larger fungi diversity of ombrotrophic bogs (west siberia)

advertisement
THE DIVERSITY OF LARGER FUNGI OF OMBROTROPHIC BOGS (WEST
SIBERIA)
Filippova N.V.1,
1
Yugra State University, (Khanty-Mansiysk, Russia)
filippova.courlee.nina@gmail.com
KEY WORDS: Ombrotrophic bog, boreal peatlands, larger fungi, bog fungal community, West Siberia
Abstract
Ombrotrophic bogs are ecosystems with specific characteristics of biogeochemical cycling and
biological community. Understanding element balance in these ecosystems is arresting question.
Ombrotrophic bogs make large part of peatlands in boreal zone and substantial part of these
ecosystems in West Siberia. Functioning of ecosystem depends of it’s living components, which
grouped in producing and decomposing blocks. The last includes such groups as bacteria,
archea, protists, fungi and others; fungi considered to be main decomposers in intrinsic to bogs
acidic environment.
Larger fungi is an arbitrary devition, outlined generally by visible by naked eye fruitbodies,
inventory of this group implies methods of direct observation. They are mainly basidiomycota,
some aso- and zygomycota, from functional perspective they include EM fungi, saprotrophs of
plant litter, and wood, rarely parasites; saprotrophic species include many operating with
complex polymers.
Fungal community of peatlands described in many publications, all groups of fungi and fungilike organisms in different peatland types were under investigation, though methods of
cultivation prevailed and since lack of knowledge on not-cultivated species.
Our research revealed about 50 species of larger fungi at ombrotrophic bogs by opportunistic
inspection over the course of 5 years. Locations are spread in the radius of 150 km through
middle and north taiga subzones of West Siberia. Absence of singular findings may say about
stability of the list; this number close with other published lists of bog macrofungi. There are two
distinguished habitats which correspond to different macromycetes species in ombrotrophic
bogs: treed bogs with low water level, and graminoid – sphagnum watered hollows. There are
three general functional groups: ectomycorrhizal species (15 from the list), saprotrophs of
sphagnum (30), parasites (6), and some miscellaneous (wood inhabiting fungi were not included
in this report). Species could be previously assigned to oligotrophic and mesotrophic on the
basis of occurrence in central and lag-zone of bog. Analysis of ecology of species from the
literature shows that only several species registered only from bogs; most of species occur in
related (in any factors) bogged habitats with sphagnum. Most of species found reported from
some of European fungal Red Lists, and three species were included in Red List of KhantyMansiysk region (KHMAO).
Peatland background
Ombrotrophic bog considered one of three general classes of peatlands (others two – poor and
rich fens). These clases devided by the difference in water supply. Ombrotrophic bogs form at
the relatively pour mineral substrates overlapped by impermeable layers of peat at the bottom,
since main source of water and dissolved minerals is from the atmosphere. Ombrotrophic bogs
reported as being relatively similar across the boreal zone with relative unity among the national
systems of classification (Vitt, 2006; Charman, 2002). Their range cover “southern part of
Scandinavia, Britain and the plains of the Netherlands, Germany and through into Poland, and
the Baltic states. In North America, the Atlantic provinces of Canada, the north-eastern part of
the USA and parts of the Pacific coast” (Charman, 2002).
Bogs characterizes by sphagnum – dominated vegetation, as opposed to brown moss dominated
in rich fens; between these two types are poor fens where mesotrophic sphagna prevail in ground
layer. Since ground layer is quite similar across continents, it helps to the unification of national
classifications. Several species of dwarf shrub – plant layer occur elsewhere as well, such as
Chamaedaphne calyculata, Ledum palustre, Vaccinium oxycoccus, V. vitis-idaea, Rubus
chamaemorus. Though many plants occur with more restricted ranges, such as: Picea mariana
dominates in North America, Pinus sylvestris in boreal Europe and Western Asia, and species of
Larix in eastern Asia. In dwarf shrub – plant layer, Ledum groenlandicum characterizes bogs of
North America, and Calluna vulgaris and Erica tetralix of Eurasia (Vitt, 2006).
Ombrotrophic bogs are most common peatland type in the fSU (former Soviet Union). Bog area
cover about 45 % of the total fSU peatland (Botch et al., 1995). They described as “very uniform
in vegetation pattern and ecological conditions throughout the enormous area from the Baltic
coast in the west to the continental part of Central Siberia, where bogs are replaced by very
similar larch bogs on permafrost. Southward, bogs are covered with more or less dense stands of
pines, while the shrub and grass layers change very little” (Botch, Masing, Läänelaid, 2010).
In West Siberia, peatlands have quite long research history, first publications being appeared at
the beginning of twentieth century (detailed description of the history of research done by Liss et
al., 2001). One of the generalizations was a “Wetland typology map” (Romanova et al., 1977),
where the whole of Western Siberia is divided into 20 wetland types and complexes at a 1:2500
000 scale. Recently, this map was refined to estimate areas covered by different types, their
biomass and NPP (Peregon et al., 2009). According to this estimation, area covered by wetlands
in taiga zone are about 30%.
Detailed description of peatlands of West Siberia provided by Liss et al. (2001). Three - levelled
classification was applied based on hydrochemistry, dominated plant forms (trees, plants,
mosses) and dominated species of plants. The smallest unit of classification is “biogeocoenosis
type” (corresponds to microlandscape by Ivanov, 1981). For oligotrophic peatlands Liss et al.
describe 20 types of biogeocoenoses in West Siberia. We will not provide description of all them
since our study was not intended to trace how (if only) macrofungal community corresponds to
these types (see methods). From our data, macrofungal community corresponds to the presence
of tree layer for mycorrhizal species and the level of water for sphagnum saprotrophs. Since we
provide two generalized descriptions, one for treed habitats and the other for graminoid –
sphagnum hollows.
Treed oligotrophic bogs are widespread, they have local name “ryam”. They form at most dry
surfaces of bogs, at the edges or at slopes and convex parts, and differ slightly in north and south
subzones by dominated plants. Well presented acrotelm layer provides conditions for tree growth
(two species, Pinus sylvestris and Pinus sibirica dominate). Pinus sibirica grows up to 3-10 m
high, with trunk diameter 8-10 (20) cm; Pinus sylvestris grows in stunted form (Pinus sylvestris
f. litwinowii has single trunk about 3-7 cm in diameter, and 4-5 m high). Dwarf shrubs layer
development depends of saturation level, and species composition changes depending on water
too. Ledum palustre dominate in dry conditions, more rarely with Chamaedaphne calyculata.
Andromeda polifolia is indicator of high water level. Betula nana occurs rarely, Vaccinium
uliginosum and V. myrtillus, V. vitis-idaea grow in shadowed spots. Herbs common there:
Oxicoccus microcarpus, Eriophorum vaginatum and Rubus chamaemorus. In moss layer
dominates S. fuscum and S. angustifolium and S. magellanicum dominate at lower positions.
Under tree trunks there are common spots of green mosses (Pleurosium schreberi, Aulacomnium
palustre, Dicranum bonjeanii). Some types may include as well spots of lichens which develop
at concave locations.
Graminoid – sphagnum hollows have conditions of high water level (0-15 cm) and they are
treeless. Dwarf-shrubs layer depressed and scarce, formed by Andromeda polifolia and
Oxycoccus palustris. Graminoids and plants common there: Scheuchzeria palustris, Carex
limosa, Rhynchospora alba, Drosera anglica; Menyanthes trifoliata and Carex rostrata appear
in locations where the water flow is faster. Widespread carpet of moss developed by hydrophilic
sphagna: S. majus, S. jensenii, S. lindbergii, S. balticum and S. fallax. Small hummocks covered
by S. angustifolium and S. magellanicum.
Landscape of bogs often built by above-listed types in mosaic combination, as low hummock –
hollow complex, high hummock – hollow complex, and quite widespread ridge – hollow
complex. The last is formed at slopes and it’s structure depends of hydrology and surrounding
elements of landscape.
Fungi background
Larger fungi (synonymous macrofungi, macromycetes) is an arbitrary division, which have some
differences between authors. Macrofungi distinguished by having fruiting structures (sporocarps)
visible by the naked eye (Lodge et al., 2004, Mueller et al., 2007). The group united by being
studied with the same method (direct observation) in contrast to the microfungi, to which
cultivation on different substrates is needed (Arnolds, 1981). Though Rossman et al. (1998)
specify that direct observations occur at two scales: in the field, and microscopically under a lens
in the laboratory. Such, Nordic Macromycetes issue includes groups among ascomycetes which
could be collected rather by lens than by naked eye (such as Hyaloscyphaceae, Dermateaceae
etc.). So, to refine the term when embarking to work with macrofungi, one should list the goups
involved, as Arnolds (1981) exampled.
