Lichens

DC-1, Semester-2
Paper-3: Mycology and Phytopathology
Lesson: Lichens
Lesson Developer: Dr. Surinder Kaur Walia and Dr. Charu Kalra
College/ Department: Sri Guru Tegh Bahadur Khalsa
College/Department of Botany and Deen Dayal Upadhyaya
College/ Department of Botany, University of Delhi

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Table of Contents

Chapter: Lichens

 Introduction
 General Structure
 Different types of associations
 The Lichen Partners
 Lichenicolous fungi
 Parasymbiosis
 Lichenicolous fungi
 Nature of Association

 Nomenclature
 Distribution of Lichens
 Ecology of Lichens

 Classification of Lichens on the basis of habitat

 Comparative view of Lichens

 Classification of lichens based on morphology of the

Lichen Thallus
 Endolithic Lichens
 Epilithic Lichens
 Different types of epilithic lichens
 Crustose lichens
 Foliose lichens
 Fruticose lichens
 Squamulose or placodioid lichens
 Filamentous lichens
 Leprose lichens
 Intermediate forms

 Structure of the lichen thallus

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 Comparative view of homiomerous and

heteromerous thallus
 Structure of heteromerous lichen thallus
 Growth of lichen thallus

 Reproduction
 Reproduction involving both partners
 Fragmetation
 Soredia
 Isidia
 Squamules
 Reproduction involving only the mycobiont
 Pycnidia
 Apothecia
 Types of apothecia
 Leceidine type
 Lecanorine type
 Perthecia
 Some peculiar structures of lichens
 Cephalodia
 Cyphellae and Pseudocyphellae

 Dispersal
 Chemical composition of Lichens
 Primary metabolites
 Secondary metabolites
 Reasons for different coloration in lichens

 Economic Importance of lichens

 Human uses of Lichens
 Lichens for perfume
 Lichens for dye
 Lichens for medicine
 Antibiotic properties
 Herbal remedies
 Lichens for food
 Problems often encountered with eating
lichens

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 Miscellaneous uses
 Sources for nest insulation
 Nitrogen fixation
 Pollution indicators

 Lichenometry and its uses

 Common names of some lichens
 Exercises
 Key to Exercises
 Glossary
 References
 Web Links

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Introduction

The word lichen is derived from a Greek word (Gr leikhēn = to lick). In Latin līchēn, means
a kind of plant. These are unique organisms in many ways. Lichens represent an intimate
and long-term symbiosis of algae and fungi. The fungus partner is called mycobiont and a
photosynthetic algal partner, the photobiont (earlier known as phycobiont). A stable
thallus with a distinct morphology is formed when two (or more) partners come together,
leading to a long-term symbiotic relationship. But usually, in many lichens, this association
breaks down, with the partners re-establishing the relationship from the separately
dispersed fungal spores and photosynthetic cells.

General Structure

The fungal partner usually contributes more to the structure and the algal partner produces
food for the lichen by photosynthesis. As the external surface is formed by fungal
partner it is knowns as the exhabitant, whereas, the unicellular or filamentous
photobiont cells are termed as inhabitants as located inside the lichen thallus.

Source: [Link]

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Table: Different types of associations

[Link]. Type Photobiont Mycobiont Special feature Example(s)

of lichen
1. One-membered One One Typical ?
2. Three-membered One Two Presence of Pestalotiopsis
Parasymbiotic maculans
fungus
3. Three-membered Two One Presence of Peltigera
Cephalodia leucophlebia,
Lobaria
pulmonaria
4. Four-membered Three One Presence of two ?
different
photobionts in
Cephalodia
5. Four-membered Two Two Presence of Dimelaena
Lichenicolous lichenicola
Lichen

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Figure: Types of associations in lichens (a) Two membered 1 photobiont+1 mycobiont

Source:author

The Lichen Partners

There are more than 13,500 species of lichens described to date. There are one hundred
fifty species belonging to 40 genera of photobionts, are known which are usually either
green algae or cyanobacteria.

Did you think?

How only 150 species (40 genera) of photobionts can take the number of lichens
more than 13,500?

The reason is; a single species of photobiont may have multiple fungal partners.e.g. all
species of Cladonia have photobionts in a single green algal genus Trebouxia.

Most common green algal partners in lichens are:

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PHOTOBIONTS MYCOBIONTS
Algal Partners Cyanobacterial Partners Fungal Partners
(in case of three
organisms’ symbiosis)
Cephaleuros Calothrix Chroococcus  Hymenoascomycetes
Coccomyxa Gloeocapsa  Discomycetes (Discolichens)
Myremesia Nostoc  Lecanorales (inoperculate
Trentepohlia Scytonema Stigonema Discomycetes)
Trebouxia
N.B. Basidiomycetes do not readily
form lichens (reason yet not
identified). e.g. Cora and
Dictyonema.

A: Trebouxia B: Trentepohlia

Source: A. [Link]

B.[Link]
[Link]

Lichenicolous fungi- These are formed when a fungus dwells on or in lichens as parasites
(pathogens), commensals or saprobes. These are also known as lichen-inhabiting fungi. So
it represents association of two fungi with a single photobiont. There are about 300
genera and 1000 species of obligately lichenicolous fungi now known.

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These are known to

 induce galls on the thallus (e.g. Polycoccum galligenum),

 occur as saprophytes on dead or decaying lichen thalli (e.g. Niesslia
cladoniicola),
 occur as parasites causing necrotic patches (e.g. Lichenoconium lecanarae)
killing parts of thalli.

Parasymbiosis- It is a stable situation of Lichenicolous fungi in which the host lichen

is totally unaffected from outward appearances, thus representing symbiotic association
with an already existing symbiosis e.g. Arthonia glaucomaria on Lecanora rupicola.

Development of wart-like structures termed cephalodia (sing. cephalodium. Gr. kephale =

head) occurs when a single mycobiont forms an association with two or rarely three
different photobionts. These outgrowths form integral part of the lichen.

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Figure: Cephalodia are pockets of cyanobacteria. Cephalodia either grow on the outside
surface of the thallus lobe; on the outer surface of a pseudopodetium; or inside the thallus
between the outer fungal layers. Cephalodia only occur in lichens that have a green alga as
a primary algae in the main thallus. Cyanobacteria (also called blue green algae) are very
important to us all because they make the nitrogen in the air usable by plants.

Source: [Link]
[Link]

Lichenicolous lichens-These are formed when lichens grow on or in other lichens either as
commensals or parasites. The algae of the lichenicolous lichens occur inside the host (e.g.
Buellia on Physconia) or as discrete thalli on its surface (e.g. Caloplaca on several lichens).
These represent four-membered symbiosis. In these, each mycobiont has an
independent photobiont.

