FUNGAL ECOLOGY
(sometimes with special regard to macromycetes)
Fungi and their environment • Life strategies and interactions of fungi • Ecological groups of
fungi, saprotrophs (terrestrial fungi, litter and plant debris, wood substrate, etc.) • Fungal
symbioses (ectomycorrhiza, endomycorrhiza, endophytism, lichenism, bacteria, animal relationships)
• Parasitism (parasites of animals and fungi, phytopathogenic fungi, types of parasitic relations)
• Fungi in various habitats (coniferous forests, broadleaf forests, birch stands and non-forest
habitats, fungal communities)
• Fungal dispersal and distribution • Threat and protection of fungi
(the study material has not been corrected by native speaker)
PF_72_100_grey_tr ubz_cz_black_transparent
MASARYK UNIVERSITY, FACULTY OF SCIENCE
DEPARTMENT OF BOTANY AND ZOOLOGY

Wood decomposition (decay) is performed by lignicolous fungi – mainly Basidiomycota, mostly
non-gilled fungi (former order Aphyllophorales s. l.). The first in succession re saproparasites,
already colonising living trees and then able to live in dead wood; they are followed by primary
saprotrophs (they directly decompose wood mass, e.g. Fomitopsis pinicola) and secondary saprotrophs
(they take nutrients from the matter decomposed by primary saprotrophs, e.g. Pycnoporellus
fulgens).
Specific conditions can prevail in wood – still standing trunks or branches on the trees are drying
out (they have lower water potential than later, when they fall into litter) and inside the trunks
there can be up to 20 times more CO2 than O2.
The course of wood decomposition can be divided into three stages: substrate colonisation, decay
process and incorporation of its products into the soil.
fomitopsis_pinicola Pycnoporellus
Fomitopsis pinicola
http://www.mykonet.ch/images/Porlinge/fomitopsis_pinicola_rotrandiger_baumschwamm300.jpg
Pycnoporellus fulgens
http://grzyby.strefa.pl/Pycnoporellus_fulgens.html

Wood substrate contains cellulose, hemicelluloses (mixed polymers with other carbohydrates -
xylose, etc.) and lignocellulose (cellulose microfibrils coated with hemicelluloses and embedded in
lignin) – it is a rich source of carbon, but poor in other basic nutrients (C/N ratio 300:1 up to
1000:1). Lignicolous fungi are adapted to this – the degradation of lignin may be important not so
much for the nutrient recovery itself,
C:\Documents and
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but for release of covalently bound nitrogen (its deficiency inhibits proteosynthesis and hyphal
growth => degradation of cellulose is alternated by ligninolysis => nitrogen and cellulose
components are released, which are reused until storage substances are depleted around the hypha =>
further degradation of lignin and so on). Lignicolous fungi must also deal with presence of
tannins, terpenes or flavo-noids, which are toxic to fungi.
Taken from http://botany.natur.cuni.cz
/koukol/ekologiehub/EkoHub_4.ppt
Structure of wood and its components
Top: Photo of cellulosic fibrils

Drevo_strukt
Source: Anke & Weber 2006; taken from http://botany.natur.cuni.cz/koukol/ekologiehub/EkoHub_4.ppt
Like cellulose
and hemicellu-
loses, pectins are cleaved at oxygen bridge sites.

The mechanisms of lignocellulose degradation are best studied on the model species Phanerochaete
chrysosporium (see photo). Lignin has a strongly irregular structure that does not allow direct
contact with the active sites of enzymes – the oxidation of lignin is therefore allowed by free
radicals: veratryl alcohol produced by lignin peroxidase and Mn2+ produced by manganese peroxidase;
these reactions require H2O2 as an electron donor. Furthermore, lignin fragments can be directly
cleaved by laccase (interestingly: this enzyme is also secreted by plant roots) or the fragments
(e.g. syringyl, guaiacyl, p-hydroxyphenyl)
Stepeni_lig phanerochaete_chrysosporium
can become radicals capable of further cleavage of lignin. Other enzymes also participate in the
degradation of ligno-cellulose – hydrolases (cellulases, cellobiases), cutinase, pectinase,
amylase, lipase, ...
Source: Anke & Weber 2006;
taken from http://
botany.natur.cuni.cz/koukol
/ekologiehub/EkoHub_4.ppt

