PRINCIPLES OF SYSTEMATIC ZOOLOGY

(AEB 413)

CHAPTER 1

Introduction

1.1 TAXONOMY

This term is derived from the Greek words ―taxis‖ (arrangement) and nomos (law) and
was first proposed in its French form by de Candolle (1813). Mayr (1971) defined
taxonomy as the theory and practice of classifying organisms. Taxonomy, as defined by
Simpson (1961) is the theoretical study of classification, including its basis, principles,
procedures and rules. Organisms form the subjects of classification while the subject of
taxonomy is classifications (Gregg, 1954).

The Science and Philosophy of Taxonomy Classification, in biology involves the

identification, naming, and grouping of organisms into a formal system. The amount of
diversity in the living world is staggering. The variety of living things identified include in a
million species of animals, ½ a million species of plants, ½ a billion of fossil, and 3 – 10
million undescribed species. Differences in sex, age classes, seasonal forms and
polymorphic forms constitute varieties among organisms. There is a strong need for the
ordering of organisms which reflects the natural order of things involved in classification
methods. The vast numbers of living forms must be named and arranged in an orderly
manner so that biologists all over the world can be sure they know the exact organism
that is being examined and discussed. Groups of organisms must be defined by the
selection of important characteristics, or shared traits, that make the members of each
group similar to one another and unlike members of other groups. Modern classification
schemes also attempt to place groups into categories that will reflect an understanding of
the evolutionary processes underlying the similarities and differences among organisms.
Such categories form a kind of pyramid, or hierarchy, in which the different levels should
represent the different degrees of evolutionary relationship. The hierarchy extends
upwards from several million species, each made up of individual organisms that are
closely related, to a few kingdoms, each containing large assemblages of organisms,
many of which are only distantly related.
To construct classification schemes that correspond as closely as possible to the natural
world, biologists examine and compare the anatomy, functions, genetic systems,
behaviour, ecology, and fossil histories of as many organisms as possible. More than 1.5
million different groups have been identified and at least partly described, and many more
remain to be studied. All branches of biology contribute to such studies; the specialties
that are immediately concerned with the problems of classification are taxonomy and
systematics. Although the two disciplines overlap considerably, taxonomy is more
involved with nomenclature (naming) and with constructing hierarchical systems, and
systematics with uncovering evolutionary relationships.

The idea of classification has been with man from the start of history. The ancients
attempted to discover an explanation for the bewildering organic diversity by approaching
classification of organisms as inanimate objects and living beings based on procedures
of logic. Their aim of classification was to serve as an identification key. This objective
shifted and the interests of taxonomists broadened, when organic diversity after 1859 was
interpreted as the result of evolutionary divergence. In addition to providing identification
keys, the taxonomist now interpreted groups of organisms as descendants of common
ancestors. He developed interests in the pathways and causations responsible for
evolutionary changes. Since this has dealings with living organisms, the taxonomist
became interested in field studies in relation to organisms. Such studies revealed that
animal behaviour and ecology supply more important taxonomic characters of species
than mere observation of morphological differences of preserved specimens.

Contributions of Taxonomy to Biology

Considering the enormous diversity of organisms, the pattern which emerges from
taxonomic studies, adds credence to the theory of evolution. The role taxonomy in biology
is multiple and can be briefly summarized as follows:

1. A vivid picture of the existing organic diversity in nature has been worked out for us by
taxonomy.

2. It provides much of the needed information permitting a reconstruction of the phylogeny

of life.

3. It reveals many evolutionary phenomena available for causal study by other branches
of biology.
4. Almost exclusively, it supplies the needed information for entire branches of biology
(e.g. biogeography).

5. It supplies classifications which are of great explanatory value in evolutionary

biochemistry, immunology, ecology, genetics, ethology and historical geology to mention
but a few.

6. Indispensable in the study of economically and medically important organisms.

7. Taxonomist has made greatest contributions to our understanding of the structure of

species and of the evolutionary role of peripheral populations.

It thus, contributes significantly to a broadening of biology and to a healthy balance in

biological science.

1.2 SYSTEMATICS

Simpson (1961) defined systematics as the scientific study of the kinds and diversity of
organisms and of any and all relationships among them. It can be simply defined as the
science of the diversity of organisms. It deals with the kinds of animals, their distinction,
classification and evolution. (Blackwelder & Boyden, 1952). The word relationship
includes all biological relationships among organisms such as their phylogeny, ecology,
behaviour, physiology, biochemistry and even genetics.

Place of Systematics in Biology

It is dominantly concerned with diversity and this makes it unique among the biological
sciences. The major preoccupations of systematics are:

• To determine by comparison, the distinguishing properties of every species and higher

taxon.

• To determine what properties, certain taxa have in common with each other, and what
the biological causes for the observed differences or similarities are, and

• Finally, it concerns itself with variations within taxa.

Systematics deals with populations, species and higher taxa. No other branch of biology
handles this task of integration at similar level in the highly diverse organic world. It
supplies urgently needed information about these levels and cultivates a way of thinking,
a way of approaching biological problems. This is of tremendous importance for the
balance and well-being of biology in its entirety (Mayr 1968).

Contribution of Systematics to Biology

The contribution of systematics to other branches of biology and to mankind, when

considered, adds to an appreciation of its scope.

Leaders in different fields of biology have acknowledged their unreserved dependence

on taxonomy. Elton (1947) emphasized that without the knowledge of systematics, the
ecologist is helpless and the whole of his works may be rendered useless (Mayr 1991).

Scientific ecological survey cannot be carried out successfully without the most
painstaking identification of species of ecological importance or significance. A similar
dependence on this subject is true for other areas of science. The experimental biologist
without sound taxonomic knowledge is a failure. In comparative biochemistry, it is needed
for the identification and classification of molecular components. The evolution of
molecules (important area of research in molecular biology) can be understood against a
sound background of classification. With this knowledge, the biochemist can determine
what organism might supply the key to important steps in the evolution of molecules.

1.3 TERMS AND DEFINITIONS

1.3.1 Zoological classification

This is the designation and ordering of animals into groups on the basis of their
relationships i.e. of association by contiguity, similarity or both (Simpson, 1961). This is
done after assemblage of data or characters that can be applied to the unidentified
species. Classification deals with populations and aggregates of populations. In
classification, the ordering of populations and groups of populations at all levels is
undertaken by inductive procedures (i.e. a posteriori form of reasoning from effects to
causes).

For example,

All animals with feathers and beak = Birds (tentatively labelled as Group 1)

All animals with scales = Reptiles (tentatively labelled Group 2)

 All animals with hairs (furs) and mammary glands = Mammals (tentatively labelled
Group 3)

1.3.2 Identification

This is the determination of the taxonomic identity of an individual and its placement in
the previously established taxa or classes.

The procedure for identification is based on deductive reasoning (i.e. a priori – from
causes to effects) and deals with individuals and few characters (or a single one) that
throw the given specimen into an already established taxa.

For example, if a given specimen is found to possess a feature that describes a particular
taxon in the identification key, then the specimen is so identified. From the previous
example, if Specimen A to be identified has feathers and beak; then it must be a bird.

1.3.3 Zoological Nomenclature

This is the application of distinctive names to each of the groups recognized in the
zoological classification. Nomenclature is thus an essential adjunct or secondary outcome
of classification.

For example, Groups 1, 2 and 3 in the previous classification can be given the names
Aves, Reptilia and Mammalia respectively.

1.3.4 Taxon (plural = Taxa)

A taxon is a taxonomic group of any rank that is sufficiently distinct to be worthy of being
labeled by a name and assigned to a definite category in the hierarchy of classification.
According to Simpson, it is a group of real organisms recognized as a formal unit of any
level of a hierarchic classification. For example, Porifera (a Phylum), Crustacea (a Class),
Chelonia (an Order), Baetidae (a Family) and Homo sapiens (a Species) are all taxa. A
taxon always refers to a concrete zoological object. The species is a concept and not a
taxon because it cannot be described, but a given species such as the Lion (Panthera
leo) is.

Two aspects that need emphasis:

1. A taxon refers to concrete zoological objects (i.e. can be described).

2. Thus the species is not a taxon, but a given species such as Homo sapiens is.
3. Taxon must be formally recognized by the taxonomist.

Demes and geographical isolates become taxa only when formally recognized as
subspecies.

