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

THREAT AND PROTECTION OF FUNGI
CHANGES OF FUNGAL OCCURRENCE
Human influence on the occurrence of fungi begins already in the Neolithic period with
deforestation of the land (today, forests cover ca 30% of our territory). However, the greatest
changes came during the 20th century, when, in addition to mechanical changes, there are
significant changes in the chemistry of eco-systems, which as a result lead to changes in species
representation (including disappearance of some species) of previously unseen dimensions. Since
around the 1970s, the decline of not only individual species, but also entire systematic and
ecological groups in forest and non-forest habitats has been recorded.
Causes of fungal decline can be distinguished to natural and anthropogenous.
Natural causes are mainly two:
• climate change and weather fluctuations – uneven precipitation, more pronounced and longer dry
periods (rainfall then occurs suddenly, in torrents), sudden temperature changes and longer periods
of heat or frost, recently more frequent in Europe („continentalisation“, resp. „Mediterranisation“
of Europe) => fluctuations acting as a stress factor lead to selection => reduction of the mycelia
vitality up to disappearance of stenoecic species; there is also a distribution shift, recently
thermophilic and xerotolerant species are spreading here, and on the contrary, psychrophilous
species which require balanced humidity are receding;

• habitat loss and/or vegetation changes:
– natural succession in forest stands, change in tree species composition => can lead to
disappearance of accompanying mycorrhizal fungi (see below); such changes can be observed even in
the „most natural“ forests (in the Žofín or Boubín virgin forests, spruce and beech
gradually dominate at the expense of fir);
=> lignicolous fungi with a narrow
substrate specificity are also threatened
(for example Rhodotus palmatus bound
to dead wood of elms, which in recent
decades have been widely affected by
tracheomycosis – when the currently
dead trunks decay, Rhodotus will have
nothing to grow on and will disappear);
– overgrowth of non-forest habitats with tall herbs (grasses) and trees or shrubs (which can be a
part of natural forest regeneration), affecting steppe habitats (with occurrence of a number of
gasteroid species), already unmown meadows or ungrazed pastures (affecting, for example, fungi of
the genera Hygrocybe, Entoloma, family Clavariaceae) or overgrown mires (Geoglossaceae) => in such
cases human intervention is a suitable solution to maintain the desired conditions.
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\16_rhodotus_palmatus_eva_collakova_2006_4
8181.jpg
http://www.nahuby.sk/obrazok_detail.php?obrazok_id=48181&poradie=23
Rhodotus palmatus
Photo Eva Čolláková

The most serious anthropogenic influences which are manifested today are:
• direct destruction of localities during the construction of roads, buildings, waterworks,
conversion of meadows into fields, etc. – it acts on a local level, but in specific cases the
locality of a species, which does not occur a long way off, may be destroyed;
• changes in forest management:
– replacement of natural stands by others => change of accompanying species of fungi (there are
known cases of elimination of single localities of rare species), or change of the life strategy of
fungi (originally saprotrophs => parasites of trees weakened in conditions unnatural for them);
– "generation change", felling of mature trees and complete replacement with new planting => it has
an effect on the mycorrhizal partners or (sapro-)parasites of mature trees (more than in the forest
stand, it is manifested in places of isolated or scattered tree growth – in alleys, parks,
orchards, on dams of ponds);
– fragmentation of stands (clear-cuts) => change in climatic and vegetation conditions, a
fragmented forest holds water more difficult, possibility of fungal dispersal in the stand is also
limited (especially important for mycorrhizal species);
– use of heavy mechanisation => disruption of the surface layer of the soil and/or its compression
(„damped“ soil will „rise“ back in 4–8 years), water erosion easily follows on slopes;