Lignin, a three-dimensional polymer formed by alcohol units (three types of cinnamic acid alcohols
– coumaryl alcohol, sinapyl alcohol and coniferyl alcohol, see scheme above) with admixture of
nonstructural phenolic substances (vanilla, gallic, ferulic acid; flavonols, catechin, catechol,
pyrogallol), terpenes, resins, etc., occurs not only in wood, but also in herbal parts of plant
bodies; each plant family has a specific ratio of building blocks.
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Cellulose consists of glucose units linked by a
β-(1-4)-glycosidic bond; it may be in crystalline or amorphous form.
Its breakdown involves endoglucanase, cellobiohydrolase (exocellulase) and β-glucosidase.
http://www.sigmaaldrich.com/life-science/metabolomics
/enzyme-explorer/learning-center/lysing-enzymes.html
Taken from http://
botany.natur.cuni.cz/koukol
/ekologiehub/EkoHub_4.ppt

http://forestpathology.coafes.umn.edu/microbes.htm; taken from http://botany.natur.cuni.cz/koukol
/ekologiehub/EkoHub_4.ppt
Lignicolous fungi are divided into two groups according to enzymatic equipment
– the first causes soft and brown rot, the second white rot.
• The first fungi colonising wood, especially very moist wood, cause soft rot (SR) – in principle,
it is a „steaming“ of wood at higher temperature and humidity, during which cellulose and
hemicelluloses are degraded, while lignin is only slightly disturbed (side chains are split off);
the wood breaks in a cube structure. Deuteromycota (Acremonium, Phialophora, Cephalosporium,
Doratomyces, even aquatic Tricladium) are mainly involved here, also teleomorphic Ascomycota
(Chaetomium), to a lesser extent Basidiomycota (Mucidula mucida).
C:\Documents and
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comycota_Ascomycota-html.jpg soft
http://palaeos.com/fungi/ascomycota/ascomycota.html

Top right: brown-red rot caused by Laetiporus sulphureus; bottom: brown rot caused by
Leucogyrophana mollusca.
• Pectins, cellulose and hemicelluloses are degraded by fungi which have (similarly to the previous
case) cellulase and cellobiase and cause destructive decomposition; the wood is coloured with
released lignin (which is not degraded or only minimally), resulting in brown rot (BR; also the
term red rot may be used).
brown C:\Documents and
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na_mollusca.jpg
Wood decreases in weight and volume, breaks and crumbles, often in cube structure. Brown rot is
caused e.g. by Gloeophyllum, Fomitopsis, Piptoporus, Postia, Laetiporus, Phaeolus or
Conio-phoraceae, total about 10% of lignicolous basidiomycetes.
http://forestpathology.coafes.umn.edu/microbes.htm; taken from http://botany.natur.cuni.cz/koukol
/ekologiehub/EkoHub_4.ppt
   Photo A. Lepšová, http://botanika
.bf.jcu.cz/mykologie/galerie/Basidio/Aphyllophorales/Laetiporussulphireushnil1.jpg
..../mykologie/galerie/Basidio/Aphyllophorales/Leucohygrophanamolluscahnil1.jpg

• In contrast, fungi which have the strongest enzymatic equipment, including polyphenol oxidases
(ligninolytic enzymes; lignin is degraded together with cellulose or hemicellulose), cause
corrosive decay. The wood tissue is completely decomposed, it does not disintegrate in a cube
(holes can be formed in the spring wood of the annual ring => white pocket or honeycomb rot), the
wood loses colour and white rot (WR) is formed. It is caused
mainly by basidiomycetes (Pleurotus, Phanerochaete,
Stereum, Onnia, Phellinus, Ganoderma, Daedalea,
Fomes, Trametes, Polyporus squam.), and ascomycete
Xylariaceae (Xylaria, Hypoxylon, Ustulina, Daldinia).
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etinum.jpg C:\Documents and
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olomitatus.jpg
Anna Lepšová,
http://botanika.bf.jcu.cz/mykologie/galerie/Basidio/Aphyllophorales/Fomfomentariushnil.jpg
http://botanika.bf.jcu.cz/mykologie/galerie/Basidio/Aphyllophorales/Trichaptumabietinumhnil.jpg
http://botanika.bf.jcu.cz/mykologie/galerie/Basidio/Aphyllophorales/Phellinusnigrolimitatushnil.jpg
Left to right: white rot of Fomes fomentarius, white pocket rots of Trichaptum abietinum and
Phellinus nigrolimitatus.

