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Each Lichen is a miniature ecosystem, consisting of a fungus and one or more photosynthetic partners. Lichenologist William Purvis explores their unusual biology; amazing diversity and ecological importance. He explains how the rewards of understanding lichen biodiversity are great and may lead to technological developments in medicine, metal prospecting and pollution control.
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100% found this document useful (9 votes)
2K viewsLichens
Each Lichen is a miniature ecosystem, consisting of a fungus and one or more photosynthetic partners. Lichenologist William Purvis explores their unusual biology; amazing diversity and ecological importance. He explains how the rewards of understanding lichen biodiversity are great and may lead to technological developments in medicine, metal prospecting and pollution control.
Uploaded by
flocky23905Copyright
© Attribution Non-Commercial (BY-NC)
Available Formats
Download as PDF, TXT or read online on Scribd
/ 113
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9 Lichens
William Purvis
From rocky coasts and urban churchyards to rain forests and!
ountain summits, lichens are a familiar part of the landscape
Fach lichen isa miniature ecosystem, consisting of a fungus
and one or more photosynitietic partners. This delicate balance
‘between partners is easily upset by environmental disturbance,
providing, us with practical tools to assess the effects of air
pollution, radioactive falltout and climate change.
Lichenologise William Purvis explores their unusual biology,
amazing diversity and ecological importance. He explains
how therewards of understanding lichen Biodiversity are
Great and may lend) x0 technological developments in medicine,
mecal prospecting and pollution control, This unique book
includes new information on economic uses and outlines
practical project ideas. Packed with stunning photoeraphs
Tichens reveals the varied and intriguing, world of these
fascinating organisms.
The Life Series
‘The Natural History Muscum’s Life Series explores the
parlorama of the natural world. Fach book covers a single
pplant or animal group, or ecosystem. The series presents an
accessible introduction to the wonders of the natural world
suitable for all narure enthusiasts Lichens
William Purvis
PUBLISHED ay
“THe Naural History Museum, LONDON Dedicated to Peter James who has done so
much to stimulate lichenology worldwide
First published by The Natural History Museum,
Cromwell Road, London SW7 SBD
© The Natural History Museum, London, 2000
ISBN 0-565-09153-0
All rights reserved. No part of this publi
bbe transmitted in any form or by any means
prior permission from the British publisher.
tion may
chou
A catalogue record for this book is available from
the Bricish Library
Edited by Jacqui Morris,
Designed by David Ro
Reproduction and printing by Craft Print, Singapore
DisrrmuTioN
North America, South America,
Central America, and the Caribbean
Smithsonian Institution Press
470 L'Enfant Plaza
‘Washington, D.C. 20560-0950
USA
Australia and New Zealand
CSIRO Publishing
PO Box 1139
Collingwood, Vietoria 3066
Australia
UK and rest of the world
Plymbridge Distributors Ltd.
Plymbridge House, Estover Road
Plymouth, Devon PL6 7PY
UK Contents
Preface
What is a lichen?
How lichens grow, multiply and disperse
Lichen biodiversity
Evolution, classification and naming
Ecological role
Lichens in forests
Lichens in extreme environments
Biomonitoring
Prospecting and dating
Economic uses
Practical projects
Glossary
Index
Further information
Acknowledgements
19
33
46
49
56
62
76
88
92
99
108
109
11
112 Preface
This book explores the fascinating world
of lichens. Lichens occur almost everywhere
and the lichen symbiosis is one of the most
successful examples of two or more
organisms living together. How lichens are
formed is one of the greacese puzzles in
biology. Just as salt is dissimilar to sodium
‘and chlorine, so is a lichen dissimilar to the
‘organisms that create it. Lichen morphology,
physiology and biochemistry are very
different to chat of the isolated fungus and
alga in culture, Lichens occur in some of the
most extreme environments on Earth
tundra, hor deserts, rain forests, rocky coasts
and toxic spoil heaps. Many of these
environments are also among the most
sensitive to destruction and pollution, Our
etic
The author
William Purvis geaduated in botany from the
University of Sheffield in 1981 and was
awarded a Natural History Museum (NHM)
Centenary studentship funded by British
Pecroleum ar the Royal School of Mines,
Imperial College and Botany Department,
The Natural History Museum, London. He
took up his post as a lichenologist at The
Natural History Museum in 1988, He is
rich and varied lichen heritage is incredibly
useful in allowing us to assess what we are
doing to our environment.