Macrofungi include most (but not all) basidiomycetes, but also ascomycetes, and some
zygomycetes (Kirk, 2008; Lodge et al., 2004). As reported by Mueller et al. (2007), macrofungi
putatively comprise 10% of total fungal diversity and ratio of macromycetes species to vascular
species predicted 5:1 for tropical and 2:1 for temperate regions. Most terrestrial macrofungi are
saprobes or mycorrhizal symbionts, but some are pathogens of plants, animals, or fungi (Lodge
et al., 2004).
Inventory of fungi (and macrofungi) in taiga zone of West Siberia is ongoing subject. About 50
works have been published since 50 year period and about 500 species listed. There are no
published checklist or flora for the area. For the inventory work first reference books are Nordic
Macromycetes (Hansen et al., 1992, 1997, 2000), which describes flora of regions with similar
climate (through temperate to alpine zones of North Europe). Area enclosed in this publication
also includes high bogged regions (such as Finland, Sweden).
Research on fungal community of bogs (at wider scale – peatlands) has grown considerably over
past two decades (Thormann, 2007). Different aspects of community organization were revealed:
decomposition rates of substrates across ecological variables; successions of species through
decomposition; species richness of microhabitats and others (Thormann, 2006). It is noticed
repeatedly, that the group of fungi linked to the degradation of the complex structural polymers
(species of basidiomycetes and selected groups of ascomycetes) is underrepresented. The reason
likely it is methodical limitation, since isolation of many groups of fungi is media dependent. As
Dighton et al. (2005) summarizes, there are now three wide classes of methods helping to
characterize fungal communities and they are mutually complementary (classical methods of
direct observation, culturing and molecular/biochemical).
In addition, to collect species from sources not concerned directly with peatlands, Thorman et al.
(2007) compiled the list on the basis of reviewed mycological and ecological journals plus
selected systematic monographs, taxonomical and herbaria data bases where fungi associated
with sphagnum were reported. Total list summed up to 600 species (macrofungi and microfungi
from four fungal phyla). About a third of species (225) from this list are macromycetes, from
them 22 were recorded in this publication as well. The rest 35 species were new and could
supplement the list of fungi from peatlands.
Macromycetes communities of peatlands were described in several publications as well (Salo,
1993; Favre, 1948; Augustin and Runge, 1968; Lange, 1948; Sotek and Stasinska, 2010;
Stasinska et al., 2005; Roberts et al., 2004; Chinan, 2010; Chinan, 2011, Kalamees, 1982;
Kalamees and Raitviir, 1982). Only part of these publications were available to us yet. Analysis
shows weak similarity between all lists apparently due to the difference of ecosystems.
Roberts et al. (2004) describes macrofungi from sphagnum bogs from Vouncouver island. The
trophic state of these bogs must be higher due to the oceanic influence, tree species are different
as well. From 73 species presented in Roberts’ list only 3 common with list in the paper. Chinan
(2010, 2011) describes macromycetes flora of two sites in Eastern Carpathians, where 41 and 69
species (12 and 14 are shared with our list) reported. These bogs have tree layer from Picea
obovata and Betula pendula, since trophic state must be richer there as well. Sotek (2010) and
Stasinska et al. (2005) revealed macromycetes lists from two bogs in Poland, with 78 and 58
species (the lists were not available to us).
As said before, one of generalised publications about macromycetes of boreal countries is Nordic
Macromycetes (and Funga Nordica, next edition). It could be representative source of bog
inhabiting species, given high peatland areas of main countries included. We used key words
(bog, sphagnum, peat, and names of common plants) to draw species list. All list included
about100 species (but only 35 with “bog” key word). Total list shares 30 species with list in this
report.
In Russia many works were done to reveal micromycetes community of different peatlands (only
several examples being: Golovchenko, 2002; Grum-Grzhimaylo, 2012; Polyakova et al, 2001).
To our knowledge, there was not research on macrofungi of bogs in Russia before.
Study site
West Siberian plain covers about 2,5 million km² , placed in central part of Asian continent from
the Ural Mountains in the west to the Yenisey river in the east. Climate and vegetation changes
here from steppe to tundra forming near latitudinal zones. Taiga zone cover most of the area in
central part, being subdivided into three sub-zones (sought, middle, and north taiga). Area has
characteristic flat relief, with differences not more then 100 m under see level, geological surface
presented mostly by alluvial deposits. Climate subarctic, with wet warm summer and quite
severe snowy winters. General climate characteristics near Khaty-Mansiysk: average
temperature of coldest month (January) about -20° C (from 3 to -53), warmest month (July) 17°
C (from 37 to 1). Winter lasts for 26 weeks, and snow lays for 210 days, depth of snow being
about 80 cm. Total year sun radiation near 3300 KDj/m2. Mean sum of precipitation about 500
mm, this prevail sum of evaporation for about 100-150 mm, so precipitation-evaporation ratio
about 1,5. When comparing different climate characteristics, taiga zone of WS have most
analogy with several territories in Canada (Manitoba, Saskachevan, Alberta) (Dikunez et al.,
2004).
The Western Siberian plain is an extremely paludified area. Complex factors were influencing
peatland development here over the coarse of Holocene period (5000-10000 years). Peatlands
cover 50–75% of its area now and contain 40% of the global peat deposits by estimation of
Peregon et al. (2009).
Points of the study lay in taiga zone (north and middle subzones), being separated from each
other in the radius about 150 km. They are in vicinities of towns (approximate geographical
coordinates of visited locations in parenthesis): Noviy Urengoy (N 65,76 ° E 75,52°), Noyabrsk
(N63,19° E76,38°; N 63,09° E 74,3°; N 63,79° E 75,54°), Berezovskiy (N 64,23 ° E 65,79 °),
Poykovskiy (N 60,85° E 71,5°), Khanty-Mansiysk (N61,06° E69,45°; N60,88° E68,66°),
Sovetskiy (N 60,85° E 63,52°), which are in the administrative borders of Tyumen region.
Methods
Term “larger fungi” varies from author to author. Next artificial groups considered under this
term in this report: agaricoid fungi, boletoid fungi, clavarioid fungi, cup-shaped fungi, and some
miscellaneous. They are from following families: amanitaceae (1), boletaceae (1), clavariaceae
(1), cortinariaceae (9), endogonaceae (1), entolomataceae (1), exobasidiaceae (2), Geoglossaceae
(1), hydnangiaceae (1), hygrophoraceae (2), lyophyllaceae (1), marasmiaceae (2), mycenaceae
(4), physalacriaceae (1), rhytismataceae (1), russulaceae (3), sarcosomataceae (1), sclerotiniaceae
(3), strophariaceae (15), suillaceae (1), thelephoraceae (1), tricholomataceae (2).
Embarking to work with macromycetes should consider several items, which define methods of
work and restrict the final results: 1) fruit bodies are only part of fungus, which get indirect
knowledge about distribution and functioning of vegetative part; 2) species may fruit at different
seasons, so monitoring through all season must be done; 3) species may fruit not every year,
what appoint continuous monitoring; 4) identification of larger fungi depends largely on vivite
characteristics, which must be described in the field (Lodge et al., 2004). The methods of
inventory work could differ from so called “opportunistic inspection” to using protocols which
provide quantitative statistical information (Mueller et al., 2004). Opportunistic sampling it what
probably must be done at first in any case, yeilding list of species which may be envolved in
further elaboration.
Field trips to collect fungi from bogs were done over the coarse of 5 years in about 10 locations
of north and middle taiga zones of West Siberia. Most locations were forayed during several
days at earlier years, and two locations near Khanty-Mansiysk (middle taiga) were visited
regularly in vegatation season over the coarse of last two years (since there is field station of
Yugra State University).
Specimens were processed according to protocols for larger fungi collection (Lodge et al., 2004).
Type of vegetation, microhabitats and substrates were recorded for each collection. Rare and
single findings and cryptic species were collected in herbarium every time; common species
were collected in several replicas. Pictures prepared in the field and under the microscope are
available online: http://www.flickr.com/photos/bog-fun/collections/. Herbarium specimens are
stored in biodiversity collection of Yugra State University field station (not registered
collection).
To express some information about structure of diversity, speculative 4-graded scale was used: 1.
common species may be found always in related community, 2. regular species suppose to be in
community after some time of search, 3. rare species were collected several times, 4. single
findings collected once. This scale can only approximately estimate relative quantity. It is
urgently recommended to use plots and long-term monitoring to get precise quantitive
information (Arnolds, 1981).