Milestones in Lichenology

Some of the important discoveries/contributions made in the study of lichens, known as

lichenology is as follows:

Table: Milestones in Lichenology

Year Investigator Significant Event

1729 Pier Antonio Micheli Published Nova Plantarum Genera, in

which he enumerated about 300
species within 38 ‘orders’, illustrated
asci for the first time from soredia.
1757- Erik Acharius Founder of systemic study of lichens,
1819 introduced many of the descriptive
terms and generic names now used,
and synthesized the extant world
literature, Methodus Lichenum, 1803;
Lichenographia universalis, 1810; and
Synopsis Methodica Lichenum, 1814.
1866 Anton de Bary Hinted about the dual nature of lichen
thallus.

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1867 Simon Schwendener A Swiss botanist, in 1867 explained the

dual nature of lichen thallus. He
announced to the scientific world his
hypothesis that lichen was formed by
two separate organisms, a fungus and
an alga. At the time his theory was
largely rejected, but afterwards proven
to be factual.
1877 DA Stahl Achieved the synthesis of lichen
experimentally for the first time.
1961 David C. Smith In work on physiological studies, he
demonstrated the transfer of carbon
between the lichen partners for the
first time.
1968 Oliver L. Gilbert Demonstrated the use of lichens in the
monitoring of pollutants; published
first zonal scale calibrated according to
sulphur dioxide levels.

1970 V Ahmadjian & H Accomplished first spore to spore

Heikkila synthesis of lichens with the lichen
Endocarpon pusillum.

Nature of Association

The relationship between the fungal partner and the photosynthetic partner of lichens has
been described in various ways.

 Some describe it as mutualism in which both partners are being benefitted.

The fungal partner benefits by getting carbohydrates, from the algae which being
green synthesizes them through photosynthesis. The photosynthetic partner gets
protection from dessication in adverse weather thus resulting in the algae having a
more stable and constant environment to live in allowing it to grow better.

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The fungi collect the carbohydrates by means of special hyphae called appressoria
or haustoria which contact the wall of the algal cells. The fungi may produce a
substance which increases the permeability of the algal cell walls so that they lose as
much as 80% of the sugars that they produce. The carbohydrates pass into the
fungal hyphae through diffusion.

Figure: Diagram showing appresorium and haustorium

[Link]
[Link]

 Alternatively, the association is considered to be a controlled form of parasitism

of the algal cells in which the fungal partner benefits and the photobiont
loses by showing controlled growth inside the thallus by the hyphae and are also
prevented from forming zoospores.

However, in some cases, the algal cells are killed sooner or later by the fungus or it
cannot survive alone in the habitat occupied by the lichen.

From an ecological perspective, the relationship is usually mutualistic because the

composite lichen thallus survives in more habitats than either partner when alone.

Nomenclature

 Because lichens are "dual organisms" composed of at least two separate species,
their nomenclature is not easy. According to recent rules lichens are formally
assigned the name based on their fungal partner. e.g. the common orange-yellow

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coloured lichen Xanthoria parietina growing on rocks in coastal areas, is classified as

a fungus (Xanthoria) which has Trebouxia, a green alga as photosynthetic partner.

Figure: Xanthoria parietina

Source:[Link]
[Link]

 Recent nomenclatural trend in fact considers that lichens are a nutritional and not a
taxonomic group.
 Earlier, some lichen taxonomists placed lichens in their own division, the
Mycophycophyta, but this practice is no longer accepted because the components
viz. fungal and algal, belong to separate lineages.

Distribution of Lichens

Lichens are spread over large geographical areas e.g. species of Parmelia sulcata, Xanthoria
elegans are worldwide in distribution. Moreover, their geographical range is more extended
than that of any other class of plants as from hot valleys of tropical climates, semiarid
deserts to the shores of arctic and Antarctic seas.

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Figur[Link] Anholt Desert showing patchwork of lichens and heaths; B: Antarctic lichen.

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Source:[Link]
Anholt_desert.jpg,[Link]

DID YOU KNOW?

 World’s richest areas appear to be Southern temperate rain forests, Northern
temperate forests, high latitude zones for lichen-forming fungal species but however
these are not present in humid tropics.
 The maximum lichen species have been reported from North Mexico followed by
Australia.
 Lecidea geographica occur on projecting rocks even far above the snow line.

The distribution of lichens over the surface of the globe is regulated, not only by the
presence of suitable substrata, but more especially by climatic conditions.

To assist their survival in such inhospitable conditions, lichens are able to shut
down metabolically during periods of unfavourable conditions and with the
appropriate amount of light and moisture, clean air, and freedom from
competition, lichens will continue to grow.

Ecology of Lichens

Because of their unique symbiotic association, lichens are known to be present in almost
every terrestrial habitat, capable of supporting photosynthesis. They compete with plants
for sunlight, space and other resources.

Major factors affecting the presence and distribution of lichens are the availability of light,
moisture, temperature and substratum.

Lichens do not have roots and do not need to tap continuous reservoirs of water like higher
plants, thus they can grow in locations impossible for most plants, growing on cooled lava
flows and bare rocks surfaces, sterile soil or sand, the woody bark of trees, wood, and
mosses.

They are poor competitors of vascular plants but they compete well with
bryophytes. Because of their small size and slow growth; they thrive in places where
higher plants have difficulty growing. They can produce a battery of more than 500 unique

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biochemical compounds that help them not only to discourage competition from plants but
also control light exposure, repel herbivores, and kill attacking microbes.

As a rule lichens grow commonly in open exposed habitats, though some are found
only or chiefly in shady situations; while, scarcely any occur where the atmosphere is
polluted with smoke. Many species also prefer growing in moist places by streams, lakes
and the sea.

Xanthoria is common in places enriched by manure e.g. dust from cattle yards or from
birds. So it becomes apparent that in many cases lichens are quite indifferent to the
substrata on which they grow.

Thus it can be inferred that the preference for a certain substrata depends upon the
temperature of the locality or that of the special habitat.

DID YOU KNOW?

Various lichens occur abnormally in some unexpected habitats like
 dried dung of sheep,
 bleached bones of reindeer and whales,
 old leather iron glass and
 dead organic matter in environment with low productivity.

Classification of Lichens on the basis of habitat:

They have been divided into the following categories on the basis of their habitat:

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Figure: Different habitats of Lichens

Source:[Link]
dri_shilapushpa/[Link]

Table: Comparative View of Lichens:

Corticoulous Saxicolous Terricolous Muscicolous

fungi /terrestrial/ar
boreal
Substrata Bark e.g. Parmelia rocks and stones, On different growing with /
(on old trees) and ancient monuments, dry types of soil e.g. on decayed
(grow on) Graphidei spp.(on stone walls, roof tiles Lecanora crassa mosses and
young trees) and farm buildings near on sandy soil. on the
the sea. ground, trees,
rocks or
amongst or
moss e.g.
Cladonia and
Peltigera spp.