– chemical treatment of stands (herbi-, insecti-, fungicides – these substances affect not only
occurrence of fungi, but potentially also humans – „false poisonings“ were reported after eating
edible mushrooms from stands treated with herbicides);
– fertilization => higher nitrogen content in the soil, changes in pH;
– the removal of litter (bedding for farm animals) is practically no longer done => it has a partly
negative effect – removal of the „food“ of fungi (limiting growth of saprotrophic fungi in
particular, which participate in its decomposition), changes in the soil structure, physical (the
litter helps to regulate moisture, reduces evaporation) and chemical properties (loss of some
elements) – but, on the contrary, it can be better for ectomycorrhizal species preferring mineral
soils;
– removal of dead wood (trunks, branches) from the forest is good from the forest protection point
of view (restriction of the „food“ for potential parasites of trees), but it leads to a reduction
of fungi – not only directly (elimination of lignicolous saprotrophs bound to wood at a certain
stage of decay), but also by worsening the humidity conditions in the growth (rotting wood retains
water, which evaporates in dry season and humidifies the air);
• management changes, especially in cases of long-term human-maintained habitats suddenly left to
natural succession:
– pastures, meadows, fields, gardens represent artificially maintained forest-free areas => if they
remain fallow, there are initial changes in the herb vegetation and then spontaneous overgrowth of
woody plants until onset of the forest => elimination of non-forest species (see above);

• environmental pollution:
– dust fallout, especially in the vicinity of industrial plants (exhalation of toxic substances,
ash, lime dust => affecting the soil pH and physical properties);
– air pollution by smoke emissions (1971, annual total for Czechoslovakia: 35 million tons, of
which 3 million tons of SO2, nitrogen oxides also have a significant share) then falling in the
form of „acidic“ precipitation (up to around pH = 2, for comparison, normal rain has pH 5.6–5.7) =>
decrease in pH and change in ratio of cations in the soil => affects fungi both directly and
through weakening of mycorrhizal partners
=> it turns out that impact on the ecosystem can be reversible, after a significant reduction in
emissions of sulphur and nitrogen oxides in the 1990s, the situation in the affected areas improved
– occurrence of some mycorrhizal species, missing in previous decades, has been again recorded in
the last three decades;
– accumulation of nitrogen compounds in the environment (nitrates from fertilizers, NOx oxides from
exhaust gases) => eutrophication => reduction of diversity, sensitive oligotrophic species
disappear (e.g. of the genera Cortinarius, Inocybe, Lactarius, Russula);
– accumulation of heavy metals (e.g. 14–36 ppm of lead measured in boletes 10 m from the road,
while the WHO norm is 0.5 ppm), radioactive isotopes, etc.;
– around salted roads, salinity also increases (NaCl, KCl, MgCl2) up to ca 50 m;

• change in the water regime (especially the groundwater level) – melioration (=> drainage),
irrigation, connection with construction activity => in particular wet habitats are threatened
(mires, springs, banks of ponds and lakes, floodplains, wet meadows);
• excessive „pressure“ of mushroom pickers in some places – not only the collection of fruitbodies
(restriction of spore dispersal), but especially massive trampling, damaging the mycelium in the
litter and surface layers of the soil (it can lead to 2–3x higher compactness of the soil in 10
years).
Although human interventions can in some cases lead to enrichment of the mycobiota in a given area
(meadow or steppe species in long-term deforested habitats, in some cases also replacement of
original forests by other stands – for example, rare finds from planted spruce stands in vicinity
of Karlštejn are known), to a greater extent they limit the occurrence of popular edible mushrooms
(chanterelles, boletes, Tricholoma species etc., see below) and many others – for example, in
Germany, around a third of macromycete species are currently considered threatened, and about a
quarter of our species are in the Red List of Macromycetes of the Czech Republic.

