the stem once, then dying ; fruiting once .
living for many years, bearing Perennial, Polycarpic. Living more than
flowers and fru its . two years; fruiting more than once.
Cladoptosic. Shedding of branche s , Persistent. Remain ing attached; applied
st,erns and leaves simultaneously, to individual parts.
as in Taxodium . Pliestesial or Nultiperennial. Hono-
Deciduous . Persistent for one g r ow- carpic but living several to many
ing sea son. years before flol,ering , as in Agave.
Deliquescent. Soften i ng and wasti ng Rhizocarpic. Plants having the roots
away . living for many years with the
Ephemeral. Ge rmina ting , grO\ving , stems dying annua lly .
flowering a nd fr uiting in a short Seasonally Deciduous . Falling af ter
peri.od, at.: 1:.(1'O!;" JLf'p rt herbs . one growing season .
Evanescent. Passing away, disappear- Summer Annual. Germinating in spring
ing early . or early summer and flO\~ering and
Evergreen . Persistent two or more fruiting in late summer or early
growing seasons . fall , then dying .
 .,....
lit;>,: (;.;';'

6 147

Winter Annual . Living less than and usually floHering and fruiting
one year but through the winter; in early spring.
germination usually in late fall ,

IX . GROlITH AND DEVELOP}lENT

A. GroHth Regions
Adaxial or Abaxial . GroHth region Intercalary. GroHth region near the
localized on top or bottom of a base of an internode or base of
leaf part or any dorsi-ventral blade .
structure. Interst itial . Growth all-over in an
Apical or Terminal. Growth region organ, no localized meristems, as
at the apex of the structure. i n some fruits.
Basal. Gro\"th region at the base Narginal. Grm"th region near the edge
of a blade , as in grasses . of a leaf .

B. Growth Per iodicity

Accrescent . GroHing after flo\o,e r Indeterminate or Evergrm"ing . Cont i n-
ing or bud development has oc- ual groHth of plant parts, not lim-
curred , as the sepals in Hyperi- ited by a cessation of meristematic
cum and bud scales in Carya . activity .
Determinate . Growth of plan t parts, Intermittent. A renewal and cessation
the size of Hhich is limited by of meristematic activity which pro-
cessation of meristematic acti- duces c l usters of stems and/or
vity during the year. leaves along an axis .

C. General Development
Acropetal. Developing upHard, tOl"ard Centrifugal . Developing from the in-
apex. side out\"ard , or from top dOHIlHard .
Basipetal. Developing dm"m"ard, Determinate . Central flower develops
toward base . first , arresting development o f
Centripetal . Developing from the primary axis.
outside inward, or from bottom Indeterminate . Latera l f lowers develop
upward. first with primary axis continuing
to elongate or develop .

D. Developmental Shoot Types

(Classification based on oT1g1n and development of shoots)
Dwarf Shoots or Spurs . Shoots that more shoots and a terminal bud
develop from preformed buds Hhich Hhich Hill develop etc. - -several
have very short internodal lengths times in a season with several
or intervals . flushes of growth .
Epicormic Shoots or Water Sprouts. Heterophyllous Shoots. Shoots that
Shoots that develop f rom dormant develop from winter buds which do
lateral buds on the trunk which not contain the primordia of a l l
have very long and frequently the leaves to develop during the
variable internodal lengths or year .
intervals. -.~... Lammas Shoots. Abnormal late season
Flushing Shoots. Shoots that shoots that develop from the ter-
develop fr om mature terminal minal bud , not a recurring phenome -
buds several times during a non as in flush ing shoots .
season . Terminal bud will Long Bud Shoots . Abnormal buds or
develop shoot with new termi- shoots which elongate, then have
nal bud which Hill develop arrested grm.,th without the
148 6

development of leaves and lateral the lateral buds immediately

branches . beneath the terminal.
Long Shoots . Normal shoots that Sucker Shoots . Shoot s that
develop from terminal or axillary develop from adventitious
buds which have normal internodal buds on old stumps or roots,
lengths or intervals. usu.i11Y after cutting or
Preformed Shoots . Normal shoots that ~nJ ury , whi ch have elongate
develop from \.,Iinter buds which con - internodal l eng ths and intervals.
tain primordia of a l l l eaves that Syleptic Shoots . Abnormal shoots
will expand during the season . that deve l op from lateral buds
Proleptic Shoots. Abnormal late before they have reached
season shoots that develop from maturity .

IX . PATTERNS

A. Symmetry
Actinomorphic or Radial. Hith flora l Irregular . \.Jith flo ral parts Hithin
parts radiate from center like a Hhorl dissimilar in shape and!
spokes on Hheel. or size.
Asymmetric . Hitho u t regularity in Regular . Hith floral parts within
any dimension . a whorl simi la r in shape and
Dorsiventral. Pl anate and having size .
distinct dorsal and ventral sur- Spherical. With multi-dimen-
faces, the t\W usually different . sional radial symmetry .
Equilateral . With halves or sides Zygomorphic or Bilateral . Hith
equal in shape and size. floral parts in two symmetri-
Inequilateral. I·lith halves or sides cal halves .
unequal i n shape and size .
B. Arrangement Systems
(Based on Leppik [1961])
Anthotaxis . Arrangement of sporo- 1/3, 2/5 , 3/8 , 5/13, 8/21, etc .
phy1ls, primarily reproduct i ve rhe numerator rep r esents the num-
in function . ber of turns or spirals around a
Carpotaxis . Arrangement of fruits , stem before one leaf is directly
reproductive in function. above another and the denominator
Phyllotaxis . Arrangement of leaves, r epresents the number of l eaves
pr imarily photosynthetic in func - i n the turns or spirals before one
tion . is direct l y above the other. 2/5
Rhizotaxis . Arrangement of roots. phyll otaxy would mean two twists
Sematax is. Arrangement of semaphyl ls and f ive leaves before one leaf
(petals , sepals, tepa1s) . pri- is directly above the other or
marily advertising (pollinator an angle of divergence of 144 0
attracting) in function . betHeen succeeding leaves in the
xxxxxxxxxxxxxxxxxxxxxxxxxxxxxxxxxxxxx stem (2/5 of 360 0 ) . According
Fibonacc i Phyllotaxis . A fundamen- to Leppik ant hot axis and sema-
tal type of leaf arrangement ex- taxis do not necessarily follow
pressed as a fraction in t"hich the same pattern, \"ith anthotaxis
each succeeding f r action is the in Mi chelia cited as being in 2/7,
sum of the two previous numera- 3/7 , 3/8 , and 4/10 systems of
tors and the sum of the two pre- arrangement .
vious denominators , i . e ., 1/2 ,
6 149

C. Branching Patterns (Figure 6- 17, p . 141)

(Adapted from Hall e [1971])
Dichotomous . Branc hing into two f. I~ith spi ral or whorle d main
equa l par ts. l aterals and dichotomous
Flabella t e . Fan-shaped branching. branches; e . g ., Phyllanthus .
Honopodial. Branching with a main g . I.]ith tiered or clustered
axis and r educed or missing lateral branches . Hain axis
laterals; ~xcu rr en t . wi th periodic lateral branch
a . !,'ithou t lateral branches . production ; e . g ., Bombax .
Main axi s ending in a clus- Sympodial. Branching without a main
ter of l arge l eaves ; e . g . , axis but with many , mor e or less ,
Cari ca papaya. eq ual late rals .
b. With c on tinuou s la ter al a . Leader displacement . Succes-
branches . Naio axis bese t sive l aterals take over l eader
\,,1th laterals; e . g . J Trema grow t h producing umbre lla -
lamarckiana. shaped trees; e.g . , Acacia .
c. Wi th continuous but se lf- ~~l ia; del iquescent .
pruning lateral bra nches b. Leader dis placement by t wo
(excurrent). Nain axi s with equal latera ls (dichotomous
upper laterals pr esen t and subs titution) . Produc tion of
lower having fal l en off; a fo r ked branching system; e . g .,
e . g .• Pinus . Nan i hot .
d. Hi th short- lived l ateral s . c . Leader displacement as a short-
lolain axis beset with phyllo- shoot (appo sit i on) . Usua lly a
rno r phic branches , appearing single lateral becomes domi-
as compound l eaves that may nant; e . g . , Terminalia .
be deciduous ; e . g ., some Note : Other types of s tem growth and
t r opical Rubiaceae, Pycnanthus . branching pa t terns oc cur , parti-
e. \.,lith s piral or whorled main c ularly in tropical woody plants
laterals on main axis and dis- but no ad equate class ific ation ha s
ticho us branches ; e.g ., Pvcnanthus . been devised .

X. SECRETORY AND EXCRETORY STRUCTURES

(Ne c taries, Di gestive Glan ds, Hydathod es , Tanniferous a nd Ethereal Oil Glands)

Clas sification based on St ru ctu re Note: No sa ti sfactor y c l assification scheme

based on structure is available . Some gl and s are single-ce lle d , most are multi-
cellul a r ; some a r e ses sile , some stipitate; some a re cap itate, some are vari -
ously s hap ed ; some are highly mod ified st ru c tures , some are so s i mple tha t they
simply secrete the substance through the cuticle.
A. Classifica tion based on Function .
Digestive . Enzyme secreting Salt Excreting . Glands that
glands found mos t ly on l e aves excret e o r secre te sal t s olu-
of carnivorou s and ins ectivo- tions which usua lly dry during
rous plan ts . t he day and de li quesce at night .
Ethereal Oil Producing . Tannin Bearing . Tann i n produc i ng
Mo s tly aromatic compound pro- glands fo und in va rious parts of
ducing gl ands fou nd on var i ous the plant, pre sumably protec tive
par ts of the plant , without in s ome s tr uctures .
de fi ni te ly known functions. Ha t e r Excre ting . Specia lized vein
Nectar Se cret ing . Sugary com- openings (hydathodes) through
pound producin g gl ands fo und \o,fhich wate r is lo st under ce rtain
mostly on f lora l parts that a tmospheric conditions .
produce attractant s for po l-
linators.
150 6

B. Classification based on Pos i t i on .

1 . Floral Nectaries
a . Petaliferous . Most fre d . Stamina! Disc . Entire disc
quently at the base of frequently nectariferOllS.
petal s . e . S taminal . Host frequently
h . Sepaliferous . Most fre- at . . .base of filament.
quently at the base of f . Staminodial. Frequently
sepals . entire structure is nectari-
c. Septal. At the junction of fe rous.
the septa i n the ovary.
2. Extraf l oral Ne ctaries
a . Bracteal and b. Laminar. c . Nodal. At the nodes.
Hay be lo calized or found
over entire structure .
3. Etherea l Oil Glands.
a. Cauline . Nay be a l lover c . Petiolar . May be allover
(general) or along the ribs (general) or near apex
(costal), or in the grooves (acrope tiolar) or near
(canalicula t e) . base (basipetiolar) .
b . Laminar . Hay be allover
( general) or near apex
(acrolaminar) or near base
(bas ilamina r ) o r marginal .

XI . COLOR

A. Standardized Colors
Use standardized color charts, such as the Ridgeway for solid color descrip-
tions .

Distribution of Colors (Variegation)

B.
(Adapted from Lindley [1848 ])
Banded . Transverse stripes of one Ocellated. A broad spot of some color
color crossing another . has another spot of a different
Blotched . The color disposed i n co l or \"ith in it.
broad, irregular blotches . Painted . Colors disposed in
Bordered. One color is surrounded of unequal intensity .
by an edging of anothe r. Spotted. The color disposed in small
Clouded. Colors are unequally spots.
bl ended together . Striped . Longitudinal stripes of one
Discoidal . A single large spo t of color crossing anothe r.
color in the center of another . Tessellated . Color arranged in small
Dotted . The col or dispos e d in very squares , so as to have some resec-
small round spots . blance to a checkered pavement.
Edged. One color is surrounded by Variega ted . The color disposed in
a very narrm.J rim of another. various irregular, sinuous ,
Marbled. A surface traversed by Zoned . The same as ocellated , but
irregular veins of c olor ; as a concent r ic bands more numerous.
bl ock of marble often is .

