UNIVERSITAS NEGERI JAKARTA

FAKULTAS MATEMATIKA DAN ILMU PENGETAHUAN ALAM

PROGRAM STUDI BIOLOGI

MATA KULIAH
BIODIVERSITAS
DAN SISTEMATIKA
KRIPTOGAM

LICHEN

OLEH :
AGUNG SEDAYU, M.Sc.
RIZAL KOEN ASHARO, M.Si.
INTRODUCTION

At the beginning of the nineteenth

century botanists were making their first
attempts to classify the lichens, and in
order to systematize the immense variety
of growth habit they first divided the
class on the basis of several major
growth forms.

For example, lichens covering rocks,

trees, or soil with a thin, more or less
well-developed crust were separated
from leaflike species which adhere more
or less firmly to the substrate and from
upright, branching, bushlike forms.
A. Caloplaca ceracea
(Teloschistaceae);
crustose endolithic
growth-form. B.
Teloschistes
chrysophthalmus
(Teloschistaceae);
fruticose. C. Xanthoria
elegans (Teloschistaceae);
foliose. D. Megalospora
tuberculosa
(Megalosporaceae);
crustose. E. Letrouitia
domingensis
(Letrouitiaceae); crustose.
F. Brigantiaea leucoxantha
(Brigantiaeaceae);
crustose. G. Calicium
salicinum (Caliciaceae);
ascomata close-up,
crustose thallus. H.
Physcia phaea
(Physciaceae); foliose. I.
Rinodina mniaraea
(Physciaceae); crustose.
Certainly the growth form of lichen thalli cannot be considered as a principal characteristic on
which taxonomy can be based. The foliose lichens, for example, do not form a taxonomic group
of related species but only a morphological unity. The lichens of one family, even of one genus,
may belong to the crustose, foliose, and fruticose growth form.
A theory proposed by Reinke (1894-1896) suggests that the lichens of different
taxonomic groups have developed independently from crustose species to
foliose forms and finally to fruticose plants. Fruticose lichens are regarded as
the most highly developed peaks of several parallel lines of evolution.

This theory, although attractive, is not acceptable.

As will be seen, the structure of foliose thalli is not
less complicated than that of fruticose thalli. On the
contrary, some highly differentiated organs such as
cyphellae are confined to foliose genera. For this
reason, the foliose and fruticose lichens must be
regarded as forms equal in their level of development
in a line of evolution leading from a crustose
organization to a more highly differentiated habit.
Crustose lichens can, with some certainty, be
considered as primitive or secondarily derived.
In spite of the above-mentioned objections it is still convenient to divide the
lichens into growth forms. The habit being often the most obvious
characteristic for distinguishing lichen species, the growth form is the most
useful starting point in the construction of artificial keys for their
determination.

The habit of a lichen is due

not only to the overall
growth form, but is often
the result of special
anatomical characteristics.
For example, the shape of
the thallus surface depends
on the anatomy of the
cortex. External appearance
and internal structure are
interdependent.

Cross-section of a foliose lichen

General Structure
Lichens do not have
a waxy cuticle like
plants have on their
leaves, nor do they
have vascular tissue
such as xylem and
phloem to move
nutrients and water
around their thalli as a
plant does. Everything
in the lichen’s
environment is
absorbed into the
lichen's structure.
Lichens get their
water and nutrients
from their
surrounding
environment via air
and rain.
Cortex
The cortex is the outer layer of the lichen thallus. These cells are thicker and more closely
packed than the other fungal cells in the lichen. This layer provides some small measure of
protection, as well as provides color in some species.
Algal Layer
You can usually tell what kind of alga a lichen has just by color alone. When a lichen is dry, its color is
usually gray or colored like the fungal cells on the upper cortex. When a lichen is wet, those cells
become transparent, and the algal cells underneath get a chance to show their vibrancy.

Green algae generally give the lichen a bright

green color when wet, although there are exceptions
of pigmented lichens with green algae due to the
fungal partner showing its colors.

Cyanobacteria can be a layer under the upper

cortex or in tiny pockets on top of the upper cortex if
there is a green algal layer already present.
Cyanobacteria will give the lichen a dark green, brown,
or black color.

In some lichens, however, there are no layers of fungus

Xanthomendoza mendozae close-up. Notice the
and alga. The individual components are mixed green algal layer underneath the orange cortex.
together in one big uniform layer and the resulting
growth form is gelatinous. These types of lichens are
called jelly lichens.
Medulla
The majority of the lichen thallus is comprised of fungal filaments called
the medulla. It is made of fungal cells that are loosely packed in the
middle of the lichen thallus, have thin cell walls, and are threadlike. The
result is a cotton-like substance underneath the outer cortex.