Ectomycorrhizal species are disappearing most markedly – the occurrence of these fungi is
completely bound to partner trees (conversely not), and it depends not only on the presence of
trees at the site, but also on their age and vitality => damage or weakening of symbiotic tree
species can reduce occurrence of „their“ fungi => changes in the mycobiota can thus signalise later
changes in the ecosystem, which may not be visible at the level of vascular plants. A really
„killer“ combination for mycorrhizal fungi, which has become a reality in many places during the
20th century, is represented by a changed spectrum of tree species and atmospheric emissions
changing the soil pH (increasing acidity).
Examples: fungi of the genera Boletus, Suillus, Gomphidius, Sarcodon, Cantharellus, Ramaria,
Gomphus, some species of other genera – Tricholoma equestre, Amanita caesarea or Lactarius volemus;
significant decline of hypogeous fungi (Tuber, Elaphomyces) was also recorded.
37_tuber_aestivum 38_elaphomyces_granulatus
Left:
Tuber aestivum; right: Elapho-myces granulatus.
http://botany.upol.cz
/atlasy/system/gallery.php
?entry=Elaphomyces

C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\31_sarcodon_imbricatus.JPG C:\Documents
and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\34_cantharellus_cibarius.JPG
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\35_tricholoma_flavovirens.JPG
C:\Documents and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\36_lactarius_volemus.JPG
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\33_gomphidius_glutinosus.JPG
Top left: Sarcodon imbricatus; centre: Boletus pinophilus; right: Tricholoma equstre;
bottom left: Cantharellus cibarius; centre: Gomphidius glutinosus; right: Lactarius volemus.
C:\Documents and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\32_boletus
pinophilus.jpg
http://
www.houbar.cz/default.aspx?show=3&text=171

Availability of a suitable substrate and its condition (wood/detritus/humus, raw/fine humus, tree
species, trunk/branches/twigs/litter, chemistry, water content) are essential for saprotrophic
species, but the overall composition of the community in the substrate, succession and mutual
interactions between organisms can also play a key role.
Saprotrophic species are not affected by changes
in ecosystems so fundamentally as mycorrhizal fungi,
but decline of the genera Morchella, Verpa, Helvella,
meadow fungi of the genus Hygrocybe, the polypore
Polyporus umbellatus, Albatrellus species or montane
fungi of the genera Hericium and Bondarzewia was
also recorded.
44_ramaria_botrytis 45_hygrocybe_miniata
MycorrhizalRamaria botrytis (left) and Gomphus clavatus (right).
Hygrocybe miniata
39_gomphus_clavatus
Photo Dan Dvořák

C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\41_morchella_esculenta.JPG C:\Documents
and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\42_verpa_bohemica.JPG C:\Documents
and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\43_helvella_lacunosa.JPG C:\Documents
and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\46_polyporus_umbellatus.JPG
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\47_albatrellus_ovinus.JPG C:\Documents
and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\48_hericium_flagellum.JPG
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\49_bondarzewia_mesenterica.JPG
Top left: Morchella esculenta; centre Helvella lacunosa; right Albatrellus ovinus; bottom left
Verpa bohemica; centre Hericium flagellum; right Bondarzewia mesenterica and Polyporus umbellatus.

Abiotic factors of the environment also play a role, for example acid rain decreases the pH => a
change in the soil reaction leads to influence on the occurrence of calciphilous species, such as
mycorrhizal Boletus satanas, Inocybe erubescens (= I. patouillardii) or saprotrophic Entoloma
sinuatum.
C:\Documents and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\51_boletus_satanas.JPG
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\52_inocybe_erubescens.JPG C:\Documents
and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\53_entoloma_sinuatum.JPG
Situation of
parasitic fungi is quite simple, especially obligate parasites (facultative ones usually with a
wider „scope“ are not so threatened) – the presence and vitality of the host organism is essential
(contrary to mycorrhizal symbionts, saproparasites commonly „welcome“ weakening of the host), and
environmental conditions and interspecies relationships also play a role (similar to previous
groups).
Top right: Entoloma sinuatum; bottom left: Boletus satanas; right: Inocybe erubescens,

Other species expand into the niches which become vacant – mycorrhizal Imleria badia (= Boletus
badius), Tylopilus felleus, Paxillus involutus, Russula ochroleuca, Lactarius turpis or Amanita
phalloides, terrestrial saprotrophic Clitocybe species, Lepista nuda, Gymnopus dryophilus and
Agaricus species, lignicolous Auricularia auricula-judae, Pycnoporus cinnabarinus and Stropharia
aeruginosa or parasitic Armillaria, Heterobasidion or Fomes species.
C:\Documents and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\55_tylopilus_felleus.JPG
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\56_paxillus_involutus.JPG
Left: Tylopilus felleus; right: Paxillus involutus.

