Lichen PDF

Lichen
A lichen is a composite organism that arises from algae or cyanobacteria living

among filaments of multiple fungi[1] in a symbiotic relationship.[2][3][4] The
combined lichen has properties different from those of its component organisms.
Lichens come in many colours, sizes, and forms. The properties are sometimes
plant-like, but lichens are not plants. Lichens may have tiny, leafless branches
(fruticose), flat leaf-like structures (foliose), flakes that lie on the surface like peeling
paint (crustose),[5] or other growth forms.[6]
A macrolichen is a lichen that is either bush-like or leafy; all other lichens are
termed microlichens.[2] Here, "macro" and "micro" do not refer to size, but to the
growth form.[2] Common names for lichens may contain the word moss (e.g.,
"reindeer moss", "Iceland moss"), and lichens may superficially look like and grow
with mosses, but lichens are not related to mosses or any plant.[4]:3 Lichens do not
have roots that absorb water and nutrients as plants do,[7]:2 but like plants, they
produce their own food by photosynthesis.[8] When they grow on plants, they do not
live as parasites, but instead use the plants as asubstrate.
Lichens occur from sea level to high alpine elevations, in many environmental
A tree covered with leafy foliose
conditions, and can grow on almost any surface.[8] Lichens are abundant growing on lichens and shrubby fruticose lichens
bark, leaves, mosses, on other lichens,[7] and hanging from branches "living on thin
air" (epiphytes) in rain forests and in temperate woodland. They grow on rock,
walls, gravestones, roofs, exposed soil surfaces, and in the soil as part of a biological soil crust. Different kinds of lichens have
adapted to survive in some of the most extreme environments on Earth: arctic tundra, hot dry deserts, rocky coasts, and toxic slag
heaps. They can even live inside solid rock, growing between the grains.
It is estimated that 6% of Earth's land surface is covered by lichen.[9][10]:2 There are about 20,000 known species of lichens.[11]
Some lichens have lost the ability to reproduce sexually, yet continue to speciate.[7][12] Lichens can be seen as being relatively self-
contained miniature ecosystems, where the fungi, algae, or cyanobacteria have the potential to engage with other microorganisms in a
functioning system that may evolve as an even more complexcomposite organism.[13][14][15][16]
[4][17] They are among the first living things to

Lichens may be long-lived, with some considered to be among the oldest living things.
grow on fresh rock exposed after an event such as a landslide. The long life-span and slow and regular growth rate of some lichens
can be used to date events (lichenometry).
Contents
Pronunciation
Growth forms
Color
Internal structure and growth forms
Physiology
Symbiotic relation
Ecology
Miniature ecosystem and holobiont theory
Lichenicolous fungi
Reaction to water
Common lichen growth forms
Metabolites, metabolite structures and bioactivity
Growth rate
Life span
Response to environmental stress
Reproduction and dispersal
Vegetative reproduction
Sexual reproduction
Taxonomy and classification
Fungi
Photobionts
Controversy over classification method and species names Letharia vulpina, wolf lichen, grows
Diversity like a multiple-branched tuft or
Identification methods leafless mini-shrub, so it has a
Evolution and paleontology fruticose growth form.
Ecology and interactions with environment
Substrates and habitats
Lichens and soils
Ecological interactions
Effects of air pollution
Human use
Food
Lichenometry
Biodegradation
Flavoparmelia caperatahas leaf-like
As dyes
structures, so it is foliose.
Traditional medicine and research
Aesthetic appeal
In literature
History
Gallery
See also
Notes
References
External links Caloplaca marina grows like an
Identification and classification orange crust coating the rock, so it is
Internal structure crustose.
Miscellanea
Societies
By region
Images
Cultivation
Historic
Pronunciation Caloplaca thallincola grows like a

In American English, "lichen" is pronounced the same as the verb "liken" crust, and in a pattern that radiates
outward from the center, so it has a
(/ˈlaɪkən/). In British English, both this pronunciation and one rhyming with
crustose placodioid growth form.
"kitchen" /ˈlɪtʃən/) are used.[18][19][20][21]
Growth forms
Lichens grow in a wide range of shapes and forms (morphologies). The shape of a
lichen is usually determined by the organization of the fungal filaments.[22] The
nonreproductive tissues, or vegetative body parts, is called the thallus. Lichens are
grouped by thallus type, since the thallus is usually the most visually prominent part
of the lichen. Thallus growth forms typically correspond to a few basic internal
structure types. Common names for lichensoften come from a growth form or color
that is typical of a lichengenus.
Pannaria lurida forms small leaf-like
Common groupings of lichen thallus growth forms are: scales crustose below but free at the
tips, so it is squamulose.
1. fruticose[23][24][25] – growing like a tuft or multiple-branched leafless
mini-shrub, upright or hanging down, 3-dimensional branches with
nearly round cross section (terete) or flattened
2. foliose[23][24] – growing in 2-dimensional, flat, leaf-like lobes
3. crustose[5][23][24] – crust-like, adhering tightly to a surface substrate)
(
like a thick coat of paint
4. squamulose[25] - formed of small leaf-like scales crustose below but
free at the tips
5. leprose[26] – powdery
6. gelatinous – jelly like Chrysothrix chlorina grows like
7. filamentous – stringy or like matted hair powder dusted on the rock so it is
8. byssoid – wispy, like teased wool leprose.
9. structureless
There are variations in growth types in a single lichen species, grey areas between
the growth type descriptions, and overlapping between growth types, so some
authors might describe lichens using different growth type descriptions.
When a crustose lichen gets old, the center may start to crack up like old-dried
paint, old-broken asphalt paving, or like the polygonal "islands" of cracked-up mud
in a dried lakebed. This is called being rimose or areolate, and the "island" pieces
separated by the cracks are called areolas.[23] The areolas appear separated, but are
Collema nigrescens is gelatinous,
(or were) connected by an underlying "prothallus" or "hypothallus".[26] When a
without internal structure for its parts.
crustose lichen grows from a center and appears to radiate out, it is called crustose
placodioid. When the edges of the areolas lift up from the substrate, it is called
squamulose.[27]:159[25]
These growth form groups are not precisely defined. Foliose lichens may sometimes branch and appear to be fruticose. Fruticose
lichens may have flattened branching parts and appear leafy. Squamulose lichens may appear where the edges lift up. Gelatinous
lichens may appear leafy when dry.[27]:159 Means of telling them apart in these cases are in the sections below
.
Structures involved in reproduction often appear as discs, bumps, or squiggly lines on the surface of the thallus.[7]:4 The thallus is not
always the part of the lichen that is most visually noticeable. Some lichens can grow inside solid rock between the grains (endolithic
lichens), with only the sexual fruiting part visible growing outside the rock.[23] These may be dramatic in color or appearance.[23]
Forms of these sexual parts are not in the above growth form categories.[23] The most visually noticeable reproductive parts are often
circular, raised, plate-like or disc-like outgrowths, with crinkly edges, and are described in sections
below.
Color
Lichens come in many colors.[7]:4 Coloration is usually determined by the photosynthetic component.[22] Special pigments, such as
yellow usnic acid, give lichens a variety of colors, including reds, oranges, yellows, and browns, especially in exposed, dry
habitats.[28] In the absence of special pigments, lichens are usually bright green to olive gray when wet, gray or grayish-green to
brown when dry.[28] This is because moisture causes the surface skin (cortex) to become more transparent, exposing the green
photobiont layer.[28] Different colored lichens covering large areas of exposed rock surfaces, or lichens covering or hanging from
bark can be a spectacular display when the patches of diverse colors "come to life" or "glow" in brilliant displays following rain.
Different colored lichens may inhabit different adjacent sections of a rock face, depending on the angle of exposure to light.[28]
Colonies of lichens may be spectacular in appearance, dominating much of the surface of the visual landscape in forests and natural
places, such as the vertical "paint" covering the vast rock faces ofYosemite National Park.[10]
Color is used in identification.[29]:4 Color changes depending on when a lichen is wet or dry.[29] Color descriptions when used for
.[29] Dry lichens with a cyanobacterium as the photosynthetic partner tend to be dark
identification are based on when the lichen is dry
grey, brown, or black.[29]
The underside of the leaf-like lobes of foliose lichens is a different color from the top side (dorsiventral), often brown or black,
sometimes white. A fruticose lichen may have flattened "branches", appearing similar to a foiliose lichen, but the underside of a leaf-
like structure on a fruticose lichen is the same color as the top side. The leaf-like lobes of a foliose lichen may branch, giving the
appearance of a fruticose lichen, but the underside will be adifferent color from the top side.[26]
The sheen on some jelly-like gelatinous lichens is frommucilaginous secretions.[22]
Internal structure and growth forms