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Mainly In large forests In Temperate and cold In all kinds of growing with /
found in (e.g. Usnea; zones of the world. soil, from the on decayed
sands of the sea- mosses
A few on trees Rare in tropics. shore to the
by roadsides granitic detritus
(e.g. Physcia of mountains.
parietina)
No spp. found
on cultivated
ground
Special Lichens growing on Divided into two groups
term/grou decaying wood of
p (if any) trees and on old  Calcicolous-

pales are known as growing on

lignicole (e.g. calcareous and

Lecidea) cretaceous rocks

e.g. Lecanora
calcarea
 Calcifugous.
Rest all belong
to this group.

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Figure: Corticolous fungi [Link] sulcata; B. Usnae scrabida

Source: [Link]
[Link]
[Link]

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Figure: Saxicolous fungi Lecanora allophana growing on rocks.

Source: [Link]

Figure: Folliocolous lichen Strigula spp. growing on leaf.

Source: [Link]

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Figure: Terricolous lichen Bryoria bicolor

Source: [Link]

Fact finder
 Some ground-dwelling lichens, such as members of genus Cladina rangiferina
(reindeer lichens), however, produce chemicals which leach into the soil and inhibit
the germination of plant seeds and growth of young plants.

Cladonia rangiferina

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Source: [Link]

 Terricolous lichens are often an important part of soil stabilization; indeed, in some
desert ecosystems, vascular (higher) plant seeds cannot become established except
in places where lichen crusts stabilize the sand and help retain water.

Recently another type has been elucidated:

Follicolous or Epiphyllous These are very peculiar as grow on perennial leaves of

certain trees and shrubs, whose vitality is not at all affected by their presence as it is
by that of fungi, e.g. Felhanera bouteillei, Lecidea and Strigula.

Figure: Fellhanera bouteillei a epiphyllous lichen.

Source: [Link]

Many lichens also grow as epiphytes (epi- on the surface, phyte- plant) on other
plants, particularly on the trunks and branches of trees. When growing on other
plants, lichens are not parasites; they do not consume any part of the plant nor
poison it.

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A major ecophysiological advantage of lichens is that they are

poikilohydric (poikilo= variable; hydric= relating to water), meaning that
though they have little control over the status of their hydration, they can
tolerate irregular and extended periods of severe desiccation.

Like some mosses, liverworts, ferns, and a few "resurrection plants",

upon desiccation, lichens enter a metabolic suspension or stasis
(cryptobiosis) in which the cells of the lichen symbionts are dehydrated to
a degree that halts most biochemical activity. In this cryptobiotic state,
lichens can survive wider extremes of temperature, radiation and drought in
the harsh environments they often inhabit.

Classification of lichens based on morphology of the Lichen thallus

On the basis of growth forms lichens are informally classified into:

1. Endolithic Lichens –
 Lichens growing inside substrates where rocks save algae from strong
sunlight, which otherwise can kill algal partner hence leading to the
death of lichens
 They are not visible with naked eye as they show high degree of
camouflage.
 With time these break down the rock they dwell in.
2. Epilithic Lichens –
 Lichens growing on substrate surfaces like insects, rocks, leaves,
bark, soil, under water and man-made substrates like glass.
 They are visible to the naked eye.

Different Types of Epilithic Lichens:

In epilithic lichens mycobionts and photobionts vary in their association patterns giving
rise to six morphologically distinguishable types or growth forms as discussed below can
associate in a variety of ways to form several different morphological.

 Crustose - crust-like
 Foliose - leaf-like

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 Fruticose - shrub or beard-like

 Squamulose or placodioid - scale-like
 Filamentous and
 Leprose

1. Crustose lichens

 Thallus forms a thin and flattened crust

 Crust is firmly attached to the surface of the substratum by its entire lower
surface

A distinct layer of fungal tissue (upper cortex) covers the photobiont cells due to
which surface appears smooth and less powdery as seen in leprose type.

However thalli surface can be continuous, dissected by cracks, warts or special

polygonal areas termed as areolae as in Rhizocarpon.

Figure: Crustose lichen Rhizocarpon geographicum

Source: [Link] (CC)

Areolae are adaptations to alternate wetting and drying regions, when thalli are
moist the tissues swell and the cracks close. Between areolae and the growing

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regions of crustose thalli may be present a white or dark brown to black marginal
prothallus which contains only the mycobiont and no photobiont, e.g. Caloplaca
flavescens.

A B

Figure:A.[Link]
[Link]

2. Foliose lichens.

 Thallus dorsiventrally flattened leaf-like thallus.

 The upper cortex of the thallus is protective, gelatinized mycelial layer
with a pseudoparenchymatous structure followed by a photobiont
layer, and a usually thick medulla.
 Further, a well differentiated lower cortex is present which is attached
to the substratum by puckering of the underside (e.g. Hypogymnia), a
fine felted tomentum (e.g. Lobaria) or coarse bundles of compacted
hyphae or root-like structures called rhizines (e.g. Parmelia).

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Figure:A. Hyphogamia physodes; B. Lobaria pulmonaria; C., D. Parmelia squarrosa

Source:[Link]
_geograph.org.uk_-_1049273.jpg;
[Link]

 The development of the tissues on lower surface enables foliose thalli to be peeled from
their substratum much more readily (without getting damaged) than crustose or
squamulose lichens e.g. Hypogymnia physodes.

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A: Thallus overview B:In Cross Section

A: [Link]

B: [Link]

3. Fruticose lichens.

 Most complex and largest of the lichens.

 Attached to their substrate from a single point by a disc-like holdfast or
clusters of rhizoids.
 The thallus may be erect or pendulous, hair or strap-like with profuse
branching with usually a radial symmetry arranged around a central cavity
(e,g. Bryoria) or a tough central strand (e.g. Usnea).

A. B.

Figure: A. Bryoria furcellata .B. Usnea flammea, thallus branch with soralia and
isidiomorphs. Collection on bark of Betula pubescens, Argyll, Scotland.

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Source:[Link] ,
[Link] (CC)

 Some Usnea spp. may attain a length of 10 m.

 Some foliose lichens may appear shrubby like fruticose lichens; but a close
examination clearly demarcates two with alga existing only on one side of the
flattish thallus in foliose and as a ring around the thallus in fruticose types.