57_russula_ochroleuca 58_amanita_phalloides 59_clitocybe_nebularis 60_lepista_nuda
61_gymnopus_dryophilus 62_agaricus_xanthoderma
Mycorrhizal fungi at the left – top: Russula ochroleuca; bottom: Amanita phalloides; other species
are saprotrophic – top centre: Clitocybe nebularis; right: Gymnopus dryophilus; bottom centre:
Lepista nuda; right: Agaricus xanthodermus.

C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\63_hirneola_auricula-judae.JPG
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\64_pycnoporus_cinnabarinus.JPG
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\65_stropharia_aeruginosa.JPG C:\Documents
and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\66_armillaria_ostoyae.JPG
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\67_heterobasidion_annosus.JPG
C:\Documents and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\68_fomes_fomentarius.JPG
Lignicolous saprotrophs or saproparasites – top left: Auricularia auricula-judae, centre:
Heterobasidion annosum; right: Pycnoporus cinnabarinus; bottom left: Fomes fomentarius; centre:
Armillaria ostoyae; right: Stropharia aeruginosa.

OPTIONS OF PROTECTION OF FUNGI
• Regulation of mushroom picking means restriction of the fruitbody collection for a certain period
of time, possibly on certain days of the week, or weight limit of collections for one mushroom
picker per day (e.g. in Switzerland – here both these methods of regulation are applied in addition
to „total protection“, the ban on the collection of all mushrooms in certain protected areas) –
this approach is only relevant for rare species (restriction of fruitbody collection => greater
chance of spore dispersal) and to some extent for reduction of trampling.
• Much more effective way is overall protection of the site (protected area – National Nature
Reserve, National Nature Monument) and related restrictions (access restricted to paths, prohibited
collection of nature specimens).
• The most effective way of protection of fungi is the protection of entire habitats – appropriate
land management is crucial (in particular, limited mechanical interventions and chemical changes,
leaving dead wood as a substrate for lignicolous fungi).

Protection of fungi in the Czech Republic was an empty concept until roughly the 1960s. In the
1970s, professional mycologists began to „sound the alarm“in connection with marked decline of some
species (especially mycorrhizal ones). Fungi are no longer perceived only as food and the object of
mushroom picking as a „national sport“, research concerning distribution of disappearing species
and inventorying in a number of protected areas has started.
In the 1990s and the 2000s, concrete steps were taken at the professional level (issue of the Red
Book and the Red List, ongoing inventory in protected areas) and practically applied (legal
protection of selected species, protection of myco-logically valuable sites, consideration of fungi
in management of some areas).
C:\Documents and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\90_chranene_houby_cr.JPG
Introduction of mushroom picking regulation (see above) would probably have a hard time in our
country, not to mention any effective control. The picking ban applies only to selected protected
species, since the issuance of the Nature Protection Act (114/92 Coll.) and the decree (395/92
Coll.) in which these species are listed; the law defines the protection of plant species (their
underground and above-ground parts) including their habitats, whereby plants are meant here
including fungi.
This publication provides an overview of protected species of Czech fungi with descriptions and
distribution maps.

According to which criteria should the species deserving protection at local or national level be
selected? At first, rare species (with a minimum absolute number of localities), relict species
(destruction of the locality would probably mean the irreversible end of its occurrence here),
endemic species (if they disappear from here, so...), or species having edge of the distribution
range or extrazonal occurrence here; conspicuous edible mushrooms are under greater threat (this
is why Amanita caesarea and Butyriboletus regius are protected in our country, even though among
other species they have significantly more localities here).
Species whose habitats are acutely threatened by economic activities deserve increased attention.
C:\Documents and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\91_boletus_regius.JPG
C:\Documents and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\92_amanita_caesarea.JPG
Left: Butyriboletus regius; right: Amanita caesarea.