A lichen consists of a simple photosynthesizing organism, usually green algae or
cyanobacteria, surrounded by filaments of a fungus. Generally, most of a lichen's
bulk is made of interwoven fungal filaments,[30] although in filamentous and
gelatinous lichens[22] this is not the case. The fungus is called a mycobiont. The
photosynthesizing organism is called a photobiont. Algal photobionts are called
phycobionts.[31] Cyanobacteria photobionts are calledcyanobionts.[31]
The part of a lichen that is not involved in reproduction, the "body" or "vegetative
tissue" of a lichen, is called the thallus. The thallus form is very different from any
The cyanobacterium Hyella
form where the fungus or alga are growing separately. The thallus is made up of
caespitosa with fungal hyphae in the
filaments of the fungus called hyphae. The filaments grow by branching then
lichen Pyrenocollema halodytes
rejoining to create a mesh, which is called being "anastomose". The mesh of fungal
filaments may be dense or loose.
Generally, the fungal mesh surrounds the algal or cyanobacterial cells, often enclosing them within complex fungal tissues that are
unique to lichen associations. The thallus may or may not have a protective "skin" of densely packed fungal filaments, often
containing a second fungal species,[1] which is called a cortex. Fruticose lichens have one cortex layer wrapping around the
"branches". Foliose lichens have an upper cortex on the top side of the "leaf", and a separate lower cortex on the bottom side.
Crustose and squamulose lichens have only an upper cortex, with the "inside" of the lichen in direct contact with the surface they
grow on (the substrate). Even if the edges peel up from the substrate and appear flat and leaf-like, they lack a lower cortex, unlike
foliose lichens. Filamentous, byssoid, leprose,[26] gelatinous, and other lichens do not have a cortex, which is called being
ecorticate.[32]
Fruticose, foliose, crustose, and squamulose lichens generally have up to three different types of tissue, differentiated by having
different densities of fungal filaments.[30] The top layer, where the lichen contacts the environment, is called a cortex.[30] The cortex
is made of densely tightly woven, packed, and glued togetheragglutinated)
( fungal filaments.[30] The dense packing makes the cortex
act like a protective "skin", keeping other organisms out, and reducing the intensity of sunlight on the layers below.[30] The cortex
layer can be up to several hundred micrometers (μm) in thickness (less than a millimeter).[33] The cortex may be further topped by an
epicortex of secretions, not cells, 0.6–1 μm thick insome lichens.[33] This secretion layer may or may not have pores.[33]
Below the cortex layer is a layer called the photobiontic layer or symbiont
layer.[24][30] The symbiont layer has less densely packed fungal filaments,
with the photosynthetic partner embedded in them.[30] The less dense
packing allows air circulation during photosynthesis, similar to the anatomy
of a leaf.[30] Each cell or group of cells of the photobiont is usually
individually wrapped by hyphae, and in some cases penetrated by an
haustorium.[22] In crustose and foliose lichens, algae in the photobiontic
layer are diffuse among the fungal filaments, decreasing in gradation into the
Schematic cross section offoliose lichen:
layer below. In fruticose lichens, the photobiontic layer is sharply distinct
1. The cortex is the outer layer of tightly
woven fungus filaments (hyphae) 2. This from the layer below.[22]
photobiont layer has photosynthesizing
green algae The layer beneath the symbiont layer called is called the medulla. The
3. Loosely packed hyphae in the medulla medulla is less densely packed with fungal filaments than the layers above.
4. A tightly woven lower cortex, with In foliose lichens, there is usually, as in Peltigera,[27]:159 another densely
anchoring hyphae calledrhizines where the packed layer of fungal filaments called the lower cortex.[26][30] Root-like
fungus attaches to the substrate. fungal structures called rhizines (usually)[27]:159 grow from the lower
cortex to attach or anchor the lichen to the substrate.[2][26] Fruticose lichens
have a single cortex wrapping all the way around the "stems" and
"branches".[27] The medulla is the lowest layer, and may form a cottony white inner core for the branchlike thallus, or it may be
hollow.[27]:159 Crustose and squamulose lichens lack a lower cortex, and the medulla is in direct contact with the substrate that the
lichen grows on.
In crustose areolate lichens, the edges of the areolas peel up from the substrate and appear leafy. In squamulose lichens the part of the
lichen thallus that is not attached to the substrate may also appear leafy. But these leafy parts lack a lower cortex, which distinguishes
[30] Conversely, foliose lichens may appear flattened against the substrate like a
crustose and squamulose lichens from foliose lichens.
crustose lichen, but most of the leaf-like lobes can be lifted up from the substrate because it is separated from it by a tightly packed
lower cortex.[26]
Gelatinous,[27]:159 byssoid, and leprose lichens lack a cortex (are ecorticate), and generally have only undifferentiated tissue, similar
to only having a symbiont layer.
In lichens that include both green algaland cyanobacterial symbionts, the cyanobacteria may be held on the upper or lower surface in
small pustules called cephalodia.
Pruinia is a whitish coating on top of an upper surface.[34] An epinecral layer is "a layer of horny dead fungal hyphae with
indistinct lumina in or near the cortex above the algal layer".[34]
[1]
In August 2016, it was reported that macrolichens have more than one species of fungus in their tissues.
Physiology
Symbiotic relation
"Lichens are fungi that have discovered agriculture" —Trevor Goward[35]
A lichen is a composite organism that emerges from algae or cyanobacteria living among the filaments (hyphae) of two fungi in a
mutually beneficial symbiotic relationship. The fungi benefit from the carbohydrates produced by the algae or cyanobacteria via
photosynthesis. The algae or cyanobacteria benefit by being protected from the environment by the filaments of the fungi, which also
gather moisture and nutrients from the environment, and (usually) provide an anchor to it. Although some photosynthetic partners in
a lichen can survive outside the lichen, the lichen symbiotic association extends the ecological range of both partners, whereby most
descriptions of lichen associations describe them as symbiotic. However, while symbiotic, the relationship is probably not
mutualistic, since the algae give up a disproportionate amount of their sugars (see below). Both partners gain water and mineral
nutrients mainly from the atmosphere, through rain and dust. The fungal partner protects the alga by retaining water, serving as a
larger capture area for mineral nutrients and, in some cases, provides minerals obtained from the substrate. If a cyanobacterium is
present, as a primary partner or another symbiont in addition to a green alga as in certain tripartite lichens, they can fix atmospheric
nitrogen, complementing the activities of the green alga.
The algal or cyanobacterial cells are photosynthetic and, as in plants, they reduce atmospheric carbon dioxide into organic carbon
sugars to feed both symbionts. Phycobionts (algae) produce sugar alcohols (ribitol, sorbitol, and erythritol), which are absorbed by
the mycobiont (fungus).[31] Cyanobionts produce glucose.[31] Lichenized fungal cells can make the photobiont "leak" out the
[7]:5
products of photosynthesis, where they can then be absorbed by the fungus.
The lichen combination of alga or cyanobacterium with a fungus has a very different form (morphology), physiology, and
biochemistry than the component fungus, alga, or cyanobacterium growing by itself, naturally or in culture. The body (thallus) of
most lichens is different from those of either the fungus or alga growing separately. When grown in the laboratory in the absence of
its photobiont, a lichen fungus develops as a structureless, undifferentiated mass of fungal filaments (hyphae). If combined with its
photobiont under appropriate conditions, its characteristic form associated with the photobiont emerges, in the process called
morphogenesis.[4] In a few remarkable cases, a single lichen fungus can develop into two very different lichen forms when
associating with either a green algal or a cyanobacterial symbiont. Quite naturally, these alternative forms were at first considered to
be different species, until they were found growing in a conjoined manner.
Evidence that lichens are examples of successful symbiosis is the fact that lichens can be found in almost every habitat and
geographic area on the planet.[13] Two species in two genera of green algae are found in over 35% of all lichens, but can only rarely
be found living on their own outside of a lichen.[36]
In a case where one fungal partner simultaneously had two green algae partners that outperform each other in different climates, this
might indicate having more than one photosynthetic partner at the same time might enable the lichen to exist in a wider range of
habitats and geographic locations.[13]
Phycobionts can have a net output of sugars with only water vapor.[31] The thallus must be saturated with liquid water for
cyanobionts to photosynthesize.[31]
Algae produce sugars that are absorbed by the fungus by diffusion into special fungal hyphae called appressoria or haustoria in
contact with the wall of the algal cells.[37] The appressoria or haustoria may produce a substance that increases permeability of the
algal cell walls, and may penetrate the walls.[37] The algae may lose up to 80% of their sugar production to the fungus.
[37]
Ecology
Lichen associations may be examples of mutualism, commensalism or even parasitism, depending on the species. There is evidence
to suggest that the lichen symbiosis is parasitic or commensalistic, rather than mutualistic.[38] The photosynthetic partner can exist in
nature independently of the fungal partner, but not vice versa. Photobiont cells are routinely destroyed in the course of nutrient
exchange. The association is able to continue because reproduction of the photobiont cells matches the rate at which they are
destroyed.[38] The fungus surrounds the algal cells,[8] often enclosing them within complex fungal tissues unique to lichen
associations. In many species the fungus penetrates the algal cell wall,[8] forming penetration pegs (haustoria) similar to those
produced by fungi that feed on a host (pathogenic fungi).[25][39] Cyanobacteria in laboratory settings can grow faster when they are
alone rather than when they are part of a lichen.
Miniature ecosystem and holobiont theory