1.3.5 Ranking

This is the placement of a taxon in the appropriate category in the hierarchy of categories.
The rank of a taxon, therefore, is that of the category of which it is a member. The rank of
a category is its absolute position in a given hierarchic sequences of categories.

1.3.6 Category

This designates rank or level in the hierarchic classification. It is a class, the members of
which are all the taxa assigned a given rank. Such terms as phylum, class, order, family,
genus and species designate categories. It is thus an abstract term and a class name
while the taxa placed in these categories are concrete zoological objects. Category can
be defined but taxa are things and can only be described or delimited.

1.3.7. Phenon (Plural = Phena)

This is a sample of phenotypically similar or homogenous specimens at the species level.

For example, male and female, miracidium and cercaria belong to different phena, while
in the case of sibling species (i.e. morphologically identical but reproductively isolated
pairs or groups of closely related species), it is possible that several species belong to a
single phenon.

1.3.8 Taxonomic Character

This refers to any attribute of a member of a taxon by which it differs or may differ from a
member of a different taxon. It is in simple terms the characteristics of a taxon. Taxonomic
characters are not only morphological but could be functional in some aspects.
CHAPTER 2

Categories in Classification

The categories used in classification by taxonomists fall into three major groups:

1. Species category: The species is the lowest category in taxonomy and represents the
starting point in classification.

2. Infraspecific categories: These include groupings of distinguishable populations within

species

3. Higher categories (collective categories for higher taxa, used for grouping of species)

2.1 The Species Category

The nature of species is controversial in biology and philosophy. Biologists disagree on

the definition of the term "species." Philosophers disagree over the ontological status of
species. A proper understanding of species is important for a number of reasons. Species
are the fundamental taxonomic units of biological classification. Environmental laws are
framed in terms of species. Even our conception of human nature is affected by our
understanding of species.

2.1.1 Species Concepts

The concept of species plays an important role both in and outside of biology. Within
biology, species are the fundamental units of biological classification. Species are also
units of evolution — groups of organisms that evolve in a unified way. Outside of biology,
the concept of species plays a role in debates over environmental law and ecological
preservation. Our conception of species even affects our understanding of human nature.

Biologists offer various definitions of the term "species" (Claridge, Dawah, and Wilson
1997). Biologists call these different definitions ‗species concepts." The Biological
Species Concept defines a species as a group of organisms that can successfully
interbreed and produce fertile offspring. The Phylogenetic Species Concept (which itself
has multiple versions) defines a species as a group of organisms bound by a unique
ancestry. The Ecological Species Concept defines a species as a group of organisms that
share a distinct ecological niche. Others include the Typological Species Concept and the
Nominalistic Species Concept to mention a few. These species concepts are just few of
a dozen prominent species concepts in the biological literature.
What are we to make of this variety of species concepts? Monists believe that an aim of
biological taxonomy is to identify the single correct species concept. Perhaps that concept
is among the species concepts currently proposed and we need to determine which
concept is the right one. Or perhaps we have not yet found the correct species concept
and we need to wait for further progress in biology. Pluralists take a different stand. They
do not believe that there is a single correct species concept. Biology, they argue, contains
a number of legitimate species concepts. Pluralists believe that the monist's goal of a
single correct species concept should be abandoned. The controversy rages on and on!

One of the most elementary urges of man is the need to identify things and name them.
The prehistoric men even have names for kinds of birds, reptiles, fishes, flowers and
plants. The diversity of the world is discontinuous and consists of more or less well defined
―Kinds‖ of animals (called species) in any local fauna. The species problem has been
made much difficult than it is by confusion of the concepts underlying the terms phenon,
taxon and category.

Philosophically, all species concepts fall into three groups; the first being mainly of
historical significance although still upheld by few contemporary authors.

Typological Species Concept

The Typologists (believers of this concept) assert that the observed diversity of the
universe is a reflection of the existence of a limited number of underlying types or
universals. These types were supposed to be an equivalent of species. There is no
special relationship between the individuals which are believed to be mere expressions
of the same type. A type is seen as a sacred entity or a species. The observed variations
in nature were considered to be the result of imperfect manifestation of the implicit
characteristics in each species. This happens to be the concept of Linnaeus and his
followers, going back to the philosophies of Plato and Aristotle. Since Aristotle, species
have been paradigmatic examples of natural kinds with essences. An essentialist
approach to species makes perfect sense in a pre-Darwinian context. God created
species and an eternal essence for each species. After God's initial creation, each species
is a static, non-evolving group of organisms. Darwinism offers a different view of species.
Species are the result of speciation. No qualitative feature — morphological, genetic, or
behavioral — is considered essential for membership in a species. Despite this change
in biological thinking, many philosophers still believe that species are natural kinds with
essences. We can start with a brief introduction to kind essentialism and then turn to the
biological reasons why species fail to have essences.

Kind essentialism has a number of tenets. One tenet is that all and only the members of
a kind have a common essence. A second tenet is that the essence of a kind is
responsible for the traits typically associated with the members of that kind. For example,
gold's atomic structure is responsible for gold's disposition to melt at certain temperatures.
Third, knowing a kind's essence helps us explain and predict those properties typically
associated with a kind. The application of any of these tenets to species is problematic.
But to see the failure of essentialism we need only consider the first tenet. Biologists have
had a hard time finding biological traits that occur in all and only the members of a species.
Even such pre-Darwinian essentialists as Linnaeus could not locate the essences of
species (Ereshefsky 2001). Evolutionary theory explains why. A number of forces
conspire against the universality and uniqueness of a trait in a species (Hull 1965).
Suppose a genetically based trait were found in all the members of a species. The forces
of mutation, recombination and random drift can cause the disappearance of that trait in
a future member of the species. All it takes is the disappearance of a trait in one member
of a species to show that it is not essential. The universality of a biological trait in a species
is fragile.

Since this philosophical concept is sometimes referred to as essentialism, the typological

definition is also called the essentialist species definition at times. It must be emphasized
that basing one‘s species on morphology is quite different from using morphological
evidence as reference for the application of a biological species concept. (Simpson 1961).

Why the Typological Species Concept has been universally rejected

There are two practical reasons:

1. In nature, some individuals which are closely related (conspecific) may have widely
different structural characters (e.g. different phena existing in one species due to sexual
dimorphism, age differences, polymorphism and other forms of individual variation).
Different phena that belong to the same breeding population cannot be called different
species, regardless of the degree of morphological difference.

2. This concept does not support the existence of sibling species because sibling species
differ hardly at all morphologically, yet they are good biological species. From the
foregoing one can conclude that the degree of morphological difference is not a reliable
criterion for determining species.

Nominalistic Species Concept

The Nominalists deny the existence of ―real‖ universals (types) but recognize the
existence of only individuals, while species are man-made abstractions for convenience.
As Bessey (1908), put it ―Nature produces individual and nothing more… Species have
no actual existence in nature. They are mental concepts and nothing more… Species
have been invented in order that we may refer to great number of individuals collectively‖.
Species is just a name given for convenience. "I look at the term species, as one arbitrarily
given for the sake of convenience to a set of individuals closely resembling each other…
Only individuals exist, not universal classes. Taxonomic categories are arbitrary logical
sets with definitions.

This is not true because species of animals are not human constructs, nor are they types
in the sense of Plato and Aristotle, but they are something for which there is no equivalent
in the realm of inanimate objects.

Biological Species Concept

The two medieval species concepts discussed above lost their validity in the late 18th
century when it became clear that neither of them was applicable to biological species.
Historically, the Biological Species Concept is the most widely used concept among
ecologists. This concept combines the elements of the typological and nominalistic
concepts by stating that species have independent reality and are typified by statistics of
populations of individuals. It differs from both by stressing the populational aspect and
genetic cohesion of the species.

Basic Properties:

The members of a species therefore form:

1. A reproductive Unit. As a reproductive community, members recognize each other as

potential mates and seek each other for the purpose of reproduction. They are
reproductively isolated through the development of Reproductive Isolating Mechanisms
(RIMs) (i.e. features that prevent mating outside the species). They are also armed with
Species Recognition Mechanisms (SRMs) (i.e. features that allow recognition of potential
mates).

2. An ecological unit of Natural Populations, which regardless of the individuals

composing it, interacts as a unit with other species with which it shares the environment.
Organisms therefore derive their properties from the group.

3. A genetic unit which consists of a large intercommunicating gene pool. The species
form the gene pool, whereas the individual is merely a contemporary vessel holding a
small portion of the gene pool contents for a short duration. Conspecifics (members of
the same species) resemble each other because they are related (have common
ancestors).