At this stage, we were not able to work in the fungal systematics laboratory (work with herbaria
and molecular methods). Since some taxons (marked with * in checklist) must be refined in the
future. The genus Galerina presents hight diversity in bogs, but it’s systematics not worked in
contemporary literature. The same the genus Cortinarius, where several species quite reliable,
but one species was not identified, and one group (C. huronensis and similar species) also must
be verified. Several species reported in the literature as forest inhabitants, and indeed, they occur
in nearby forests, but with larger fruitbodies. Since the substrate differ as well (in bogs they are
sphagnum saprotrophs, as Gymnopus dryophilus, Gymnopilus penetrans), they probably could be
described as intraspecific categories.
In the checklist information for every species is organized in following order. Frequency and
habitat (community types) described first, then locations of samples reported. Then substrates
described according to our observations. And finaly data about ecology of species in North
Europe from the literature reported.
Checklist with annotations
Amanita porphyria Alb. & Schwein.
Rare in pine-dwarf shrubs – sphagnum communities, at hummocks and in depressions, among common sphagna and
plants. Collected three times in the north and middle taiga zones.
Forming mycorrhiza with Pinus. In North Europe reported in Pinus and Picea forests (Hansen et al., 1992). In
Switzerland in coniferous forests, primarily under Picea, Pinus, or Larix, on acid soils (Breitenbach, Kränzlin,
1995).
Armillaria ectypa (Fr.) Lamoure
Rare species, and exactly it was not collected at ombrotrophic bogs, but two times at mesotrophic communities
(poor fens) in vicinity of primary lakes (middle taiga, Sovetskiy), among S. angustifolium, S. magellanicum, Betula
pubescens and Salix lapponum. We suppose that species may be found in ombrotrophic bogs around secondary
lakes or bog streams as well.
Saprotroph of sphagnum and litter (?). In North Europe reported among fen plants and bryophytes in marshes
(Hansen et al., 1992).
Arrhenia sphagnicola (Berk.) Redhead, Lutzoni, Moncalvo & Vilgalys
Common species, it occurs in all types of bog communities in central part of bog and at lag - zone, in hollows and
hummocks, often among S. fuscum and (more rare) hydrophilic sphagna. It collected regularly in all visited locations
(north and middle zones).
Saprotroph of sphagnum. In North Europe reported on Sphagnum (Hansen et al., 1992).
Ascocoryne sarcoides (Jacq.:Fr.) Groves & D.E.Wilson
Common species in pine-dwarf shrubs – sphagnum habitats on buried in sphagnum decorticated logs of Pinus
sylvestris, where it grows nearby with it’s anamorphic stage Coryne dubia S.F.gray. Collected regularly at two
macrotops in middle taiga.
Saprotroph of wood. Reported from North Europe on recently cut stumps (Hanse, Khudsen, 2000).
Ascocoryne turficola (Boud.) Korf
Rare species, inhabits sedge – sphagnum hollows, among S. jensenii, S. majus, S. papillosum, Warnstorfia fluitans,
stems attached to buried leaves of Carex limosa (4 sightings) and branches of dwarf shrubs (once), in some locations
substrate was not registered (leave – sphagnum decaying litter). Most probably that hyphae spread in decaying
sphagnum layer, but litter of sedge and other plants initiates fructification. Found in four locations, in north taiga
(Noyabrsk) and middle taiga (Kogalym, Natural Reserve Uganskiy, Khanty-Mansiysk), in the last location several
collections from one population were done over the coarse of 3 years.
Trophic preference of species remains open to question, since substrate of collections differ, probably sphagnum –
litter saprotroph. Species has occasional reports in Europe and North America, from different types of peatlands
(fens and bogs) (Bunyard et al., 2008), in Russia reported from Leningrad and Moscow regions.
Since rare findings and preference to threatened communities, species included in Red lists of countries where was
reported; and in Red list of KHMAO region as well.
Clavaria sphagnicola Boud.
Regular species, grows in pine-dwarf shrubs – sphagnum communities, at hummocks, among S. fuscum, S.
magellanicum, S. angustifolium. Collected in north taiga (Noyabrsk), and at two macrotops in middle taiga(KhantyMansiysk).
Saptortoph of sphagnum. In North Europe reported from sphagnum bogs (Hansen, Knudsen, 1997).
*Cortinarius bataillei J. Favre
Regular species, inhabits treed bogs (pine-dwarf shrubs – sphagnum communities), at different positions in micro
relief, among common sphagna and plants, and at lag-zone. Collected from two macrotops in middle taiga, but
probably it was confused with quite similar C. huronensis in other locations.
Forming mycorrhiza with Pinus. In North Europe among mosses, often Sphagnum, on moist soil in coniferous
forests, seldom on drier soil, mycorrhizal with Picea, Pinus or Betula (Niskanen et al., 2008).
*Cortinarius flexipes (Pers.) Fr.
Regular species, grows in pine-dwarf shrubs – sphagnum communities, at hummocks and in depressions, among
common sphagna and plants. Collected many times in two macrotops in middle taiga and in one location in north
taiga.
Forming mycorrhiza with Pinus. In North Europe with deciduous trees, e.g. Fagus, Betula and Quercus, and with
conifers, preferably in moist and acid habitats (Niskanen et al., 2008).
*Cortinarius flos-paludis Melot
Regular species, grows in pne-dwarf shrubs – sphagnum communities, at hummocks and in depressions, among
common sphagna and plants. Collected many times in two macrotops in middle taiga.
Forming mycorrhiza with Pinus. In North Europe with Picea and Pinus, often in Sphagnum (Niskanen et al., 2008).
*Cortinarius huronensis Ammirati & A.H. Sm.
[C. huronensis and several similar species: C. tubarius, C. croceus, C. bataillei, all inhabiting the same
communities, need more thorough taxonomic work.]
Common species, grows in pine-dwarf shrubs – sphagnum communities, at different positions in micro relief,
among common sphagna and plants. Collected regularly in the middle and north taiga.
Forming mycorrhiza with Pinus. In North Europe among Sphagnum in fens, bogs or swamp forests, most frequent
under Pinus, more seldom Picea or Betula (Niskanen et al., 2008).
*Cortinarius obtusus (Fr.) Fr.
Common species in pine-dwarf shrubs – sphagnum communities, at hummocks and in depressions, among common
sphagna and plants. Collected regularly in the middle and north taiga.
Forming mycorrhiza with Pinus. In North Europe in mesic to damp coniferous forests with Picea, sometimes with
Pinus, Betula and Salix dwarf shrubs, in needle litter or mosses (Niskanen et al., 2008).
Cortinarius scaurus Fr. (Fr.)
Regular species in pine-dwarf shrubs – sphagnum communities, at hummocks and in depressions, among common
sphagna and plants. Collected many times in three macrotops in middle taiga and at two location in the north.
Forming mycorrhiza with Pinus. In North Europe in all kinds of coniferous forests or among Sphagnum in bogs,
rarely in deciduous forests (Hansen et al., 1992). In coniferous forests on moist, silicious soil, even in ombrotrophic
bogs (Niskanen et al., 2008).
Cortinarius semisanguineus (Fr.) Gillet
Regular species in pine-dwarf shrubs – sphagnum communities, at hummocks and in depressions, among common
sphagna and plants. Collected regularly at three macrotops in middle taiga and at one location in the north.
Forming mycorrhiza with Pinus. In North Europe on humus, needle litter or among mosses in poor coniferous
forests, often on sandy soil in open Pinus forests, rarely in other forest types, occasionally with Picea, rarely with
Betula or Fagus (Niskanen et al., 2008).
*Cortinarius sp.
Regular species in Pine-dwarf shrubs – sphagnum communities, at hummocks and in depressions, among common
sphagna and plants. Collected many times in two locations in middle taiga.
Forming mycorrhiza with Pinus.
Cortinarius stillatitius Fr.
Pine-dwarf shrubs – sphagnum communities, at hummocks and in depressions, among common sphagna and plants.
Collected many times in three macrotops in middle taiga and at one in the north.
Forming mycorrhiza with Pinus. In North Europe in Vaccinium myrtillus - Picea forests, more rarely with Pinus, in
mountain Betula forests or in Fagus forests (Niskanen et al., 2008).
Endogone pisiformis Link
Single finding comes from sedge – sphagnum hollow, in ridge – hollow complex mesotop, at the surface of
sphagnum and plant litter. Middle taiga zone.
Saprotroph on sphagnum and litter and probably forming VAM with some plants. Typically among Sphagnum, but
also at diverse plant debris in moist habitats (Berch abd Fortin, 1983).
Exobasidium cassandrae Peck
Rare in all communities where Chamaedaphne calyculata grows. Registered several times in middle taiga.
Parasite on Chamaedaphne calyculata, species parasitizes only on the one host-species (Hansen, Knudse, 1997).