Symbiosis in lichens is so well-balanced that lichens have been considered to be relatively self-contained miniature ecosystems in
and of themselves.[13][14] It is thought that lichens may be even more complex symbiotic systems that include non-photosynthetic
bacterial communities performing other functions as partners in aholobiont.[15][16]
Many lichens are very sensitive to environmental disturbances and can be used in cheaply[8] assessing air pollution,[40][41][42] ozone
depletion, and metal contamination. Lichens have been used in making dyes, perfumes,[43] and in traditional medicines. A few lichen
species are eaten by insects[8] or larger animals, such as reindeer.[44] Lichens are widely used as environmental indicators or bio-
indicators. If air is very badly polluted with sulphur dioxide there may be no lichens present, just green algae may be found. If the air
is clean, shrubby, hairy and leafy lichens become abundant. A few lichen species can tolerate quite high levels of pollution and are
commonly found on pavements, walls and tree bark in urban areas. The most sensitive lichens are shrubby and leafy while the most
tolerant lichens are all crusty in appearance. Since industrialisation many of the shrubby and leafy lichens such as Ramalina, Usnea
and Lobaria species have very limited ranges, often being confined to the parts with the purest air
.
Lichenicolous fungi
Some fungi can only be found living on lichens as obligate parasites. These are referred to as lichenicolous fungi, and are a different
[45]
species from the fungus living inside the lichen; thus they are not considered to be part of the lichen.
Reaction to water
Moisture makes the cortex become more transparent.[7]:4 This way, the algae can conduct photosynthesis when moisture is available,
and is protected at other times. When the cortex is more transparent, the algae show more clearly and the lichen looks greener
.
Metabolites, metabolite structures and bioactivity

Lichens can show intense antioxidant activity.[46][47] Secondary metabolites are often deposited as crystals in the apoplast.[48]
[48]
Secondary metabolites are thought to play a role in preference for some substrates over others.
Sometimes lichens contain structures made from fungal metabolites, for example crustose lichens sometimes have a polysaccharide
layer in the cortex.
Growth rate
Lichens often have a regular but very slow growth rate of less than a millimeter per year.[35] Different lichen species have been
.[49]
measured to grow as slowly as 0.5 mm, and as fast as 0.5 meter per year
In crustose lichens, the area along the margin is where the most active growth is taking place.[27]:159 Most crustose lichens grow
only 1–2 mm in diameter per year.
Life span
Lichens may be long-lived (longevity), with some considered to be among the oldest living things.[4][17] Lifespan is difficult to
measure because the definition of what constitutes the "same" individual lichen is not precise:[49] lichens grow by vegetatively
having a piece break off, which may or may not be considered to be the "same" lichen; moreover, two lichens can grow into each
other and then become the "same" lichen.[49]
Response to environmental stress