These three properties took the species beyond the typological interpretation of a ‗class
of objects‘ and led to the definition of species.

"Species are groups of interbreeding natural populations that are reproductively isolated
from other such groups" (Mayr, 1969).

The development of the biological concept of species brought to an end the inappropriate
philosophy based on the phenomena of inanimate nature. This species concept is called
biological not because it deals with biological taxa, but because its definition is biological.

The criteria employed here are meaningless as far as the inanimate world is concerned.
A species is a protected gene pool, with its own devices (i.e. isolating mechanisms) which
protect it against harmful gene flow from other gene pools. The criterion of reproductive
isolation can be tested, observed, and /or inferred: A and B interbreed, but neither with
C; therefore, A and B are conspecific and C is a separate species. Populations maintain
identity when sympatric (occurring in the same place) and synchronic (occurring at the
same time). The more distant two populations are in space and time, the more difficult it
becomes to test their species status in relation to each other, but the more relevant
biologically this also becomes.

How different is the Biological Species Concept from the Typological Species
Concept?

The crucial difference between the reasoning of the Typologists and the adherents of the
Biological Species Concept is a follows:
1. The Typologist focuses essentially on morphological differences. He says if there is a
clear-cut morphological difference between samples A and B, they are by definition, two
morphospecies (i.e. two species). Any list of synonyms will show how often this
philosophy has led to the description of phena as species.

2. The Biological Concept is a means to an end because it provides a set of data with
which inferences can be drawn with regard to the nature of species. The biological
taxonomist uses the amount and kind of morphological difference only as an indication of
reproductive isolation, only as evidence to draw an inference. His species are usually
confirmed by subsequent researches in the area of behaviour, studies of breeding biology
(to sort out isolated types), life history and physiology or biochemistry.

Naming of Species (Binominal or Binomial System)

The scientific name of a species (binomen) consists of two words - the generic name and
the specific name. The former begins with a capital letter while the latter begins with a
small letter, and both are always underlined or italicized when in lower case. Hence Rana
temporaria, Bufo regularis, Periplaneta americana, Homo sapiens or preferably Rana
temporaria, Bufo regularis, Periplaneta americana, Homo sapiens. When all are in capital
letters, there is no need to underline or italicize the scientific names, as in RANA
TEMPORARIA, BUFO REGULARIS, PERIPLANETA AMERICANA, HOMO SAPIENS.

Difficulties in the application of the Biological Species Concept

Difficulties sometimes arise when the biological species concept is applied to natural taxa.
However, this does not make the concept invalid because many generally accepted
concepts face similar difficulties when they have to be applied in a particular situation or
to a specific example.

The three most serious difficulties in the application of this concept are those caused by
lack of pertinent information, those caused by uniparental reproduction and those caused
by evolutionary intermediacy.

1. Limited information: This is often encountered when handling preserved specimens

of either fossils or living forms. As a result of individual variation in all of its forms, it is
often difficult to tell whether a certain morphotype is a separate species or only a phenon
within a variable population. When sufficient information is obtained from a study of life
histories and through population analysis, individual variation resulting from sexual
dimorphism, polymorphism, age difference, e.t.c. can be unmasked. Only then can the
species status be fully established. The neontologist who works with preserved
specimens encounters the same problem as the palaeontologist who works with fossils;
they are likely to assign phena (morphotypes) to species.

2. Uniparental Reproduction: There are some organisms which do not reproduce

sexually but through one of the following ways: self-fertilization, parthenogenesis,
pseudogamy, vegetative reproduction, e.t.c. which are all forms of uniparental
reproduction. As defined in evolutionary biology, a population is an interbreeding group.
This makes asexual biological population a contradiction. The problem here is that we
cannot explain the grouping into species on grounds of interbreeding. Therefore, the
biological species concept based on the presence or absence of interbreeding between
populations is inappropriate for organisms reproducing uniparentally. Fortunately, there
are usually well-defined morphological discontinuities among kinds of uniparentally
reproducing organism produced by natural selection. It is customary to utilize the
existence of such discontinuities, and the amount of morphological difference among
them, to delimit species among uniparentally reproducing types.

3. Evolutionary Intermediacy: The species exists in full classical distinctness only in a

local fauna (i.e. when found in the same locality). The stage is set for incipient speciation
as soon as the dimensions of space (spatial) and time (temporal) are added. A species of
organism may be scattered all over Nigeria with different population densities. It can be
possible to observe some morphological differences between the extreme, of their
location. Under these circumstances (spatial and temporal distribution), populations will
be found which are in the process of becoming separate species (i.e. undergoing gradual
changes). This may impede classification into groups or species if the evolutionary
knowledge is lacking. In particular, the acquisition of morphological distinctness is not
always correlated with the acquisition of reproductive isolation. There could still be
interaction and interbreeding in which case they will be called some species but with the
presence of reproductive barrier among various subgroups, the application of the
biological concept becomes difficult. The following difficulties may result from evolutionary
intermediacy.
a. Acquisition of reproductive isolation without equivalent morphological change. Such
reproductively isolated groups (species) without (or with slight) morphological differences
are called sibling species.

b. Acquisition of strong morphological difference without reproductive isolation. This is

well illustrated by the West Indian Snail genus Cerion

c. Hybridization - occasional breakdown of Isolating Mechanisms (IM) between two

species. This leads to the production of occasional hybrids which may either be sterile or
of reduced viability. This does not cause any taxonomic problem. Hybrid individuals are
sometimes described as species before their hybrid nature is discovered. Such names
lose their validity as soon as the hybridism is established. Hybrid individuals are not
population, and are therefore not recognized as taxa. Taxonomic difficulty arises in
situations where new populations are formed as a result of hybridization. This is one sign
of the continuous nature of the process of speciation (the evolutionary formation of new
kinds of organisms). If hybrids are formed between two populations that are barely
differentiated, they may remain undetected, since their features may fall within the range
of variability of one or both of the populations. One would expect, if those populations
were to remain in contact, that they would blend together and lose their distinctness. On
the other hand, two populations may each have diverged so far from their common
ancestor that their individuals no longer recognize each other as potential mates. In that
case, it is agreed that the two populations should be considered separate species. They
will not fuse back into a single species.

It is between those two extremes of complete blending and total distinctness that
hybridization can provide glimpses of the complex process of differentiation, of evolution
in action. Speciation normally occurs in geographic isolation, but the distributions of birds
are complicated and ever changing. Populations once isolated often come into contact,
and when they do, the amount, duration, and results of hybridization will vary from
instance to instance.

There are different forms of hybridization:

i. Sympatric hybridization: When parental species maintain their genetic integrity over
a more or less wide area in which they occur together, it is advisable to uphold their
species status in spite of breakdown of isolating mechanism in a portion of their range.
ii. Amphiploidy: Hybridization in plants may lead to production of an allopolyploid, which
combines the chromosome sets of two parental species. Such hybrids may give rise
by uniparental reproduction to populations reproductively isolated from both parents
and behave like new species (if able to reproduce and occupy a new ecological niche).
The establishment of this in animals is yet to receive light.
iii. Semi species: Intermediate status between subspecies and species occur
occasionally in geographical isolates. They may or may not be considered species on
the basis of some criteria. It becomes more convenient for taxonomist to attach such
doubtful populations to the species with which they are most nearly allied. Other
instances of evolutionary intermediacy are circular overlaps and other borderline
cases (Mayr‘63).
CHAPTER 3

Polytypic Species and Infraspecific Categories

3.1 Polytypic Species

Our understanding of species is based on local populations without relating it to
changes in time and in space. This is the species of the local naturalist which is called
the nondimensional species. It is nondimensional because it lacks the dimension of
space and time. In a given locality, it is possible to have more than one species of
animals. These different species within the same locality are described as sympatric
species.
Every species consists of numerous local populations, and some of these may be
visibly different from each other. A population which is quite district from the population
of the original type locality of the species is describe as a subspecies.
A subspecies is therefore a geographically defined aggregate of local populations
which differ taxonomically from other such subdivisions of the species.

Fig. 3.1: Formation of Subspecies

These different populations, since they belong to the same species, are described as
conspecific (populations or individuals of the same species). Species which contain
two or more subspecies are called Polytypic species. Species which are not divided
into subspecies are called monotypic species.