Lichenology is at an exciting stage
of development, attracting great interest
from biologists, chemists, geologists,
ceavironmental scientists and consultants,
and industry. Lichens are used successfully
as bioindicators of air quality by school
children and students throughout the
world. The rewards of understanding
lichen biodiversity are great and in the
future may lead co important technological
developments, with the discovery of new
‘medicines and the clean-up of contaminated
sites, as well as monitoring the health of
principal author of the Lichen Flora of
Great Britain and Ireland and bas carried
‘out pioneering research into why different
lichen species accumulate metals, and
the implications for biomonitoring and
classification. He is actively involved in using
lichens as bioindicators of enviconmental
health and is carrying out taxonomic research
on critical groups. Many people think that lichens are simple
organisms rather like mosses. In fac, they
are mini-ecosystems, consisting of atleast
two organisms: a fungus (mycobiont) and a
photosynthetic partner (photobiont). The
photobiont, which contains chlorophyll, may
be cither a green alga or belong to an entirely
different kingdom —a cyanobacterium
(a bacterium that contains a blue-green
photosynthetic pigment). Even where only
two partners are present, the association or
symbiosis is not usually a simple mixture,
Instead, it results in a distinct lichen body
Lit Lchen-eavered fence aaove Boulders with
post, with leafy orange ‘lichens (Steeocaufon
anthoria sp shrubby sp. and reindeer icens,
Usnea sp. (green thal) __Clodania spp.) mosses
and Hypogymmia sp. (Rhacomitium, Andreaea
(pale-groy, lft) and a and Plytrichum spp.)
crustke Thelomma sp. and shrub. Pare Grandes
(laclcand-wnite, op). Jardins, Laurens,
Coastal hills of Norhorn Quebec.
Califia, hati a iehen?
‘ae Section through
Heppia adgutinata
showing algal cals (rom
Schwendener 1869).
(thallus) being formed. It looks very different
from the individual partners when they are
separated and grown in laboratory cultures.
‘The names we call lichens refer to the fungal
partner because every lichen has a unique
fangus. The algae and cyanobacteria have
their own names and most of them occur in
many different lichens.
Fungi, and algae and/or cyanobacteria
associate in many different ways in lichens.
‘At one end of the scale fungal filaments and
algae coexist, as in the tropical lichen genus
Coenogonium (see p. 45). At the other end of
the seale are the majority of lichens, which
are highly organized and, without derailed
examination, appear to be a single entity. The
International Association of Lichenology in
1982 defined a lichen as an “association of a
fangus and a photosynthetic symbiont
resulting in a stable thallus of specific
structure”. However, where several partners
‘occur in the same thallus, the precise role
played by each individual partner is by no
‘means clear
An intriguing question is whether the
mycobiont and the photobiont benefit from
the association, Many biologists consider
lichens to be one of the finest examples of
symbiosis (two or more organisms living
together) because lichens are widespread and
the algal cells appear to be healthy inside the
lichens. Others believe that the lichen
telationship is one of controlled parasitism,
that the photobiont cells are victims rather
than partners of the mycobiont. Simon
Schwendener, a German, was the First 0
discover the dual nature of lichens. In 1869,
he wrote: As the result of my researches, the lichens
are not simple plants, not ordinary
individuals in the ordinary sense of the
words they are, rather, colonies, which
consist of hundreds of thousands of
individuals, of which, however, one alone
plays the master, while the rest, forever
imprisoned, prepare the nutriment for
themselves and their master, This fungus
is a fungus of the class Ascomycetes, a
parasite which is accustomed to live
upon others’ work. Its slaves are green
algae, which it has sought out, or indeed
caught hold of, and compelled into its
service. Ir surrounds chem, as a spider
its prey, with a fibrous nec of narrow
meshes, which is gradually converted
into an impenetrable covering, but while
the spider sucks its prey and leaves it
dead, the fangus incites the algae found
in its net to more rapid activity, even to
‘more vigorous increase.