XII. GENERAL
(Unclassified descriptors applicable to any character. from Porter et al .
[1973])
6 151

Compound . Composed of two or more Polyan thous. Having dif ferent states
anatomically or morphologically in several to many (more than
equivalent units, whether sub- three) different sets of flowers,
divided into them or an aggre- only one state present in each
gate of them . set.
Dimorphic . Having two different Polyheterophytous . Having dif ferent
sizes andlor shapes \<1ithio the states in several to many (more
same species. than three) different individuals
Evident . Clearly visible macro- or sets of plants , only one state
scopically . present in each set .
Heteranthous. Having different Polymorphic . Having s everal to many
states in two different sets (more than three) different shapes
of flowe r s, only one state and/or sizes within the same
present in each set . species.
Heterophytous. Having different Same . Same as in description of next
states in tHO different sets higher taxon.
of plants , only one state Sepaloid. Sepal-like in shape , tex-
present in each set . ture and/or color.
Homanthous. Having more than Simple . Not composed of more than
one state within each indi- one anatomically or morphologi-
vidual flot...e r . all flm.;rers cally equivalent unit.
the same. Triheteranthous . Having different
Homophytous. Having more than one states in three different sets
state within each individual of flowers, only one state present
plant , all plants the same . in each set.
Honomorphic. All of the same shape Triheterophytous. Having different
and size . states in three di ffe rent sets o f
Not Applicable . Not relevant to plants, only one state present in
the taxon at hand; passive null each set.
value . Trimorphic. Having three different
Obscure. Not clearly visible shapes and/or sizes t.,rithin the
macroscopically , usually owing same species .
to incomplete differentiation. Unknown. Relevant to taxon at hand
Petaloid. Petal-like in shape, but status not knotffi; information
texture and/or color . unavailable .

Section C. GYHNOSPERH GLOSSARY*

A. Vegetative Structures
Acicular Leaf. Needle-like , l ong Fascicle Sheath. Closely imbricated
and slender; e . g ., Pinus . bud scales at the base of the fas-
Awl-shaped Leaf. Subulate ; narrow , cicle of needles ; e . g ., Pinus .
flat , stiff , sharp-pointed, usu- Linear Leaf. Narrow , flattened ,
all y less than 1/2 in . long; e.g . , triangular, or quadrangular
Juniperus . ""' - ..... leaf usually 1/2-2 in . long;
Fascicle . Cluster of needles borne e.g ., Taxus, Picea.
on a minute determinate short Long Shoo~longated inte rnodes,
shoot in the axil of a primary rapid annual gro\.,rth.
leaf (bract) ; e . g .• Pinus .

*Contributed by James W. Hardin , North Carolina State University , Raleigh.

152 6

Multinodal Shoot . Spring shoot devel- Short Shoot. Very s hort or inconspi-
oping from the terminal winter bud cuous internodes and grOl~th very
and producing 2 or more Hhorls of slow if at a l l.
branches; the cones are pa r tly Determinate . No continued growth;
lateral in the middle of the shoot with l eaves drops as a
shoot ; e . g . • Pinus echinata . unit ; ve. g., Pinus fascicle .
Needle . Acicular ; slender, e l ongated Indeterminate. Continued growth
l eaf, usually over 2 in long; e . g ., by an apical meristem ; spur-
Pinus . branch; e . g ., Larix, Cedrus ,
Peg (ster i gmata) . Lateral stem pro- Ginkgo .
jection to which leaf is attached Uninodal Shoot. Spring shoot
and persistent after leaf dehis- developing fro m the terminal
cence ; i . e. , abscission layer be- winter bud and prodUCing only
t~"een peg and leaf . Leaf may be one internode with one who r l of
sessil e ; e . g .• Picea; or petio- branches at the end; the cones
late; e.g . , Tsuga . on the peg. are subterminal at the end of
Scale Leaf. Small, usual l y appressed the shoot ; e . g . , Pinus resi -
and imbricate ; e . g . , Jun iperus , nasa .
Thuja .

B. Reproductive Structures
Apophysis . Exposed outer surface Epimatium . Fleshy covering of the
of eithe r an ovuliferous scale seed and more or less fused ,,;ith
or megasporophyll as seen when the integument : arising from the
the cone is closed. chalazal end of the ovule like an
Aril. An outgrowth from the stem additional integument; e.g ., Pod 0-
forming a fl es hy covering of carpus.
the seed; e. g., Taxus , Torreya; Megasporophyll. Modified leaf bear-
or on l y rudimentary at base of ing ovules; e . g . , Zamia .
the f l eshy seed; e .g., Cephalo- Hicrosporophyll. l-Iodified leaf bear-
taxus . ing microsporangia or pollen sacs .
Bract. Hodified leaf sub tending Ovuliferous Scale . Highly modified
the ovuliferous scale; may be lateral branch i n the axil of a
distinct or fused to the scale . leaf (bract) , and bearing ovules.
Cone (strobilus). Aggregat i on of l-~y be flat or peltate, woody or
sporangia-bearing structur es fleshy; e . g. , Pinaceae .
at tip of th e stem (either Receptaculum . A fleshy structure
sporophylls or scales in the below the seed formed from the
Gymnosperms) . bases of bracts and the slwllen
Female Cone (megasporangiate receptacle or cone axis; e.g . ,
strobilus) . Bearing ovu l es Acmopyle, and some Podocarpu s
or seeds. spp .
Hale Cone (microsporangiate stro- Umbo . Projection, with or without
bilus) . Bearing pollen sacs spine or prickle, on th~ apophy-
(microsporangia). sis of the cone scale .
6 U3

Section D. LOWER VASCULAR PLANT GLOSSARY*

I. SPOROPHYTE

Stems

l. Branching
Axial. With branches arising from Monopodial . Having one main axis of
buds in leaf axil. grm.,rth .
Dichotomous. Hith branches forking Random . With branches arising from
into two more or less equal buds without relation to leaves.
parts . Sympodial . With branches more or
Epipetiolar. '.Jith branches aris- less equal without a main axis .
ing from buds on the petiole.

2. Phyllotaxis
Dis tichous. \-lith leaves in two Polystichous . With leaves in several
rm"s. rows.

3. External Features
Aerial Stem. An erect stem aris- J ointed . With stems that can be
ing from a horizontal rhizome. pulled apart easily at the nodes.
Articulate. Generally meaning as in Equisetum .
having a joint as in leaves, Node . Point on the stem where
leaflets or stems, as in hetero- leaves are attached; or the point
phyllous species of Selaginella; of branching of the stem.
or having a swollen area. often Rameal Sheath. Leaf sheath on the
discolored. at the point of stem joints. as in Equisetum.
branching of the stem. Tubercules. Silica deposits on the
stem ridges. as in Eguisetum .

4.
Internal Features
Canals. As in Eguisetum . Dictyostele . A dissected soleno-
Central Canal. The large cen- stele with each individual bun-
trally located air space dle a meristele.
in the stem . Eustele . A dissected siphonostele
Carinal Canal. A canal be- with phloem only to the outside
neath a stem ridge asso- of the xylem.
ciated with a vascular Plectostele. A protostele dis-
bundle . sected into anastomosing plate-
Vallecular Canal. A canal like units.
beneath a stem groove . Protostele . Stele having a solid
Stele . The central primary vascu- column of vascular tissue with
lar system of the stem and xylem centrally located.
associated tissues . De- Siphonostele . A stele having vas-
limited fro m the cortex by cular tissue in the form of a
endodermis and pericycle. hollow cylinder. with a central
Actinostele. A protostele pith .
having a xylem core ~ in the Solenostele . A siphonostele hav-
form of radiating ribs . as ing phloem both i nt ernal and
viewed in transverse sec- external to the xylem.
tion.

*Contributed by John T. Mickel. New York Botanical Garden .

 154 6

B. Leav es

1. Duration
Evergreen . Bearing g reen leaves Narcescent . The leaves of short
through the Hinter . duratio~~ dying at the end of
the grOlving season .

2. Megaphylls
(Leaves Hith a branching vein system \."hich are associated \.,ith nodal leaf gap)

a. General
Crozier . The coiled developing Frond. The leaf of a fern .
leaf of a fern .

b . Parts of a Leaf
Blade . The expanded por ticn of a Rachis . The axis of a compound fern
leaf . blade .
Costa . The midvein of a minor divi- Segment . The ultimate division or
sion of a fern leaf . unit of a dissected fern leaf.
Lam i na . The leaf t i ssue other than Stipe . The petiole of a fern
the veins or axes . leaf .
Pinna . A primary divis i on of a fern Stipe Bundles . The vascular bun-
leaf . dles of the fern petiole .
Pinnule . A secondary division of a
fern leaf .

, c . Venation
Areoles . The spaces fo rmed by a
Patterns
Free . No veins uniting to form a
vein netlvork. netHork .
False Veins . Small vein-like areas Included Veinlets . Veins ending
of thick-,valled cells in the inside areoles .
leaves of some IOHer vascular Net. Veins uniting to for m a net-
plants. Hark .

d . Leaf Division
Bipinnate . 1.\.,rice pinna te. Pinnate-pinnat ifid . Pinnate Hith
Pectinate . Pinna tifid Hith closely pinnatifid pinnae .
set segments ; comb- like . Pinllatifid . Pinna tely cut, more
Pinnate . Compound , \.Ji th the leaflets than half way to the mid vein .
arranged on both sides of a cornman Simple. Undivided .
axis .

e . Blade Architecture
Anadromous. Having the first l obe or Pa lmate . Radiately lobed or di-
s egment of a pinna arising basi- vided .
scopically in compound leaves. Pedate . Palmately cut or divided
Catadromous . Havi ng the first lobe Hith the Imler pair basiscopi-
or segment of a pinna arising cally exaggerated .
acroscopically in compound leaves . Pinnate. Compound , \.,rith the leaf-
Imparipinnate . Pinnate \.,rith a con- lets a r ranged on both sides of
form terminal leaf let . a common axis.
 155
6

f . Dimorphi sm (Figure 6-18 . p . 142)

(tUth leaves of two t.ypes in ferns, t he ferti l e being of different size ,
shape or dissection t han the vegetative)
Complete Dimorphi sm . Leaves com- Slight Dimo r phism . Fertile and
pl etely fertil e or vegetat i ve . vegetative leaves only s l igh tl y
Partial Dimorphism . Leaves \-lith different in appearance .
on ly a part mod i fied as fe rtile . Vegetative Frond . Frond lacking
sporangia .

3. Nicrophylls
(Leaves with only one vein and no leaf gap . )
Axillary leaves . Leaves borne in the Ligule . A small membranous outgrm.. th
axil s of branches . a s in hetero- or projection at the base of the
sporous species of Sel asinella. leaf. appearing above the sporan-
Bas t b und l es (periphe ral st rand s) . glum in fertile l eaves. as in
Bundles of thick- walled ce ll s S~lagin e ll a and Isoetes.
parallel t o the midrib , as in Ned ian leaf. Leaf on top of s tem,
Isoe tes. as in heterophyllous species of
Lacuna . Chambe r or internal air Se laginella.
s pace. Se t a . A hair like extension of the
Lateral leaf. Leaf on the side of leaf . as in homoph yllous specie s
the stem, as in heterophyllous of Sel aginella.
spec ies of Selaginella.