Basal Attachment
Lichens attach to their substrate by different means.

Rhizines are fungal filaments that extend from the

medulla and attach the lichen to its substrate. Rhizines
have no vascular capabilities like the roots in plants.
They do not move water or nutrients to the lichen;
they simply hold the lichen down to whatever it is
sitting on.
Umbilicaria phaea, rock tripe, on a rock in
southern California. It has one central holdfast,
like an umbilical cord, hence the name. Holdfast is an extension of the lichen thallus. Instead
of many rhizines, some lichens have a central peg or
holdfast that attach to the substrate, generally a rock.
These types of foliose lichens are called umbilicate
lichens, since the central holdfast is like an umbilical
cord.
Anatomy of the Thallus
 Hyphae with hyphae with branched hyphae
hyphae with multi- globose cells ellipsoid cells with cylindrical
angular cells cells

interwoven parallel oriented

hyphae in parallel oriented hyphae in
anticlinal interwoven hyphae in hyphae in periclinal anticlinal
arrangement periclinal arrangement arrangement arrangement
prosoplectenchymatous pseudoparenchymatous branched hyphae
tissue developed from tissue formed from thin-
fan-shaped
anticlinal hyphae with walled anticlinal hyphae arrangement of in a netlike
strongly gelatinized walls hyphae arrangement;

Prosoplectenchymatous tissue
cell lumina of netlike formed by hyphae in a netlike
pseudoparenchymatous
hyphae lying in the arrangement with gelatinized tissue
homogeneous substance of walls
the gelatinized walls
hyphae from the medulla of Parmelia cetrarioides
with bone-shaped cross-septa perforated by pores
hyphae from a rhizine of Peltigera praetextata
showing anastomoses and pores
palisade tissue in the cortex of Roccella phycopsis
ascus and paraphyses of Coenogonium with
club-shaped cells
longitudinal and vertical sections through the thallus of Darbishirella gracillima showing the algal
layer and arrangement of cortical hyphae
 Leptogium

Physma
Types of cortex in the Collemataceae. Above: cortex of
isodiametric cells in Leptogium sinuatum seen in cross
section (left) and from above (right). Middle: primitive
cortex of Leptogium apalachense; the cell lumen lies
inside a gelatinous substance and forms an irregular
pattern when seen from above (right). Below: cortex
of Physma byrsinwn which is several cells deep in
some places.
 Trentepohlia

Physolinum

hairlike thalli of Coenogoniwn sp. with hairlike or granular thallus of Coenogoniwn

Trentepohlia as phycobiont; in two places the moniliforme with Physolinum as phycobiont
hyphae gather to form a fruiting body
 Cystocoleus Porina nucula

hairlike thallus of Cystocoleus niger with young perithecium of Porina nucula within a thallus
Trentepohlia as phycobiont; the alga is granule; the thallus contains large crystals of calcium
completely covered by hyphae oxalate
 cross section of the
thallus of Sphaerophorus
melanocarpus showing
cortical hyphae with
part of a cross gelatinized walls in a
section of the netlike arrangement
radial thallus of
Sphaerophorus
globosus
showing a
strongly
gelatinized
cortex

cross section of the thallus

of Letharia vulpina with a
central cord formed by
several smaller
strands
cross section of the isolateral thallus of
the double cortex of Ramalina siliquosa.
Ramalina siliquosa. The algal cells are either
situated in the medulla or in the cortex
Development of the Thallus
The thalli of Lichen thalli tend to This type of
frutescent lichens fuse, especially in the connection should
grow at the top and dense cushions of be clearly
die away at the base. certain fruticose lichens distinguished from
As a result branched such as Cladonia. In the complete fusion
thalli may split and most lichen species achieved by other
become separate only the branches of lichens (P).
plants (N) different plants grow
together at points of
contact (O)
 Growth Forms
Crustose Lichens
Crustose lichens are just that,
crusts. They form a crust over a
surface, like a boulder, the soil, a
car, or your roof shingles. They can
come in many bright, vibrant
colors like sunny yellow, orange, Foliose Lichens
and red, as well as grays and Foliose lichens have two easily distinguishable sides. In other
greens. Crustose lichens are words, there is a top side and there is a bottom side. They can
pressed against their substrate. be very flat, leafy like lettuce, or convoluted and full of ridges
and bumps.