A: Thallus overview B: In Cross section

A: [Link]

B: [Link]

4. Squamulose or placodiod lichens.

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 Portion of their thallus partially lifted off the substrate to form

'squamules' (small scales).
 However, otherwise they are similar to crustose lichens in having an
upper cortex but no lower cortex.
 The best example of a squamulose lichen thallus is Cladonia. Its
primary thallus is squamulose and secondary thallus is upright and
cylindrical with a hollow stalk having apothecium at its tip.

Figure: Cladonia fimbriata

Source: [Link]

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Figure: A: Thallus overview;B: In Cross section

Source:[Link]
m/content/kingdoms-living-world/[Link]

This vertical thallus arising out from primary thallus, bearing apothecium is known as
podetium and is typical of this genus. The podetium opens out into a cup and the
apothecia ultimately develop at the rim of the podetium.

5. Leprose lichens.

 An odd group of crustose type lichens where the fungal hyphae are loosely
associated with photobiont cells.
 They usually have a highly hydrophobic surface.
 Simplest type of the thallus without any protective upper cortex, inner cortex
and only an algal cell layer and sometimes a weakly defined medulla.
 Because they do not produce fruiting bodies which is essential for the
identification of fungi therefore these lichens have not yet been identified
properly or at least not yet have been given full scientific names.

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A. B.

Figure: Lepraria inacana a leprose lichen.

Source: A. [Link] (copyright Alan

Silverside); B. [Link]

6. Filamentous lichens.

 There are a few filamentous lichens (e.g. Racodium) in which the

filamentous form of the alga is predominant.
 The fungus forms the outer structure and gives the lichen its shape.

Figure: Racodium rupestre a filamentous lichen

Source: [Link]

7. Intermediate forms:

 There are several intermediate forms of lichen thalli observed.

 In a few genera, such as Cladonia, Pilophorus and Stereocaulon, there is a
combination of different growth forms: like the basal part of Cladonia is
squamulose, from which arise fruticose, hollow, simple or branched
structures.

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Figure: Cladonia chlorophaea – with basal squamulose and upper fruticose thallus.

Source: [Link] (CC)

Structure of the lichen thallus

On the basis of distribution of photobiont among the fungal hyphae, there are two general
types of lichen thalli.

 Homiomerous Lichens
 Heteromerous Lichens

Compartive view of Homiomerous and Heteromerous Lichens

CHARACTER  HOMIOMEROUS/  HETEROMEROUSL

unstratified or unlayered stratified or layered
thallus thallus

DEFINITION When photobiont is more or less The photobiont is found as a

evenly distributed throughout the definite layer or layers embedded
thallus in a pseudoparenchymatous mass
of fungal hyphae
DESCRIPTION The alga is usually gelatinous and is
a cyanobacterium. The fungal
hyphae are intermixed with algal
components and form a thin outer

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protective layer of thallus.

EXAMPLES Collema - jelly lichen Parmelia, Usnea
PICTURE

A B
[Link]
es/226/Lichens/Lichens_files/image [Link]

[Link] es/226/Lichens/Lichens_files/imag
[Link]

Structure of Heteromerous Lichens’ Thallus

The vertical section of foliose lichen which is a typical example of heteromerous type, the
following layers may be distinguished from top to bottom.

 Upper Cortex. Usually composed of tightly interwoven mycelium, that gives it a

cellular appearance. This cellular appearance is referred to as
pseudoparenchymatous.
 Algal Layer. Composed of interwoven hyphae with the algal cells. This is the ideal
location for the algal cells below the upper cortex where it receives the optimal
amount of solar radiation, for photosynthesis, but not direct solar radiation which
would be harmful.
 Medullary Layer or medulla. Composed of loosely interwoven mycelium. This layer
is entirely fungal.

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 Lower Cortex. Usually same composition as the upper cortex and attached to the
substratum by special attachment organs as explained earlier. These structures are
entirely fungal in origin, and serve to anchor the thallus to the substratum.

Growth of Lichen Thalli

Though lichens lack roots, they are usually firmly attached to the substrata by various kinds
of attachment structures explained above. Lichens grow extremely slowly; with a few
millimeters increase their length or diameter per year. In case of the fastest growing
varieties not more than 30 mm are added to their length per year.

How Lichens Are Capable Of Growing In Harsh Environments?

During prolonged periods of dessication, for their survival lichens retain a small amount of
water and reduce their growth and metabolic processes. After a rain, the fungal partner is
able to soak up water like a sponge, absorbing two to three times its weight in water. In
moist internal environment is critical for the alga, requiring water, along with carbon dioxide
and sunlight, to manufacture food through photosynthesis. Further, during dry periods in
scarcity of other food, the fungus benefits by absorbing sugars and nitrogen-rich compounds
produced by the alga during photosynthesis.

Reproduction

There are two components to lichen reproduction. That portion which reproduces the
individual entities and that portion which reproduces both the bionts. It can be studied
under two separate headings:

 Reproduction involving: both partners

o only mycobiont
o only photobiont

Reproduction involving both partners

Reproduction involving both partners can take place vegetatively or asexually by a wide
range of methods, such as:

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 Lichens

1. Fragmentation. Small units (propagules or fragments), having portions of both the

alga and fungus, break off from the main organism (lichen) and begin to develop on
their own. This mode of reproduction is of special importance in fruticose lichens
which break off easily when trampled due to their fragile nature, as in Cladonia
stellaris and C. uncialis.
2. Soredia. These specialized minute propagules are formed when one or more algal
cells got surrounded by small clumps of fungal hyphae.

[Link]

 These become noticeable on the surface of the thallus by arising in the algal
layer either as individual units forming a powder over the entire surface of the
thallus, or as well defined small clusters known as soralia (sing. soralium)
and break through the upper cortex,.
 This has been reported in about 30% of the lichens.
 Soredia are usually hydrophobic and are dispersed by wind, perhaps following
their initial detachment by the impact of a rain drops.
 The position, shape and sometimes the colour of the soralia are the features
used for the identification of genera.

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 Lichens

A B

Showing white soralia white soralia and brownish apothecia on

margins

Source A: [Link]
[Link];B:[Link]

Figure: Cross section through the thallus of a typical foliose lichen.

Source: [Link]

3. Isidia. These structures appear as tiny outgrowths on the upper or outer cortex of
foliose and fruticose lichens respectively.

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 Lichens

 They contain cortical, algal and medullary layers and appear as

cylindrical finger-like, simple or branched growths, or flattened pads.
 They are found in 15% of the lichens.
 They increase the surface area of the lichen thallus but can also
become detached and then function as vegetative propagules.
 Like soredia, the position of the isidia is used to identify genera.

Figure: Leptogium cyanescens

Source:[Link]
[Link]

4. Squamules: In some lichens such as Cladonia, squamules break off from a

vegetative thallus and serve to establish a new thallus.
5. Lobules. These are small outgrowths, usually formed from the edge or margin of the
lichen thallus. They are much like isidia, but more fragile. Lobules function in
reproduction just like isidia and soredia, dispersing the alga and fungus together.