Importance of Polytypic Species

The recognition of polytypic species is one of the most important developments in
taxonomy. It has led to considerable simplification of the classification of well-known
groups of animals like birds, mammals, butterflies and snails. In 1910, 19,000
monotypic species of birds were listed in the literature. In 1969 when the recognition
of polytypic species came to light, the number of bird species became reduced to
about 8,600. Polytypic species have also been identified in mammals, insects
(especially butterflies) and in molluscs (especial land snails). They show variety of
subspecies and similar simplification has also been reported in these groups.
Of much greater significance is the restoration of a definite biological meaning and
homogeneity to the species category which was greatly abused by the awarding of
species rank to every local population no matter how insignificant the difference. This
completely destroyed the biological significance of the species category.
Some difficulties are encountered by taxonomists when trying to establish polytypic
species. The first difficulty is that polytypic species are composed of (a) allopatric (i.e.
in terms of geographical spread, in which case different populations occupy different
geographic zones which are close together) and (b) allochronic (i.e. in terms of time,
in which case species do not occur at same time level). These populations differ from
each other. However, all populations of sexually reproducing organisms also differ
slightly from each other. Therefore, certain standards must be met before subspecies
are recognized.
The second problem is that occasionally, closely related species replace each other
geographically and this is possible where they have similar ecological requirements.
Yet they are recognized as full species and not subspecies.

Nomenclatural Problems
A polytypic species is often a compound of several species originally proposed as
monotypic. It differs from the Linnaean species by no longer being the lowest category
(which is now the subspecies) and by being a collective category.
What scientific name should be given to this new collective taxon, and who should be
the author? Subspecies are named according to the trinominal or trinomial system.
Thus, we have Sarotherodon melanotheron melanotheron Rupell - a tilapia fish,
Brachionus patulus patulus (rotifer), Osteolaemus tatraspis tetraspis (dwarf crocodile),
Passer domesticus domesticus. Osteolaemus tetraspis was the original species
described by the author in West Africa. This species was divided into two subspecies
(when another related pop was discovered in Congo). The specific name of the West
African species was now duplicated, thus Osteolaemus tetraspis tetraspis (the
nominate subspecies), while the Congolian group became Osteolaemus tetraspis
osborni.

3.2 Infraspecific Categories and Terms

1. The Variety
This is one of the commonest and most abused terms in taxonomy. Morphological
differences in the past constitute variety as used by Linnaeus in his classification. On
typological principles, each species has a fixed pattern. Anything that did not fit the
idealized pattern was a ‘variety‘, a conceptual extension of the scholastic ‗accident‘.
This term has become obsolete in systematics.
2. The subspecies
This was a replacement for the term variety in some point in history. This was when
variety was also applicable to geographic race. The subspecies may be defined as an
aggregate of phenotypically similar populations of a species, inhabiting a geographic
subdivision of the range of a species, and differing taxonomically from other
populations of the species.
From this definition, it is clear that a subspecies may consist of many local populations
all of which, though very similar, are slightly different from each other genetically and
phenotypically. It is therefore a collective category. Secondly, subspecies are to be
named only if they differ taxonomically, i.e. by sufficient diagnostic morphological
characters.
Difficulties in the application of the subspecies category

The following aspects of the subspecies have reduced its usefulness.

1. Different characters may show independent trends of geographic variation, i.e.

there may be no correlation between the varying characters.
2. There could be independent reoccurrence of similar or phenotypically
indistinguishable populations in geographically separated area. The tendency is to
associate the subspecies with certain geographic zones.
3. The arbitrariness of the degree of distinction considered by different specialists as
justifying the subspecies status of slightly differentiated local populations. There are
no laid down rules or uniform guidelines for the separation of subspecies.
Various aspects of geographic variations cause difficulties. Indeed, the subspecies
have been misused in many ways. Some authors applied the term to individual
variants and sibling species; many authors named insignificant local populations as
subspecies and finally, some authors considered the subspecies as a unit of evolution
rather than as on arbitrary device to facilitate intraspecific classification.

Determination of sub species

This process is largely based on morphological features.

1. To determine the overlap in the range of the features, i.e. to plot linear overlap of
observed samples

The parameter for range determination in this case = Head size ÷ body length
The question is "How meaningful is such overlap?"

This method was later found to be of limited use because there could be some minor
differences in characters. It gives information on the overlap of the sample but does
not give much information about the size of the populations. It exaggerates the
importance of the end point of the range, i.e. tails of the overlapping curves. Linear
overlap is therefore a very unsatisfactory way of describing the degree of difference
between two populations.

2. Race
When two populations of a single species show considerable genetic differences and
where they occupy ecologically distinct or geographically well separated habitats, they
may be called races.

a. Geographic race: The term subspecies and geographic race are frequently used
interchangeably by taxonomists of mammals, birds and insects. Other taxonomists
apply the word race to local populations within subspecies.
b.
c. Ecological race: The nature of ecological races among animals is still
controversial in that it is based on the existence of ecological differences. Since no
two localities are exactly identical with respect to their environment, every
subspecies could also be considered, at least theoretically as an ecological race.
It is however possible for some populations to differ in their ecological requirement
without necessarily acquiring taxonomically significant differences.

The characteristics of a race can easily be determined with reference to man. Human
races are subdivisions of mankind based on geographical localities, and
morphological differences like skin colour, hair colour, shape of the nose etc. These
differences are genetically based and do not reflect differences in ecological
requirements.

Inheritance of characters may be polygenic (i.e. controlled by several genes) e.g. as

in skin colour for the black, white and the Negro race. Racial differences may be
temporary or permanent depending on the degree of isolation.

d. Host Races: These apply to parasites and species-specific plant feeders. If gene
flow between populations on different hosts is drastically reduced, such host races
are the equivalent of geographic races in free - living animals, and often develop
subspecific characters.

3. Cline: This term is used to represent a character gradient. It is not a taxonomic

category, but it is so important in systematics because it refers to the existence of
small morphological and genetic changes over the distribution range of the
species. In most cases, the population is not broken into distinguishable groups,
but shows gradual continuous change in characteristics along a gradient such as
north to south, lowlands to highlands or dry to moist climates. Such cases (clines)
 are exemplified by species of birds and mammals that are larger in cooler climates
(Bergmann‘s rule) or darker in warm humid regions.

4. Intrapopulation Variants (variants within a population): The existences of phena

are examples of intrapopulation variants. The morphs (different forms of the same
stage) found in polymorphic populations are also examples of such variants.
5. Asexual entities: In most lower invertebrates, reproduction is uniparental through
parthenogenesis or vegetative budding. Since interbreeding is the ultimate test of
conspecificity in animals, and since this criterion is available only in sexually
reproducing organisms, the determination of categorical rank for taxa of
uniparentally reproducing organisms is difficult. It is always difficult for taxonomists
to treat clones, strains or stocks of such organisms.

6. Neutral terms: These are terms which a taxonomist may assign informally to
phena, particularly for the purpose of convenience. The ones most frequently used
are "form" for a single unit, "group" or "complex" for a number of units.
Form is used when one is not sure whether the phenon in question is a full species
or a subspecies. Seasonal and polymorphic variants are referred to as forms. The
term group is applied to an assemblage of closely related taxa which the
taxonomist does not want to place in a separate category.

3.3 Speciation and Isolation

This is the splitting of a phyletic line, the process of multiplication of species, or the origin
of discontinuities between populations caused by the development of reproductive
isolating mechanisms. Put simply, speciation is the process by which new species are
formed and isolation is usually the essential step in the formation of new species.

Reproductive isolation is on the one hand, a basis of our definition of species and on the
other hand a major factor in creating one species from another.