C. Reproductive St r uct ure s (Figure 6-18, p . 142)

Ann ulus . Thi ck-walled ring of cell s Inferior indusium. An indusium
on the sporangium. attached beneath the so rus \dth
Elater . One o f fo ur elongate appen- t he spor a ng i a appea r ing above
dages on the spores , as in it .
Eguise tum. Leptosporangiate. Having the en tire
Eusporangiate . Having the sporan- sporangium develop from a peri-
gium develop from a great clina l divis ion of a super ficial
amount of leaf tissue as cell or smal l gro up of ce lls.
opposed to only one or a few Lip Ce lls . The line of cells between
cells . which the sporangium deh i s ces.
Exospor e or Exine . Outer spore Hassula. A c lump of mic r ospo re s , as
wall layer . in AzalIa .
Fovea . Pit or depression con- Negasporangium. The sporangi um i n
taining the sporangium in the which megaspores are pr oduced .
leaf base of Isoetes . Megaspo r e . A spore that gives rise
Heterosporou s . Having two kinds of to a female gametophyte .
spores , usually differing in MegasporophylL A l eaf bearing one
s ize . or more megaspo r angia .
Homosporous . Having spores of only Hicrosporangium . The sporangium in
one kind. which microspores are produced .
Indusium . A f l ap of tissue c over- Nicrospore . A spor e t hat gives rise
ing a sorus . to a male game top hyte .
False indusium . A folded leaf Nic r ospo rophyll. A l eaf bearing one
margin protecting t he so rus. or mor e microsporangia .
True indusium . An epidermal Paraphyses . Hair s or hai r-l ike
outgrowth protecting the st ruc tures in the so rus .
sorus .
 156 6

Peri spore or Perine . An outer cover- Trilete . Basically tetrahedral,

ing of some fern spores, with dif- but often appearing round or
ferent configuration than that of triangular, with three scar
the exospore. lines forming a Y.
Receptacle . Point on a leaf where Sporocarp . A hard, nut-like struc -
sporangia are attached . ture con~aining the sporangia in
Sorus . A cluster of sporangia . hetero ~po-r ous ferns.
Sporangiophore. The umbrella-shaped Sporophyll. A l eaf bearing sporangia .
sporangium- bearing unit of the Stomium. Lip cell region of a fern
strobilus, as in Equ isetum. sporan gium.
Sporangium. A spore case. Strobilus . Stem with short inter-
Spore Shapes . nodes and spore-bearing appen-
Honolete. Bean-shaped , with a dages ; a cone.
single scar line. Velum . The membranous flap covering
the sporangium, as in Isoetes.

II . GAMETOPHYTE

Antheridium . The male sex organ pro- Prothallus . Gametophyte of lower

ducing the sperm. vascular plants.
Archegonium. The female sex organ Rhizoid. A hair- like absorptive or-
producing the egg. gan on gametophytes and rarely on
sporophytes .

III. ASEXUAL REPRODUCTION

Apogamy. Producing sporophytes from Bulblet. A small. bud-like vegetative

a gametophyte without fertiliza- propagule produced on the leaves of
tion. some ferns.
Apospory. Producing gametophytes Gemma . A vegetative reproductive bud
d i rectly from a sporophyte {dth- borne on the stem , as in Lycopodium;
out produc ing spores . a multicellular reproductive propagule
on gametophytes . as in ferns.

PHYTOGRAPHIC LITERATURE

Bocquet . G. 1959. The campylotropous ovule . Phytomorphology 9: 222-227.

Bjornstad , I . N. 1970. Comparative embryology of Asparagoideae-Polygonateae.
Liliaceae. Nytt Nagasin for Botanikk 17 (3- 4): 169-207 .
Correll, D. S., and H. C. Johnston. 1971. Hanua1 of the Vascular Plants of
Texa s . Texas Research Foundation. Renner .
Cronquist . A. 1968. The Evolution and Classification of Flowering Plants.
Houghton-Hifflin Company. Boston .
Davis, P. H., and V. H. Heywood. 1963. Principl es of Angiosperm Taxonomy .
D. Van Nos trand, Inc . Princeton.
Duke, J. A. 1969. On tropical tree seedlings 1. Seeds. seedlings, systems.
and systematics . Annals Hissouri Botanical Garden 56(2): 125-161.
Faegri, K. , and van der L. Pijl . 1966 . The Principles of Pollination
Ecology . Pergamon Press . Ne{.,r York.
Featherly , H. I . 1954. Taxonomic Terminology of Higher Plants . Iowa State
College Press . Ames .
Fernald, H. L. 1950. Gray's Hanua1 of Botany. 8th ed . American Book Com-
pany. New York .
Gray, P. 1967 . The Die tionary of Biological Sciences. Reinhold Publishing
Corporation. New York.
6 157

Halle, F. 1971. Architecture and growth of tropical trees exempl i fied by

the Euphorbiaceae . Biotropica 3(1) : 56 - 62 .
Hickey, L. J . 1973 . Classification of the architecture of dicotyledonous
leaves. American Journal of Botany 60(1): 17-3 3.
Jackson, B. D. 1928. A Glossary of Botanic Terms, 4th ed. J . B. Lippincott
Company . Philadelphia .
Johnson, A. M. 1931 . Taxonomy of the Flowering Plants . The Century Company .
New York .
Lawrence, G. H. N. 1951. Taxonomy of Vascular Plants. The HacMillan Com-
pany . New York .
Leppik, E . E. 1957 . A neH system for classification of flmoJer types . Taxon
6, 64-67.
1961. Phyllotaxis. anthotaxis , and semataxis. Acta Bi otheoretica
14: 1-28 .
Lindley, J . 1848 . An Introduction to Botany , 4th ed. London .
}lartin, A. C. 1946. The comparative internal morphology of seeds . American
Hidland Naturalist 36 : 513-660.
Porter , D. N., R . H. Kiger , and J. E. Honahan. 1973 . A guide for contribu-
tors to Flora North America, Part I I : An outline and glos sary of terms
for morphological and habitat description (Provisional Edition) . FNA
Report 66. Department of Botany, Smithsonian Institution. \Jashington.
Rickett, H. H . 1944 . The classification of inflorescences . Botanical
Review 10 : 187-231.
Stearn, \~ . T . 1966. Botanical Latin. Thomas Nelson and Sons Ltd . London .
Swartz, D. 1971. Collegiate Dictionary of Botany. The Ronald Press Company .
New York .
Systematics Association Committee for Descrip tive Terminology. 1960. I . Pre-
liminary list of works relevant to Descriptive Biological Terminology .
Taxon IX(8) : 245- 257 .
1962 . II. Terminology of simple symmetrical plane shapes . Taxon
XI(5) : 145-156 , 245 .
Uphof, J . C. Th . 1938 . C1eiHtogamic flowers . Botanical Review 4(1) : 21-49.

PHYTOGRAPHIC EXERCISE SUGGESTIONS

I . General
Phytographic studies should be a combination of laboratory and field
\.;rork . The nature of the subject makes it ideal for individual and group
scavenger hunts for different structures and features . The objectives of
these studies should be to provide (1) a vocabulary for intelligent com-
munication about vascular plants, (2) an understanding of the use of relative
terms and to help the student become (3) a more critical observer , and (4)
more precise in his descriptions of vascular plants.

1. Identify the parts of the plants, stems, roots, buds, leaves, inflores -
cences , flm.;rers, fruits, and seeds in the material on display (or in
the field .
2. Identify the types or - -,habits of plants, stems, roots , buds, leaves,
inflorescences , flm.;rers, fruits, seeds in the material on display
(or in the field) .
3. Determine the position and arrangements of stems , roots. buds , leaves,
inflorescences. flowers, stamens, carpels, and cotyledons of the
material on display (or in the field) .
 158 6

4. Determine the shapes , margins, bases , and apices of the structures on

display (or in the fi eld).
5. Determine the surface, texture , vestiture fe atures for the material on
display (or in the field) .
6. Distinguish the perianthal, androecial , and gynoecial types or structural
types of the material on display . .....~ '-
7. Hake a comparative study of seeds, embryos, end osperm, and seedlings on
display . Identify parts, types , and arrangements of parts.
8. Hake a study of numbers of parts, fusion of parts, number of whorls ,
symmet ry of parts, and inserti on of parts for the material on display
(or in the field) .
9. Determine the sex , pollinator, and insect visitor--flm.,rer relationshi ps
for several species in the field.
10 . Determine the periodicity states , maturation times, and growth and develop-
ment features for several species in the field .

II. Special
Select one species and determine or describe the characte r state or
characteristic for each of the foll m.,ring:

1. Plant 11. Stipule Type

1. Type 12 . Blade Shape
2 . Duration 13 . Blade Apex
II . Roots 14 . Bla de Hargin
1. Type 15 . Blade Base
2. Structural Type 16. Laminar Secretory
II I. Stems Structure
1. Bark Type VI. I n f lores cences
2. 1·lood Ring Type 1. Type
3. Pith Type 2 . Position
4. Texture VII . FloI.,rers
5. Type or Structural Typ e 1. Sex
6 . Position 2 . Type
7 . Configuration 3 . Insertion of Parts
8. Sur f ace 4 . # Hembers in I'Thorl
9 . Vestiture 5 . II of tfuorls
I V. Buds 6 . Fusion of Parts
1. Type 7 . Perianth Type or
2. Structural Type Structural Type
3 . Scale Arrangement 8. Aestivation
4 . Shape 9 . Nectary Position
5 . Surface VIII. Androecium
6. Vestiture 1. Androecial Type
V. Leaves 2 . Stamen Structural Type
1. Type or Structural Type 3. Anther Attachment
2 . Duration 4. Anther Shape
3 . Position 5 . Anther Apex
4 . Arrangemen t 6 . Anther Base
5 . Textur e 7 . Anther Type
6. Surface 8. Stamen Arrangement
7 . Vestiture 9. Thecal Arrangement
8. Venation
9. Ptyxis
10. Petiole Type
 ;:::':~:,,,,,
;.:,::;., • •<i::;:·

6 159

I X. Gynoecium 2 . Pollination Type

1. Gynoecial Typ e 3. Fertili zation Type
2 . Carpel ~rrangement 4 . Pol lination PathHay
3 . Carpel Type 5 . Fertilization Pathway
4 . Stigma Type XI II . Periodicity & Haturation
5 . Style Type 1 . Leaf Periodicity
6 . Ovary Position 2. Flm.,rer Periodicity
7. Ovary Shape 3 . Fruit Periodicity
8 . Ovule Type 4 . Leaf Maturation
9 . Ovule Position 5. Stamen Maturation
x. Fruits 6. Carpel Haturation
1. Structural Type XI V. GroHth and Deve l opment
2. Position 1. Specialized Growth
3 . Pl acentation Regions
XI. Seeds and Seedl i ngs 2 . Grm.,rth Periodicity
1 . Seed Surface Feature (Stem and Root)
2 . Cotyledon Pos ition 3 . F1m.,rer Development
J. Seed Type 4 . Stamen Development
4. Embryo Type 5 . Flora l Symmetry
5 . Aril Structural Type 6 . Anthotaxis
6 . Eophyll # & Shape 7. Carp otaxis
7. Seedling Type 8. Serna tax i s
XII. Sex and Pollination 9 . Phyllotaxis
1 . Plant Sex 10 . Stem Branching Pattern