Reproduction involving only the mycobiont

1. Pycnidia: The thallus of some lichens is dotted with black pits called pycnidia, which
are lined with minute, asexual conidia. These bacilliform spores (also called
pycnidiospores) are strikingly similar in appearance to the male spermatia produced
by certain non-lichenized fungi such as wheat rust.

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 Lichens

A; showing pycnidia on margins B: pycnidia in section

Source:A.[Link]
[Link]/html/p1/[Link]

The second method of reproduction involves only the mycobiont. The fungual partner
reproduces only sexually by the production of ascospores or basidiospores in most of the
lichens. Conidial states if produced are often pycnidial and are flask-shaped structures
producing characteristically shaped conidia. The exact functional role of these conidia is not
yet fully understood and may in some species be sexual.

Since majority of the fungi in lichen associations belong to Ascomycota, the

reproduction normally found within this group will be explained. As in the non-
lichenized groups both homothallic and heterothallic mating systems have been recognized
in them.

Apothecia with inoperculate asci are especially common with perithecia and
pseudothecia in few are observed in non-lichenized groups.

On the contrary, no apothecia with operculate asci or cleistothecia are observed among
lichenized fungi.

Discharge of ascospores varies in most lichens, either forcibly or by a pore (in

unitunicate asci); by rostrate dehiscence (in lecanoralean asci);by fissitunicate dehiscence
(in bitunicate asci).

2. Apothecia: These are saucer-shaped, 2-3 mm, small structures, which may be raised
above the surface of the thallus, level with it, or sunk below it.

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 Lichens

Figure: magnified view of a section through a apothecia

Source: [Link]

In some genera the apothecia look like slits, i.e. Graphis sp.

Figure: Graphis elegans with slit like apothecia

Source: [Link]

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 Lichens

The asci are held within a mass of special sterile hairs called paraphyses. These paraphyses
are composed of fungal hyphae and often have coloured tips. These then give the apothecia
their coloured centres. They may be reddish-brown (e.g. Peltigera) or yellow (e.g. Cladonia,
Lecanora, Xanthoria). Differences in the colours of apothecia can be important in
identification. Apothecia are mainly perennial with serial development of asci.

Figure: A. Peltigera horizontalis; B. Cladonia coccifera with its red apothecia

Source:[Link]
[Link]

Types of apothecia:

The apothecia are of two types depending upon the presence or absence of photobiont cells
in the tissues supporting the margins of discs.

a) Lecideine type. Here the apothecia are covered only by the fungal hyphae which
form the proper thalline margin. The photobiont is not present in the apothecial
margin and thus this part differs in colour from the main thallus e.g. Cladonia,
Lecidea and Peltigera.
b) Lecanorine type. In this type, apothecia are more developed as the photobiont
component also takes part along with the fungal hyphae in the formation of
thallic margin, which is of the same colour as the main thallus e.g. Lecanora and
Parmelia.

3. Perithecia usually arise singly, but in a few genera are combined into a common
stroma as in Melanotheca. The wall surrounding the generative locule (centrum) is

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 Lichens

referred to as the exciple or peridium. Asci in perithecia are discharged by way of

an apical ostiole.

Source:[Link]
a)_(4502463061).jpg

Perithecia are often buried almost completely in the substrate on which the lichen is
growing. In species of Endocarpon, Gonohymenia, Staurothele and Thelenidia, photobiont
cells occur in the ascomatal cavity or amongst the asci.

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 Lichens

Figure: Endocarpon pusillum. A. The lichen E. pusillum. B. The isolated mycobiont and
phycobiont. C. Cross section of a perithecium with hymenial algal cells inside. D. Cross
section of a thallus under scanning electron microscopy (SEM). E. The algal layer (SEM). F.
An algal cell is clasped and surrounded by some hyphae (SEM).

Source: [Link]

Some Peculiar Structures of lichens

Cephalodia

These appear as small, hard, dark, wart-like outgrowths generally on the upper cortex of
some lichens e.g. Peltigera spp.

Figure:Showing cephalodium on the upper cortex

Source:[Link]
[Link]/220px-Peltigera_aphthosa-[Link]

 These outgrowths cannot be removed easily.

 The cephalodia contain cyanobacterium with the mycobiont but the major
part of the lichen thallus is composed of a green alga (primary photobiont).
 Lichens with these structures can fix atmospheric nitrogen and supply some
essential substances or food to the fungus which is not supplied by the green
alga, rather the lichen may get considerably benefitted from especially in
extremely ologotrophic habitats.
 Such lichens with more than one photobiont have been called chimeras or
joined thalli. These thalli have two morphologies within a single thallus, one
in the region of each of the photobionts.

Cyphellae and Pseudocyphellae

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 Lichens

Some foliose lichens bear regularly arranged aerating structures, visible to the naked eye
as small white spots or cracks on the upper or lower cortical surfaces. These open into a
depression lined by specialized rounded cells and fungal hyphae in these pores are loosely
interwoven. They are represented by organised breaks in the lower cortex where the
medulla is exposed. If they have a definite border formed by the edge of the cortex they are
called cyphellae;

But if they lack any border and loose hyphal medullary tissue comes to the surface from
which cortex is absent, they are called pseudocyphellae.

Source: [Link]

These differ from one another in following manner:

 Cyphellae are bigger and more complex structures than pseudocyphellae.

 Cyphellae have globular terminal cells while pseudocyphellae do not show any
such cells.
 Cyphellae are only known from the genus Stricta and pseudocyphellae have
been observed from a wider number of genera such Alectoria, Bryoria,
Parmelia sulcata and Pseudocyphellaria etc.

However these two structures are thought to lower the gas diffusion resistance of the
cortex and thus act as pathways for gas diffusion into thalli.

Dispersal

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 Lichens

Wind and rain drops are two very common methods of dispersal of asexual propagules
(soredia, isidia, conidia etc.) or sexual spores (ascospores).

Animals can also play a role in dispersing lichens. It has been shown for Xanthoria parietina
that mites feeding on lichen thalli can spread both the mycobiont and photobiont via their
faecal pellets.

Since some lichens do not produce soredia or isidia, dispersal by invertebrates could be
significant in these. Lichens are also dispersed by birds that use them in their nests.

Thallus of Lecanora esculenta rolls up from substratum and become detached upon maturity
and is blown about by wind.