Exobasidium karstenii Sacc. & Trotter
Regular in all communities where Andromeda polifolia grows. Registered several times in middle taiga.
Parasite on Andromeda polifolia, species parasitizes only on the one host-species (Hansen, Knudse, 1997).
Entoloma fuscomarginatum (Fr.) P. Kumm.
Regular species, collected at three macrotops in middle taiga zone; 14 collections near Khanty-Mansiysk (20112012); one collection comes from Natural Reserve Uganskiy in September 2012, and at three locations in the north.
Saprotroph of Sphagnum. Common and abundant in the atlantic peat-bogs of the United Kingdom, rare in central
European mountain bogs (Noordeloos, 1992).
The species included in Red List of fungi in Denmark, Netherlands (Combined European Red List), and was
included in Red List of KHMAO as well.
Galerina allospora A.H. Sm. & Singer
Regular species, collected in pine-dwarf shrubs – sphagnum communities at the edge of bog (lag - zone), rare in
central part, in small pits and hollows, among S. angustifolium, S. magellanicum, collected once on pine bark (Pinus
sylvestris). Found at two macrotops in middle taiga (Khanty-Mansiysk).
Saprotroph of Sphagnum and wood. In North Europe on mosses (Sphagnum, Dicranum, etc.), rottening stumps and
litter of Picea (Hansen et al., 1992). In North America on debris and dead sphagnum under spruce or on lightly
burned over areas in bogs (Smith and Singer, 1964).
Galerina paludosa (Fr.) Kühner
Regular species, collected in pine-dwarf shrubs – sphagnum communities at the edge of bog (lag - zone), rare in
central part, in small pits and hollows, among S. angustifolium. Also collected in mesotrophic sphagnum bogs (more
common there), and bogged forests with Sphagnum. Collected regularly in the north and middle taiga zones.
Saprotroph of Sphagnum. In North Europe in oligotrophic mires, bogs, swampy forests (Hansen et al., 1992). Smith
and Singer (1964) report it being “the most common species in the sphagnum bogs throughout the United States and
Canada, also well known in Europe, and is very likely circumpolar in distribution”.
*Galerina sphagnicola (G.F. Atk.) A.H. Sm. & Singer
[There are several similar species with calyptrate spores inhabiting in bogs, but more thorough taxonomical work
must be done with this group. Now we describe most common species from them (G. sphagnicola), differing in
larger carpophores and inhabiting in groups at wet sphagnum hollows where it presents quite constant features. In
other cases, when single fruitbodies form at dry hummocks, features become more blurred and these collections will
be analyzed later, as well as more scarce collections of potentially other species with calyptrate spores.]
Common species inhabiting in graminoid – sphagnum hollows, low hummocks, among hydrophilic common
sphagna species. Collected regularly in the north and middle taiga zone.
Saprotroph of Sphagnum. In North Europe in bogs (Hansen et al., 1992). In North America scattered to gregarious
on Sphagnum in bogs (Smith and Singer, 1964).
*Galerina sphagnorum (Pers.) Kühner
Common species in graminoid – sphagnum hollows, low hummocks, rare in pine-dwarf shrubs – sphagnum
communities among S. fuscum, S. angustifolium. Collected regularly in the north and middle taiga zone.
Saprotroph of sphagnum. Smith and Singer (1964) describe it growing on various species of Sphagnum and widely
distributed.
Galerina stagnina (Fr.) Kühner
Regular species in lag – zone between bog and forest, bogged forests, among mesotrophic sphagna, doesn’t sighted
at ombrotrophic communities. Collected in the north (3 locations) and middle taiga (4 locations) zone.
Saprotroph of Sphagnum. In North Europe on Sphagnum and other mosses (Hansen et al., 1992). Smith and Singer
(1964) describe it growing on wet mossy stream beds, often among Sphagnum riparium in North America.
Galerina tibiicystis (G.F. Atk.) Kühner
Common species in graminoid – sphagnum hollows, low hummocks, among hydrophilic common sphagna species.
Rare in pine-dwarf shrubs – sphagnum communities among S. fuscum, S. angustifolium. Rarely collected from lag –
zones and mesotrophic sphagnum bogs. Noticed in north and middle taiga zone at many locations.
Saprotroph of Sphagnum. In North Europe in mires, bogs, mossy banks of streams and lakes (Hansen et al., 1992).
Smith and Singer (1964) report that species common in sphagnum bogs of northern and mountainous regions of the
United States, Canada, Central, Eastern and Northern Europe, and Japan.
Gymnopilus fulgens (J. Favre & Maire) Singer
Rare species, collected in one location from low hummocks in pine - dwarf shrubs – sphagnum habitat, among S.
magellanicum, S. papillosum. One collection gcomes from eutrophic fen at sedge peat.
Saprotroph of Sphagnum and sedge peat. In North Europe in Sphagnum, moist peat (Hansen et al., 1992).
Gymnopilus penetrans (Fr.) Murrill
Regularly collected in pine - dwarf shrubs – sphagnum communities, at high hummocks, among S. fuscum. Several
findings in middle taiga zone.
Saprotroph of Sphagnum. In North Europe on wood of coniferous, more rarely deciduous trees (Hansen et al., 1992).
Gymnopus androsaceus (L.) J.L. Mata & R.H. Petersen
Common species, collected in pine – dwarf shrubs – sphagnum communities and in lag zones, on different litter:
Pinus sylvestris needles, branches; leaves and branches of dwarf shrubs; leaves of Eriophorum vaginatum; leaves of
Rubus chamaemorus. Black rhizomorphs were seen spreading widely at sphagnum surface and attached to buried in
sphagnum litter. Regularly collected in north and middle taiga zone.
Saprotroph of different plant litter, coniferous needles and branches, leaves and branches of dwarf shrubs. Reported
in North Europe on sticks and needles of coniferous and deciduous trees and shrubs in oligotrophic localities
(Hansen et al., 1992).
Gymnopus dryophilus (Bull.) Murrill
Common species in pine-dwarf shrubs – sphagnum communities, at hummocks, on S. fuscum, S. angustifolium, S.
magellanicum. Collected in north taiga (Beresovskiy, Noyabrsk) and in middle taiga (Sovetskiy, Khanty-Manskiysk
– many collections around these locations).
Saprotroph of Sphagnum. In North Europe in deciduous forests and alpine heaths (Hansen et al., 1992).
Hebeloma incarnatulum A.H. Sm.
Common species in pine-dwarf shrubs – sphagnum communities, at hummocks and in depressions, among common
sphagna and plants. Collected regularly in north and middle taiga zone.
Forming mycorrhiza with Pinus. Vesterholt (2005) reports the species one of the most common agaric in
Fennoscandia, among mosses, sometimes in Sphagnum, in association with Picea or Pinus in the spruce forests.
Hygrocybe cinerella (Kühner) Arnolds
Rare species, grows at low hummocks in graminoid – sphagnum hollows, among S. papillosum, S. jensenii.
Collected twice at one macrotop (middle taiga).
Saprotroph of Sphagnum. Species with rare reports in the world, recorded only from Scandinavia, Iceland,
Greenland and Siberia. In Greenland it occurs in marshes or on peat or Sphagnum (Boertmann, 1995).
Hypholoma capnoides (Fr.) P. Kumm.
Rare findings in pine-dwarf shrubs – sphagnum communities, at logs of Pinus sylvestris buried in sphagnum (in wet
conditions). Collected twice at one macrotop (middle taiga).
Saprotroph of wood. In North Europe on roots of Picea and Pinus (Hansen et al., 1992).
Hypholoma elongatum (Pers.) Ricken
Common species, grows from graminoid – sphagnum hollows in central part to lag – zone, among hydrophilic
oligotrophic and mesotrophic sphagna species. Collected regularly in middle and north taiga zone.
Saprotroph of Sphagnum. In North Europe in Sphagnum, on peaty soil, rarely in Polytrichum (Hansen et al., 1992).
Hypholoma eximium (C. Laest.) Rald
Singular finding in graminoid – sphagnum pool in central part of bog massif, location was dried after long dry
summer, since bare peat was the substrate, in middle taiga zone (Khanty-Mansiysk).
Saprotroph of Sphagnum. From North Europe collected on naked soil in snow beds (Hansen et al., 1992).
Hypholoma myosotis (Fr.) M. Lange
Common species in lag – zone between bog and forest, among mesotrophic sphagna, doesn’t sighted in
ombrotrophic communities. Was collected as well at palsa bogs in mesotrophic hollows. Collected from north and
middle taiga zone.
Saprotroph of Sphagnum and litter. In North Europe reported in Sphagnum, on naked soil in bogs and swamps,
moist acid grasslands and woodland (Hansen et al., 1992).