Unlike simple dehydration in plants and animals, lichens may experience a complete loss of body water in dry periods.[8] Lichens are
capable of surviving extremely low levels of water content (poikilohydric).[50]:5–6 They quickly absorb water when it becomes
available again, becoming soft and fleshy.[8] Reconfiguration of membranes following a period of dehydration requires several
minutes or more.
In tests, lichen survived and showed remarkable results on the adaptation capacity of photosynthetic activity within the simulation
time of 34 days under Martian conditions in the Mars Simulation Laboratory (MSL) maintained by the German Aerospace Center
(DLR).[51]
The European Space Agencyhas 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 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
[52][53]
lichens were brought back to earth and were found to be in full health with no discernible damage from their time in orbit.
Reproduction and dispersal
Vegetative reproduction
Many lichens reproduce asexually, either by a piece breaking off and growing on its
own (vegetative reproduction) or through the dispersal of diaspores containing a few
algal cells surrounded by fungal cells.[2] Because of the relative lack of
differentiation in the thallus, the line between diaspore formation and vegetative
reproduction is often blurred. Fruticose lichens can easily fragment, and new lichens
can grow from the fragment (vegetative reproduction). Many lichens break up into
fragments when they dry, dispersing themselves by wind action, to resume growth
when moisture returns.[54][55] Soredia (singular: "soredium") are small groups of
algal cells surrounded by fungal filaments that form in structures called soralia, from
which the soredia can be dispersed by wind.[2] Isidia (singular: "isidium") are
branched, spiny, elongated, outgrowths from the thallus that break off for
mechanical dispersal.[2] Lichen propagules (diaspores) typically contain cells from
both partners, although the fungal components of so-called "fringe species" rely Xanthoparmelia sp. with dark-colored
[56]
instead on algal cells dispersed by the "core species". reproductive structures (disc-like
apothecia) at center, surrounded by a
pale coloured vegetativethallus.
Sexual reproduction
Structures involved in reproduction
often appear as discs, bumps, or squiggly lines on the surface of the thallus.[7]:4
Only the fungal partner in a lichen reproduces sexually.[2] Many lichen fungi
reproduce sexually like other fungi, producing spores formed by meiosis and fusion
of gametes. Following dispersal, such fungal spores must meet with a compatible
algal partner before a functional lichen can form.
Some lichen fungi belong to Basidiomycetes (basidiolichens) and produce

mushroom-like reproductive structures resembling those of their nonlichenized
Disc-like apothecia (left) and thallus relatives.
(right) on a foliose lichen
Most lichen fungi belong to Ascomycetes (ascolichens). Among the ascolichens,
spores are produced in spore-producing structures called ascomata.[7] The most
common types of ascomata are the apothecium (plural: apothecia) and perithecium (plural: perithecia).[7]:14 Apothecia are usually
cups or plate-like discs located on the top surface of the lichen thallus. When apothecia are shaped like squiggly line segments instead
of like discs, they are called lirellae.[7]:14 Perithecia are shaped like flasks that are immersed in the lichen thallus tissue, which has a
[7]:14
small hole for the spores to escape the flask, and appear like black dots on the lichen surface.
The three most common spore body types are raised discs called apothecia (singular: apothecium), bottle-like cups with a small hole
at the top called perithecia (singular: perithecium), and pycnidia (singular: pycnidium), shaped like perithecia but without asci (an
ascus is the structure that contains and releases the sexual spores in fungi of theAscomycota).[57]
The apothecium has a layer of exposed spore-producing cells called asci (singular: ascus), and is usually a different color from the
thallus tissue.[7]:14 When the apothecium has an outer margin, the margin is called the exciple.[7]:14 When the exciple has a color
similar to colored thallus tissue the apothecium or lichen is called lecanorine, meaning similar to members of the genus
Lecanora.[7]:14 When the exciple is blackened like carbon it is called lecideine meaning similar to members of the genus
Lecidea.[7]:14 When the margin is pale or colorless it is calledbiatorine.[7]:14
A "podetium" (plural:podetia) is a lichenized stalk-like structure of the fruiting body

rising from the thallus, associated with some fungi that produce a fungal
apothecium.[24] Since it is part of the reproductive tissue, podetia are not considered
part of the main body (thallus), but may be visually prominent.[24] The podetium
may be branched, and sometimes cup-like. They usually bear the fungal pycnidia or
apothecia or both.[24] Many lichens have apothecia that are visible to the naked
eye.[2]
Most lichens produce abundant sexual structures.[58] Many species appear to

disperse only by sexual spores.[58] For example, the crustose lichensGraphis scripta
and Ochrolechia parella produce no symbiotic vegetative propagules. Instead, the
Crust-like thallus with pseudopodetia
lichen-forming fungi of these species reproduce sexually by self-fertilization (i.e.
they are homothallic). This breeding system may enable successful reproduction in
harsh environments.[58]
Mazaedia (singular: mazaedium) are apothecia shaped like a dressmaker's pin in (pin lichen)s, where the fruiting body is a brown or
[7]:15
black mass of loose ascospores enclosed by a cup-shaped exciple, which sits on top of a tiny stalk.
Taxonomy and classification