Sapwood rot (right) spreads from sap- to heartwood (i.e. towards trunk inside), while heartwood rot
(left) begins in the centre and can lead to cavity formation.
Even slower than in wood, bark decay takes many years; pyrenomycetes are mainly involved,
mechanically tearing the bark by their stromata.
C:\Documents and
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http://www.biolib.cz/DOC/horak-proc-je-dulezite-mrtve-drevo.pdf
Luminiscence
So far, the phenomenon of bioluminescence is only known in lignicolous basidiomycetes (Panellus,
Pleurotus, Armillaria) – it occurs when oxygen reacts with luciferin with the participation of the
enzyme luciferase. It activates DNA repair and detoxification of radicals, which are formed, for
example, during the lignin degradation. The possible ecological significance has not yet been
clarified – there are hypotheses of attracting invertebrates (to spread spores) or, conversely, a
warning for nocturnal fungicolous invertebrates.
Source: Weitz 2004;
taken from http://botany.natur.cuni.cz/koukol/ekologiehub/EkoHub_2.ppt
Bioluminiscence of Armillaria mellea cultures in the dark (expozition 16 hours).

A special case is coprophilic fungi, growing mainly on the feces of herbivores. Excrement is a
fairly ideal nutrient medium, at least initially sufficiently moist and with a neutral pH
(coprophilic fungi, unlike others, prefer a pH higher than 6), which contains carbohydrates
(oligosaccharides and structural polysaccharides), nitrogenous substances, minerals, vitamins,
fatty acids, ...
The community composition on excrement is influenced by the animal species and its food (food
selection and digestion), environmental humidity, contact with soil and surrounding vegetation,
coprophilic insects, interactions with other microorganisms. With the changing composition of the
substrate, a succession similar to that of soil saprophytes is applied here – first zygomycetes
(Mucor, Pilobolus, Piptocephalis /this genus may rather contain parasites on coprofiles/) – fungi
capable to quickly utilise simple sugars, „ending“ after their depletion), ...
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Piptocephalis3
Left: Piptocephalis (Zoopagales),
right: Pilobolus (Mucorales),
photos of sporangia.
http://www.mycolog.com
/chapter11a.htm;
http://www.uoguelph.ca
/~gbarron/MISCELLANEOUS
/pilobolu.htm (George Barron)

... followed by ascomycetes (Ascobolus, Podospora, Sordaria, Chaetomium) and basidiomycetes (first
Coprinus, later Bolbitius, Psathyrella, Panaeolus, Nidularia).
C:\Documents and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_saprofyt\51_chaetomium.jpg
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Ascobolus Coprinus Kravinec
Top left: Chaetomium, bottom left: Sordaria (both perithecia); centre: Ascobolus (bottom:
apothecia, top: cross section of apoth.);
bottom right: primordia of Coprinus,
top right: Panaeolus ovatus.
http://www.mycolog.com/chapter11a.htm
http://www.ms.mff.cuni.cz/~jhum8111/hory/all_in_one.php?path=schladminger_tauern_2006/big_pictures
http://www.idsystem.cz/mushrooms/houbymesice/2003_11/houbymesice.htm
http://www.mold-help.org/content/view/412/  .../chapter11a.htm
http://www.uoguelph.ca/~gbarron/MISCE2002/sordaria.htm

Their mycelia probably develop continuously, but fructify gradually; later-stage fungi prevail over
time due to stronger enzymatic equipment and often antibiotic action => suppression of earlier
stages (hyphal interference, known in basidio-mycetes – for example, degeneration of Pilobolus
cells and stopping the formation of sporangiophores was observed after contact with Coprinus
mycelium).
For comparison: Pilobolus was noted on horse manure after 2 days, Ascobolus after 7 days and
Coprinus after 2 weeks after defecation; the total decomposition of excrement takes about 2 months.
C:\user\Houba\Škola-obrázky\clemencon\09-78_pseudorhizae-from-mole-latrine.jpg
Pretty life of Hebeloma radicosum – in the soil it grows into mole latrines, draws nutrients from
excrements and shares them with mycorrhizal partner.
Heinz Clémençon: Cytology and Plectology of the Hymenomycetes. Bibliotheca Mycologica, vol. 199. J.
Cramer, Berlin-Stuttgart, 2004.