PHYTOGRAPHI C INDEX
Amorphic 102 Anis osty lous 126
Abaxial 123, 147 Adnate 127, 128 IInnotinal 14 5
Adnat e Stipu1es 97 AmphicarpouH 116 , 117
Abllent 97, 128 Ann ual 146
Adpre"scd 122 Amph iflo rou s 117
Acarpous 1 26 ,\nn ual Ring 91
Adventltious Bf! Amphi s arca III
Accessory Bud 93 Annular 133
Aerating 88 Amp hitrop ous 109
Accessory Organs 99 Annulus 155
Aeria l 116 Amplexicaul 134
Accrescent Ift7 Anom alicidal
Aer ial Root 88 AmpI iate 126
Accurnhent 11 7 IInadromous 15ft Capsule 110
Acerose 133 Aerial Stem 153 Anterior
Aerocaulous 116 Anandrous 126
Achene 109 Gr ooves 102
Aerophy11ous 116 ,\natropous 109
Achenecetum III Ante ri or Lines 102
AesUval 1 45 Ancipital 12 5
Achlamydeous 103, 126 Ant e r io r Lobes 102
Aestivation 85 And rHgamocephHlou.; Iftft
Acicular 130 Ante rior Ridges 102
Agamandrocep halous Ift3 Andragamous 144
Acicu!.'!r Leaf 151 Anther 104
Agamandrous Ift3 Androdiocciou" 144
Acicu1ate 138 Anthe ridium 156
Agamogynecephalous 143 Androeclum 99
Acrocaulous 117 , 124 Anthesis 1 46
Agamogynou;; lft3 Androgynecandrous 143
Acrocida1 Capsule 110 AnChoearpous 127
Agamohermaphrodi - Androg ynecepha lous 1 ft 3
Ac ro dromous 138 ,\nthotaxis 148
cephalous 1ft4 Androgynophore 99
Acrolarninar 150 Anth ropophi1y 1 45
Agamohcrmaphroditic Iftft Androgynous 1ft)
Acropetal 147 Ant l.pe talous 118
Agamous 143. 144 Anclrohermap hro-
Acropetiolar 150 ,\ nti sepal ous 118
Agglomerate 119, 122 dicepha1ou,; lft4
Acroramaus 117 Anti t ropous 118
Aggregat e 119, 122 Androhermaphrodlt iC lftft
Acroscopic 122 Antro r se 122
Aianthaus 145 Andromono ecious 11,1,
Acti nodromous 138 Apetalous 103. 126
Alate 11 5 , 13 8 Anemophi.ly 1 45
Actinomorpilic 102 , ,\ pe x 86
Alhumi1mus 114 .Angus tate 126
103, 148 Aphyll opodic 117
Allagos t emonous 118 Anisocarpous 126
Actinostele 153 Aphyllous 126
Allautogamy Ift 5 An isocotylou5 126
Aculeate 137, 140 ,\pica1 116, 118.
Allogm'lY Ift 5 Anisolateral 126
Acumi nate 134 119, 123 , 147
Alternate 119. 124 Anisomcrous 127
Acute 13ft ,\ pi culate 13ft
Alveol.'!tc 138 ,\r,i,;opetalollS 126
Adaxial 12 3 . 147 Apocarpous 107, 127
Ament 98 Anisophyllous 126
Adherent 127
160 6

Apogamy 156 llasifixcd 107 Bullate 1)8 Chaff <)t,

ApopetaloU5 10 3, 127 B...,,;) 1amin<l.r 150 Hur J 11 Cha.L lza lOR , 114
Apoph::,,;h; 15 2 lla si pet<11 11,7 But tress 88 ehaJ a zogamy 1.45
Aposcpalo lls ]03 , 127 IlHs~petiolar 124 , Chamhered Pith 91
Apos pory 156 150 Cha n nel l ed 97
Apostemon ous !04, 12i llasira tTlOus 117 c Ch;.ntaceous 1"3
Ap pendiculill" Stamen 104 Basis copic 122 Chasmantheric
Applanate 123 Ilast Ilundles 1 55 Caducous 146 Pollinat ion 145
APl'oC'ition 1(.9 Beak 109 Ca1carate 103 Che i ropteroph i ly 145
AppresseJ 122 Be ard 102 CalceoLlte 10) Ch1amydeolls 103 , 126
Arac hnoid 140 He arded 11. 0 Callosity 102 Chor i p etalous 10) , 127
Arbo r escen t 92 Ilent Emll r yo 114 Callus 102 Chorisepalous 103, 127
ArchegoniuT:l 156 ller r y 111 Calyh J. um 109 Cil i at e 1)7, 1/,0
ArcuHte 123 . 124 , 13 3 lIihacca III Calycle 97 C1liolate 1 40
Areolate 138 Ilicarpellate 125 Calyx 99 Cincinnus 98
Areoles 154 Hicrena t e 137 Campanulate 103 Circinate 1 24
Arhizous 126 Biden tat e 125 Campy1odromous 139 Circular 128
Ar11 114. 15 2 Bid uou s .1/.6 Campylotropous 109 Circumalate 115
Arilla t e 11 5 Iliennial 1 46 Canal 1 53 Circumferentia.l 1 2)
Aristate IJt, Bife rous 1 45 Canalicula. te 97, Circumscissile
Arrallljement 85 Bi fid 128 138 , 150 Capsule 110
Arran gc!Ilcnt Systems 87 Rif1orous 125 , 11.6 Cance l late 138 Ci rrhou s 134
Ar ticulate 15 3 Bifoliatc 125 Can esce nt 1 40 Clad ode n
Ascending 92 , 122 IlifolioIate 94 Cant harop hi ly 145 Cladodromous 1)9
Asepal ous 103 , 1 211 Bifu rcate 128 Capilla te 133 Cladoptosic l46
Asperous 139 fligem inate 9/j Capitate 108 , 133 Clamhering 92
Assurgen t 122 Ilijup,ate 94 Capita t e Emhryo 114 Clasping 134
Astemo nollS 12 6 lIi1ahiate 10 3 , 125 Capitulum 98 Clathrate 138
Astylocarpel lous 108 llilatc ra.l .\ 48 Capsule 1 1r) Clavate 108 , 13)
Asty lo ca r pe podic 108 lIi10cular 12 5 Capsule, Inde hi s- Clm,' 102
Astylo u& 108 Himes t r ia 1 1 46 cent 109 Cleft 128, 134, 137
Asyr.lmetric 148 g inate 111) , 12.1 Carina 1 02 C1eistantheric
Atropous 109 Hi palmate 128 Carinal Canal 153 Pollination 145
At tenua te 134 Ri palmatc1 v Ca rinate 10 ) C":limbing 92
Au riculate 134 Compound 94 Ca rnose 143 Clinan thiu m 99
Au r iculiform 130 Bipinnat e 154 Carpe l 99, 107 Clouded 150
Autogamy 145 Bil'i nnate 1y Cilrpellate 1 4 ) , 144 Clustered 11')
Autumnal 145 Compound 9t. Car pophore 109 Coalesced 127
AI! I- shap ed Leaf L51 Bi.seriat e 125 Car pop o d i um 107, 108 Coch le ate 124 , 1)3
Axin l IS ) Bi serrate 137 110 Coenoc ar pi um II I
A.-dIe 118 Bise xll<ll 125 , 1 /,) C"rpot" xi s 148 Coetaneous 146
Axillary 116, 11 7 IHtt' rna te 9/, Cartilag ino us 143 Coherent 127
Axi l la r y Bud q ] IHade 93 , 1. 54 Car uneulate 115 Coleo pt He 114
A:dl1ary Lea ve s 155 IHa stocarp ous 1M, Caryopsis 109 Coleorhiza 11"
Blotched 150 Cas i ng 103 Coll ateral Bud 93
Hordered 1 50 Catadromous 154 Collet 11 5
!\ J\os try x 9 8 Ca t nph yll 115 Column 128
ll o t uli fo rm 133 Catkin 98 Co1unUlar 92
Haccacetum III Bractea1 Ne<:tarv 150 Caudate 1)/. Comose 115 , 140
llacc;He 14) !l rac teoles 9" , 97 Caudex 92 Complanate 133
Balausta 10C) Bra ctlet 97 Cauline 11 7 Complete 127
Banded 150 Br<1cts 94 , 97 , 152 Cauline Ethereal Oil Complete Dimorphism 155
Banne r 10) t;r an chi ng Patterns 87 Gl and 150 Comnle te Leaf 94
Barbilte 140 Bristles 102 Caulocarp ic 146 Co mpoun d 94 , 151
Barbed 140 llr is tl y ])9 , 14(1 Caulous 117 Co mpound Corymb 99
B<l r be l Late 140 Ilrodd Emhr yo 11" Central 12 1 Compound Cvme 99
KHrk 91 Broc hidod r omous 139 Cen t ral Canal IS ) Compound Umhel 99
Basal ll6 , 118 , 12) , Bud 88 , 91 Centri f uga l 147 Compressed 133
147 Bud P ri.mordium 9) Centripet al 1"7 Conduplicate 108, 123 ,
Bas;ll Stipu1es 97 1Iu lh 92 Centroramous 12 1, 124
Base 86 , 102 , I n) !\,,]hel 92 Cer.1 ceo us 143 Co n e 1 5 2
Basl cau 10us 117 Bulbil 92 Ce rnuous 123 Cone, Female 152
llasicida1 Capsule 11 0 Il tllhlet 92 , 156 Cespi t ose 92 Cone, Nale 152
6 161

Configuration 86 Cyme 98 Dipterous 125 Ephemeral 146

Conglomerate 119, 122 Cymule 98 Disc 99 Epicalyx 97
Con ical 13) Cynarrhodion III Discoid lOB , 133 Epico rmic
Conjugate 128 Cypsela 109 Discoidal 150 Shoots 147
Connate 127, 134 Discontinuous 116, 119 Epicotyl 114
Connate-perfoliare 134 Dissected 128 Epigeous 116
Connective 107 D Dissepiment 109 Epigyny 118
Connivent 122 Distal 123 Epihyperigyny 118
Conocarpium III Deciduous 146 Distant 119 Epihypogyny 118
Contiguous 127 De c1inate 122 Distichous 119, 153 Epima ti um 152
Continuous 116 , 119 Decompound 94 Distinct 127 Epiperigyny 118
Continuous Pith 91 Decumbent 92 Distribution of Epipeta10us 119
Contorted 122 , 124 Decurrent 108, 134 co loration 87 Epipetio1a r 153
Contortup l icate 124 Decussate 11 9 Diurnal 146 Epipetric 116
Contractile Root 88 Deflexed 122 Divaricate 122 Epiphy 110us 117
Convolute 124 Deliquescent 146 Divergent 122 Epiphytic 116
Corda te 134 Deltate 130 Divided 12B , 137 Epirhizous 116
Cordiform 130 Deltoid 130 Division 86 Episepa10us 119
Coriaceous 143 Dentate 137 Do1abriform 133 Epitropous 117
Corm 92 Dcnticidal Dorsal 123 Equilateral 148
Carmel 92 Capsule 110 Dorsal Side 102 Equinoctial 146
Corneous 143 Denticulate 137 Dorsifixed 107 Equitant 119
Corniculate 103 Depauperate 126 Dorsilaminar 124 Eramous 92
Corolla 99 Depressed 122 Dorsiventra1 148 Erec t 92
Corona 102 Descending 122 Dotted 150 Erose 137
Coronate 103, 115 Determinate 147 Douhly-crenate 137 Essential Organs 99
Cotani f orm 133 Determina te Doubly-serrate 137 Etaerio 111
Corrugate 124, 138 Inflorescences 98 Downy 140 Ethereal Oil Gland
Corymb 9 8 Developmental Shoot Drupe 111 149 , 150
Types 87 Drupecetum III Eucamptodromous 139
Costa 1 54
Costal 150 Dewy 140 Drupelet 111 Eusporangiate 155
Costate 138 Dextrorse 122 Duration B7 Euste1e 153
Cotyledon 94 , 114 Diadelphous 104, 125 ))warf 126 Evanescent 146
CotylespermOlls 114 128 DHarf Embryo 115 Evergreen 146, 154
Cotyliform 133 Dlandrous 125 Dwarf Shoots 147 Evergrowing 147
Craspedodromalls 139 Diaphonous 143 Dyads 107 , 125 Ev ident 151
Crateriform 133 Diaphragmed Pith 91 Excentric 133
Cremocarp 110 Dichasium 98 , 125 Excurren t 149
Crenate 137 Dich1amydeous 103 , 125 E Exfoljating Bark 91
Crenulate 137 127 Ex ine ISS
Crested 108, 115 Dic hogamous 1t,6 Eccentric 108 Exocarp 109
Crisp ate 137 Dichotomous 92, 128, Echinate 139 Exospore ISS
Crisrate 108 139 , It,9, 153 Echma 109 F.xplanate 122
Cro\~ded 119, 122 Didesium 111 Ectocarp 109 Exstipellat e 97
Crozier 154 Dic1inous 125 , 144 Edged 150 Exstipulate 97
eruciate 103, 133 Dicotyledonous 125 E1aminate 94 Extrafloral
Cruciform 133 Dictyostele 153 Elater ISS Nec tary 150
Crustaceous 143 Dicyclic 127 Ellipsoid 128 Extrorse
Cryptantherous 118 Didymous 119 Elliptic 128 Dehiscence 107
Cryptocoty1ar Didynamous 119 Emarginate 134
Seedling 116 Diffuse 108 Embryo 114 F
Cubical 129 Diffuse Porous Hood 91 Embryo Sac lOB
Cucu11a te 103 Digestive Gland 149 Embryonic Position 85 Falcate DO , 133
Culm 92 Digitately Net ted 139 Emergent 1 16 False Veins 154
Cuneate 130 , 134 Dilated 126 Emersifolious 116 Farinaceous 139
Cuneiform 130 Dimerous 127 Endocarp 109 Fasciateu 127
Cupu1e 97 Dimidiate 128 , 130 Endosperm 114 Fascicle 151
Curvative 124 Dimoxphic 151 Endospermous 114 Fascicle Sheath 151
Cuspidate 134 Dimorphism 155 Entire 137 Fascicled Root 88
Cyathium 99 Dioecious 125, 144 Entomophily 145 Fasciculate 119
Cydy 85 Diplec01oba1 124 Eo phy11 115 Fas tigiate 92
Cylindric 129, 133 Diplostemonous lIB Epet i olate 94 Faucal Area 102
Cymbiform 133 Dip10tegium 110 Epetiolulate 97 Felted 140
 162 6