Lichens are classified by the fungal component. Lichen species are given the same scientific name (binomial name) as the fungus
species in the lichen. Lichens are being integrated into the classification schemes for fungi. The alga bears its own scientific name,
which bears no relationship to that of the lichen or fungus.[59] There are about 13,500–17,000 identified lichen species.[37] Nearly
[37]
20% of known fungal species are associated with lichens.
"Lichenized fungus" may refer to the entire lichen, or to just the fungus. This may cause confusion without context. A particular
fungus species may form lichens with different algae species, giving rise to what appear to be different lichen species, but which are
[60]
still classified (as of 2014) as the same lichen species.
Formerly, some lichen taxonomists placed lichens in their own division, theMycophycophyta, but this practice is no longer accepted
because the components belong to separate lineages. Neither the ascolichens nor the basidiolichens form monophyletic lineages in
their respective fungal phyla, but they do form several major solely or primarily lichen-forming groups within each phylum.[61] Even
more unusual than basidiolichens is the fungus Geosiphon pyriforme, a member of the Glomeromycota that is unique in that it
encloses a cyanobacterial symbiont inside its cells. Geosiphon is not usually considered to be a lichen, and its peculiar symbiosis was
not recognized for many years. The genus is more closely allied toendomycorrhizal genera.
.[62]
Lichens independently emerged from fungi associating with algae and cyanobacteria multiple times throughout history
Fungi
The fungal component of a lichen is called the mycobiont. The mycobiont may be an Ascomycete or Basidiomycete.[11] The
associated lichens are called either ascolichens or basidiolichens, respectively. Living as a symbiont in a lichen appears to be a
[63]
successful way for a fungus to derive essential nutrients since about 20% of all fungal species have acquired this mode of life.
Thalli produced by a given fungal symbiont with its differing partners may be similar, and the secondary metabolites identical,
indicating that the fungus has the dominant role in determining the morphology of the lichen. But the same mycobiont with different
photobionts may also produce very different growth forms.[60] Lichens are known in which there is one fungus associated with two
or even three algal species.
Although each lichen thallus generally appears homogeneous, some evidence seems to suggest that the fungal component may
consist of more than one genetic individual of that species.
[64]
Two or more fungal species can interact to form the same lichen.
The following table lists theorders and families of fungi that include lichen-forming species.
Photobionts
The photosynthetic partner in a lichen is called a photobiont. The photobionts in lichens come from a variety of simple prokaryotic
and eukaryotic organisms. In the majority of lichens the photobiont is a green alga (Chlorophyta) or a cyanobacterium. In some
lichens both types are present. Algal photobionts are called phycobionts, while cyanobacterial photobionts are called
cyanobionts.[31] According to one source, about 90% of all known lichens have phycobionts, and about 10% have cyanobionts,[31]
while another source states that two thirds of lichens have green algae as phycobiont, and about one third have a cyanobiont.[25]
Approximately 100 species of photosynthetic partners from 40[31] genera and five distinct classes (prokaryotic: Cyanophyceae;
[65]
eukaryotic: Trebouxiophyceae, Phaeophyceae, Chlorophyceae) have been found to associate with the lichen-forming fungi.
Common algal photobionts are from the genus Trebouxia, Trentepohlia, Pseudotrebouxia, or [[Myrmecia (alga)]|Myrmecia].
Trebouxia is the most common genus of green algae in lichens, occurring in about 40% of all lichens. "Trebouxioid" means either a
photobiont that is in the genus Trebouxia, or resembles a member of that genus, and is therefore presumably a member of the class
Trebouxiophyceae.[24] The second most commonly represented green alga genus is Trentepohlia.[25] Overall, about 100 species of
eukaryotes are known to occur as photobionts in lichens. All the algae are probably able to exist independently in nature as well as in
the lichen.[64]
A "cyanolichen" is a lichen with a cyanobacterium as its main photosynthetic component (photobiont).[66] The most commonly
occurring cyanobacterium genus is Nostoc.[64] Other[25] common cyanobacterium photobionts are from Scytonema.[11] Many
cyanolichens are small and black, and have limestone as the substrate. Another cyanolichen group, the jelly lichens of the genera
Collema or Leptogium are gelatinous and live on moist soils. Another group of lar
ge and foliose species including Peltigera, Lobaria,
and Degelia are grey-blue, especially when dampened or wet. Many of these characterize the
Lobarion communities of higher rainfall
areas in western Britain, e.g., in the Celtic rain forest. Strains of cyanobacteria found in various cyanolichens are often closely related
to one another.[67] They differ from the most closely related free-living strains.[67]
The lichen association is a close symbiosis. It extends the ecological range of both partners but is not always obligatory for their
growth and reproduction in natural environments, since many of the algal symbionts can live independently. A prominent example is
the alga Trentepohlia, which forms orange-coloured populations on tree trunks and suitable rock faces. Lichen propagules (diaspores)
typically contain cells from both partners, although the fungal components of so-called "fringe species" rely instead on algal cells
dispersed by the "core species".[56]
The same cyanobiont species can occur in association with different fungal species as lichen partners.[68] The same phycobiont
species can occur in association with different fungal species as lichen partners.[31] More than one phycobiont may be present in a
single thallus.[31]
Although each lichen thallus generally appears homogeneous, some evidence seems to suggest that the photobiont component may
consist of more than one genetic individual of that species. A single lichen may contain several algal genotypes.[69][70] These
multiple genotypes may better enable response to adaptation to environmental changes, and enable the lichen to inhabit a wider range
of environments.[71]
Controversy over classification method and species names

There are about 20,000 known lichen species.[11] But what is meant by "species" is different from what is meant by biological
species in plants, animals, or fungi, where being the same species implies that there is a common ancestral lineage.[11] Because
lichens are combinations of members of two or even three different biological kingdoms, these components must have a different
ancestral lineage from each other. By convention, lichens are still called "species" anyway, and are classified according to the species
of their fungus, not the species of the algae or cyanobacteria. Lichens are given the same scientific name (binomial name) as the
fungus in them, which may cause some confusion. The alga bears its own scientific name, which has no relationship to the name of
the lichen or fungus.[59]
Depending on context, "lichenized fungus" may refer to the entire lichen, or to the fungus when it is in the lichen, which can be
grown in culture in isolation from the algae or cyanobacteria. Some algae and cyanobacteria are found naturally living outside of the
lichen. The fungal, algal, or cyanobacterial component of a lichen can be grown by itself in culture. When growing by themselves,
the fungus, algae, or cyanobacteria have very different properties than those of the lichen. Lichen properties such as growth form,
physiology, and biochemistry, are very different from the combination of the properties of the fungus and the algae or cyanobacteria.
The same fungus growing in combination with different algae or cyanobacteria, can produce lichens that are very different in most
properties, meeting non-DNA criteria for being different "species". Historically, these different combinations were classified as
different species. When the fungus is identified as being the same using modern DNA methods, these apparently different species get
reclassified as the same species under the current (2014) convention for classification by fungal component. This has led to debate
about this classification convention. These apparently dif .[2][60]
ferent "species" have their own independent evolutionary history
There is also debate as to the appropriateness of giving the same binomial name to the fungus, and to the lichen that combines that
fungus with an alga or cyanobacterium (synecdoche). This is especially the case when combining the same fungus with different
algae or cyanobacteria produces dramatically different lichen organisms, which would be considered different species by any measure
other than the DNA of the fungal component. If the whole lichen produced by the same fungus growing in association with different
algae or cyanobacteria, were to be classified as dif
ferent "species", the number of "lichen species" would be greater
.
Diversity
[59] Some of
The largest number of lichenized fungi occur in the Ascomycota, with about 40% of species forming such an association.
these lichenized fungi occur in orders with nonlichenized fungi that live as saprotrophs or plant parasites (for example, the Leotiales,
Dothideales, and Pezizales). Other lichen fungi occur in only five orders in which all members are engaged in this habit (Orders
Graphidales, Gyalectales, Peltigerales, Pertusariales, and Teloschistales). Lichenized and nonlichenized fungi can even be found in
the same genus or species. Overall, about 98% of lichens have an ascomycetous mycobiont. Next to the Ascomycota, the largest
number of lichenized fungi occur in the unassignedfungi imperfecti, a catch-all category for fungi whose sexual form of reproduction
has never been observed. Comparatively few Basidiomycetes are lichenized, but these include agarics, such as species of
Lichenomphalia, clavarioid fungi, such as species of Multiclavula, and corticioid fungi, such as species of Dictyonema.
Identification methods
Lichen identification uses growth form and reactions to chemical tests.
"Pd" refers to the outcome of the Pd test or is used as an abbreviation for the chemical used in the test, para-phenylenediamine.[24] If
putting a drop on a lichen turns an area bright yellow to orange, this helps identify it as belonging to either the genus Cladonia or
Lecanora.[24]
Evolution and paleontology
The fossil record for lichens is poor.[72] The extreme habitats that lichens dominate, such as tundra, mountains, and deserts, are not
ordinarily conducive to producing fossils.[72][73] There are fossilized lichens embedded in amber. The fossilized Anzia is found in
pieces of amber in northern Europe and dates back approximately 40 million years.[74] Lichen fragments are also found in fossil leaf
Miocene.[75]
beds, such as Lobaria from Trinity County in northern California, USA, dating back to the early to middle
The oldest fossil lichens in which both symbiotic partners have been recovered date to the Early Devonian Rhynie chert, about 400
million years old.[76] The slightly older fossil Spongiophyton has also been interpreted as a lichen on morphological[77] and
isotopic[78] grounds, although the isotopic basis is decidedly shaky.[79] It has been demonstrated that Silurian-Devonian fossils
Nematothallus[80] and Prototaxites[81] were lichenized. Thus lichenized Ascomycota and Basidiomycota were a component of early
Silurian-Devonian terrestrial ecosystems[82][83]
The ancestral ecological state of both Ascomycota and Basidiomycota was probably saprobism, and independent lichenization events
may have occurred multiple times.[84] In 1995, Gargas and colleagues proposed that there were at least five independent origins of
lichenization; three in the basidiomycetes and at least two in the Ascomycetes.[85] However, Lutzoni et al. (2001) indicate that
lichenization probably evolved earlier and was followed by multiple independent losses. Some non-lichen-forming fungi may have
secondarily lost the ability to form a lichen association. As a result, lichenization has been viewed as a highly successful nutritional
strategy.[86][87]
Lichenized Glomeromycota may extend well back into the Precambrian. Winfrenatia, an early zygomycetous (Glomeromycota)
lichen symbiosis that may have involved controlled parasitism, is permineralized in the Rhynie Chert of Scotland, of early Devonian
age.[88] Lichen-like fossils consisting of coccoid cells (cyanobacteria?) and thin filaments (mucoromycotinan Glomeromycota?) are
permineralized in marine phosphorite of the Doushantuo Formation in southern China. These fossils are thought to be 551 to 635
million years old or Ediacaran.[89] Ediacaran acritarchs also have many similarities with Glomeromycotan vesicles and spores.[90] It
has also been claimed that Ediacaran fossils including Dickinsonia,[91] were lichens,[92] although this claim is controversial.[93]
Endosymbiotic Glomeromycota comparable with living Geosiphon may extend back into the Proterozoic in the form of 1500 million
year old Horodyskia[94] and 2200 million year old Diskagma.[95] Discovery of these fossils suggest that fungi developed symbiotic
partnerships with photoautotrophs long before the evolution of vascular plants.
Ecology and interactions with environment
Substrates and habitats