Hypholoma udum (Pers.) Quél.
Common species, grows from graminoid – sphagnum hollows in central part to lag – zone, among oligotrophic and
mesotrophic sphagna, in hollows and pits of pine – dwarf shrubs – sphagnum habitats. Collected regularly in middle
taiga zone.
Saprotroph of Sphagnum. In North Europe in sphagnum or peaty soil (Hansen et al., 1992).
Laccaria proxima (Boud.) Pat.
Rare species in hollows in pine – dwarf shrubs – sphagnum communities among S. angustifolium, in altered places
(along tracks) on bare peaty substrate. Collected several times at two macrotops in middle taiga zone.
Mycorrhizal (possibly facultative). In North Europe reported in moist, oligotrophic sites, often with Sphagnum
(Hansen et al., 1992).
Lactarius helvus (Fr.) Fr.
Regular species in pine – dwarf shrubs – sphagnum communities, in hollows among S. angustifolium and at
hummocks (S. fuscum). It fruits often more abundant in altered places (along bog tracks). Sighted several times at
two macrotops in middle taiga.
Forming mycorrhiza with Pinus. In North Europe on moist to wet ground under conifers and Betula, often on
disturbed soil (Hansen et al., 1992). On sandy and peaty soil, typically among Sphagnum (Heilmann-Clausen,
1998).
Lactarius rufus (Scop.) Fr.
Common species in pine – dwarf shrubs – sphagnum communities, in hollows among S. angustifolium and at
hummocks (S. fuscum). Collected regularly in middle and north taiga zone.
Forming mycorrhiza with Pinus. In North Europe mainly in coniferous forests, particularly Pinus, also in Betula
forests and arctic/alpine areas with Betula nana (Hansen et al., 1992).
Leccinum holopus (Rostk.) Watling
Rare species at ombrotrophic bogs, in Pine – dwarf shrubs – sphagnum communities, at hummocks among S.
magellanicum, S. fuscum. Forming mycorrhiza with Betula nana. But more often met in lag-zone wmon
mesotrophic sphagna and in bogged forests with Betula pubescens. From ombrotrophic bogs collected several times
at one macrotop (middle taiga).
Forming mycorrhiza with Betula nana (in central part of bog) and Betula pubescens (at lag-zone). Associated with
betula in Sphagnum bogs or among grass and mosses, on humid, peaty soil (den Bakker, 2005).
Lichenomphalia umbellifera (L.) Redhead, Lutzoni, Moncalvo & Vilgalys
Regular species in pine – dwarf shrubs – sphagnum communities, at disturbed sphagnum surface (often along bog
tracks), on S. fuscum or peaty substrates (covered with liverworts and algae). Collected several times at one location
in middle taiga.
Saprotroph of sphagnum. Reported on moss, Sphagnum, or on peat or rotten wood in North Europe, lichenised
(Hansen et al., 1992).
Monilinia oxycocci (Woronin) Honey
Regular species at all types of communities where Oxycoccus palustris grows. Sighted several times in middle taiga
zone, probably more often since regular observations of whitened affected berries.
Parasite on berries, fruiting in early spring. Forming macroconidial state on wilting host hoots (Hansen, Khudsen,
2000).
Mycena concolor (J.E. Lange) Kühner
Regular species in hollows in pine – dwarf – shrubs – sphagnum communities, among S. angustifolium, often in
altered places on peat (along bog tracks). Collected regularly in north and middle taiga.
Saprotroph of Sphagnum and peat. In North Europe reported in peatbogs, among Sphagnum (Hansen et al., 1992).
Mycena galopus (Pers.) P. Kumm.
Rare at ombrotrophic bogs, sighted in hollows in pine – dwarf – shrubs – sphagnum communities, among S.
angustifolium. Collected twice at one macrotop (middle taiga).
Saprotroph of Sphagnum. In North Europe reported from variable habitats, on dead leaves, in Sphagnum, or no burnt
places (Hansen et al., 1992).
Mycena megaspora Kauffman
Regular species, in pine – dwarf shrubs – sphagnum communities among S. angustifolium, S. fuscum, also (more
often) in lag – zone between forest and bog, among mesotrophic sphagna. Collected regularly in north and middle
taiga zone.
Saprotroph of sphagnum and ? litter. In North Europe on burnt places, on peaty ground or in Sphagnum (Hansen et
al., 1992).
Myriosclerotinia caricis-ampullaceae (Nyberg) N.F. Buchw.
Exactly, this species was not collected at ombrotrophic bogs, where C. rostrata also occurs, but at lag – zone
between bog and forest, also collected from marches (salix – sedge wetlands near lake). Collected several times in
north and middle taiga zone.
Parasite or saprotroph on Carex rostrata. Reported from North Europe on Carex rostrata and Carex aquatilis.
Conidial state produced within host culms (Hansen, Khudsen, 2000).
Myriosclerotinia dennisii (Svrček) J. Schwegler
Regular species in all communities where Eriophorum vaginatum grows. Sclerotia form inside of flower stalks,
apothecia arise in early summer. Sclerotia collected more often then fruiting was sighted, several times at two
macrotops in middle taiga.
Parasite or saprotroph on Eriophorum vaginatum. Reported from North Europe on Eriophorum, Baeothrion
caespitosum, or Eleocharis uniglumis, conidial state not observed (Hansen, Khudsen, 2000).
Omphaliaster borealis (M. Lange & Skifte) Lamoure
Regular species in all types of communities in central part of bog and at lag - zone, in hollows and hummocks,
among S. fuscum and hydrophilic sphagna. More often in pine – dwarfshrubs communities on S. fuscum. Collected
regularly in north and middle taiga zone.
Saprotroph of Sphagnum. For North Europe reported in moist, acid, boreal heaths (Hansen et al., 1992).
Pseudoplectania sphagnophila (Pers.) Kreisel
Rare species in pine – dwarf shrubs – sphagnum communities, on hummocks, among S. fuscum, S. magellanicum.
Collected several times in north and middle taiga.
Saprotroph of Sphagnum. Reported from North Europe on S. fuscum (Hansen, Knudsen, 2000).
Psilocybe atrobrunnea (Lasch) Gillet
Rare (?) species in graminoid – sphagnum hollows and pools, among hydrophilic sphagna. But there was abundant
fructification in the autumn after dry summer 2012 at two locations near Khanty-Mansiysk. Before that was
collected only once at one of these locations.
Saprotroph of Sphagnum. In North Europe in mires or other wet places (Hansen et al., 1992).
Rhytisma andromedae (Pers.) Fr.
Common species, in all types where Andromeda polifolia grows. Recorded regularly in middle taiga, epidemic was
recorded in autumn after dry summer 2012. Anamorph stage of fungus appeared abundantly on Andromeda at two
macrotops near Khanty-Mansiysk.
Parasitic on leaves of Andromeda polifolia.
Russula paludosa Britzelm.
Regular species in pine – dwarf shrubs – sphagnum communities, at hummocks and in depressions, among common
sphagna species. Sighted regularly in middle and north taiga zone.
Forming mycorrhiza with Pinus. In North Europe in coniferous forests (Hansen et al., 1992).
Sphagnomphalia brevibasidiata (Singer) Redhead, Moncalvo, Vilgalys & Lutzoni
Rare species in pine – dwarf shrubs – sphagnum communities at S. fuscum hummocks. Collected several times in
middle taiga.
Saprotroph of Sphagnum. Reported in Sphagnum in in bogs for North Europe (Hansen et al., 1992).
Suillus variegatus (Sw.) Kuntze
Regular species in pine – dwarf shrubs – sphagnum communities, at hummocks and in depressions, among common
sphagna species. Collected several times in middle taiga zone.
Forming mycorrhiza with Pinus. In North Europe in dry Pinus forests (Hansen et al., 1992).
Tephrocybe palustris (Peck) Donk
Common species of graminoid-sphagnum hollows, from central part to lag-zone, also occurs in mesotrophic bogs
and in bogged forests among mesotrophic sphagna. Collected regularly in north and middle taiga zone.
Parasite of sphagnum. Growing in sphagnum bogs (Hansen et al., 1992).
Thelephora terrestris Ehrh.
Regular species in pine – dwarf shrubs – sphagnum communities, at hummocks and in depressions, grows at S.
fuscum surface, Eriophorum vaginatum tussocks, Rubus chamaemorus stems. Collected regularly in north and
middle taiga zone.
Saprotroph of sphagnum, probably other litter and probably mycorrhizal with Pinus. In North Europe reported on
humus, roots, or among mosses in coniferous forests, saprophytic, mycorrhizal, or semi-parasitic (Hansen, Khunsen,
1997).