ve .Hde 143 , 14" Gen,:ral 87 liermaphrodandrO\lS t4/. IIv"oco L" I eSilc rl'IOUS
Fenio'~;trate 138 Ge neral lIermapli rod igyne·· IIYl'ogeolis II f1
FerLilization De V,!.l opmCll t B7 cc pha10us 144 Hy po),,}," i 11m 101
Pathvav 87 r,eneral St nlC tll ral Ilermaphrodigynous 144 Ilvpogvnv IIR
Fertilization Position 85 Hermaphroditic 144 Hypo p hy11olls 126
Tvpe 86 GenicuLate 92 , 108, Hesperidium 111 HYl'otropous 118
Fihonacci Phyllo- 121 lleterandrous 126 I1vsteranthous 146
taxis 148 Ceocaroous 117 Ilete ranthous 151 ~
Fibrous 115, 1 43 (;eoflorol1s 11li, 1 ,],7 Heteroblasty 126
Fibrous Root 88 Gib bou" 103 lIeterocarpou~ J26
Filament 104 Gif!.antic 126 Heteroceph<llous 141,
Filamentose 137 (;labrate 139 , I t,1) lIete r ocladous 126 Imbricate 119 , 124
Filantherous Glabrescent 139 Ileteroeymous 144 Impar ipin nat e 1St.
Stamen 104 Glabrous 139 Heteromerous 127 1mpa r ir inna te 1y
Filifcrous 137 Glandular 139, 140 Il ete ropetalou~ 126 Compound 9t.
Filiform 107 , 130 Glans HI lleterophyllol.ls 126 i mperfect lid
Fimbriate 108 , 137 Gl auc esce nt 139 lleteroph yl 1ous Shoots Implicate 125
Flmbriolate 137 (~laucolls 119 11,7 Incano"s 11,1)
Fissur<:!J nark 91 Globose ]03 IIctcrophytous 151 rncis e d 128 , 1 3 7
Fis tulose 133 Glocloidi<lte If,O Heterosepdlous 126 l nclina te 125
Flabellatc 108, 149 Clomerule 98 Heterosexual 144 Tnclined 122
Fbbe lliform 133 Giume 97 !leterospicollS 11,4 Included Veinlets 15t.
Flaccid 143 (;1utinous 139 Heterosporous 155 IncompLete 97, 127
!'lcshy Root 91 {:rain 109 Heterostichou~ 12 6 Incrassate lid
Flexuous 123 , 138 {:ranular 139 !leterosty1olls 108 Incumbent J17
Floating 116 Cre,1sy 139 Heterotropous 118 I ncurv ed 1.2)
Floccose 140 Gro!!th Pe riodicity 87 lIiberllal 146 Indeh'i.s cent Capsule
Floral Nectary 150 Crmlth Region 87 ](jemal 146 Indeterminate 11,7
Flotophyliou.'> ll6 Cynagamocep hal ous It,4 lIil urn 114 I nd ete rminat e
F101,'c[ 88, 93 , 97 Gynagamo\ls 1M, Hip LJ 1 Inflorc~cence 98
Flot..'cr Primordium 93 Gyna n drium lOt" 128 Hippocr e pi for m 13 3 In dument 86
Flower Sex 86 r:ynecandrous 144 Hirsute 140 I ndtlplicilt e 123 , 12t.
Flushing Shoots 147 Gynehermapll rocl i -· Hi.rsutul1ous 140 Indu r ate 11, 1
Fly Trap 97 cepllalous 144 Hirtellous 140 I ndusi.um 155
Folded Embryo 115 Gynehermaphrodi- Hisp id l!.0 fndus tum, False 155
Follicetum 111 tie 144 !lispirlu10us It,o Indu s ium, Inferio r 155
Follicle 110 Cy nobasi c 108 , 119 HolJOh' Pi th 91 f ndu sium , Trtl e 155
Fovea 155 Cynodioe eious 11,4 110manurous 126 Ineq\li1atera l 126 , It.8
Foveolate 138 Gynoeciulfl 99 Homanthous 151 lnermous 1M)
Free 127, 123 Cynomonoecio us V I4 llomocar pous 126 InferLer 11 7
Fr ee-adnate 128 Gynophore 99 liomoce phalolls It.t. I nflated 97
Free-cent ral 118 Gynostemium 104, 128 Ilomoch.lamyrleous 103 , Inf1exe d 122
Free Veins 154 1 27 i nf lorescence ,~f'X 8f,
Fringe 102 1I0mocymous 11,4 In f lorescence
Frond 154 ilomogamous l 46 Types 9R , qg
Fruit 88 Homop hy tous 151 Tnfru folia r 117
Fruticos<o 92 flalf-inferior 117 Homosexual 144 InfrupeL i o1.ar Blid 9)
Fugacious 146 Half-terete lJ3 I!omospicou~ 11.1, TnfUildihll la r 10)
Functional T~!p c s 87 lIilpaxanthic 11,6 Homosporous 155 I n tegume nts L08
Funi cular 108, US I!a p lomorpllic 102 Ilomostylous 108 Lnte r calarv It.7
Funiculus 108 , 109 ]i,1state 111, lIood 102 lnterfoliar 117
Furcate 128 Hastiform 133 Ho r izontal lv 122 In termit t ent 147
Fus ifo rm 133 lIaustoriilJ <)1 110m 102 lnternudl' 9J
Fusion 86 lIead 98 Humistrate 92 Interretiolar StipuJes
!lelicoid Cyme <)8 Hyaline lI.) 07
Ilemianatropous 109 Ilyda thodc It.9 Inlerru[lted 11q
G Ilemirrnpous 10'1 Ilvdrophilv It.5 Int er rupt edly Pinn aL",ly
r:erh 88 lI"menopterophily 145 f:ompollnd 94
Galeate 103 Herr.ac eo ll~ 143 Iiyuanepigvny 118 Int e rstitial 147
Geitonogamy 1 45 ile rmdphrodagamo - I!ypa n thium 99, 128 fntrorse 107
Gelatinous 143 <.:ephalous 141, lIypanthodiurl 98 Investing Emb l'yo 115
Geminate 94 , 119 llermiJphrodagnmous 11,t, l!yphodromous 139
lnvo1uc el q8
Gemma 156 lIermaphrodandro- ! 1~'l'o c ot yl 114 Il1vol\1cr ~ 93
cephalous 144
6 163

Leptophy11ous 126 ~'ericarp 109 Nectar Sec r e tin ~ Glands

Involute 108, 123,
125 . 137 Leptosporan giate 15 5 Herosity 86 149
irregular 148 Liana 88 Hesoca rp 109 nectary 149
I!locotyloU5 126 Ligneous 143 Netaphyll 115 Needle 152
lsodynamo us 12 6 Ligulate 103 , 137 Mic r o Embryo 115 Net Veins 154
Isomerous 127 Ligule 93 , 102, 155 Nic r omellitophily 145 Netted 139
lsopetalous 126 Limb 102 , 103 Micropyle 108, 114 Neuter 143 , 144
lsophyllous 126 Linear 129 Hicrosporang I n te Ni tid 11.0
lsosepalous 126 l.inear Embryo 115 Stroh·Uu s 152 Nocturnal 146
lsostichous 126 Linear Leaf 151 11icrosporanglum 155 Nod a l Nectary 1 50
L'lnea t e 108 Hicrospore 155 Node, 9 1, 153
Lingulate 1)) Hicrosporophyl1 152, Nod'lf o rm 133
J Lip 102 155 Nodulose 133
Lip Cells 155 ~!idrib 93 Non-porous \o.'ood 91
Jaculator 109 Lobate 1)4 md vein 93 Not applicable 151
Jointed 153 Lobe 102 lUnor 126 ;lotorhiza 1 117
Jugate 94 Lobed 1.08, 128 , 137 ~1inute 126 Nucellus L08
Locarion 85 ~tixe d Hud 93 Numher 85
Locule 107, 108 }!onad 107, 125 Nut 109
l.oculicidal Capsule }!onade1phous 104 , Nutlet 110
110 125 , 128
Keel 102 I.ocusta 98 Honilifo r m 91
Knee 88 Lodicule 102 Honocarpel1ate 125 o
Loment 110 Honocarpic 146
L Long Bud Shoots 147 }lonocephalous 125 Obcon i c 133
Longitudinal ~lonocha!::lium 99 , 125 Ohcordate 134
Labellum 102 Dehiscence 107 ~'onochtamydeous 125 Obcordiform 133
Lacerate 128, 137 Longit udinal tlonoc1inous 14/. Ohdel t ate 130
Laciniate 128 , 137 PORtu r e 85 11onocotyledonous 125 Obdeltoid 130
Lacuna 155 Lon~ Shoots 148 , Honocyclic 127 Ohdlp lostemonous 119
l.;l.cvigate 140 151 Honoecio us 125, 144 Oblan ceo ]ate 129
Lamina 154 Loose 119 Nonolete 156 Ob lan ceoloid 129
Laminar ethereal Oil Lo r ate 123 Nonomerous 127 Oblate 128
Gland ISO Lunate 133 Nonomorphlc 151. Ohli'lue 122 , 137
Laminar Nectary 150 Lyrate 133 ~!on o)lhi1y 145 Oh1oi.d 128
Laminar Stamen 101, )\ollophyllo us 125 Ob l ong 129
~lollopodial 11>9, 153 Obovate 129
Laminate 118
Lamrnas Shoots 147 Nuc1].1gino1..:s 140 Obovo id 129
Lanate 140 Hucronate 13/, Ohpyram idal 130
Lanceolatc 129 Hacropodia1 114 Hucronul:lce 13/, Obscure 151
Lanccoloid 129 Najor 126 ~!ultlcellu1.nr 125 Obtdangu 1a r ])0
Lanuginose 140 Halacophi1y 145 Nul t icipit a 1 125 Obtru11.J.te 130
Hale 143 , 144 ~!ulticostal 125 OhtruIloi d 130
La t eral 107, 116 ,
!larhled 150 r·!u1tilocu l ar 125 Obtuse 134
119
Lateral Bud 93 Narcescent 146, 154 ..!u1tinodal Shoot 1 52 Ocellated 150
Lateral Embryo 115 Hargin 86 ~' ultjperennin1 11,6 f)create 137
Late r al Leaf 155 ~I.lrginal 118, 123, Nu l tiseriate 125 Oc r eo l ate 137
Lateral Stipu lcs 97 147 Nultist r iate 125 01 igomerous 1 27
Laterospcrmous 124 Hassu1a 155 Hudcate 140 Oligotaxy 127
Latiflorous 126 Haturation 87 Nuriculate 11,0 Opaque 140
tatTarse nehiscence Hatutinal 146 Huticous 134 Operculate Capsul e 110
107 ~Iedial 123 Hy iophlly 145 Opposite 119
~!edian Leaf 155 ~ly rmecophily 145 Orientation 85
Leaf 88 , 93
Leaflet 93 ~Iedian Stipu1es 97 Ornithophily 145
Leaf-opposed 117 Hegasporangiate Ort hotropous 109
Leaf Primordium 93 Strob ilu s 152 Osseo us 143
Megasporangium 155 Naked Bud 93 Ovarv 107, 108
Leaf Scar 91
Hegaspore 155 NanophyIlous 126 Ovate 129
Legume 110
Hegasporophyll 152, Na piform 133 Ovoid 129
Lemma 97
:i. 55 Navicular 133 Ovule 108
Lenetee! 91
Lenticular 128. 133 Nel1ttophily 11.5 Neck 103 Ov uli fc rous Scale 152
~\embranoufl 143 Nccroco1eop teroph ll.y
Lcntininous 140
Lepidote 140 Neniscoidal 133 145
164 6