Lichens grow in a wide range of substrates and habitats, including some of the most extreme conditions on earth. They are abundant
growing on bark, leaves, and hanging from branches "living on thin air" (epiphytes) in rain forests and in temperate woodland. They
grow on bare rock, walls, gravestones, roofs, exposed soil surfaces. They can survive in some of the most extreme environments on
Earth: arctic tundra, hot dry deserts, rocky coasts, and toxic slag heaps. They can even live inside solid rock, growing between the
grains, and in the soil as part of a biological soil crust in arid habitats such as deserts. Some lichens do not grow on anything, living
out their lives blowing about the environment.[2]
When growing on mineral surfaces, some lichens slowly decompose their substrate by chemically degrading and physically
disrupting the minerals, contributing to the process of weathering by which rocks are gradually turned into soil. While this
contribution to weathering is usually benign, it can cause problems for artificial stone structures. For example, there is an ongoing
lichen growth problem on Mount Rushmore National Memorial that requires the employment of mountain-climbing conservators to
clean the monument.
Lichens are not parasites on the plants they grow on, but only use them as a substrate to grow on. The fungi of some lichen species
may "take over" the algae of other lichen species.[8][96] Lichens make their own food from their photosynthetic parts and by
absorbing minerals from the environment.[8] Lichens growing on leaves may have the appearance of being parasites on the leaves,
but they are not. However, some lichens, notably those of the genus Diploschistes
are known to parasitise other lichens. Diploschistes muscorumstarts its development
in the tissue of a host Cladonia species.[39]:30[25]:171
In the arctic tundra, lichens, together with mosses and liverworts, make up the
majority of the ground cover, which helps insulate the ground and may provide
forage for grazing animals. An example is "Reindeer moss", which is a lichen, not a
moss.[8]
A crustose lichen that grows on rock is called a saxicolous lichen.[24][27]:159

Crustose lichens that grow on the rock are epilithic, and those that grow immersed
inside rock, growing between the crystals with only their fruiting bodies exposed to
the air, are called endolithic lichens.[23][27]:159[66] A crustose lichen that grows on
bark is called a corticolous lichen.[27]:159 A lichen that grows on wood from which
the bark has been stripped is called a lignicolous lichen.[32] Lichens that grow
immersed inside plant tissues are called endophloidic lichens or endophloidal
Lichens on a statue made of
lichens.[23][27]:159 Lichens that use leaves as substrates, whether the leaf is still on limestone on a tower ofRegensburg
the tree or on the ground, are called epiphyllous or foliicolous.[31] A terricolous Cathedral
lichen grows on the soil as a substrate. Many squamulous lichens are
terricolous.[27]:159 Umbillicate lichens are foliose lichens that are attached to the
substrate at only one point.[23] A vagrant lichen is not attached to a substrate at all, and lives its life being blown around by the
wind.
Lichens and soils

In addition to distinct physical mechanisms by which lichens break down raw stone, recent studies indicate lichens attack stone
chemically, entering newly chelated minerals into the ecology
.
The lichen exudates, which have powerful chelating capacity, the widespread occurrence of mineral neoformation,
particularly metal oxalates, together with the characteristics of weathered substrates, all confirm the significance of
lichens as chemical weathering agents.[97]
Over time, this activity creates new fertile soil from lifeless stone.
Lichens may be important in contributing nitrogen to soils in some deserts through being eaten, along with their rock substrate, by
snails, which then defecate, putting the nitrogen into the soils.[98] Lichens help bind and stabilize soil sand in dunes.[2] In deserts and
semi-arid areas, lichens are part of extensive, living biological soil crusts, essential for maintaining the soil structure.[2] Lichens have
[95]
a long fossil record in soils dating back 2.2 billion years.
Ecological interactions
Lichens are pioneer species, among the first living things to grow on bare rock or areas denuded of life by a disaster.[2] Lichens may
have to compete with plants for access to sunlight, but because of their small size and slow growth, they thrive in places where higher
plants have difficulty growing. Lichens are often the first to settle in places lacking soil, constituting the sole vegetation in some
[99] Some survive in the tough conditions
extreme environments such as those found at high mountain elevations and at high latitudes.
[100]
of deserts, and others on frozen soil of the Arctic regions.
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 (known as 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.
Lichens do not have roots and do not need to tap continuous reservoirs of water like
most higher plants, thus they can grow in locations impossible for most plants, such
as bare rock, sterile soil or sand, and various artificial structures such as walls, roofs
and monuments. Many lichens also grow as epiphytes (epi- on the surface, phyte-
plant) on plants, particularly on the trunks and branches of trees. When growing on
plants, lichens are not parasites; they do not consume any part of the plant nor
poison it. Lichens produce allelopathic chemicals that inhibit the growth of mosses.
Lichens suppress the growth of
Some ground-dwelling lichens, such as members of the subgenus Cladina (reindeer
mosses and higher plants around
lichens), produce allelopathic chemicals that leach into the soil and inhibit the
them
germination of seeds, spruce and other plants.[101] Stability (that is, longevity) of
their substrate is a major factor of lichen habitats. Most lichens grow on stable rock
surfaces or the bark of old trees, but many others grow on soil and sand. In these latter cases, 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
.
Lichens may be eaten by some animals, such as reindeer, living in arctic regions. The larvae of a number of Lepidoptera species feed
exclusively on lichens. These include Common Footman and Marbled Beauty. However, lichens are very low in protein and high in
carbohydrates, making them unsuitable for some animals. Lichens are also used by the Northern Flying Squirrel for nesting, food,
and a water source during winter.
Effects of air pollution