Chemical Composition of Lichens

Lichens produce a wide range of primary (intracellular) and secondary (extracellular)

compounds.

i. Primary metabolites include

 amino acids,
 polyols,
 carotenoids,
 polysaccharides, and
 vitamins.

a. Some, like the polysaccharide cell wall compounds lichenin and isolichenin,
have taxonomic significance.
b. Carotenoid compounds have also been intensely studied for clues to
evolutionary relationships.
c. For example in Cetraria islandica the wall contains lichenin (C6H1005), a
gummy substance, which swells in cold water and dissolves in hot.
d. Another substance, isolichenin is also found which is distinguished from
lichenin by the fact that it dissolves in cold water and turns blue under the
reaction of iodine.

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 Lichens

e. Calcium oxalate is a very common substance, especially in crustaceous

lichens; fatty oil in the form of drops or as an infiltration in the membrane is
also common.

ii. Secondary metabolites are often called lichen substances and since they have
an acid nature they are generally known as or lichen acids.
a. They are produced primarily by the mycobiont, and are deposited externally
on the hyphae of the cortex and/or medulla.
b. Common products include steroids and triterpenes like zeorin found in
species of Cladonia.
c. Pulvinic acid derivatives, derivatives of orcinol or ß-orcinol units such as
depsides, depsidones, depsones, anthraquinones, xanthones, as well
as aliphatic fatty acids are the most common.

Reason for different coloration in lichens

The main reason behind the vibrant colouring of many lichens is the incrustation of the
hyphae with the crystalline excretory products.

Lichen thalli growing at higher altitudes or on surfaces facing the sun often contain higher
pigment concentrations than those growing in shade. An example of a light-screen pigment
is polyketide usnic acid which is also toxic against bacteria, fungi and several other
organisms. This substance has not been found in non-lichenized fungi.

Another example is vulpinic acid produced by the wolf’s lichen, Letharia vulpina. This
species is extremely toxic. Its thalli have been used to poison foxes and wolves. Lichens
containing toxic acids are therefore avoided by lichen-grazing animals.

Figure:Letharia vulpine

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 Lichens

Source:[Link]

A large number of these acids (mostly benzene derivatives) have been isolated and are
generally characterized by their insolubility or little solubility in water; for example,
xanthone lichexanthone, usnic acid and derivatives of pulvinic acid in Lecanora spp.
give it bright yellow colouration at exposed surfaces at high altitudes.

Figure: Lecanora muralis

Source: [Link]
[Link]/220px-Lecanora_muralis-[Link]

Similarly anthraquinone parietin found in Xanthoria parietina gives yellow colour to the
thalli. There are some other acids found in various other lichens e.g. evernic acid in
species of Evernia, usnic acid in Cladonia spp.

Economic importance of lichens

Human Uses of Lichen Substances

Lichens have and are being used for many different purposes by human cultures on every
continent, with the possible exception of Australia. This study of the relationship between
lichens and people is known as ethnolichenology.

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 Lichens

Lichen substances have been used by humans (both historically and today) for dyes,
medicines, food, decoration, etc. Different human cultures across the world have also
found many other more novel uses for lichens.

Commercial products such as litmus, dyes and essential oils for perfumes have been
obtained from lichens but now have been replaced by synthetic products.

Lichens for perfume:

Evernia prunastri (oakmoss) and other species are harvested from nature to be used in
perfume industry as a fixative.

Lichens for dye: Lichens are a common source of natural dyes.

 The lichen dye is usually extracted by either boiling water or ammonia fermentation
(to steep lichen in ammonia for at least 2 to 3 weeks.

 A yellow dye is obtained by boiling Letharia vulpina in water.

 Traditional dyes in Scotland were very important since ages as mentioned under:

 Brown lichen dyes (crottle) and red lichen dyes (corkir) were used extensively to
produce tartans.

 Purple dyes from lichens were historically very important throughout Europe from the
15th to 17th centuries. They were generally extracted from Roccella spp. which was
imported from the Canary Islands, Cape Verde Islands, Madagascar, or India. These
lichens, and the dye extracted from them, are called (variants archil, orchilla).

 Orchil was also produced from Ochrolechia spp., lichens available in Britain and was
called cudbear.

 Both Roccella spp. and Ochrolechia spp. contain the lichen substance orcin, which
converts into the purple dye orcein in the ammonia fermentation process.

Lichens for medicine: Many lichens have been used medicinally across the world because
of the abundance of unique secondary metabolites present in most lichen thalli.

 Antibiotic properties- One of the most potent lichen antibiotics used in

traditional medicines is usnic acid, obtained from Usnea spp. These are not

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 Lichens

only active against Gram positive bacteria and streptomycetes but also have
tumour-inhibiting, antihistamine, spasmolytic and virucidal properties.

 Herbal remedies- Cetraria islandica (Iceland moss) was used in the treatment
of tuberculosis. Lobaria pulmonaria (Lungwort) is still sold, for the relief of lung
diseases and catarrh in Europe.

Lichens for food: Lichens are eaten by people in North America, Europe, Asia, and Africa,
and perhaps elsewhere.

Often lichens are merely famine foods eaten in times of dire needs, but in some cultures
lichens are a staple food or even a delicacy.

Problems often encountered with eating lichens

 Presence of slightly toxic secondary compounds,

 Polysaccharides present in them are usually indigestible to humans.

Therefore to overcome these problems, lichens are often thoroughly washed, boiled, or
soaked in ash water to remove secondary compounds.

In Northern Europe and Scandinavia Cetraria islandica (Iceland moss) was an important
human food. It was used in different preparations like porridge, pudding, soup, or salad.

Figure:Cetraria islandica (Iceland moss)

Source:[Link]
etraria_islandica)_6101.JPG

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 Lichens

Cladina rangiferina, or reindeer lichen is a staple food of reindeer and caribou

in the arctic. These lichens act as important component of the ground cover grazed
by animals.

Figure: Cladina rangiferina (Reindeer lichen)

Source:[Link]
437180).jpg

Northern peoples in North America and Siberia traditionally eat the partially digested lichen
after they remove it from the rumen of caribou that have been killed. It is often called
'stomach icecream'. In India, and other centers of curry powder production, garam
masala sauce contains certain lichens used as bulking agents.

Many invertebrates (insects, mites and sluges) also use lichens as food. They may also be
eaten by a number of large hoofed mammals.

Miscellaneous uses

Source of nest insulation

Flying squirrels build nests that contain upto 98% lichen thalli.

Certain insects (African bush cricket) hide in lichens and a few have mysterious colouration
and morphology that give them the appearance of leaves covered with foliicolous lichens.

Nitrogen fixation

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 Lichens

Lichens with cyanobacterial photobionts are important in certain ecosystems such as,
deserts, tundra, and certain new and old temperate and tropical forests, where they provide
the bulk of fixed nitrogen as lichens along with bryophytes and free-living cyanobacteria are
present in the crust of soil.