There are 46 protected species of fungi in the Czech Republic, while the number of more or less
threatened ones is in the hundreds, and most of the protected species cannot be recognised by the
layman in the field (however, they are not usually picked, with the exception of a few species, as
the already mentioned Amanita caesarea, Butyriboletus regius) => the main impact of the legal
protection of fungi is „educational“ (fungi also deserve protection) and especially as a basis for
justifying the declaration of protected areas – since the 1980s, some protected areas have been
declared directly for mycological purpose (dam of the Luční pond near Tábor) or protected mainly
due to occurrence of rare fungi (Rendezvous near Valtice, Velký vrch near Vršovice in Louny
District; all three areas are today classified as National Nature Monument).
Maps taken from http://geoportal.cenia.cz/mapmaker/cenia/portal
C:\Documents and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\93_lucni_u_tabora.png
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\93_lucni_u_tabora_detail.png

C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\94_rendezvous_u_valtic.png
Maps taken from http://geoportal.cenia.cz/mapmaker/cenia/portal
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\95_velky_vrch_u_vrsovic.png C:\Documents
and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\94_rendezvous_u_valtic_detail.png
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\95_velky_vrch_u_vrsovic_detail.png

The ideal constellation for enforcing the protection of fungi and their habitats (can be applied in
general, not only for fungi) is the cooperation of „local experts“ (regular monitoring of
localities => knowledge of the occurrence of species which no „externalist“ can have),
mycologists-specialists (with knowledge, on the basis of which they can put the data on the species
occurrence into a wider context) and politicians (who apply the results of the work of experts at
the national or local level => legal regulations, protection of the localities).
As a guide for interested persons in the state administration, experts and the public, Methodology
for Species Protection of Fungi was published in 2022. It presents the fungi of important habitats
in the Czech Republic, analyses the causes of their threat and, above all, is focused on practical
options of their protection with an emphasis on appropriate management in various habitats.
The methodology is available on the website of the Ministry of Environment of the Czech Republic as
a ZIP file (60 MB), partial files and appendices (see below) on the website of the Czech Scientific
Society for Mycology.

C:\Documents and Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\96_cervena_kniha_4.JPG
A complete overview of the entire issue of protection of fungi is also provided by an excellently
prepared page on the website of the Czech Mycological Society.
In the Red Data Book, fungi were elaborated in the 4th volume (cyanobacteria, algae, fungi, lichens
and bryophytes); it was completed in 1991 (still as „federal“ Red Data Book of the Slovak Republic
and the Czech Republic), published in 1995 (Príroda publishing house, Bratislava).
A proposal of 87 species, suitable for amending the list of protected fungi in the aforementioned
decree 395/92 Coll., was prepared at the same time. These are presented in appendices of the
methodology with a detailed description, data on their ecology, current distribution, main
threatening factors and proposed level of protection.

At the pan-European level, 33 species of threatened fungi (representing their occurrence in various
habitats and different climatic conditions) were selected for inclusion in the appendix of the Bern
Convention (orig. Convention on the Conservation of European Wildlife and Natural Habitats).
C:\Documents and
Settings\Hrouda\Dokumenty\S_Vyuka\stazeno\ekolhub_ochrana\97_cerveny_seznam_makromycetu_cr.jpg
The Red List of Macromycetes of the Czech Republic was published in 2006 and, unlike the Decree
395/92 Coll. (list of 46 species) and the Red Data Book (114 species), it contains 904 species of
Czech fungi, which can be considered potentially threatened => thus it represents a valuable basis
for declaring new protected areas, but also a tool for routine decision-making about the value of
particular localities.
Note: With the exception of species of lichenised Agaricomycetes, lichenised fungi are not included
in the Red List - a separate Red List of Lichens of the Czech Republic was published for them in
2010.
Most European countries currently have their Red Lists of fungi (in varying quality and with
different amounts of species included).