Xeromphalina cornui (Quél.) J. Favre
Rare species in pine – dwarf shrubs – sphagnum communities, at hummocks, among S. fuscum, stems attaches to
sphagnum at middle part of shoots. Collected several times at two locations in middle taiga.
Saprotroph of Sphagnum (stems attachéd to live sphagnum shoots 2-3 cm below the capitula). For North Europe
reported on conifer debris among Sphagnum (Hansen et al., 1992).
Results and discussions
Opportunistic inspection of ombrotrophic bogs in taiga zone of West Siberia (about 10 locations
were visited over a coarse of 5 years, two locations were visited repeatedly during several years)
revealed 57 species in these communities. Ratio of macromycetes species (in our conception) to
number of vascular plants in ombrotrophic bogs is about 2:1 as reported generally for temperate
regions (Mueller et al., 2004).
Two main functional groups (guilds) may be outlined: sphagnum saprotrophs and EM fungi
(about half species (32) are saprotrophs of sphagnum, 13 species forming mycorrhiza with trees,
2 species wood and plant litter saprotrophs, 7 species parasites of plants and Sphagnum).
Structure of functional groups among larger fungi resemble those of boreal forest’s except
litter/humus saprotrophs are replaced on sphagnum saprotrophs; litter saprotrophs almost absent
among larger fungi (there are great number of ascomycetes inhabiting litter, but they have small
sizes); wood saprotrophs are present, but we not included them in this analysis as well (we
decided to consider them in another report since there only about 3 species forming conspicuous
fruitbodies, and others are corticioid fungi).
It is interesting to register large number of EM fungi (13 species) given only two species of trees
supported them. Exactly, Pinus sylvestris prevails and Pinus sibirica is in minor participation.
Since probably all 13 species could be assigned to one mycorrhizal host.
Opportunistic inspection doesn’t suppose to yield quantitative characteristics of community /
biodiversity. Since long forays to cumulate new species were done, we could suppose indirectry
that cumulative curve of species diversity was achieved. In related publications about larger
fungi of bogs similar numbers of species reported. Though the full lists not coincide, there is
some part of shared species. Exact comparison could not be done since differences of trophic
status of involved bogs and their tree layer. Analysis of data in a floristic publication from
similar climatic region (Nordic Macromycetes and Funga Nordica, next edition) revealed 35
species reported from bogs there, and about 100 species with relation to sphagna, peat, bogged
habitats (from which about 30 shared species with list from this report). Thereby, there are
number of species preferring bogs, though large number with wider range of habitats. And for
some species bog habitats were not registered before (as reflected in Nordic Macromycetes).
For presentation of structure of diversity we used speculative scale, where species were assigned
to one of 4 grades; we allot 14 species being common, 22 regular, 23 rare; only 2 species were
with singular finding. Small number of singular finding may say about stability of the list.
There are two general habitats, responding to different lists of larger fungi in ombrotrophic bogs.
Treed (pine – dwarfshrubs – sphagnum) communities inhabited by about 35 species. They are
forming mycorrhiza with trees (Amanita porphyria, Cortinarius bataillei, C. flexipes, C. flospaludis, C. huronensis, C. obtusus, C. scaurus, C. semisanguineus, C. stillatitius, Cortinarius
sp., Lactarius helvus, L. rufus, Russula paludosa, Suillus variegatus, ), or wood saprotrophs (as
Hypholoma capnoides, Ascocoryne sarcoides), or parasites of dwarf shrubs (Exobasidium
cassandrae). Several saprotrophs of Sphagnum were collected only from this habitat (e.g. they
prefer high hummocks of S. fuscum to watered hollows with hydrophilic sphagna): Arrhenia
sphagnicola, Clavaria sphagnicola, Entoloma fuscomarginatum, Gymnopilus fulgens, G.
penetrans, Gymnopus dryophilus, Lichenomphalia umbellifera, Mycena concolor, Mycena
megaspora, Omphaliaster borealis, Pseudoplectania sphagnophila, Sphagnomphalia
brevibasidiata, Xeromphalina cornui.
Other general habitat, graminoid sphagnum hollows, characterised by following list of
Sphagnum saprotrophs: Ascocoryne turficola, Galerina sphagnicola, G. sphagnorum, G.
tibiicystis, Hypholoma elongatum, H. eximium, H. udum, Psilocybe atrobrunnea, Tephrocybe
palustris. There several species related to hollow plants as well: Rhytisma andromedae,
Monilinia oxycoci, Myriosclerotinia dennisii.
The research was aimed to ombrotrophic bogs, habitats with specific environment and
biochemistry. Data about extreme habitats could be useful in general ecology of fungi. Visits of
lag-zone between bogs and mineral forests with mesotrophic communities occationally were
done as well. Some interesting information received from visiting this zone (mesotrophic
sphagnum communities). Hypholoma myosotis was collected only from mesotrophic habitats, it
never occurs in central parts of bogs. On the contrary, H. elongatum, H. udum and H. eximium
could be sighted in both habitats, though in mesotrophic conditions they have larger fruit bodies.
Similar differences in trophic preferences between species in the genus Galerina. Galerina
stagnina could be collected only in lag zone and G. paludosa seems prefer more trophic habitats
to central part of bogs. On the contrary, G. sphagnicola, G. sphagnurom, G. tibiicystis occur in
central part of bogs in oligotrophic hollows and abundant there. These examples shows that
among sphagnum saprotrophs even from one genera there are preferences in trophic status of
community, and there are species specialized to oligotrophic conditions more than others. But
only thorough mycosociological investigation with analysis on plots may show real differences
in fungal communities in response to ecological factors at bogs.
Larger fungi are more often considered by conservation activities than other fungal groups.
Although all fungi are threatened by human activity, more information about extinction of larger
fungi available up to date, and they are easier to manipulate in conservation programms. Red
lists include mostly macromycetes (Moore, 2001). There is Red Book of endangered species of
KHMAO region (Vasin, 2003; second edition in 2013, in press), which lists about 50 species of
fungi as well. Two species from ombrotrophic bogs were included in the next edition of the
Book, Ascocoryne turficola and Entoloma fuscomarginatum, two are characteristic for
ombrotrophic bogs and collected rare and regular in KHMAO. Analysis of Combined European
fungal Red List (European Red List of endangered macrofungi) showed that most of species (44
from 57) from ombrotrophic bogs are included in some countries lists. Since ombrotrophic bogs
are widespread and almost pristine in the area, the derived annotated list may probably serve as
etalon for altered areas where these ecosystems are threatened.
Table 1. Species list with related data about number of collections, frequency of occurence, ecological groups, substrates and presence in European
fungal Red Lists (species which taxonomy needs elaboration marked *).
Taxons
need
elaboration
Total 11
Species (alphabetical
order)
Total 57 species (50
basidiomycetes, 6
ascomycetes, 1
zygomycetes).
Amanita porphyria
Alb. & Schwein.
Armillaria ectypa (Fr.)
Lamoure
Arrhenia sphagnicola
(Berk.) Redhead,
Lutzoni, Moncalvo &
Vilgalys
Ascocoryne sarcoides
(Jacq.:Fr.) Groves &
D.E.Wilson
Number of
collections/years/subzones
About 450 collections in a
coarse of 5 years (N – north, M
– middle taiga subzones).
Frequency
Community
type
Ecological
group
Substrate
Species in European
Red Lists
Common
species - 14,
regular - 22,
rare - 19,
singular
findings - 2.
30 species occur
constantly in p-ds (pine – dwarf
shrubs –
sphagnum)
habitats, 10
species in g-s
(graminoid –
sphagnum).
Sphagnum
saprotrophs - 30,
EM - 15, parasites
- 6.
(from our observations)
Total 44 species
3 coll/07,12/N,M taiga
rare
p-d-s
pine
mycorrhiza
2 coll/07,09/M taiga
rare
g-s
(mesotrophic)
sphagnum
saprotroph
among S. angustifolium, S.
magellanicum, stunted Betula
pubescens
19 coll/06-12/N, M taiga
common
all types
sphagnum
saprotroph
among S. fuscum and (more
rare) hydrophilic sphagna
4 coll/10-12/M taiga
common
p-d-s
wood
saprotroph
decorticated logs and stumps
of Pinus sylvestris (probably P.
sibirica as well)
Austria, Denmark,
Flandern, France,
Hungary, Latvia,
Netherlands, Norway,
Turkey
Austria, Czech Republic,
Denmark, Finland,
France, Germany, Great
Britain, Italy, Latvia,
Netherlands, Sweden,
Switzerland
Austria, Czech Republic,
France, Germany, Latvia,
Norway, Poland, Serbia,
Switzerland
Taxons
need
elaboration
Species (alphabetical
order)
Number of
collections/years/subzones
Frequency
Community
type
Ecological
group
Substrate
Species in European
Red Lists
Ascocoryne turficola
(Boud.) Korf
7 coll /08,09,12/N, M taiga
rare
g-s
sphagnum
saprotroph
among S. jensenii, S. majus, S.
papillosum, Warnstorfia
fluitans, stems attached to
buried leaves of Carex limosa
(4 sightings) and branches of
dwarf shrubs (once), in some
locations substrate was not
registered (leave – sphagnum
decaying litter)
Clavaria sphagnicola
Boud.