There are two types of speciation

1. Allopatric speciation – formation of species during geographical isolation

2. Sympatric speciation – Speciation without geographical isolation, which is the

acquisition of isolating mechanisms within a deme (a local population of a species).
Formations of isolative Mechanisms (Reproductive barriers)

1. Geographic isolation:

This has to do with spatial separation which may prevent physical contact between
different populations. 2. Reproductive isolation

a. Prezygotic barriers (preventing formation of zygote).

i. Seasonal differences: If we have populations which differ according to the breeding

season, the chances of having cross-breeding between the two populations are slim.

ii. Mechanical differences: This is another prezygotic barrier which relates to differences
in the copulatory organs. If there are differences in the copulatory organs of closely related
species, such as to prevent copulation, a barrier is created.

iv.Ethological differences: This is relevant when considering species which show

courtship behaviour before mating. It is observed that some animals show slight
differences in courtship behaviour which constitute a barrier. Example is found in two
closely related species of Drosophilia, D. melanogaster and D. percincitis. The female
of D. melanogaster is highly discriminatory in its response to the courtship behaviour
of the males. The difference in the courtship behaviour of these related species is a
barrier to breeding and therefore constitutes reinforcement to the separation of species.
v. Gametic incompatibility: Gametes for one reason or the other do not fuse to form a
zygote. This is attributed to the existence of physiological differences.

b. Postzygotic barriers

i. Hybrid inviability - where hybrid zygote fails to develop to adult.

ii. Hybrid sterility - where hybrids do not reproduce at all, even if adult. The existence
of this barrier implies that fertilization has taken place but zygote fails to develop.
Example is the cross between the Horse and the Donkey to produce a mule which is
sterile. The same is true of crosses between fowls and turkeys or pheasants.
CHAPTER 4
Higher Taxa

A higher taxa can be defined as an aggregate of related species separated from others
by a discontinuity. The groups of species found in nature are higher taxa. Cats,
carnivores, mammals, and vertebrates are higher taxa of different rank.
In classification, the first step is to determine which species show similarities indicating
that they belong to the same group, and secondly to determine what separates one
group from the other (to locate the discontinuity).
Suppose we have the lizards and snakes, both of which are examples of reptiles.
Reptiles are Gnathostomes → vertebrates → chordates. The cats are mammals →
gnathostomes → vertebrates→ chordates. The different animals can be regrouped into
closely related families, and arranged in an ascending order. This grouping is possible
because the members of the family share common structural characteristics.
Here, the difference between similarity and relationship (affinity) needs clarification.
They do not belong to the same family because they share common characteristics,
but they share common character because they belong to the same family, because
they originated from a common ancestor. In order words, members of a taxon are
similar because they share in a common heritage, they do not belong to the taxon
because they are similar. It is like the identical twins; two brothers are not identical
twins because they look similar, but they are similar because they share the same
origin (they are both derived from a single zygote), i.e. because they are identical twins.
One fatal weakness of the nominalist thinking when applied to classification of
organisms, is the reversal of an existing causal relation between similarity and affinity.
Groups of animals sharing a common ancestry clearly qualify as a monophyletic group.
This common ancestry seems to be the central theme for classification. It is known that
members of a monophyletic group may occupy the same adaptive zone or a well-
defined niche, but undergo adaptive radiation in several directions, while maintaining
considerable amount of structural unity. The groups in a taxon evolve from a founder
species i.e. the first colonizers of a virgin land. Examples are the finches of Darwin.
Phylogenetic as well as ecological relationship must be associated with the discussion
of higher taxa. They have same origin and occupy the same adaptive zone.
The Linnaean Hierarchy
The taxa of animals and plants are ranked in a hierarchy of categories. Aristotle was
the first to attempt classification but his method could not stand the test of time because
of its arbitrariness. Linnaeus, a Swedish naturalist introduced his system of
classification in 1758 in his popular book ‘Systema Naturae‘ (which literality means
Natural System). It was based on some biological relationship of the organisms within
the animal kingdom. The highest regularly used category is the phylum and the lowest,
the species. Linnaeus, the first taxonomist to establish a definite hierarchy of taxonomic
categories, recognized only five categories – Class, Order, Genus, Species and
Variety.
The term variety was later replaced by the subspecies. More categories were created
with time. These were the phylum and family. Any given species belong thus to seven
obligatory categories as follows:
1. Kingdom
2. Phylum
3. Class
4. Order
5. Family
6. Genus
7. Species

Question: Attempt the classification of 1. Housefly, 2. Rat, 3. Man, 4. Cobra

With the discovery of more animals and more relationships, it became necessary to
create more categories. This involved the splitting of some of the existing categories
by using prefixes as –sub, or super - thus we have:
1. Kingdom
2. Sub- kingdom
3. Super phylum
4. Phylum
5. Sub phylum
6. Super class
7. Class
8. Sub class
9. Super order
10. Order
11. Sub order
12. Super family (oidea)
13. Family (-idea)
14. Sub family (-inae)
15. Genus
16. Sub genus
17. Species
18. Sub species

The sub divisions are a reflection of groupings of closely related organisms. Although
Linnaeus created categories on the basis of affinities, he could not explain the
significance. He has thus been described as a pre-evolutionary taxonomist. Though
his affinities have some evolutionary relationships, he couldn‘t recognize these before
the dawn of evolution by Darwin.

How different is the concept of the species from the concepts of the higher
categories?
A higher category is a class into which all the taxa that rank at the same level in a
hierarchic classification are placed.
The category selected for a given taxa indicates its rank in the hierarchy. Taxa as earlier
pointed out, are based on zoological realities, but categories are based on concepts.
 In that respect, there is no difference between the category species and the higher
categories from the genus up. Examples like species, genus family etc are all
categories and are therefore conceptual.
In many other respects there is a great deal of difference between the concept of
species and the concepts of the higher categories.
1. The category species is self-operationally defined by the testing of isolating
mechanism in nature, but definition for the supra specific (higher) categories are all
arbitrary.
2. Species category signifies singularity, distinctness and difference, while the higher
categories have the function of grouping and ordering by emphasizing affinities among
groups of species rather than differences between species. They are therefore
collective concepts.
3. Finally, evidence used for the delimitation of higher taxa and their ranking into
categories are furnished by comparative data while interbreeding is the criterion used
for ranking at the species level.

Higher category includes genus, family, order, class, phylum.

The genus (pl genera)

This is the lowest higher category. The genus is not different in concept from family, order
or other higher categories.

It is defined pragmatically as a taxonomic category containing a single species or a

monophyletic group of species, which is separated from other related genera by a decided
gap (or major differences). The characteristic of a genus are not static. In the course of
evolution, some differences may occur making it necessary for the genus to be separated.
E.g. All Tilapia belong to one genus but as more important differences were discovered
in some of the populations, it becomes imperative to introduce more generic names.
The Family

The definition of the family is also pragmatic, since it cannot be defined operationally. The
definition is equivalent to that of the genus. A family is a taxonomic category containing a
single genus or a monophyletic group of genus, which is separated from other families by
a decided gap. E.g. the ladybird beetles (Coccinelidae), the long-horned beetles
(Cerambycidae), the woodpeckers (Picidae) etc. Family characteristics are supposed to
adapt the individuals to certain adaptive zones or to certain life style. The association of
families with adaptive zones or specific niches is much stronger than a similar relationship
with the genera. Families are more widespread in distribution than the genera.

Illustrating with local examples of the family of freshwater fishes (Cichlidae): They are
found in America, Africa (West, East, Central and South). The genus Cichlosoma is found
only in Central America. In West Africa, we have the genera Tilapia, Sarotherodon,
Hemichromis. In East/central and south Africa, Tilapia, Sarotherodon, while in East
African Lakes, Haplochromis.

The family is a very useful category to the general zoologist because each family usually
presents a general feature which is recognizable at a glance.

Orders, Classes and Phyla (Upper categories)

Each of these can be defined in a similar way as in the genus and family. The upper
categories are very well defined and represent the main branches of the phylogenetic
tree. They are characterized by a basic structural pattern laid down in evolution.
Superimposed on it is adaptive modification resulting from series of adaptive radiations
that have taken place in the classes and phyla.
It should be noted that classification and phylogeny are based on natural groupings.
Secondly, classification seems to have a lot of meaning when related to phylogenetic
relationships. This type of classification based on phylogenetic relationships is called
―Evolutionary taxonomy‖. The relationships are based on homologous structures
(having similar origins but different functions; appearance may/may not be similar).
Analogous features are similar in appearance but cannot be traced back to the same
origin in the common ancestor.

Similarity is not part of the definition in the case of homology because homologous
structures are by no means necessarily similar (e.g. ear ossicles of mammals and the
corresponding jaw bones in the lower vertebrates). Similarity must be referred to in the
definition of analogy, because non-homologous features that are not similar are not
considered analogous.
CHAPTER 5

Methods of Zoological Classification

5.1 Taxonomic collection and the process of identification

Taxonomic collections may be borrowed from museums/which are repositories of

systematic collections or collected by the specialist himself.