Fruticose Lichens
Fruticose lichens can be pendant and hair-like, upright and
shrubby, or upright and cup-like. Many fruticose lichens
have round branches that have a central core and others
are hollow in the middle. Other fruticose lichens have flat
branches that tangle up with each other.
Reproduction
Lichens are different. Unlike plants that can produce seeds that grow into new plants, lichens do not
have a straightforward way to grow more lichen. Since the fungus is the dominant partner in the
relationship, it gets to develop its fruiting bodies and produce spores. These spores can produce another
fungus, but unfortunately, for the alga, it does not get the opportunity to reproduce at all. Either the
new fungus has to find an algal partner or it perishes.
 APOTHECIA CONTAIN SPORE-PRODUCING ASCI
Spores are microscopic, single cells that can develop into entire new multicellular organisms. The fungus
group to which the great majority of lichens belong, the phylum Ascomycota, are sometimes called “cup fungi,”
because the specialized spore-producing hyphae (the asci) are packed together in a neat little button-shaped
cup, the “apothecium.” (Ascomycetes are also called “sac fungi,: after the sac-like asci withing which the
spores are produced.) Spores released into the air can be blown to a new place where they germinate and start
a new fungus. This new fungus must immediately find the correct species of alga to form a new lichen, or it
dies. Some lichen algae can live independently but no lichen fungi can. This may seem haphazard, but some
lichens are quite successful with this method of reproduction.

Below, see a photo of a apothecia on a typical foliose lichen, Physcia stellaris.

 ASEXUAL REPRODUCTION BY SOREDIA
When lichens reproduce by fragmentation, a small portion of the thallus containing both partners breaks
off and is transported to a new location where it grows to a new thallus. Many lichens have specialized
structures that enhance fragmentation. The most common asexual propagule (small particle that can be
transported and form a new organism) is a soredium. Soredia are very tiny (barely visible to the
unaided eye) dust-like or granular balls of hyphae along with a few intertwined algal cells. They are borne
through breaks in the upper cortex that are usually aggregated into easily-seen rough patches called
“soralia.” The size of soredia ranges from dust-like to coarsely granular, and they may be located on the
flat surface of the lichen, or along its margin. These are distinguishing features that can help in telling
lichens apart

Soredia are powdery masses of fungal

hyphae mixed with algal cells that burst
through openings in the upper cortex.
 ASEXUAL REPRODUCTION BY ISIDIA

A somewhat less common, but by no means rare, asexual propagule is formed from isidia. Isidia are
small buds that form on the upper surface of lichens that contain both the upper cortex (fungus) and algal
layer. These generally tubular or stump-shaped projections eventually break off from the lichen’s surface,
leaving a microscopic scar. Using a hand lens, a lichen with isidia can usually be distinguished from one
with soredia by having a bumpy yet shiny surface, (shiny because the cortex is intact).

Isidia are small surface outgrowths of the upper

surface of a lichen which break off to form new
lichens.
 Lichen Classification
If lichenization had evolved just once and, once gained, had never been lost by descendents, all current lichens would
form a monophyletic group and could then be treated as a self-contained group of organisms. Lichens do not form a
monophyletic group. For a start, there are both ascomycete and basidiomycete lichenized fungi, albeit very
few of the latter. Perhaps lichenization arose twice, once in the basidiomycetes, once in the ascomycetes and gave rise
to two monophyletic groups - one of basidiolichens and one of ascolichens. The basidiolichens do not constitute a
monophyletic group and neither do the ascolichens. The following figure shows where the lichenized ascomycetes fit
into the ascomycetes as a whole and is based on genomic research published in 2009.
The figure is in two parts. The large tree on the left shows the classes that contain lichenized fungi with
one class, Eurotiomycetes, given in more detail, the orders within that class being shown and three of
them named. If a taxon consists only of lichenized fungi then it is shown as a red line. A taxon
containing only non-lichenized fungi is shown by a blue
line and taxa in which both lichenized and
non-lichenized fungi are found are shown in red and blue.

The lines for the latter are drawn thicker, simply to show the bicolouring more easily and the line
thickness has no other significance. The class Lecanoromycetes consists almost totally of lichenized fungi
but there is a small number of non-lichenized species within the class, hence the blue dot within the
Lecanoromyctes. This class contains the majority of the lichenized fungi (containing about 14,000 of the
approximately 18,000 known lichen species). The Dothideomycetes is the largest class within the
ascomycetes and contains about 19,000 species - but only a small number of lichenized species. The
Verrucariales and Pyrenulales consist mostly of lichenized fungi. The yellow area in the small tree to the
right shows where the large tree fits into the tree of all ascomycetes. As you can see there are three
other groups of ascomycetes, all non-lichenized, and they have been labelled X, Y and Z. The first of
those groups contains about 18,000 species while Y and Z, combined, contain about 3,000.
Lichen Habitat
Lichens have specific requirements for their habitats. Although they can occur on a variety of
substrates, each substrate must have the individual components in the right amounts that growing
lichen needs. These requirements are: water, air, nutrients, light, and substrates.