Coprophilic fungi tend to have dormant spores, the germination of which is stimulated by contact
with certain enzymes as they pass through the digestive tract of the animals concerned. For better
transfer to (or into) the animal, spores of some species have specialized structures (hooks, mucous
membrane cover) or developed special mechanisms of spread (long-range firing; more in the Fungal
spread chapter).
A similar group are ammonia fungi, growing on substrates rich in alkaline nitrogen sources (urea,
feces, ...).
They belong to ascomycetes (Amblyosporium botrytis, Ascobolus denudatus) and basidiomycetes
(Hebeloma vinosophyllum, Tephrocybe tesquorum).
Tephrocybe_tesquorum
Tephrocybe tesquorum
http://users.skynet.be
/bs133881/champis/tephrocybe_tesquorum_(yd)_7873.htm

Keratinophilic fungi are group of specialists, colonising keratinised parts of dead animal bodies
(horns, hooves, bird quills) – this group includes some Chytridiomycota and Ascomycota, typically
keratinophilic are representatives of the order Onygenales). In addition
to dead body parts, bird nests are
a rich source of nutrients for these
fungi.
Compared to common soil fungi
with a wider scope, specialized
coprophilic and keratinophilic
fungi have considerable substrate
specificity. On the other hand,
non-specialized zygomycete or
imperfect saprotrophs, as well as
some basidiomycetes, are also growing on excrements or animal remnants (e.g. Marasmius oreades is a
soil saprotroph with an overlap to coprophilic nutrition). The spores of these fungi are spread
directly through the air, they are usually not bound to specific way of transfer (such as the
above-mentioned endozoochory).
Fungi of the mentioned groups and so-called "postputrefaction fungi" (colonising carcasses) often
occur at places with a higher incidence of animals.
C:\Documents and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_saprofyt\71_onygena_equina.jpg
Onygena equina
Photo Lairich Rig, http://www.geograph.org.uk/photo/920334

Anthracophilic fungi (also pyrophilic, carbonic or phoenicoid /according to Phoenix/) grow in
habitats affected by fire (it can be a regular natural fire, a volcanic eruption or an artificial
fire). Ash is a specific substrate with a high pH (pH rises even in the soil under the ash due to
rain leaking) with a large amount of minerals, which are often insoluble in water. The substrate is
sterilised by the fire => there is a lower competition of other saprotrophs, anthracophilic species
are competitively promoted; in some species, higher temperature stimulates the germination of
spores.
The environment, in which they live, usually creates optimal conditions for specialised fungi to
grow, develop and form competitive relationships; however,
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when competitors are eliminated, their optimum may be different. An example is Rhizina undulata,
a fungus associated with burnt places, not successful elsewhere in an undisturbed ecosystem; when
the ecosystem is disturbed, e.g. by surface fire, it becomes a facultative parasite of spruce.
Rhizina undulata
Photo Jaroslav Malý, http://www.nahuby.sk/obrazok
_detail.php?obrazok_id=23274&next_img_type=gallery