Schwendener’s views were quite revolutionary
for the rime, and his dual hypothesis, as ic
was called, was not accepred universally,
Indeed, it was rejected with scorn by some
of the most prominent lichenologists of the
errs the lichen thallus
a contralid frm of
paras?
1M, myeobiont;
,photiont.
What i a lichen?
day. One of these, the Scottish Reverend
James Crombie, described it as “a romance
of lichenology, or the unnatural union
between a captive algal damsel and tyrant
fangal master”, It is easy to see how the
lichen fungus benefits from the association ~
the photobionts photosynthesize and provide
icwith carbohydrate. This is in the form
of sugar alcohols in the case of lichens
containing green algae and glucose in lichens
containing cyanobacteria. The mycobiont
may also obrain nitrogen compounds when
the phocobiont is a cyanobacterium because
cyanobacteria can capture and fix nitrogen
from the atmosphere.
We do not have any evidence that the
fungus passes any nutrients to the phorobiont,
but it does modify che algae and cyanobacteria
that it contains. The photobionts lose their
cell walls and do not reproduce sexually.
However, both partners share one benefit.
Together they are able to colonize many
habitats that would otherwise be ou of their
reach. The fungus can live in places lacking
the organic matter that they would normally
need a$ a source of nutrients. Algae and
‘cyanobacteria, which usually live in aquatic
cor moist habitats, can live in drier places.
They can also be affected adversely by high
light intensity and, given the protection of the
fangus, they can expand into environments
where light intensity is high.
Symbiosis is a very successful way of
living, but itis not unique to lichens.
Other examples are
+ Mycorthizas — associations of fungi with
higher plants, in which ehe fungus lives on
the plant ross. What is a ener?
+ Actinorshizas ~ where bacteria are
associated with the coots of leguminous
plants.
* Feeshwater and marine invertebrates —
‘Hydra and some other coelenterates include
algae in their bodies, as do some flatworms.
Lichen fungi
Fungi, unlike green plants, contain no
chlorophyll and cannot produce their own
carbohydrates. They have developed various
‘ways of obtaining organic carbon compounds
from both living and dead sources
(heterotrophy). Fungi form many different
symbiotic celaionships with vascular plants
(mycorrhizas). For instance, fungi may be
associated with forest tees, where they assist
in the nutrition of their host. The process of
lichenization, involving obtaining carbon
from photobiont cells, is a common mode of
fangal nutrition; around 20% of all Fungi are
lichenized. Many lichen fungi produce special
microscopic branches (haustoria), which
penetrate che photobiont. The lichen-forming
fungi are a very mixed group. Some belong to
taxonomic groups in which all the members
are lichenized, while other groups may
contain non-lichenized relatives. By far the
largest proportion of lichen-forming fungi
(over 40%) belong to the ascomycetes, whieh
produce spores in a sac-shaped cell (ascus).
Ti nacure lichen partners exist only in
symbiosis, apart from as a reproductive
spore. In laboratory conditions, however, itis
possible to take the lichen apart and grow the
two partners separately. Scientists in Europe,
er Orange-pee fungus,
‘Aleuria auranti—a non
lichenied ascemyeste
fungus. Epping Forest,
London, UK. Japan and North America have made
significant progress in culturing the fungus
from spores or extracts of fungal eels. In
pure culture, lichen mycobionts form slow-
‘growing amorphous colonies, which are quite
different in appearance from the whole
(lichenized) thallus. It is also possible to
culture some lichens and to synthesize
new ones by inoculating spores with algae,
but they are difficule co maintain for long
periods. Suecess depends partly on avoiding
contaminant organisms killing the lichen and
also on creating the right conditions, which
What is a fichen?
do not favour one or other ofthe symbionts.