If lichens are exposed to air pollutants at all times, without any deciduous parts, they
are unable to avoid the accumulation of pollutants. Also lacking stomata and a
cuticle, lichens may absorb aerosols and gases over the entire thallus surface from
which they may readily diffuse to the photobiont layer.[102] Because lichens do not
possess roots, their primary source of most elements is the air, and therefore
elemental levels in lichens often reflect the accumulated composition of ambient air.
The processes by which atmospheric deposition occurs include fog and dew,
gaseous absorption, and dry deposition.[103] Consequently, many environmental
studies with lichens emphasize their feasibility as effective biomonitors of Some lichens, like the folioseLobaria
atmospheric quality.[102][104][105][106][107] pulmonaria, are sensitive to air
pollution.
Not all lichens are equally sensitive toair pollutants, so different lichen species show
different levels of sensitivity to specific atmospheric pollutants.[108] The sensitivity
of a lichen to air pollution is directly related to the energy needs of the mycobiont, so that the stronger the dependency of the
[109] Upon exposure to air pollution, the photobiont may
mycobiont on the photobiont, the more sensitive the lichen is to air pollution.
use metabolic energy for repair of its cellular structures that would otherwise be used for maintenance of its photosynthetic activity,
therefore leaving less metabolic energy available for the mycobiont. The alteration of the balance between the photobiont and
mycobiont can lead to the breakdown of the symbiotic association. Therefore, lichen decline may result not only from the
.[102]
accumulation of toxic substances, but also from altered nutrient supplies that favor one symbiont over the other
This interaction between lichens and air pollution has been used as a means of monitoring air quality since 1859, with more
systematic methods developed byWilliam Nylander in 1866.[2]
Human use
Food
Lichens are eaten by many different cultures across the world. Although some
lichens are only eaten in times of famine, others are a staple food or even a delicacy.
Two obstacles are often encountered when eating lichens: lichen polysaccharides are
generally indigestible to humans, and lichens usually contain mildly toxic secondary
compounds that should be removed before eating. Very few lichens are poisonous,
but those high in vulpinic acid or usnic acid are toxic.[110] Most poisonous lichens
are yellow.
In the past Iceland moss (Cetraria islandica) was an important human food in
northern Europe, and was cooked as a bread, porridge, pudding, soup, or salad. Wila
(Bryoria fremontii) was an important food in parts of North America, where it was
usually pitcooked. Northern peoples in North America and Siberia traditionally eat
the partially digested reindeer lichen (Cladina spp.) after they remove it from the
rumen of caribou or reindeer that have been killed. Rock tripe (Umbilicaria spp. and
Lasalia spp.) is a lichen that has frequently been used as an emer
gency food in North
America, and one species, Umbilicaria esculenta, is used in a variety of traditional
Korean and Japanese foods.
Iwatake (Umbilicaria esculenta)
gathering at Kumano in Kishū, by
Lichenometry Hiroshige II
Lichenometry is a technique used to determine the age of exposed rock surfaces

based on the size of lichen thalli. Introduced by Beschel in the 1950s,[111] the technique has found many applications. it is used in
archaeology, palaeontology, and geomorphology. It uses the presumed regular but slow rate of lichen growth to determine the age of
exposed rock.[10]:9[112] Measuring the diameter (or other size measurement) of the largest lichen of a species on a rock surface
indicates the length of time since the rock surface was first exposed. Lichen can be preserved on old rock faces for up to 10,000
years, providing the maximum age limit of the technique, though it is most accurate (within 10% error) when applied to surfaces that
have been exposed for less than 1,000 years.[113] Lichenometry is especially useful for dating surfaces less than 500 years old, as
radiocarbon dating techniques are less accurate over this period.[114] The lichens most commonly used for lichenometry are those of
the genera Rhizocarpon (e.g. the species Rhizocarpon geographicum) and Xanthoria.
Biodegradation
Lichens have been shown to degrade polyester resins, as can be seen in archaeological sites in the Roman city of Baelo Claudia in
Spain.[115] Lichens can accumulate several environmental pollutants such as lead, copper
, and radionuclides.[9]
As dyes
Many lichens produce secondary compounds, including pigments that reduce harmful amounts of sunlight and powerful toxins that
reduce herbivory or kill bacteria. These compounds are very useful for lichen identification, and have had economic importance as
dyes such as cudbear or primitive antibiotics.
The pH indicator (indicated acidic or basic) in thelitmus test is a dye extracted from the lichenRoccella tinctoria by boiling.
In the Highlands of Scotland, traditional dyes for Harris tweed[2] and other traditional cloths were made from lichens including the
orange Xanthoria parietina and the grey foliaceousParmelia saxatilis common on rocks known as "crottle".
There are reports dating almost 2000 years old of lichens being used to make purple and red dyes.[116] Of great historical and
commercial significance are lichens belonging to the family Roccellaceae, commonly called orchella weed or orchil. Orcein and
other lichen dyes have largely been replaced by synthetic versions.
Traditional medicine and research
Historically in traditional medicine of Europe, Lobaria pulmonaria was collected in large quantities as "Lungwort", due to its lung-
like appearance (the doctrine of signatures suggesting that herbs can treat body parts that they physically resemble). Similarly,
Peltigera leucophlebia was used as a supposed cure for thrush, due to the resemblance of its cephalodia to the appearance of the
disease.[25]
Lichens produce metabolites in research for their potential therapeutic or diagnostic value.[117] Some metabolites produced by
lichens are structurally and functionally similar to broad-spectrum antibiotics while few are associated respectively to antiseptic
similarities.[118] Usnic acid is the most commonly studied metabolite produced by lichens.[118] It is also under research as an
bactericidal agent againstEscherichia coli and Staphylococcus aureus.[119]
Aesthetic appeal
Colonies of lichens may be spectacular in appearance, dominating the surface of the
visual landscape as part of the aesthetic appeal to paying visitors of Yosemite
National Park and Sequoia National Park.[10]:2 Orange and yellow lichens add to the
ambience of desert trees, rock faces, tundras, and rocky seashores. Intricate webs of
lichens hanging from tree branches add a mysterious aspect to forests. Fruticose
lichens are used in model railroading[120] and other modeling hobbies as a material
for making miniature trees and shrubs.
Pine forest with Cladonia lichen