6 coll/ 8,10,12/N, M taiga
rare
p-d-s
sphagnum
saprotroph
among S. fuscum, S.
magellanicum, S.
angustifolium
*
Cortinarius bataillei J.
Favre
4 coll/08-10,12/M taiga
regular
p-d-s
pine
mycorrhiza
*
Cortinarius flexipes
(Pers.) Fr.
7 coll/07,09,10,12/N,M
taiga
regular
p-d-s
pine
mycorrhiza
*
Cortinarius flospaludis Melot
8 coll/09,10,12/M taiga
regular
p-d-s
pine
mycorrhiza
France
*
Cortinarius huronensis
Ammirati & A.H. Sm.
13 coll/12/M taiga
common
p-d-s
pine
mycorrhiza
Czech Republic, Italy,
Latvia, Netherlands
*
Cortinarius obtusus
(Fr.) Fr.
15 coll/07-12/N,M taiga
common
p-d-s
pine
mycorrhiza
Netherlands, Norway
Cortinarius scaurus
Fr. (Fr.)
7 coll/07-09,12/N,M taiga
regular
p-d-s
pine
mycorrhiza
Czech Republic, France,
Germany, Latvia,
Netherlands, Norway
Cortinarius
semisanguineus (Fr.)
Gillet
7 coll/07,09,12/M taiga
common
p-d-s
pine
mycorrhiza
France, Latvia,
Netherlands, Norway
Cortinarius sp.
9 coll/09,12/M taiga
regular
p-d-s
pine
mycorrhiza
*
Denmark, Netherlands,
Norway, Slowakia,
Sweden
France, Germany
Norway, Poland
Taxons
need
elaboration
*
Species (alphabetical
order)
Number of
collections/years/subzones
Frequency
Cortinarius stillatitius
Fr.
6 coll/07,09,12/N,M taiga
regular
p-d-s
pine
mycorrhiza
Endogone pisiformis
Link
1 coll/12/M taiga
singular
g-s
?
among S. angustifolium and
plant litter
*
Ecological
group
among S. fuscum, S.
angustifolium, several times on
tracks with altered surface and
bare peat exposed
Substrate
Species in European
Red Lists
Austria, France, Latvia,
Norway
Entoloma
fuscomarginatum (Fr.)
P. Kumm.
13 coll/07-09,12/N,M taiga
regular
p-d-s
sphagnum
saprotroph
Exobasidium
cassandrae Peck
3 coll/10-12/M taiga
rare
p-d-s
parasite on
Chamaedaphne
calyculata
Exobasidium karstenii
Sacc. & Trotter
5 coll/10-12/M taiga
regular
g-s
parasite on
Andromeda
polifolia
Galerina allospora
A.H. Sm. & Singer
9 coll/12/M taiga
regular
lag-zone
sphagnum
saprotroph
among S. angustifolium, S.
magellanicum, once collected
on pine (Pinus sylvestris) bark
Austria, Norway
lag-zone
sphagnum
saprotroph
among S. angustifolium in
small pits and hollows, also
collected in mesotrophic
sphagnum bogs (more
common there), and bogged
forests with sphagnum
Croatia, Denmark,
France, Hungary, Italy,
Latvia, Norway, Poland,
Serbia, Spain & Portugal
g-s
sphagnum
saprotroph
among hydrophilic common
sphagna species, rare in p-d-s
communities among S. fuscum,
S. angustifolium
Austria, France, Norway,
Russia
among S. fuscum, S.
angustifolium
Austria, Denmark,
France, Germany,
Hungary, Latvia,
Netherlands, Norway,
Poland
Galerina paludosa
(Fr.) Kühner
*
Community
type
Galerina sphagnicola
(G.F. Atk.) A.H. Sm.
& Singer
Galerina sphagnorum
(Pers.) Kühner
18 coll/06-12/N, M taiga
25 coll/07-12/N, M taiga
27 coll/10,12/M taiga
regular
common
common
g-s, rare p-d-s
sphagnum
saprotroph
Denmark, Netherlands,
Norway
Switzerland
Taxons
need
elaboration
Species (alphabetical
order)
Galerina stagnina
(Fr.) Kühner
Galerina tibiicystis
(G.F. Atk.) Kühner
*
Number of
collections/years/subzones
8 coll/06-08,10,12/N, M
taiga
18 coll/06-12/N, M taiga
Frequency
regular
common
Community
type
Ecological
group
Substrate
Species in European
Red Lists
Austria, Czech Repoublic,
Denmark, France,
Germany, Italy,
Netherlands, Norway,
Switzerland
lag-zone
sphagnum
saprotroph
among mesotrophic sphagna,
doesn’t sighted at
ombrotrophic communities
g-s, rare p-d-s
sphagnum
saprotroph
among S. fuscum, S.
angustifolium, rarely collected
from lag – zones and
mesotrophic sphagnum bogs
Austria, Croatia,
Denmark, France,
Hungary, Italy, Latvia,
Norway
Austria, Czech Republic,
Denmark, Finland,
France, Germany, Latvia,
Netherlands, Norway,
Russia
Gymnopilus fulgens (J.
Favre & Maire)
Singer
3 coll/07,12/M taiga
rare
p-d-s, eutr
sphagnum
saprotroph
among S. magellanicum, S.
papillosum at low hummocks,
one collection from eutrophic
fen at sedge peat
Gymnopilus penetrans
(Fr.) Murrill
5 coll/08,12/M taiga
regular
p-d-s
sphagnum
saprotroph
among S. fuscum
Gymnopus
androsaceus (L.) J.L.
Mata & R.H. Petersen
15 coll/06-12/N, M taiga
common
p-d-s, lag-zone
litter
saprotroph
at different litter: Pinus
sylvestris needles, branches;
leaves and branches of dwarf
shrubs; leaves of Eriophorum
vaginatum; leaves of Rubus
chamaemorus. Black
rhizomorphs spread widely at
sphagnum surface and attache
to buried in sphagnum litter.
Gymnopus dryophilus
(Bull.) Murrill
10 coll/06-08,12/N, M
taiga
regular
p-d-s
sphagnum
saprotroph
among S. fuscum, S.
angustifolium, S.
magellanicum
Taxons
need
elaboration
*
Species (alphabetical
order)
Number of
collections/years/subzones
Frequency
Community
type
Ecological
group
Hebeloma
incarnatulum A.H.
Sm.
12 coll/06-12/N, M taiga
common
p-d-s
sphagnum
saprotroph
among common sphagna at
hummocks and in depressions
Hygrocybe cinerella
(Kühner) Arnolds
3 coll/09,10,12/M taiga
rare
g-s
sphagnum
saprotroph
among S. papillosum, S.
jensenii at low hummocks
Hypholoma capnoides
(Fr.) P. Kumm.
2 coll/12/M taiga
rare
p-d-s
wood
saprotroph
at logs of Pinus sylvestris
buried in sphagnum (wet
conditions)
France, Hungary, Latvia
Hypholoma elongatum
(Pers.) Ricken
18 coll/06-12/N, M taiga
common
g-s, lag-zone
sphagnum
saprotroph
among hydrophilic
oligotrophic and mesotrophic
sphagna
Austria, Croatia, France,
Italy, Latvia, Poland,
Romania
Hypholoma eximium
(C. Laest.) Rald
1 coll/12/M taiga
singular
g-s
sphagnum
saprotroph
among hydrophilic sphagna
and exposed peaty substrates
Hypholoma myosotis
(Fr.) M. Lange
9 coll/07,08,10,12/N, M
taiga
regular
lag-zone
sphagnum
saprotroph
among mesotrophic sphagna,
doesn’t sighted in
ombrotrophic communities
among oligotrophic and
mesotrophic sphagna species,
in hollows and pits of p-d-s
communities
among S. angustifolium, in
altered places (along tracks) on
bare peaty substrate
Hypholoma udum
(Pers.) Quél.
13 coll/09,10,12/M taiga
common
g-s, lag-zone
sphagnum
saprotroph
Laccaria proxima
(Boud.) Pat.
5 coll/12/M taiga
rare
p-d-s (pits)
? sphagnum
saprotroph
Lactarius helvus (Fr.)