Some anthracophilic species, which produce fruitbodies in burnt places, appear to be
ectomycorrhizal (Geopyxis carbonaria); it is possible that fructification in this case is a
reaction to the loss or weakening of the partner tree by fire.
Geopyxis carbonaria
Photo Tomáš Chaluš, http://www.nasehouby.cz
/houby/taxon_list.php?taxon=genus&key=Geopyxis
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Muscicolous Arrhenia retiruga
Photo Petr Bílek, http://www.nahuby.sk/obrazok_detail.php?obrazok_id=145658
In addition to the above-mentioned ones, there are a number of partial or „complemen-tary“ groups
of saprotrophic fungi specialised in certain substrates, such as dying mosses (muscicolous fungi),
Sphagnum (sphagni-colous fungi), peat (turficolous fungi) , ...
C:\Documents and
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... dead parts of herbs (herbicolous fungi), grasses and sedges (graminicolous fungi), conifer
cones (strobilicolous fungi), plant fruits (fructicolous fungi), exudates on/near the leaf surface
(already mentioned phylloplane / phyllosphere fungi) or on/near the root surface (rhizoplane /
rhizosphere fungi, or in the case of mycorrhizal roots there are mycorrhizosphere fungi).
With the involvement of molecular methodology (analysis of environmental samples, in addition to
the „classical“ approach of monitoring substrate affinity and succession based on observation of
macromycete fruitbodies, or isolation of micromycetes), unexpected species (which deviate from the
expected groups) are detected on various substrates – therefore the question arises, to what extent
can the above-mentioned traditional categories still be applied?
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Left: strobilicolous Auriscalpium vulgare
Photo Igor Kramár, http://www.nahuby.sk/obrazok_detail.php?obrazok_id=138522
Right: fructicolous Lanzia echinophila
Photo Braňo Ivčič, http://www.fotonet.sk/?idi=4138
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REPRESENTATION OF SAPROTROPHS IN SYSTEMATIC GROUPS
• Oomycetes – mainly aquatic species (Leptomitales) are saprotrophic.
• Chytridiomycetes – saprotrophic fungi are celulolytic, or some keratinophilic (stripped
snakeskin, crustacean exoskeleton).
• Zygomycetes – true saprophytes belong to Mucorales; they use proteases, lipases and amylases, few
are cellulo- or chitinolytic.
Compared to other groups, they are not so abundant, but often play a pioneering role („R“
strategists, „sugar fungi“).
Some of them are secondary saprotrophs
(e.g.Mucor hiemalis grows on decaying wood and
utilises products of basidiomycete decomposition).
Leptomitus lacteus
Photo Dirk Klos, http://protist.i.hosei.ac.jp/pdb/Galleries
/Klos/Bavaria/Leptomitus_1.html
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Mucor hiemalis
http://www.bcrc.firdi.org.tw/fungi
/fungal_detail.jsp?id=FU200802070045

• Ascomycetes and imperfect fungi – the largest number of soil fungi (species of the genera
Aspergillus, Penicillium, Trichoderma, Fusarium, etc.) are mainly cellulolytic (some of them
degrade lignin, some keratin).
Many of them compete against other fungi by producing antibiotics; tolerance to antibiotics of
other species is just as important.
Some ascomycetes also decompose wood – either they are normally soil fungi, which colonise e.g.
twigs in the soil (only cellulose component, on the surface), or they are true lignicolous fungi,
causing white rot (cellulo- and ligninolytic – e.g. Helotiaceae, Bulgaria, Xylariaceae).
Among all fungi, Ascomycota (resp. Deuteromycota, represented especially by anamorphs of sac fungi)
have probably the highest tolerance to extreme environmental factors, such
as temperature, drought,
UV radiation (melanin in the
spore walls of Alternaria or
Cladosporium).
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Left: Bulgaria inquinans, apothecia
http://houby.humlak.cz
/popis.htlm/bulgaria_inquinans_popis.htm
Right: Cladosporium herbarum
http://ww5.stlouisco.com
/doh/pollen_site/MoldInfo.html

• Basidiomycetes usually come in the final stages of succession, they are „C“ strategists with
long-lasting mycelium. In particular, they are cellulo- and ligninolytic (brown or white rot, see
above).
• A special form is represented by yeasts, which we encounter mainly on the surface of animal and
plant bodies, where they can have enough nutrients from exudates rich in carbohydrates. The
spherical shape is advantageous for survival – there is a risk of lack of water or, conversely, the
need to balance the osmotic pressure during heavy rainfall (flushing of other substances =>
hypotonic environment), exposure to considerable lighting, etc. – together with yeasts, this
environment is also used by bacteria (mostly spherical as well).
• Slime moulds feed phagotrophically, but in the plasmodial phase they are also able to absorb
nutrients (using extracellular enzymes to break down macromolecules, such as cellulose or chitin),
i.e. partially saprotrophic nutrition.
Rot_WB
Trametes hirsuta (+ white rot)
Source: Anke & Weber 2006; taken from http://
botany.natur.cuni.cz/koukol
/ekologiehub/EkoHub_4.ppt
Plasmodium of Fuligo sp.
http://www.environment
.gov.au/biodiversity
/abrs/publications/fungi
/fuligo-plasmodium.html
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