P .. r is!'ermous 11 4 P le ur otropous 118 Protected Bud 93

l'erisporo 156 Plica t e 123 , 125 , US Protective
Pachyc,l\llv 92 Persi:;tent 146 Pliestesial 14 6 Stipules 97
1'3111t(>d .1 50 Perso n ate 10l, P lumose 108 I'rot e rogynoll5 1~6
\'>a1 "t (' 10) Pe tal <)9 , 10) Plumul e 114 I'rotha11u5 156
Pet a lanthero us Pneuma to phorous 9 1 P rotostele 15)
Pide 94
~;ta m en 104 Podogyne 108 Proximal 123
P"l ea 97
Pal eaceous 1411 Petaliferous Poll e n Grain ·d 07 Pruinosc 140
l'"lcClmofohic 102 Necta r y 15 0 Poll e n Sac 107 Pseudanthillm 99
Pal 1.nac t inod ramous 139 !'e taloi d l Oll, 151 PolU.na tio n Ty pc 86 Pseudo<;.arp Ill.
Pal ma te 154 Petaloid S tamen 104 Pollination Patlway Pseudodrupe 114
Pe talos temo nous 10 1. , 8(, Pseud omonome rous
!'Hlmately Compound 94
Palmately Ne tt e d 1.39 128 Pollinia 107 127
Palmate ~ pi. nnate 9/, Petiolar Etherea l Oi l Pol y a d 107 , 125 Pseudo-terminal
Pal~atifjd 128, 1 37 r:1Ilnd 150 Pol yandrous 1 25 Bud 9)
Pa lmatisect 128 Petiolate 97 , 137 Pol ya n thous 151 Psychophily 145
Panduriform 133 Pe tiol e 9) Po l ycarpel1ate 125 Pterocauly 92
Pan icl e 99 Pe tiolul ate 97 Polycarpic 1.46 l' t yx i s 85
Pannas e 140 , 143 Pe tio1u1 e 93 l'olycepha l ous 125 Pub erulen t 140
Pnpilionaceous 104 Petrorhi z ous 116 Polycycli.c 127 Puhescent 140
:'apil los e V ,Q Pha1aenoph ily 145 Po1 y delphous 104 , Pull Root 88
Pap pus lO] Phanerantherous 119 1 25 , 128 Pulverulent 140
Parall el 122 , 139 Pll1l!l eroco tylar Seedlin g Po l ygamo -dioecious Pulvina l 97
l'arall elo Jromous 139 Il6 14l, Pulvinus 93
Pa ra physes 1 55 Photosy n thet ic Stipu 1es Po lygamo-monoeeious Punctate 138
Pa ri eta l 118 97 144 Pungent 134
Ph y l1ar y 97, 98 Pol y gamo us 14t, Pustulate 138
I'nr ipinniltely
Compound 94 Phy 110c l.ad 92 Polyher ero pl lv tous l' y ramida1 130
Pa rted 128 , 137 Ph vllo dia 1 97 151 I'yrene III
Pilrtial Dimorphism Ph y110dium 97 l'o l ymerou s 127 Pyrifo rm 133
1 55 Phv11opodic 117 Po lvmorpl,ic 151 "yxl s 110
Partly Adnatc 128 Phy 110tax i s 14 8 I'olyphore q9
" ate111 form 133 P i lUe rous 13 4 Polystichous 119, 153
Patent 122 P Uose 140 Pome I II o
Pectinate 128 , 15/, Pi nna lSI, Population Sex 86
Pedate ]28 , 154 Pin nat e I S!, Pori cidal Capsule 110 r)lIadra t e Rhombic
Pedicel 98 , 99 Pi nn ately Co mpound 9l, Poricida1 129
Peduncle 98 Pinnately Nette d 1)9 De ldscence 107 0uadrate Rhomboid
Peg 152 P i nnat e-p innatifid 1St, I'orogamy 11.5 129
Pellucid It,3 P i nna tifid 128 , lJ 7, Porrect 122 fluadrifid 1 28
Pe l tate 137 154 Pos it i on 85 f~uincuncial 124

Pelti form l} 3 Pill natisect 128 i'ositiona1 Types 87

I' e nd ulous lIS , 122 l'i nn u1e 1 54 Posterior (~roove 103
l'en ni-para Jle l 139 Pi s ifo rm 1)3 PO$terior Line 103
Pentacyc l i.e 127 Pistil 99 Pos te rior ladge 103
Pcn til go na l 125 I'istillate 143 , 144 Pouch 10) Raceme 98
I'en t<:l mcr ous 127 Pi t che r 97 Prec ocious 1M) Rach i11a 93 , 98
l' e nt andr01l5 1.25 Pith 91 Preflo rat ion 85 R,1 ch is 93 , 98, 154
Pepa 111 T' l acent,1 ]'l7 , 108 , 109 Preformed Sh oots U.8 Ra dial 148
Perennial ll,r, Plain 125 Pri c kle 91 Radiate 119
Per f ect 1 4 3 J' !d i te d 138 Pr i c k ly 140 ll. adica l 116 , 117
Pe rf ol iate 1 3 7 P la nate 125 Pr imary Root 88 Ra dicle 114
Peria n th ')') Pla n e 123 , 138 P rimocane 92 Rarnea1 Sheath 153
I'ericarp 10') PIH Il t Sex 86 l'roant hesis 1 46 Rame n tace ous 140
l'ericarpous 124 Plated Bark 91 Procumhent <)2 Ramose 92
Pe r ic l ad ial 97 P l<ltyci! n tho us 126 Projected 122 Ramous 117
Pcrigyn iu m 98 P l ecLostc l e IS} Proleptic Shoo ts 148 Random 1 53
Perigyny llil l'leiochasium <j<) Promeristem 93 Rap he lOB, 114
Perine 156 Pl ei. omerOlJ>l 1 27 Prop T\oot 9 1 Ray 98 , 10)
Pe riodi city 87 P l c io taxy 12 7 Prophyllum 94 Rec eptacle 99 , 156
Pe rip heral 123 Pleomorphic 102 Prostrat e 92 Rec e p taculum 152
PeripherIll Emhryo 11 5 P l e uroga.my 145 Pro tan droll s 146 Reclinate 122 , 125
Perisperm Ill, Pleuro rh izal 1 1 7 Pro ta n the rous 146 l\ecl ined 122
6 165

Salt Excreting Gland 149 Sheath 94 Stele 153

Reclining Stem 92 Stellate 133
Rectangular 133 Salverform 104 Sheathing 137
Samara 110 Sheathing Stipule s 97 Stem 88
Recurved 123 Ste reomorphic 102
Samaracetum 111 Shining 140
~educed 126 Sterigmata 152
Same 151 Short Shoot 152
Reflexed 122 Stigma 107 , 108
Sapromyiophily 145 Shreddy Bark 91
Regular 148 Stilt Root 91
Sarcocauly 92 Shrub 88
Reniform 133, 134 Stinging 140
Sarcous 115 , 143 Silicle 110
Repand 137 Stipe 107, 108, IS!,
Scaberulent lllO S ilique 110
Repent 92 Stipe Bundles 154
Scabridulous 140 Simple 9 4. 151 . IS!,
Replicate 125 Stipellate 97
Scabrous 140 Simple Cyme 98
Replum 109 Stipe1s 94
Scale 93, 97 , 103 Sinistrorse 122
Resinous 1 40 Stipular Scar 91
Resupinate 122 Scale Leaf 152 Sinuate 123 . 138
Scape 92 , 98 Siphonostele 153 Stipu la te 97
Retic ula te 138 Stipules 94
Reticulodromous 139 Scapose 99 Size 85
Scarious 143 Slight Dimorphism 155 Stolon 92
Retinaculum 109 Stoloniferous 92
Scarred 138 Smooth 138
Retrorsc 122 Stomium 156
Retrorsely Crenate Scattered 119 Smooth Bark 91
Soboliferous 92
Sch i;>;ocarpic Achenes 110 Sto rage Leaf 97
137 Straight 123
Schi?ocarpic Ilerries 110 So1enoste1e 153
Retrorsely Serrate Striate 138
Schizoc3rpic Carcerules Solitary 99
138 Strict 92
110 Sorosis III
Retuse 134 Strigillose 140
Schizocarpic Follicles Sorus 156
Revolute 123 , 125 , Stdgose 140
110 Spad'lx 98
138 St riped 150
Schizocarp'ic Nericarps Spathe 97 , 98
Rhipidium 98 St rob ilus 152 , 156
111 Spathulate 133
Rhizocarpic 146 Strombifor m 133
Rhi zoid 156 Schizocarpic Nutlets 111 Spatu la te 1.33
Schizocarpic Samaras 111 Spatulate !~mbryo 115 Strophio1ate 115
Rhizome 92 Sty l e 107, 108
Sclerocauly 92 Species Sex 86
Rhizotaxis 148 Stylocarpellous 108
Sclerous 143 Spherical 148
Rhombic 129 Stylocarpepodic 108
Scorp ioid Cyme 98 Spheroid 128
Rhomboid 129 StylodiOus 107
Scurfy 140 Sphingophily 145
Ribbed 138 Sty1opodic 108
Seasonally 146 Spiculate 140
Ring Porous '"toad 91 Subacrocaulous 117
Ringed 138 Seasonally Dec iduous 146 Spike 98
Secondary Root 88 Spikelet 98 Suhapical 119
Ringed Bark 91 Spine 97 Subbasal 123
Robust 126 Seculatc. 133
Secund 99, 119 Spinose 134 Subbasicaulous 117
Rolled 123 Suhbas1fixed 107
Seed 88, 109 Spiral 133
Root 88 Suberous 143
Seed Coat 114 Spiroloba1 124
Root Cap 88 Spongy 11,3 Subglahrate 140
Root Hair 88 Segmen t 154
Semataxis 148 Spongy Pi th 91 Subg10bose 10!,
Rootstock 92 Submersed 116
Semicarpous 107 Sporangiophore 156
Roridul a te 140 Submersic8ulous 116
Semic r8s pedodromous 139 Sporangium 156
Rostellum 109 Sub petio1ar Bud 93
Semperflorous 145 Spore Shapes 156
Rosulate 119 Subterminal 123
Sepal 99 , 103 Sporocarp 156
Rotate 1 04 Subterranean 116
Sepaliferolls Sporophyll 97, 156
Rounded 134 Subulate 130
Nectary 150 Spotted 150
Rudimentary Sucker 92
Sepaloid 151 Spur 92 , 103
Embryo 115 Sucker Shoots 148
Septal Nectary 150 Squamose 140
Rugose 138 Suffrutescent 143
Septate 128 Squamulose 140
Ruminate 138 Sulcate 138
Runcinate 133 Septieida1 Capsule 110 Squa re 129
Septifraga 1 Capsule 110 Squarrose 123 Summc.r Annual 146
Runne r 92 Superior 117
Septum 109 Stamen 99, 104
Rupturing Superpos ed Ilud 93
SeriCeOllS 140 Staminal Disc 104
Capsule 110 Supervo1ute 125
Serotina1 146 Staminal Di sc
Serrate 138 Nectary 150 Supine 92
Serrated 128 Staminal Nc.ct ary 150 Supra folia r 117
s Serr"ulate 138 Staminate 143. 1411 Suprarhizous 124
Sessile 97 , 137 Staminodial Necta r y Surcarpous 116
Sac 10) Surcurrent 137
Seta 155 150
Saccate 104 Surface 86
Sagittate 13/, Setaceous 140 Staminodium 104
Setose 140 Standard 103 Surficial 116
Sagittiform 133
Shape 86 Standard Coloration Suspended 118
Salient 122
87
166 6