Value of collections

Museums serve an important role as centres of documentation. They supply a permanent

record of faunas and floras particularly where the biota has since been destroyed by
natural catastrophes or human activities. Collections are reference materials, which
although not in continuous use, yet must be available when needed. Such material forms
the basis of published research. It may be needed again at a later period for verification
of the original data or for renewed study in the light of more recent knowledge.

Purpose of Scientific collection

Collection should be seen not just as an identification collection but as the sampling of
populations. Identification collection avoids duplicates, but considering the great
variability of most natural populations, an adequate sample of every population should be
collected and preserved.

The size of a collection or sample depends on the objectives of the research. For
example, more material is needed in a species with strong individual and geographical
variation than in a uniform species. When the purpose of the research is population
analysis, large samples from numerous localities are needed. In fishes for instance,
where the meristic characters of each population of a species may vary statistically, a
single specimen per locality would tell us very little. All phena of a species, should as far
as possible be represented in the collection.

Scope of collections

Most museums are largely restricted to a geographic area and to certain groups of
animals. Only very few national museums attempt worldwide coverage in all groups of
animals. However, too broad a coverage inevitably leads to shallowness and a failure to
obtain in-depth knowledge of all the groups required for monographic studies.
Faunal surveys have the best results when concentrating on a particular taxonomic group.
The study of ecology, habits and other life history parameters which may be quite
burdensome in broad purpose research becomes easy to handle when specific families
or genus are to be considered.

Where and How to collect

It is absolutely necessary to plan carefully, any collecting trip. All possible geographic
information must be obtained beforehand, including the distribution of vegetation types,
seasons, means of transportation etc. Previous collections must be carefully analysed
and existing type localities mapped out. If a species type shows seasonal variation, the
collection should be spaced seasonally.

Where should collection be made?

Owing to the rapid increase of human populations, resulting in more intensive agriculture
and in drastic deforestation, there are many areas that are in desperate need of
immediate collection before the localized faunas become extinct. Techniques for the
collection of different animal groups are numerable and are described in standard
collecting manuals. These involve the use of sampling nets, traps, baits, poisons, etc.

Preservation of Specimens

The method of preservation differs from one taxonomic group to the next. The basic rule
is to preserve specimens in such a way as to prevent them from any kind of deterioration.
There are appropriate manuals and handbooks for description of specific methods-like
slide preparation, drying, use of alcohol, formaldehyde (formalin) etc. Properly preserved
specimens are still in excellent condition 200 years after being collected.

Labelling

A specimen that is not properly labelled is worthless for most types of taxonomic research.
The jocular dictum that “the label is more important than the specimen” indeed
emphasizes the weight of a label. Information such as the exact locality of collection, date
of collection, collector’s name, etc should appear on the label.

Identification The identification of previously unidentified material can be done through

the following procedures:
Sorting of collections All material collected during any expedition or collecting trips must
be sorted and at least tentatively identified before it can be incorporated into a collection.
E.g. Samples collected from the same locality – ponds, streams, and terrestrial habitats
are separated right in the field before further attempts at identification.

The first step in the laboratory is to sort the organisms into groups on the basis of
similarities in the most visible characters.

The second step is to accumulate relevant literature that will aid in the identification. E.g.
Handbooks, manuals, identification keys, etc.

A beginner trying to identify a specimen can usually tell that it is a bird, a prawn, a
grasshopper or a butterfly. He can immediately go to identification keys and manuals
specializing in these zoological groups. The beginner is advised to seek expert advice
from qualified taxonomists whenever he encounters difficulty in using the identification
keys. The expert is able to identify the specimen down to the species level. Each
specimen identified is given a determination label which contains the following
information: generic and specific name and author name of the determiner and year in
which the identification was made. This information establishes the authenticity of the
determination.

How does the identification key look like?

E.g. group-vertebrates

5.3 TAXONOMIC CHARACTERS

A taxonomic character is any attribute of a member of a taxon by which it differs or may
differ from a member of a different taxon. The taxonomic characters thus help to
distinguish one taxon from another. Features by which individuals of the same population
differ from each other, such as differences between the sexes and age classes, are not
taxonomic characters. However, when taxa differ from each other by the presence or
absence of sexual dimorphism, larval characteristics or characteristics of age group, then
such a difference becomes a taxonomic character.

Taxonomic characters are population characteristics. They have two functions:

a. They are diagnostic in that they help to specify a taxon.

b. They are used as indicators of relationships

Taxonomic characters are not all equally useful; some are powerful indicators of
relationship, others are not. The usefulness of a character therefore depends on its
information content i.e. ability to reveal relationship. Characters could be weighted as high
or low.

There are two types of characters – phenotypic characters and genotypic characters.
Classical taxonomy deals mainly with phenotypic characters because they are more
easily discernable genotypic characters.

There are two groups of characters, whether phenotypic or genotypic Unreliable

characters:

These are characters which are poor indicators of relationship, and are therefore said to
have low weight. They include:

a. Highly variable characters Variability refers not only to the presence or absence of
the character, but also the variation of the character in a population. Here the character
is not specific enough to help in the identification of the particular taxon. E.g. Body size,
wing venation, banding in snails (white or black band)

b. Totally invariable characters

E.g. the paired eye of vertebrates

c. Characters difficult to determine

E.g. number of hairs in mammals

d. Regressive characters

Taxonomic characters based on the loss of eyes wings, toes or other appendages or
certain teeth are often unnatural and of low taxonomic weight.

Key characters

These are characters that are:

i. Easily perceived
ii. Of very low variability and

iii. Usually present in preserved specimens. The important characters of a group to be

studied should be known before attempting the classification of such a group.

5.4 KIND OF TAXONOMIC CHARACTERS

A Morphological Characters: Basis of old taxonomy

i. General external morphology.

- Feathers (birds), hairs (mammals)

- Scale counts (fish and reptiles)
- Sutures & sclerites (arthopods)
- Exoskeleton (shells of molluscs, tests and the hand parts of some protozoa)
ii. General internal morphology (Anatomy) e.g. Foraminiferans/radiolarians, testacerus
rhizopods etc.
- Skull and teeth of mammals
iii. Special structures of genitalia (arthropods)
iv. Larval stages and embryology - Tadpoles (amphibia)
- Nymphs of odonata (dragonfly and damsel fly)
- Characteristics of the eggs (e.g schistosoma eggs
- S. hemalobium and S.mansoni)
For instance, the various sibling species of the Anopheles maculipennis complex (malaria
mosquitoes) were discovered due to differences in egg structures.
v. Karyology: number and patterns of chromosomes
The classification of animals, particularly insects have been advanced through studies of
chromosomal patterns.
B. Physiological and Biochemical characters
a. Metabolic factors
b. Serological, protein and other biochemical differences
c. Body secretions
d. Genic sterility factors

Physiological characters are not easy to define.

All structures are the products of growth processes, i.e. physiological processes, and are
therefore, ultimately physiological characters. Also, since all physiology is regulated by
enzymes and other macromolecules, it is not quite different from biochemical characters.
By physiological characters, we mean growth characteristics and temperature relations
of different populations. These characters are good indications of species differences, but
they are rarely used by taxonomists because
1.They are not present in preserved specimens
2.They usually require special apparatus and techniques for their study

Biochemical characters
This is based on the kinds of macromolecules (proteins) and metabolic processes. It is
known that there is some marked specificity in the nature of these characters at every
taxonomic level, and this is the basis of serological studies (for protein comparison).
The serological method is based on the principle that the proteins of one organism will
react more strongly with antibodies to the protein of a closely related organism than to
those of one more distantly related.
Improvements in this technique have stimulated interest in the study of antigenic
reactions. Immunogenetics (study of blood-group genes) has shed some light on
relationships between the distributions of haemoglobins, amino acid and species of
Pigeons and Primates Pvriness, Pyimidines is specific can help in establishing natural
groups. They form the basis of chemo taxonomy-devoted to the taxonomic study of
specific chemical components and macromolecules.
Paper chromatography has been used to compare the chemical composition of closely
related species (e.g. for amino acids, and peptides through minhydrin treatment, purines,
pyimidines etc). Various methods of electrophoresis reveal the molecular composition of
complex proteins.
It is also possible to consider a single complex molecule (e.g. haemoglobin) of one
species and compare its amino acid composition with that of closely or more distantly
related species, this can indicate whether or not two organisms belong to the same
phyletic line.