Solid (agar) culcure media are beeter for
‘growing lichens than liquid media
Lichen algae
Only a few cypes of algae occur within
lichens. Algae can be the dominant partner,
eg: in the ‘ely lichens’, Collemaa spp. and
Leptogivns spp, but this is rare. The algae
found most frequently are the unicellular
green algae ofthe genus Trebotsia, Other
common species belong to the orange-
Pigmented genus Trentepoblia and the
er anthorisparietina: aoe The fungus of
the orange, aisle Xantheriaparetna in
‘rating bodies ofthis culture Its amorphous
lichen are superfclaly and loaks quite uke the
Similar to those of the infact chen. Lichen fun
fungus Alewris aurania. are nat known to profiace
ftuiting bodies in culture, What i a iehen?
cyanobacterium (formerly called blue-geeen
alga) genus Nostoc. Lichen photobionts look
rather differene when they are inside rather
than outside lichens. Sexual reproduction and
the formation of filaments is suppressed and
wwe usually need to isolate and culture lichen
algae ro identify them to species level. This
has been achieved successfully for less than
2% of all lichen photobionts. Around 100
green algae (occurring in about 23 genera)
and many fewer cyanobacterial species in
about 15 genera may be involved as
symbionts. Most lichen fungi are highly
selective, forming associations only wich
compatible photobiont species. Some
notable exceptions include:
* The same lichen in different geographical
locations may contain different photobionts.
# Some lichen species contain different
photobionts during different stages of their
life cycles.
* Another fungus growing on a lichen may
share the alga of its host and later swap it for
another species (e.g. Diploschistes muscorum
‘growing on Cladonia spp.).
Some lichens contain both algae and
cyanobacteria within the thallus. The main
photobiont is normally the green alga, and
cyanobacteria are localized in distinct
structures (cephalodia). This has given rise to
cone of the greatest enigmas in lichen biology
thar the same lichen can exist in several very
‘Roi Teboula sp. 2
uniceliular green alga.
‘The algae found most
frequently in lichens
belong othe genus
Trebousia. ‘What is leben?
Fr Trentepoia =p.
Green algae ofthis
‘genus contain an orange
cearotenold pigment,
which masks the green
colour af ehlarophyl
‘Many lichens in shaded.
habitats and mest lichens
In lowland troplea ain
forests contain
Trntepotiia.
se. Chains of
Nostoe sp. —
a cyancbacterium.
‘Members ofthis genus
oceurfrequenty in
lichens, especialy in
thot of wet habitats, ‘What a lehen
rigut Steta eanariensis —
leatsike form containing
‘green algae, with large
red-brown, dise-ke
‘rating boces. different forms, depending on the photobiont
it contains. These different forms of the same
lichen ate called photomorphs.
Photomorphs
‘One of the most important lichenological
discoveries in the 20th century was that
lichen algae play a major part in determining,
what a lichen looks like, how it functions and
where it grows, Pioneering research was
cattied out by the Englishman, Peter James,
and colleagues abroad. James discovered
strange thalli in sheltered gorges in New
‘Zealand and Scotland in which different
lobes belonged to different species. For
instanes, in New Zealand he found the leafy
lichen Sticta felix (containing green algae)
growing out of the shrubby Dendriscocanlon
species (containing cyanobacteria). He also
found separate individuals of both lichens
and realized that environmental factors (light,
humidity) are, intriguingly, very important in
determining which photobiont is present in
the lichen. Because all lichen names are based
‘on the fungus alone, a single name has to be
used for what were previously considered co
be two lichens. Many examples are now
known, especially within the leaf-like lichens
of the genera Nephroma, Sticta and Lobaria.
In these groups, the same fungus may be
associated with either a green alga and
appear leaf-like or with a cyanobacterium,
when it usually looks very different, often
appearing shrub-like or as tiny granules.
‘The different phorobionts can either occur
together in the same thallus or separately
Such an example is the lichen, Sticta