Syconium 111, U4 To r ose 133 Typica l Slamen 104 V.' rna1 146
Sylcptic Shoots 148 Tortuous 123, 138 Vernat ion 85
Symmetry 87 Torus 99 u Ver ruco~e 115. 140

Sympeta l ous 103, 128 Trace Sca r 91 Ve rsallle 107

Sympodial 149, 153 Trajiing 92 Umbel 98 Ve rticillaster 99
Synantherous 11,6 Transverse 122 Ur'lbellet 99 Ve r ticilla te 119
Syncarpous 107, 128 Transverse Deh iscence Ur'lbilicate IJJ .... ~ Vespe rt ine 146
Syncotyly 128 107 Umbo 152 V<:5 tl ~lal 1}7
Syngenesious 104, 128 Transve £!;e Pos tur e 85 Umbonate 115. 131 Vest iture 86
Synovario!Ls 107 Tree 88 Ur'lb r a cul:ne 1011 "<:;:i1lat" 124
Synscpalous 103 , 128 Tr ian gular 1)0 U~bracullforn 134 Vexillu", If)]
Syntropous J 18 Triearpellate 126 Unctuous 139 Vl 1105UloU5 143
Synlltyio va r ious 107 Tric yclic 127 Underground Stolon 93 Villous 143
Tridynamous 119 Undulate 123 , lJ8 V!n<! 88
Trifid 128 Ungu l cu l nte 104 VLrgate 92
T Triflorous 126 Unicarpe l lous 107 Viscid 1{'0
IJnJfoliolate ')4
Tannin llc(!ring Gland
Tr i fol iat e 126
Trlf01io1ate 9/" 126 Unilatera l 1 19 ,.
1l.9 Trifurcate 121\ Uni locul n r 126
Tap Root 91 Tr1p,ono"s 126 Un lnoda I :;hool 152 t~ at Cr Excre ting Gl an d
Te ndrIl 93, 97 Trlheteranthous 151 Un i ser l.1tC 126 149
Tentacu lar 97 Trihete r ophy rous 151 Unisexua l 126, 1{' 3 \,'horl 102
Tepal 102, 103 Tril ete 156 Unkno~'n 151 I·Jho rl ed 119
Tcrctc 108, 133 Trimerous 127 Urceolate 10/, lling 10)
Tcrgemin.1tc 9/, Trimorphic 151 Urcnt 140 lIinr,ed 97
Te r minal 116, 117, 119 Trioec i o"'; It.t. Ut ri cle 110 \!inr,ed Kark 91
Terminal Bud 93 Tripall'ate1y Compound llinged ;:u t llt.
Terminal Bud Sca l e 94 llin ter Annua l It.7
Scar Ring 91 Tripinnately Co~pound v \/ood 91
Tcrnatcly Compound 91. 94 \~ood y 1{, )

Tcssell<ltcd 1 50 Triquetrous 1 26 Vallecl!lnr Canal 15]

Tct ra cycl1c 127 Tristichous 119 Val ville l Z) , 12{'
Tet r ad 107, 126 Tri te rnate qt. Valvular Cupsu l c J 10
Tctradynamous 119 Trullate 130 ValvuL1T Dehiscence
Tetragonal 126 Trulloid 130 10) X"lloj!;any 1/.5
Tetrahedral 126 Truncate 134 Varlel:ated 150
Tetralocular 126 Trym.... lIt. Vilscular Ilu ndic Scar
Tetrumcrous 1. 27
Tctrandrous 126
Tube 103
Tuber 93
'I
Vegetative Frond 155
y

Tctrastichous 119 Tullertu 1ate 108, 140 Velum 156 Yearly 145
Texture 86 Tubercules 153 Velutinou~1 1M)
Theca 107 Tu hero us Root 91 Venat in n Sf>
Thorn 93 Tuh ular 104 Vent r al ll R, 12) z
Thr03t 10) Turh [na t e 1 33 Ve ntr al ~;Ide 10)
Thyrsc 99 Tu rgid lJ3 Ventricose lOt, Zoned 150
Tiller 93 TLldon 93 I'entristipul ar 12t, Zonoca uious 117
Tomcntose 1/,0 Twininr. 92 , 122 Verniform 134 Zvp,omorphic 102. 148
852 Appendix A

ILLUSTRATED ANGIOS PERN FAMILIES

The 25 plates that fo l low are part of a series that Carroll E. Wood, J r.,
has sel ected fo r use in t eaching from among mor e than 350 illustrations pre-
pared for A Gen eric Flora of the Southeaster n United States . which is under
hi s direct i on. One-hund r ed t wenty of these specia ll y labeled illustrations,
including r e presentatives of 70 families of dicotyl edons in eastern North
America , a r e be i ng produc ed as a separate publication by Harper and Row, the
publishers of this book .

All of the illustra tions prepared fo r the Generic Flora have been in-
tended to il lu strate as many biological de t ails as possible , and not to be
merel y " recogn ition" draHings . Host of the illustrations have been made
either from living plants o r from material preserved in alcohol , and all have
been checked f o r accuracy by one or more bo tanists who have \wrked on t he
Flora . The original illustrations are ten inches wide ; t h e length depends on
the size and number of deta i ls that are figured ; and all have been dra\<Ill for
reduction by one-half . Fo r t eaching pu rposes these plates have been modified
by placing the labels directly on the i llust r atio n , rather tha n in a conven-
t i ona l legend below it , and t hey are pub l ished here with red uction s of on ly
1 2 to 35 per cent . The combination of larger format and labeled illust r ation
makes i t poss i bl e to pinpo in t various st r uc t ures that are difficult to explain
otherwise (e. g ., the flm"er o f Asclepias) . It has not be en feasible, however,
to give the magnification of each componen t figure .

The l abels have been prepared by Professor Hood with the collaboration
of Dr . Elizabe th A. Shaw and the help of Dr . Kenn eth R. Robertson . Of t he
25 pl ates on pages 853-877 , eight were or i ginally pub l ished (with a greater
r eduction and with conventional legends) in the Journal o f the Arnold Ar boretum,
i n which the ext ensive ser i es of papers t r eating the genera of flowe r i ng plants
i n the southeas t ern United States has been appeat'ing s i nc e 1 958 (volume 39) .
The six artists whose work i s represented on the follow ing pages are Karen
Stoutsenber ger Velmure (initials KS o r KSV on illustration s), Doroth y H.
Ha r sh (Dmn , Arnold D. Clapman (ADC) , Vir g i nia Savage (VS), Rachel A. Wheeler
(RAW) , and Laverne Trautz ( LT) . The Gener i c Flora is a project of t he Ar nold
Arboret um a n d the Gray He r barium of Harvard University that has been made pos -
sible by the ge nerosity of Dr. George R. Co ol ey and by gr ants from the National
Science Founda t ion . Hany individuals have played a part i n the product i on of
the drawings and in various aspects of the research for the Flora . Even to
name them would take up mor e than the al l o t ted space , bu t t heir help has con-
t r ibuted in uncountab l e \"ays to these il lu st r ation s . I t is the hope of all
who have worked o n the Gene r ic Flora that these detailed i llustrations \"ill
be useful to both the beginning student and the experienced botanist .

Index to Illustra tions

(Fami l y arrangeme n t is that of Diels, A. Eng l er ' s Syllabus der Pflanzen-
familien . ed. 11 . nerlin . 1936)

Amaranthaceae 858 Fagaceae 857 Halvaceae 869

Araceae 853 Geraniaceae 857 Onagraceae 8 70
Asclepiadaceae 873 , 874 lIamamelidaceae 864 Or chidaceae ass
Caryophyllaceae 859 Juglandaceae 856 Ranunculaceae 860
Compositae 877 Labiatae 875 Rosaceae 865
Cruciferae 862 Leguminosae 866 Saxifragaceae 863
Ericaceae 872 Liliaceae 854 Scrophula r iace ae 876
Euphorbiaceae 868 Nagnoliaceae 861 Umbelliferae 871
 ARACEAE : ArisacllI a. a-j, A. atrol'U bclIs; k ·m, A. siewartison ii ; II , A. draconliulll ,.
'"~
'"
~
~.' ~
",,
"
,.
,
"

" ~"

~d
c : staminate flower

'%
g: carpel1ate h: Ovule
I.s. ~ (ofthotropous)
a: A. atrorubcns,
habit
sterile par! of ~pa.
dix 'appefldi~)

i: fruit, 1.5., show-

ing seed i: unusuall y if
large
i"

i~Z~j~
," infructescence
,
( ~~ "

/.
k: spathe
I: A. steward-
staminate"
view
sonii, spathe in
inflores· \ '1:'
cence, most I ' n: A. dracontium,
back view habit
of spathe r emoved