C. Ecological characters
This is very important in establishing relationships especially at the species level, and to
some extent genera. In nature, every species has its own niche, and differs from its
nearest relatives in terms of:
Food preference
Breeding seasons
Tolerance to physical factors
Resistance to predators, competitors, pathogens and in other ecological factors.
According to the competitive exclusion principle of Gause, no two species occupying
exactly the same niche can coexist in the same place at the same time. So, when two
closely related species coexist in the same general habitat, they avoid fatal competition
by the above species-specific niche characteristics. Several sibling species have been
discovered as a result of discrepancies in food preference or habitat preference. E.g in
the Galapagos finches, the different genera have different niche requirements. Geospiza
(ground finch), feeds chiefly on seeds, camarhynchus (tree finch) feeds chiefly on insects,
and certhidea (a warbler finch (feeds mainly on small insects.
Many aspects of the life cycle, such as life span, fecundity, time or duration of breeding
season etc may differ in closely related species. Niche specificity is of importance in the
taxonomy of such groups as birds, mammals, and particularly substrate specific animals)
e.g. host=specific plant feeders, and parasites).
Many new species of insects were discovered by comparing the populations of the ‘same’
species occurring on different plant hosts. This principle was carried too far by some
taxonomist who described many new species that were later find out to be synonyms. It
has also been observed that closely related animals may harbour similar parasites, and
it is also known that these parasites evolved parallel with their host, although they are
more conservative (i.e. Rate of change slower than in the host). Further evidence in man
(Homo) and the African apes (pan) is the presence of similar external and internal
parasite.
It is also possible to separate sibling species on the basis of the parasites, which they
harbour. Ecological differences were used in the separation of sibling species in the
Anopheles maculipennis complex.

4.5 ETHOLOGICAL CHARACTERS

These relate to behaviour, which is one of the most important sources of taxonomic
characters. These characters are often superior to morphological characters in the study
of closely related species, particularly sibling species. However, behaviour as a taxonomic
character, suffers two major technical drawbacks.
- Behaviour cannot be studied in preserved specimens.
- It is intermittent in even the living specimen i.e. occur only at certain time of the year/day
etc.
It is known that certain behavioural characters of animals can serve as isolating
mechanisms in the taxonomic study of groups like the bees, wasps, Orthopterans, Frogs,
fishes, and other groups.

What are these ethological behaviours?

i. Courtship behaviour: Different courtship patterns exist for different groups of animals
some of which are stereotyped behaviours and depend on sign stimuli. The male usually
sets up the pattern and the female will respond only if it is of the same species. Courtship
pattern is species-specific and represent the most important isolating mechanism in
animals.
ii. Sound production (calls and songs). (acoustic behaviour)
Birds have different patterns of songs and frogs and toads have different patterns of calls.
The development of sound-recording devices and sonagraphs for the translation of
sounds into graphic patterns has helped to advance the study of behaviour.
iii. Other types of ethological behaviour.
a. We construction in spiders, mites, and caterpillars may be used at various level in the
classification.
b. Nest construction: This has been used in the identification of bees, caddisfly larvae,
bag worms and birds. The two bee genera Anthidium and Dianthiduim could not be
recognized easily using morphological characters, yet all known species of the former
use cottony plant fibers in nest construction and those of the latter construct theirs from
resinous plant exudations (resins exuded from plants) and sand or small pebbles.

c. Egg cases: These have a species-specific form in the praying mantids. At the level of
the higher taxa, less attention has been paid to the diagnostic value of ethological
characters in the study of systematics.

E. Geographical characters
Geographical characters have been most useful in testing taxonomic hypothesis and as
tools for clarifying confused taxonomic pictures. These characters are to some extend
related to ecological features and they have both been employed in most sound
classifications.
Taxonomically, the most important kinds of characters are
1. General biogeographic patterns i.e. distribution of faunas, flora and their fossil forms.
2. The allopatric-sympatric relationship, which determines whether or not two populations
are conspecific.
The studies of the distribution of plants and animals have revealed well defined
geographic patterns. Based upon generalized comparisons of faunas and floras,
biogeographers have divided the world into various regions, provinces, sub provinces etc.
These represent distribution centers which exist today or have existed in the past. The
units of distribution are not static, and it therefore becomes more convenient to refer to
them as faunas, floras, or biota’s rather than as zones or areas.
This observation is based on the fact that related organisms with similar requirements will
be found in the same geographical locality. The close association between the organisms
could also be due to the fact that they descended from a common ancestor.
It is important for the taxonomist to know the geographic location of specimen, the type
of habitat and time of collection. Such information will permit a much sounder
interpretation of various higher taxa.
CHAPTER 6 VARIATION

4.6 QUALITATIVE AND QUANTITATIVE ANALYSIS OF VARIATION

4.6.1 QUALITATIVE:
The whole idea of systematics is based on diversity and diversity is in turn based on
variation. Samples obtained by taxonomists are sorted into phena and these are then
sorted out into species. However, the latter poses a problem as regards the establishment
of the species owing to the great variability of the species. Evidence exists from the long
list of synonyms in many groups. Most classifications are based on similarities. Beyond
similarities, other factors like population characteristics must be considered.
First is the continuity of the genotype from the fertilized egg through all larval stages to
the adult.
Secondly, a population must be a sexually reproducing unit. The knowledge of these
biological phenomena permits the taxonomist to assign phena correctly to species.

Criteria for sampling and assigning phena to species

1. Locality. Do the samples come from the same at different locality? If they are from the
same locality, there are two possibilities:
a. They could be individual variants (different forms) of a single species or they could be
several good species. If the samples came from different localities, there is a third
possibility:
b. They are different sub species of a geographically variable species.
2. Second criterion is to establish the existence of reproductive isolation. This
becomes a difficult task when handling collected materials.
4.6.2 MAJOR TYPES OF VARIATION WITHIN A SINGLE POPULATION
There are two types of variation-the nongenetic and genetic variation.
A. NON-GENETIC VARIATION
This adapts the individual to the environment. It can be discussed according to this
order:
1. Individual variation: this can take the following forms:
1.1 Age Variation
Normally development takes the form of egg→ larva/juveniles → adult, which are all
different.
a. No larval forms/stages
These groups have juvenile stages of course E.g. Reptiles, birds and mammals. The
juvenile stages have features which differ from the adult ones. In Agama adult (reptile),
the head has dark orange colour, and the juvenile is brownish. In birds, the juvenile
feathers are different from those of the adults. In mammals, there are difference in the
hair type and cover in the juveniles and adults. To a limited extent the fish has juvenile
forms different from the adult. In the Anguilla eels, the juvenile forms were formally
called a different genus leplocephalus, until recently when the two were discovered to
belong to the same genus Anguilla. Variation is very important taxonomic character
because without the knowledge, most genera would be confused.

b. Larval stages: E.g. caterpillars of butterflies, tadpoles of frogs, larvae of

echinoderms (Trocophore), molluscan larvae, larval of stages parasites (especially in
groups when the different stages occur on different hosts) create taxonomic problems.

c. Young and old adults: For e.g. in the male deer, the antler divides as the animal
grows. The nos of points of the antler will be different in the young and old deer-the
older the dear, the more numerous the antler points.

1.2 Seasonal variation of the same individuals. There are also changes in relation
to time of the year. This is particularly so in birds where the feathers are more colorful
in the breeding season. Also, in the arctic and sub arctic birds and mammals where
they tend to have light/white coloured feather or hair (winter dress) during winter, this
colour action serves for camouflage. They usually return from the Cryptic winter dress
to a ‘normally’ coloured summer dress. These changes may be caused by the
influence of hormones. Failure to recognize these seasonal changes can lead to
misidentification of some of the groups.

1.3 Seasonal variation of consecutive generations. Many species of short-lived

invertebrates (especially insects) produce several generations in the course of a year,
and individuals of different generation may be different. In such species, the
individuals which hatch in the cool spring may be different from those produced in the
summer, or the dry-season individuals may be different from the wet-season
population.
Cyclomorphosis is a special kind of seasonal variation found in certain freshwater
organism (e.g. rotifers and cladocerans). The populations of a species undergo a cycle
of morphological changes through the seasons in connection with changes in the
temperature, turbulence and other physical and chemical properties of the water. In
the genus Daphine (Cladocera) many ‘species’ have been named, that are nothing
but mere seasonal variants. In rotifers, different kinds of food may produce different
morphs (e.g. in Keratella). Failure to recognize these seasonal changes may lead to
taxonomic confusion.