Pollution indicators

Source:[Link]

Sensitive to aerial pollutants like sulphur dioxide (SO 2) and to acid rain. Different lichens
show a differential sensitivity to SO2. Thus the presence or absence of key species can be
used as an index of the levels of air pollution.

The SO2 tolerant lichen, Lecanora conizaeoides may have evolved in SO2 polluted areas
and thus was present abundantly in parts of Northern Europe. This lichen may actually
require elevated SO2 levels for good growth as shown by its disappearance from some areas
after the implementation of legislation to curb SO2 emissions.

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 Lichens

Figure: Lecanora conizaeoides

Source:[Link]
oRJ

On the contrary, many lichen species have become extinct in large areas of lowland Britain
after industrialization, one example being the beard moss Usnea articulata. This is mainly
due to SO2 pollution, but the loss of habitat, particularly ancient woodland, may also be the
reason for the reductions in some species.

Lichens are sensitive to SO2 because their efficient absorption systems result in rapid
accumulation of sulphur when exposed to high levels of these pollutants.

Out of the two bionts, the photobiont appears to be more sensitive to the pollutants where
the chlorophyll is destroyed and photosynthesis is inhibited.

A lichen zone pattern may be observed in large towns and cities or around industrial
complexes which corresponds to the mean levels of SO 2 experienced. The most commonly
used zonal index is the Hawksworth and Rose Zonation Index, first published in 1970,
consisting of a scale of 1 (poorest air quality) to 10 (purest air).

Particular species of lichen present on tree bark can indicate the typical SO 2 levels
experienced in that area. For example if there are no lichens present, the air quality is very

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 Lichens

poor (zone 1), whilst generally only crusty lichens such as Lecanora conizaeoides or Lepraria
incana can tolerate poor air quality (zone 3).

In moderate to good air, leafy lichens such as Parmelia caperata or Evernia prunastri can
survive (zone 6) and in areas where the air is very clean, rare species such as ‘the string of
sausages’ Usnea articulata or the golden wiry lichen Teloschistes flavicans may grow (zone
10).

This zonation index applies only to areas where sulphur dioxide levels are increasing. If
sulphur dioxide concentrations are falling, lichens rarely colonize in exactly the same
sequence in a few years time as lichens are slow growing.

Lichens obtain most of their minerals from the air and rain water where they are present in
very low concentration. Therefore, they can also accumulate dissolved substances like
radionuclides which enter the food chain, lichen-reindeer-man. Lichens are also used to
monitor the radioactive pollutants from the nuclear explosion accident sites.

Lichenometry and its uses

Scientists have used lichens to get information about the places they grow. It is because of
the long-lived thalli of many species.

In archeology and paleontology, lichenometry is the study of dating a surface using

lichens age markers. Crustose lichens e.g. map lichen Rhizocarpon geographicum, are
commonly used for lichenometry, because they have the slowest growth rate.

Since lichens increase in size radially as they grow, measuring the diameter of the largest
lichen on a rock surface can thus be used to determine the time the rock has been exposed.

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 Lichens

Figure: Xanthoria elegans: One of the first lichens used for lichenometry

Source:[Link]

Lichen can be preserved on old rock faces for up to 10,000 years, providing the maximum
age limit of the technique. The use of lichenometry is of increased value for dating
deposited surfaces over the past 500 years as radiocarbon dating techniques are less
efficient over this period.

Lichenometry has also been used in many different contexts to date surfaces. The dating of
the sequences of rocks forming glacial moraines has been the most widely used
application.

Other applications of lichenometry include the dating of the stone images on Easter
Island, stone walls in England, river flooding sea level change and the occurrence of
landslides.

Common Names of some lichens

Common Name Botanical Name

Map lichen Rhizocarpon geographicum
Iceland moss Cetraria islandica
Lungwort Lobaria pulmonaria
Reindeer moss, caribou Cladina (=Cladonia) rangiferina
moss, grey reindeer
lichen
British soldiers Cladonia cristatella
Oakmoss Evernia prunastri
Wolf’s lichen Letharia vulpina
String of sausages Usnea articulata
Golden wiry lichen Teloschistes flavicens
Manna lichen Lecanora esculenta
Jelly lichen, skin lichen Collema spp.

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 Lichens

DID YOU KNOW?

 Lichenoglyph is a century old picture made by Canadian Indians by scraping
lichens off the surface of a large vertical rock face.
 Arizona's Bristle cone pine is younger if compared with some lichens. It is
estimated that some lichens in the Antarctic may be 10,000 years old. If it is
right then these lichens started growing during the Ice Age.
 Lichens have been used for packing ancient Egyptian mummies!
 British Soldiers is lichen which gets its name from the resemblance to the
uniforms worn by English soldiers during the Revolutionary war. Its
scientific name, Cladonia cristatella is derived from both Greek and Latin
languages. The generic name is from the Greek word kladon which means
"sprout" in reference to the characteristic vegetative shrub-like appearance
of the fungal body. The species name is from the Latin crista, "crest" and
the diminutive suffix ella; literally means the "small crest." This is to
describe the tufted structure at the tips or crests of the appendages.

[Link]

 The European Space Agency has discovered that lichens can survive
unprotected in space. In an experiment led by Leopoldo Sancho from the
Complutense University of Madrid, two species of lichen – Rhizocarpon
geographicum and Xanthoria elegans – were sealed in a capsule and
launched on a Russian Soyuz rocket on 31 May 2005. Once in orbit the
capsules were opened and the lichens were directly exposed to the vacuum
of space with its widely fluctuating temperatures and cosmic radiation.
After 15 days the lichens were brought back to earth and were found to be

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 Lichens

in full health with no discernible damage from their time in orbit.