/" '"
i>1!1f'''''f," '''"'',
''''II':t)
: 'iV'
,.
(IU ~
fliJ'' ,' ,~ I ,!
carpella te

~~~~
inflorescence
r "'" . d "
b!
It) · d II
~:
with most of
spathe

,~
spathe,
A. alro-
rubens, I';;
portion \.
of stam i n . "
0 ,)
.D D
removed
, .
~ffi)~
from above "h ",n;;, '<om ,bo" V ,to ",d;, " 111Ii~,
s* ( 00

"' ~
w
854 Appendix A
ULIACEAE: Smilacin3 and MaianthcllIlIm. a-g, S. raCemOS3j h_l , i\L canadcnse

c : ovary in c. s.

·
.rn·
. .'.'
:".
'.-'

I : twin embryos from

GO
o
f: embryo

e; mature fruit,
a single seed
in c . s.

partly mature
fruit
k: l -seeded fruit in c. s. g: double embryo
(cf. "c")

h: flowering st em
and rhizome

j: nearly mature
fruit
Appendix A 855

ORCHIDA CEAE : Epidclldn llll sect ion EllcycJ i:l. :10111 , E. l:lml'cllsc

an th er

~ J ~.""". o~,",
YS

s" gtna " c

\ undeveloped

cavi ty m:
c. s. sepal
d: column ova ry I

"r·\'"

I: c. S. ovary between
base o f (I: sm(lll pl(lnt
nec· in fl ower
t ary
,"d
ovuli ·

ovary swlar ca nal

b: flower f: column
wi t h anther
removed ,.
stigmatic lobes with sticky
"

,
I
h: anther from below,
e: column pollini(l removed
from the 4 locules

c: l ip, fro m above, k: c. s. upper pan of

one lateral y: top of col umn, the ovary
lobe anther lifted
lip

upper sepal

OIIary

s tigmatic cavity

s ty la< canal
;: I. s. flower
856 Appendix A

J UGLA N DACEAE: C
;1-111 • • o\'al;1 v al'. ; III S I ra , ., ~.
,. II '~,
~ C. lae irlio."':t

S!illninate
f lower
....
h .
___
i
.
/ ~7

:2
i ...... - ,..
.

g.
flower ing
branC!l let

sub tendlll!J
staminate
f l ow~r

ovuk

?T\'\
V

r: term-
inal
bud in
winter
condi - ~Iigrn~

tion
seedl ing

q: carpellate c: carpelkne
flower flower
 - A
Appen d l.X

FAGACEAE: Fagus. a-I, F. CTrallllifolia

"

g: C:
stam inate flowe r
uranchlct
fru i t

branchlc!

' nfl oresc e nce

jnvo lu cr e

~?"'~ 2 OVlJ le I in eac h

la c u l e
k h
,oedlm, ~
f-
C:5. C<lr pe llate inflorescence
of- fruit to show
j: c.s . tyledons
f olded co

~- ~~"~"~:;: :; "
coty le don

~\ '~):1"10
~~rpellate~f\I~~Il\ir
I:

\6~'(,
'wig wilh ~}

floresc ence
III ZlJ'ffFF
-il
J ,
i
\,> 'b:
wmter bud -
.7'
-'--
>

WI-", P<lrt of ~j
~ , _ Y --
involucre " ;:
., to .Jl d
removeu '" /£_ bu
'?7
flowers ~" ~,~;1{"
show :t;' scale
' scars
,...,
.
.r ""
;,n, :;"',
.(\ ',II I
858 Appendix A
AMARANTHACEAE; Amaranthus. a-h , A. spinoslIs; j-j, A. rctroflexlls

h·
seed in
section

mature 2·carpellate
indehiscent frui t
(u tricle)

"
cymu le of
f lowers

b,
node wit h
pair of tap of
axillary

mature f r uit with per- i:

sistent & enlarged upper part
perianth of small
plant w ith
flowers and
fruit
pcrianth of
tcpal$

d,
3-carpellate
flower
"
5Ja
"' mature pyxis
Appendix A 859

CARYOPHYLLACEAE: Silcnc: lI-j . S_ virginica; k, S. carolillialla; I, S. ovata;

m, 11, S. antirrhina

mo
dehisced cap·
sule surround·
ed by d ried
ca lyx

r ovary
central =
plaCM(a .;
attached ~
at base &
I.s. top of ovary
'0 dehisced
laciniate petal
"". capsule

ca IV ~~"\J

g:ovule

ko
capsule
after j: seed in
dehiscence section

i; seed

"
small
plant in do
flower petal
with stamen

c: calyx lobes
860 Appendix A

RA NU NCULACEA E: C;llIh a. a.i, C. palusl ris

c: stamen

b: 8-carpcllatc flower

f: c,s, carpel
~ DC

d,
I~teral
view of
carpel g: open fol-
licles from
/",,~_pon gy f lotation 5·carpellate
tissue flower

~ndosperm

h: seed
emuryo
ncctJr
glands
i: seed in
I.s.
Appendix A 861

1\1AG NOLlAC EAE : Magnolia. a. i, 1\1. " irgi nian:l; j .l, 1\1. gra rulifl ora; Ill, 1\1. tripetal:!;
.HI, M. acuminata

dehis-
cence
lateral

stamen,
adaxial side

q~
c .s. anther

I
"'M. acuminilta
androecium (part
removed) & 9ynoe·
cium

If sty les
deciduous

I I "gynoecium in vertical
section

\
~\
I,
stamens,
adaxial
side

j:
flower bud about
to open

k'
androecium &
gynoecium, dehiscence
half of stamens introrw
removed . " ','
I,::!!:j " ':.
fleshv oUler
seed coal ,

~'1'1~ m II
"'Ip.' I•. :, hard inner
,'I ·:. " seed coat d: c.s. stamens
,
"",
h'
seed in
~ ther"
'"
, > ··'ilament"
vertical sect.
~ "stamens, ad·
SJif.
u~

1liiM.
0 axial side
862 Appendix A
RRASS ICACEAE (CRU CIFER
~ i II''.): C 'IJscll"I"• 1JIH. S'I' .pa~loriA· j.t, fru its, sec!1s, &
,-h,1lrYO~
aUll
, ~ 0 V •"lr tOUS species'

O oo"" ",o,,,:§;'-'
"Nasturtium
seed off"ICIn~l
. c, , 5","," :~',:~~"' "~ , f,
margin of
fruit
I
(rcp1urn) and
alter fall f septum
o valves

-~:dl'
-.J1,(
b: f lower
-"

r:jiaphanus
raphanistrum
fruit .
cot yl edons ,
Incu m bent "

h: embryo

"' ,
~';i/ :::~~~t
Sinapis alba
4-seedefuit LolJularia _. ", t." j : Streptanthus
m'
Corono~
maritim a
fruit fla tt'ened
',I1 perpendicular
to the rcp lurn
maculatus
fruit .
f ,5\
0>
didyr'l1-us par;;lilel to r eplum
$lfE

"'
Appendix A 863
SA XIFRAGACEAL :-:;a ... ifra"·1
r- ' .
;t.j . S• . I "", IX Ii" '. J·II
Hill" , , S, "ir,cill '," IlSJS
f"' '
...

.',
. ..Q,,'- -
", "..
,'ot 'JekJ?t""t
t~~'17JO
," ,

Y
t lower ki
. ,corolla
actmomorph-
"
n: dehl ~ced
capsule

«~/ ) i~
i

_ ii" '. :-0 "'"

~./

'Gi"s •:

I , S. virg" ini.
h: seed
I, ensi s
'
a:
S. michaux ii

', ,"l-, v /.:

q)
( '.

<<
S!f [
",
. .,
J
e : I.s. Imm ature t rU l t

"'
864 Appendix A

HAMAMELIDACEAE: 1-lall1'lIllclis. ,,·k, H. \'erllajis; I, I-I. virgilliall:!

i: H. vernalis , branch-
Ie! in late su mmer

k: seed swmi'lodium

flowe r in I.s ., stamen

at right removed 10
show staminod ium

~
'" fl ower buds
., ' . ;i'
, c: H. vernalis,

··· )\U
e : S\ilmc n

f,
stamen
u . ,",
h:

staminodia

opening
by
val ves I.

j:
locu lici-
dally de-
hiscen!

9·
abaxial side
of stamen
flowerin\!
l>ranclllet
 --- -----

Appendix A 865

ROSACEA E subfamily B.OSOIOEA E: "ol clltillll. ;1"11 , P. ean:ulcnsis; i, '>, simplex; j, P. recta;
k· m , P. Iri{It:IILala

k : leaf, b: le~f,
P. tridentata P. c~nadensi$

m: achcne

neclm
ring

f·
fruit, 2
sefJ~ls &
2 bra cts of
epicaly x removed

"
flower in vertical section

i: ICill,
"diagram P.
of cymose j: leaf ,
in f lores- P. recta
cence
embryo

g: achenc

season

c: flower
866 Appendix A
FAGACEAE (LEGUI\1INOSAE) subfamily FABOIUEAE: Baptisia. "'Ill, D. iltlstralis;
II, B. tinctoria; 0 , B. arachnifera

'o'i '/;,·
'-I y,, " . ,
Ul er "~', I

""d
-nner
coty ledon $eed
COatS

m: seed in section
h,
calyx
"od
androecium

i: st igma

f: side view of keel

k'
fruit
B. australis

0: leaf "
inflorescence
B. arachni fera

j:
I.s , gynoecium
b.Jnner
9'
hilum keel, from
w above
fruit,
seed
" . from above
B. ti nctoria

"
c: flower in sect ion
"
wing petal, '*'
"'
inner su rface
Appendix A 867

GERANJACEAE: Geraniulll. 01'.1 , G. 1ll1lculalum; k-III , G. carolinianulll

I: f lower, recep·
~ live , anthers mostl y
~ shed

b : flower at inner whorl

of stamens shedding pollen; stigmas not
(jji1\~i70"" rece ptive

•
,l f: c.s. ovary
stogmas recep- c: stamen of
a: plan t in tive
flow"r outer
whor l

ovary

disc

~DC
ovules

e: 1.5. gynoecium

m: calyx and
h: seed mature schizocarp
before dehiscence

j: c.s. of
cotyledons
of embryo
in " i"

i : "mbryo k: t ip of flowering
fruiting branch
868 Appendix A

EVI'HORBIACEAE: 11-1, Euphorbia l:orollllla; j, E. inundll(a; k, I, E. commutala;

Ill , E. denlalil ; n-p, Cham:Jcs),ce macuiata

o
stami nat e fl.
gland

embryo
"
oriented as
in seed
k,
cyathium from
ilbovc

",m," Yl ",;,",
1b r aCte o ic

d: staminate
c : cyathiurn wi th
walls re moved

flower

"upper
part o f
small plant
with flowers
y.
co lumella
,
~
anaxial
surface

& fruit (ax is) of

i: embryo
schizocarp schizocarp

p: seed

~
h; seed

m, from~
above
E. demata,
cyathium carpcl'ate

"
f : segmen t of schizocarp
after deh iscence
,,"~~-~n ectJr
gland
j:
E. inundata, "'
part of
cyathium
prost rate
plant -
note pattern
branching
Appendix A
870 Appendix A
ONAGRACEAE: Oenolhcra. a-k, O. missouriensis

. )' I,
"
stigmas

fruit before
dehiscence
b : upper part of open
"upper port of

d\]J~
floral flower
wbe
flower to show ",
insertion of t' \I
stamens at !
tip of floral
tube k,
pollen grains embryo
connected by
visein
threads

p lacenta -
tion
aKile, _ _ +_-+_'
r,
l.s. ovary
with base of ,
floral tube "

and base of
style
j: 1.5 . seed

h,
c.s. fruit

"
part of flowerin!!
plant
H
SIH
us
Appendix A 871

APIACEAE (Ui\lDELLIFERAE): Daucus. a-m, D. carota; n, D. pusillus

sty lopod;um
I.s. staminate
flower, ovary
absent

''''"'~
'. g: staminate flower

'. ~ .
• " stamen

')
I: bud,
n: D. pusillus staminate Ilower
fruit (schiwcarp)

j: fruit (schizocarp)

C;lrpophore bet_en
2 mericarps

m: embryo

"
with flowers
young fruit
I,
diagrammatic c.s.
Iruit

b: I.s. through inflorescence

e: los . per-
lect flower
after fall of stamens 1::5V

"'
SiH

"'
872 Appendix A

13
•
.. .<
~
~
~
> E
•
0 •
e"
£
'";.;.1
<
U
<
U
Appendix A 873

ASCLEPIADAC EA E , Ascle pias. a.f, A. sy riac,l; g, A. cOllllivcns; h, A. pcdiccilat a

corona hood
tip of

pof linium

gland

d : detached anther, from

inner side
b.
from above

hood
removed

g: pair of
c: 9ynostegium wilh pollin ia w ith
COro n ~
corona removed gland and trans-
hood lillor arms

h " flower wit h 2 pet als removed 10

long·st ipitilte gynoSlegium

ovary

f : 9ynoecium
 874 Appendix A

<
 -------- ----------------
Appendix A 875

LAi\lIACEAE ( LABIATAE): iI -J, S. urlicifoli "'\' k -0, S. Iyrala

st;gm"tic d: stamen , ,ide

'" "~~~
", ~
tones view
fen ile anthe,
halves

,,,,ii,
hall I l;v~ "'\ ' filame n t

fil.1ment
~\I / ,',
sic"le am ll e, /;jj;:Z:t!i;/h
f' half
stamens seen
hom ,bo,"
and behind
A
V stamens as \
seen from
fro nt
j: cm~IYooriented " 'lVH olJas1c sty le
as m nutlet

n: distal part
of s ty le k: basal
with leaf
unequal stigmatic lobes

I- 1.5 . flower

'
i' nu t let ~ OOe"i""

0 , m. \\';',.me",
-,, ',
,.. ...;
\~
'''~;::::::::;l:':~~~~~~~:;~:'O
" " fertile stamen
-" , "
both . I.s. ca lyx
t ip of with2of4
~nlher·halves
flowering nearly mature
pol liniferou5
plant nutlets
876 Appendix A
SCRO PH ULARI ACF.:AE ; Pctl slemoll. <I-J, P. C,l nC SCC Ii S

werm

j : I.s.
seed

"
inflorescence

d: flower in sect ion

stilm;nodium

i: se~d

._~~1f- p lacenta
h.
mature c~psule f·
g: c,s. ovary with persistent style LT I.s. ovary to
show axile placentation
 - ------

Appendix A 877

AST ERAC EAE (COMPOSITA E): Inflo rescence, fl o rets, and stamens

style _ _ _ _ _ _ _ _ _"!~

, . - - - - - pappus

"'Yir,!'----- involucral bract

(phyllary I

a, Aster spectabilis;
b, c, Erigeron annuus;
d, e, Erigeron (Conyza) bona-
"
inflorescence (head o r capitu lum)
riensis; f, Amph iachyris dra'
cunculoides; g, Aster umbella-
left half in section IUS; h, Pterocau lon undulatum

e: flower with
d: flower with fili- ligulate
form corolla, corolla
style not
visible

b,
tubular
flower

h,
detail of
; "
anthers
1H with
U .'
tail- Iy stamin -
like ate tubular
g : anthers with basal flower
termina l <lppendages appendages