2. Variation in social groups (social variation) e.g of social insects-bees and wasps
especially ants and termites. These social insects have castes which are made up of
the reproductive (Queens, and males or drones), workers and soldiers which all have
different morphological features. In case of Hymenoptera, the castes are usually
modified, genetically, identical females, whereas in isoptera, may involve both male
and females. The different morphological types may result from different larval food
and also be due to hormonal influence or other controlling factors.

3. Ecological variation
3.1 Habitat variation (or ecophenotypic variation) populations of a single species
occurring in different habitats may show difference in their morphology. These can
rightly be described as ecological races or ecophonotypes. For examples in molluscs
(freshwater snails and mussels), habitat forms (ecophenotype) are common. The
head stream with cooler temperature wells and rapid current) may have different form
from the lower reaches with warmer and more stagnant waters. In the limestone areas,
the shells of molluscs are heavy and of a different shape from those which grow in
waters with low calcium (lime) content. These show the dependence of certain
taxonomic characters on environmental factors. Examples are also found in the size
of fishes of the same species which grow in different habitats.

3.2 Variation induced by Temporary Climatic Conditions

Unusual climatic conditions in a given year (e.g. drought, cold etc.) may cause some
animals to produce year classes that differ visibly from the norm.

3.3 Host-determined Variation

Most variations in parasites of plants and animals are host-determined, and they
provide a potential source of error. These variations are most commonly expressed in
size difference but may involve other morphological/physiological characters. E.g in
the parasite wasp, Trichogramma semblidis Aurivillius, the males tend to be wingless
and otherwise modified when they develop in the eggs of the older fly, Sialiis lutaria
(Fabricius) (megaloptera) but not when reared from Lepidopterous hosts.

3.4 Density dependent variation

The effects of crowding may be reflected in morphological variation, especially when
crowding leads to shortage of food materials. In overcrowded condition organisms
tend to be smaller than under low population condition. E.g is the development of the
locust nymphs under various unstable conditions. Under crowded condition, newly
hatched nymphs develop in to gregarious phase. Under less crowded conditions, into
the transitional phase, and when isolated and released separately into the solitary
phase.

3.5 Allometric variation

This is simply an asymmetrical growth which leads to disproportionate size of some
structures in relation to that of the rest of the body. If individuals of a population show
allometric, then animals of different size will show allometric or heterogonic variability.
E.g in insects, it involves such features as the heads of ants or the mandibles of stage
beetles which may grow out of proportion to the rest of the body.
Failure to recognize the nature of such variations has resulted in much synonymy. The
exact causes are unknown. It is a form of age variation in species with continuous
growth, and in some cases, it is genetic in those insects (holometalolic) where the
phenomenon is common, it is correlated with sizewhich in turn may be due to
variation in food supply.

3.6 Skin colour changes (Neurogenic or Neurohumoral)

The colour of the organism may change in relation to the background or environment.
E.g. in crustaceans, cephalopods, fishes, amphibians and reptiles. Essentially the skin
colour is due to the activity to chromatophore present in the skin. In a variety of cases,
there is a darkening of the skin colour in response to dark background. Such are
hormonally induced. In other cases, these is a permanent darkening of the colour,
where this darkening involves the deposition of the black pigment in the pores of the
skin, the colour is more permanent and the condition is described as melanogenesis.
The aforementioned are not genetic in that a change of habitat causes loss of the
particular character.

4. Traumatic Variation
4.1 Parasite induced variation
In addition to such familiar effects of parasitism as swelling, distortion, and mechanical
injury, parasites may produce conspicuous modification of structures. Some strikingly
different termite soldiers were assigned to a new genus and species, Gnathotermes
aurivillii. It was shown later that these modified soldiers are nothing but parasitized
individuals from colonies of macrotermes malaccensis.

4.2 Accidental Variation

This is usually externally induced, in most cases, such changes are easily recognized
but in those forms which undergo metamorphosis, injuries to an earlier stage may
produce later abnormalities which are not so easily recognized as such. This is
especially true when these anomalies involve major taxonomic characters in the group
in question.

5. Post-mortem changes
Post-mortem changes occur in many groups of animals during preservation. E.g in
birds, the deep orange-yellow plumes of seleacidis ignotus (twelve-wire birds of
paradise) fade to white in collections. Many species have been wrongly labelled as
synonyms owing to the comparison of freshly collected material with old museum
specimens.

B. GENETIC VARIATION
This is due primarily to differences in genetic constitutions. It adapts the population
and species to the environment. This variation is divided arbitrarily into two classes.
1. Sex associated variation
These may be sex-limited i.e. expressing themselves in one sex only or associated
with one or the other sex, or they may involve sex characters or mode of reproduction.

1.1 Primary sex-differences

These relate to differ in primary sex organs utilized in reproduction (e.g gonads,
genitalia etc). Where the two sexes are quite similar, this rarely constitute taxonomic
confusion. E.g. boundary and interstitial cells = male gonads of some fishes and
amphibians, 2 claspers may vary in structure in Elasmobranchs.

1.2 Secondary sex differences

This relates to sexual dimorphism. In most species, the differences between males
and females are often very striking. E.g. birds of paradise, duck etc different sexes
were mistakenly called different species. In the king parrot, Eclectus roratus, the male
is green with orange bill which the female is red and blue with a black bill. The two
sexes were called different species for almost 100years until recently disproved.

1.3 Difference in alternating generations

This is particularly related to Aphids where there is alternation of generation between
the Pathenogenetic and sexual forms. This pathenogenetic wingless females differ
from the winged females of the sexual generations.

2. Non-Sex associated individual variation

This does not usually involve sex characters.
2.1 Continuous Variation
This is very common and usually results from slight genetic differences between
individuals. No two individuals in a population of sexually reproducing organism are
exactly alike, genetically and morphologically, except monozygotic twins. These
differences are slight and are not easily discovered. They are usually polygenic-
controlled by several genes. E.g difference-height, weight, colour. Since these
varieties are gradual and since no one individual is typical of the characters of a
population, it is wise to have as wide a sample as possible from the population to
determine the extent of variability.

Discontinuous variation (polymorphism).

Difference between individual in a population are in general slight, but in certain spp,
members of a population can be grouped into very definite classes, due to the
presence of certain conspicuous characters. Such discontinuous variety is called
polymorphism. Such characters are determined by a single gene subject to mendelian
inheritance. E.g. in Trypanosomes, social insects etc, spotting in lady beetles, and
industrial melanism in moths. It is of great biological importance because it proves the
existence of selective differences between apparently neutral characters. Its practical
importance to the taxonomist is that it has led to the description of many
pseudospecies that are actually polymorphic variants.

6.7 ZOOLOGICAL OR BIOLOGICAL NOMENCLATURE

The world is inhabited by a variety of plants and animals. About 250,000 known
species of higher plants and between 5 and 10 million species of animals are known.
This organic diversity arose as a result of evolutionary divergence. There is therefore
strong need for the ordering of organisms and naming of varieties or types or species
which are reflected in classification methods.
Nomenclature refers to the naming of organisms. Essentially names are a means of
reference and principally a means of communication. Applying this is the use of
scientific names to organisms and it constitutes a special language in the scientific
literature. The binomial (binomen’s = two names) system of nomenclature is used in
the naming of plants and animals. Under this system, the name of a species consists
of two words, respecting the generic and the specific names.
The generic name begins with a capital letter while the specific name begins with a
small letter. The two names are always underlined or written in italics when in prints.
Hence the name Homo sapiens for man Canis familiaris for Dog, and Khaga ivorensis
for the Iroko tree. The system of communication has certain objectives:
(a) Uniqueness: Every name has to be unique because it is the key to he entire
literature relating to this species. Every name gives immediate access to all the known
information about a particular species as taxon.
(b) Universality: There is a need for a common language and this calls for some
code, which will be observed by the users on worldwide basis. Local names are of
limited use for communication since the same organism may have different names,
reflecting the multiplicity of languages.
(c) Stability: Scientific names must not be changed frequently or arbitrarily. To
preserve these objectives, there is a set of code or rule described as the international
code of Zoological Nomenclature or Botanical Nomenclature as the case maybe. The
rules are mandatory but without any legal status in national or international law.
Enforcement depends on voluntary agreement of scientist. The language used in
nomenclature is Latin in form, written in Latin alphabets and subject to the rules of
Latin grammar.