Summary

The word lichen is derived from a Greek word (Gr leikhēn = to lick). In Latin līchēn, means
a kind of plant. These are unique organisms in many ways. Lichens represent an intimate
and long-term symbiosis of algae and fungi. The fungus partner is called mycobiont and a
photosynthetic algal partner, the photobiont (earlier known as phycobiont). The fungal
partner usually contributes more to the structure and the algal partner produces food for
the lichen by photosynthesis. As the external surface is formed by fungal partner it is
knowns as the exhabitant, whereas, the unicellular or filamentous photobiont cells are
termed as inhabitants as located inside the lichen thallus. The relationship between the
fungal partner and the photosynthetic partner of lichens has been described in various
ways. Some describe it as mutualism in which both partners are being benefitted. The
fungal partner benefits by getting carbohydrates, from the algae which being green
synthesizes them through photosynthesis. The photosynthetic partner gets protection from
dessication in adverse weather thus resulting in the algae having a more stable and
constant environment to live in allowing it to grow better. The fungi collect the
carbohydrates by means of special hyphae called appressoria or haustoria which contact the
wall of the algal cells. The distribution of lichens over the surface of the globe is regulated,
not only by the presence of suitable substrata, but more especially by climatic conditions.
Lichens are spread over large geographical areas e.g. species of Parmelia sulcata, Xanthoria
elegans are worldwide in distribution. Lichens are classified based on the substrata on which
they grow viz. lichens- growing on bark of trees - Corticolous, growing on rocks, stones
etc.- Saxicolous, growing on different types of soil - terricolous, growing with / on decayed
mosses – muscicolous. On the basis of growth forms they are mainly of two types-
Endolithic (growing inside substrates) and epilitihc (growing on substrate surfaces). Epilithic
lichens usually comprises of six morphologically distinguishable types- Crustose - crust-like,
Foliose - leaf-like, Fruticose - shrub or beard-like, Squamulose or placodioid - scale-like,
Filamentous and Leprose. Further on the basis of distribution of photobiont among the
fungal hyphae, lichen thalli may be either homiomerous with photobiont more or less evenly
distributed throughout the thallus or heteromerous with the photobiont present as a
definite layer or layers embedded in a pseudoparenchymatous mass of fungal hyphae. In
heteromerous type, the layers which can be distinguished from top to bottom are -Upper

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 Lichens

cortex, algal Layer, medullary Layer or medulla and lower Cortex. Lichen may reproduce by
vegetative or asexual method involving both the partners as fragmentation, soredia, isidia
and squamules. However, reproduction involving only the mycobiont occurs by the
formation of pycnidia, apothecia, perithecia. The main reason behind the vibrant colouring
of many lichens is the incrustation of the hyphae with the crystalline excretory products.
Lichens are being used for many different purposes by human cultures on every continent,
with the possible exception of Australia. This study of the relationship between lichens and
people is known as ethnolichenology. Lichen substances have been used by humans (both
historically and today) for dyes, medicines, food, decoration, etc. Commercial products such
as litmus, dyes and essential oils for perfumes have been obtained from lichens but now
have been replaced by synthetic products. Lichens with cyanobacterial photobionts are
important in certain ecosystems such as, deserts, tundra, and certain new and old
temperate and tropical forests, where they provide the bulk of fixed nitrogen as lichens
along with bryophytes and free-living cyanobacteria are present in the crust of soil.

Exercises

Define the following: pseudoparenchymatous, Foliicolous, lichenometery, cryptobiosis,

endolithic, isidium, soredium

Fill in the blanks:

a. Litmus dye is obtained from…………….

b. The botanical name of reindeer moss is………….
c. Soredia are usually ……………….and are dispersed by wind.
d. Lichen-inhabiting fungi are termed as ……………..
e. ………achieved the synthesis of lichen experimentally for the first time.
f. Lichens growing on leaves are known as ……………
g. A microscopic, powdery mass of algal cells encircled by fungal hyphae is known as
…………….

Match the following:

a. Rhizocarpon geographicum, 1. Jelly lichen

b. Letharia vulpine 2. Wolf’s lichen
c. Calcium oxalate 3. Iceland moss

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 Lichens

d. Cladonia cristatella 4. Fruticose l;ichen

e. Cetraria islandica 5. Lichenometry
f. Parmelia 6. Crustaceous lichens
g. Usnea 7. British Soldiers
h. Collema 8. Lecanorine type

Answer the following questions:

1. Why can lichens establish themselves in the most inhospitable environments?

2. Where do lichens grow?
3. What are the differences between homoiomerous and heteromerous lichens?
4. How do lichens reproduce?
5. Write short note on economic importance of lichens.
6. What functions is being performed by the secondary metabolites present in the lichen
thallus?
7. How do lichens help to maintain ecosystems?
8. What is lichenometry? What are its applications?
9. What role do lichens play in soil formation?
10. What is the function of some of the structures peculiar to lichens - cyphellae,
pseudocyphellae and cephalodia?

Key to Exercises:

Fill in the blanks

a. Roccella tinctoria b. Cladonia rangiferina c. hydrophobic d. lichenicolous

fungi e. DA Stahl f. foliicolous g. Soredia

Match the following

a-5, b-2, c-6, d-7, e-3, f-8, g-4, h-1

Glossary

Saxicolous: growing on rock surfaces.

Terricolous: growing in or on soil.

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 Lichens

Cephalodium: internal or external swelling in the thalli of diphycophilous lichens, in which

nitrogen fixing Cyanobacteria are segregated.
Fruticose thallus: an erect or pendent, usually many branched lichen thallus with tissue
tending to form cylinders, but which may be flattened.
Isidium: a minute, corticated, more or less columnar lichen propagule consisting of both
fungal hyphae and algal cells, breaking off the thallus and dispersed for short distances by
various means.
Lichenometry: the use of lichens to determine the minimum elapsed time of exposure of a
substrate; important in dating geomorphological or archeological events of relatively recent
occurrence.
Poikilohydric: Poikilohydry is a condition in organisms that lack a mechanism to prevent
desiccation, such as the lichens and bryophytes.
Cryptobiosis: A state in which metabolic rate of the organism is reduced to an
imperceptible level.
Heteromerous: lichen thalli in which the algal cells form a distinct layer within the thallus.
Homoiomerous: lichen thalli in which the cells of the photobiont are more or less evenly
distributed throughout.
Corticolous: growing on bark.
Crustose thallus: a crustlike lichen thallus.
Discolichen: a lichen in which the fungus partner is a Discomycete i.e. an ascomycete
producing apothecium.
Endolithic: inside rock.
Foliicolous: growing on leaves.
Foliose thallus: a leaflike lichen thallus. .
Soralium: region in a lichen thallus in which soredia are produced.
Soredium: a microscopic, powdery mass of algal cells encircled by fungal hyphae and
formed in soralia on some lichen thalli; soredia act as propagative units.

References
1. Alexopoulos, C.J., Nims, C.W., Blackwell, M. (1996). Introductory Mycology, John Wiley &
th
Sons (Asia) Singapore. 4 edition.

2. Webster, J. and Weber, R. (2007). Introduction to Fungi, Cambridge University Press,

rd
Cambridge. 3 edition.

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 Lichens

3. Sethi, I.K. and Walia, S.K. (2011). Text book of Fungi and Their Allies, Macmillan
Publishers India Ltd.

4. Mehrotra, R.S. and Aneja, K.R. 1990. An Introduction to Mycology. New Age
International.

5. Kirk, P.M., P.F. Cannon, D.W. Minter, and J. A. Stalpers (eds.). 2008. Ainsworth and
Bisby's Dictionary of the Fungi, 10th edition. CAB International, Oxon, UK.

Web links
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