In literature ground-cover
In early Midrashic literature, the Hebrew word "vayilafeth" in Ruth 3:8 is explained
as referring to Ruth entwining herself around Boaz like lichen.[121] The tenth century Arab physician, Al-Tamimi, mentions lichens
[122]
dissolved in vinegar and rose water being used in his day for the treatment of skin diseases and rashes.
The plot of John Wyndham's novel Trouble with Lichen revolves around an anti-aging chemical extracted from a lichen.
History
Although lichens had been recognized as organisms for quite some time, it was not until 1867, when Swiss botanist Simon
Schwendener proposed his dual theory of lichens, that lichens are a combination of fungi with algae or cyanobacteria, whereby the
true nature of the lichen association began to emerge.[123] Schwendener's hypothesis, which at the time lacked experimental
evidence, arose from his extensive analysis of the anatomy and development in lichens, algae, and fungi using a light microscope.
Many of the leading lichenologists at the time, such as James Crombie and Nylander, rejected Schwendener's hypothesis because the
common consensus was that all living organisms were autonomous.[123]
Other prominent biologists, such as Heinrich Anton de Bary, Albert Bernhard Frank, Melchior Treub and Hermann Hellriegel were
not so quick to reject Schwendener's ideas and the concept soon spread into other areas of study, such as microbial, plant, animal and
human pathogens.[123][124] When the complex relationships between pathogenic microorganisms and their hosts were finally
identified, Schwendener's hypothesis began to gain popularity. Further experimental proof of the dual nature of lichens was obtained
[123]
when Eugen Thomas published his results in 1939 on the first successful re-synthesis experiment.
In the 2010s, a new facet of the fungi-algae partnership was discovered. Toby Spribille and colleagues found that many types of
lichen that were long thought to beascomycete-algae pairs were actually ascomycete-basidiomycete-algae trios.[1][125]
Gallery
Lobaria pulmonaria, tree Cladonia macilenta var.
lungwort, lung lichen, lung bacillaris 'Lipstick Cladonia'
moss; Upper Bavaria,
Germany
"Lichenes" fancifully drawn byErnst

Haeckel to emphasize his ideas of
symmetry in his Artforms of Nature,
1904
Usnea australis, a fruticose Hypogymnia cf. tubulosa with

form, growing on a tree Bryoria sp. and
branch Tuckermannopsis sp. in the
Canadian Rockies
Letharia sp. with Bryoria sp. Lobaria oregana, commonly

on pine branches near called 'Lettuce lichen', in the
Blackpine Lake, Washington Hoh Rainforest, Washington
State State
Xanthoparmelia cf. lavicola, a Map lichen (Rhizocarpon Physcia millegrana (a foliose
foliose lichen, on basalt. geographicum) on rock lichen), with an unlichenized
polypore fungus (bottom
right), on a fallen log.
Reindeer moss (Cladonia Crustose lichens on limestone Cladonia cf. cristatella, a

rangiferina) in Alta Murgia-Southern Italy lichen commonly referred to
as 'British Soldiers'. Notice the
red tips.
A crusty crustose lichen on a Lichen on a lilac bush Foliose lichens on rock
wall growing outward and dying in
the center. These lichens are
at least several decades old.
Xanthoria sp. lichen on Lecanora cf. muralis lichen on Microscopic view of lichen
volcanic rock in Craters of the the banks of the Bega canal in growing on a piece of
Moon National Monument Timișoara, Romania concrete dust.[a]
(Idaho, USA)
See also
Lichenology
Notes
a. This was scraped from a dry, concrete-paved section of a drainage ditch. This entire image covers a square that is
approximately 1.7 millimeters on a side. The numbered ticks on the scale represent distances of 230 micrometers, or
slightly less than 0.25 millimeter.
References
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macrolichens" (http://science.sciencemag.org/content/early/2016/07/20/science.aaf8287) . Science. 353: 488–92.
Bibcode:2016Sci...353..488S (http://adsabs.harvard.edu/abs/2016Sci...353..488S) . doi:10.1126/science.aaf8287(htt
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PMID 27445309 (https://www.ncbi.nlm.nih.gov/pubmed/27445309).
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Retrieved 10 October 2014.
3. Introduction to Lichens – An Alliance between Kingdoms(http://www.ucmp.berkeley.edu/fungi/lichens/lichens.html).
University of California Museum of Paleontology
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4. Brodo, Irwin M. and Duran Sharnoff, Sylvia (2001) Lichens of North America. ISBN 978-0300082494.
5. Galloway, D.J. (13 May 1999). "Lichen Glossary" (https://web.archive.org/web/20141206070827/http://www .anbg.go
v.au/glossary/webpubl/lichglos.htm). Australian National Botanic Gardens. Archived fromthe original (http://www.anb
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External links
Identification and classification

LIAS light: an online interactive identification key for the lichen species of the W
orld
Internal structure
University of California Museum of Paleontology microscopic image of cross section of crustose or squamulose
lichen
Earth Life Web – Schematic drawings of internal lichen structures for various growth forms
Miscellanea
University of Sydney lichen biology
Memorial University's NL Nature project, focusing primarily on lichens
Fungi that discovered agriculture
Societies
The British Lichen Society
By region
Lichens of Armenia
Lichens of Ireland
Lichens of North America
Pacific Northwest Fungi Online Journal, includes articles on lichens
Images
Pictures of Tropical Lichens
Lichen species found in Joshua Tree National Park
Very High Resolution Image of a Lichen Covered Rock
Cultivation
How To Grow Lichen (dead link)
How To Grow Lichen (archived)
Historic
"Lichens". New International Encyclopedia. 1905.
"Lichens". Encyclopædia Britannica(11th ed.). 1911.
"Lichens". Encyclopedia Americana. 1920.
Retrieved from "https://en.wikipedia.org/w/index.php?title=Lichen&oldid=827483386
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What are lichens composed of?

What are lichens composed of?

What are some common growth forms of lichens?

What are some common growth forms of lichens?

Lichen

A lichen is a composite organism that arises from algae or cyanobacteria living

[4][17] They are among the first living things to

Pronunciation Caloplaca thallincola grows like a

The sheen on some jelly-like gelatinous lichens is frommucilaginous secretions.[22]

Internal structure and growth forms

"Lichens are fungi that have discovered agriculture" —Trevor Goward[35]

Miniature ecosystem and holobiont theory

Metabolites, metabolite structures and bioactivity

Response to environmental stress

Reproduction and dispersal

Some lichen fungi belong to Basidiomycetes (basidiolichens) and produce

A "podetium" (plural:podetia) is a lichenized stalk-like structure of the fruiting body

Most lichens produce abundant sexual structures.[58] Many species appear to

Taxonomy and classification

Controversy over classification method and species names

Ecology and interactions with environment

Substrates and habitats

A crustose lichen that grows on rock is called a saxicolous lichen.[24][27]:159

Lichens and soils

Effects of air pollution

Lichenometry is a technique used to determine the age of exposed rock surfaces

Pine forest with Cladonia lichen

"Lichenes" fancifully drawn byErnst

Usnea australis, a fruticose Hypogymnia cf. tubulosa with

Letharia sp. with Bryoria sp. Lobaria oregana, commonly

Reindeer moss (Cladonia Crustose lichens on limestone Cladonia cf. cristatella, a

Identification and classification

This page was last edited on 24 February 2018, at 23:57.

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