Fr.
4 coll/09,12/M taiga
regular
p-d-s
pine
mycorrhiza
Lactarius rufus (Scop.)
Fr.
4 coll/08-12/N,M taiga
common
p-d-s
pine
mycorrhiza
Leccinum holopus
(Rostk.) Watling
2 coll/12/M taiga
rare
p-d-s
birch
mycorrhiza
Substrate
Species in European
Red Lists
Austria, Bulgaria, France,
Germany, Latvia, Poland
Denmark, Flandern,
France, Hungary, Latvia,
Netherlands, Norway,
Switzerland, Turkey
Denmark, Flandren,
Hungary, Latvia, Norway
Austria, Czech Republic,
Denmark, Flandren,
France, Hungary,
Taxons
need
elaboration
Species (alphabetical
order)
Number of
collections/years/subzones
Frequency
Community
type
Ecological
group
Substrate
Species in European
Red Lists
Norway, Switzerland
? sphagnum
saprotroph
disturbed sphagnum surface
(often along bog tracks), on S.
fuscum or peaty substrates
(covered with liverworts and
algae)
France
g-s
parasite on
Oxycoccus
palustris
on berries, fruiting in early
spring
Denmark, Germany,
Norway
regular
p-d-s
sphagnum
saprotroph
among S. angustifolium, often
in altered places on peat (along
bog tracks)
Denmark, Netherlands,
Norway, Poland
2 coll/12/M taiga
rare
p-d-s
sphagnum
saprotroph
among S. angustifolium
9 coll/06-08,10,12/N, M
taiga
regular
p-d-s, lag-zone
sphagnum
saprotroph
among S. angustifolium, S.
fuscum, also in lag – zone,
among mesotrophic sphagna
lag-zone
parasite on
Carex rostrata
at stems of Carex rostrata,
also collected from marshes
(salix – sedge wetlands near
lake)
Austria, Czech Republic
sclerotia form inside of flower
stalks of Eriophorum
vaginatum, apothecia arise in
early summer
Croatia, Germany,
Norway
among S. fuscum and
hydrophilic sphagna
Germany, Norway
Lichenomphalia
umbellifera (L.)
Redhead, Lutzoni,
Moncalvo & Vilgalys
3 coll/12/M taiga
rare
p-d-s (pits)
Monilinia oxycocci
(Woronin) Honey
3 coll/08,09/M taiga
rare ?
Mycena concolor (J.E.
Lange) Kühner
9 coll/07-10,12/N, M taiga
Mycena galopus
(Pers.) P. Kumm.
Mycena megaspora
Kauffman
Myriosclerotinia
caricis-ampullaceae
(Nyberg) N.F. Buchw.
3 coll/08,11,12/N, M taiga
rare
Myriosclerotinia
dennisii (Svrček) J.
Schwegler
3 coll/08,10,12/M taiga
regular
p-d-s, g-s
parasite on
Eriophorum
vaginatum
Omphaliaster borealis
(M. Lange & Skifte)
Lamoure
17 coll/07-10,12/N, M
taiga
regular
p-d-s, g-s
sphagnum
saprotroph
Denmark, Latvia,
Norway, Turkey
Austria, Czech Republic,
Denmark, France,
Germany, Latvia,
Netherlands, Norway,
Poland
Taxons
need
elaboration
Species (alphabetical
order)
Pseudoplectania
sphagnophila (Pers.)
Kreisel
Number of
collections/years/subzones
Frequency
4 coll/08-11/N, M taiga
rare
Community
type
p-d-s
Ecological
group
sphagnum
saprotroph
Substrate
among S. fuscum, S.
magellanicum
Czech Republic, Finland,
Germany, Norway
Czech Republic,
Germany, Norway and (as
P. turficola) Denmark,
France, Netherlands,
Poland
Psilocybe atrobrunnea
(Lasch) Gillet
4 coll/10,12/M taiga
rare ?
g-s
sphagnum
saprotroph
among hydrophilic sphagna
Rhytisma andromedae
(Pers.) Fr.
2 coll/08,12/M taiga
regular
all types
parasite on
Andromeda
polifolia
at leaves
Russula paludosa
Britzelm.
8 coll/07,09,12/M taiga
regular
p-d-s
pine
mycorrhiza
Sphagnomphalia
brevibasidiata (Singer)
Redhead, Moncalvo,
Vilgalys & Lutzoni
4 coll/08,10,12/N,M taiga
rare
p-d-s
sphagnum
saprotroph
Denmark, Flandren,
Frence, Hungary, Latvia,
Netherlands , Norway
hummocks, among S. fuscum
Austria, Denmark,
Flandren, France,
Hungary, Latvia,
Netherlands, Norway,
Turkey
Suillus variegatus
(Sw.) Kuntze
3 coll/12/M taiga
rare ?
p-d-s
pine
mycorrhiza
Tephrocybe palustris
(Peck) Donk
10/06/07/N, M taiga
common
g-s, p-d-s, lagzone
sphagnum
saprotroph,
parasite
hydrophilic, meso and
oligotrophic sphagna
at hummocks and in
depressions, grows at S.
fuscum surface, Eriophorum
vaginatum tussocks, Rubus
chamaemorus stems
among S. fuscum, stems
attaches to sphagnum at
middle part of shoots
Thelephora terrestris
Ehrh.
7 coll/07-12/N, M taiga
regular
p-d-s
sphagnum and
litter
saprotroph,
mycorrhiza ?
Xeromphalina cornui
(Quél.) J. Favre
3 coll/07,12/M taiga
rare
p-d-s
sphagnum
saprotroph
Species in European
Red Lists
France, Latvia, Norway,
Poland, Switzerland
France, Latvia
Literature
Ainsworth, G.C., Bisby, G.R., Kirk, P.M., 2008. Ainsworth & Bisby’s dictionary of the
fungi. CABI, UK.
Arnolds, E., 1981. Ecology and coenology of macrofungi in grasslands and moist
heathlands in Drenthe, the Netherlands. Part 1. Introduction and Synecology. J. Cramer, Vaduz.
Augustin, A., Runge, A., 1968. Pilze in Scheiden-Wollgras-Rasen des Emsdettener Venns.
Natur und Heimat (BRD) 8, 152–153.
Berch, S.M., Fortin, J.A., 1983. Endogone pisiformis: axenic culture and associations with
Sphagnum, Pinus sylvestris, Allium cepa, and Allium porrum. Canadian Journal of Botany 61,
899–905.
Boertmann, D., 1995. The genus Hygrocybe. Danish Mycological Society, Denmark.
Botch, M.S., Kobak, K.I., Vinson, T.S., Kolchugina, T.P., 1995. Carbon pools and
accumulation in peatlands of the former Soviet Union. Global Biogeochemical Cycles 9, 37–46.
Breitenbach, J., Kränzlin, F., 1995. Fungi of Switzerland: a contribution to the knowledge
of the fungal flora of Switzerland. Volume 4, Agarics 2nd part. Edition Mykologia, Lucerne.
Bunyard, B.A., Wang, Z., Malloch, D., Clayden, S., Voitk, A., 2008. New North American
Records for Ascocoryne turficola (Ascomycota: Helotiales). Fungi 1, 23–31.
Charman, D.J., 2002. Peatlands and environmental change. J. Wiley & Sons, England.
Chinan, V., 2010. Macrofungi identified in the peat pogs from the Naruta 2000 site “Gaina
Lucina” (Eastern Carpathians, Romania). Analele ştiinţifice ale Universităţii “Al. I. Cuza” Iaşi
LVI.
Chinan, V., 2011. Macrofungi from “Gradinita” peat bog (Eastern Carpathians, Romania).
Analele ştiinţifice ale Universităţii „Al. I. Cuza” Iaşi.
Den Bakker, H.C., 2005. Diversity in Leccinum: a molecular phylogenetic approach.
National Herbarium Nederland.
Dighton, J., White, J.F., Oudemans, P., 2005. The Fungal community: its organization and
role in the ecosystem (Introduction). Taylor & Francis group, LLC.
Dikunez, V. A., Kotova, T. V., Makeev, V. N., Tikunov, V. S., ed, 2004. Atlas of KhantyMansiyskiy Okrug - Yugra. vol 2. Natura and ecology. Moskow - Khanty-Mansiysk. (in
Russian)
European Red List of endangered macrofungi, at: European Council for the Conservation
of Fungi home page, http://www.wsl.ch/eccf/activities-en.ehtml
Favre, J., 1948. Les associations fongiques des hauts-marais jurassiens et de quelques
regions voisines. Mater. flore cryptogam. Suisse 10, 228.
Golovchenko, A.V., Semenova, T.A., Polyakova, A.V., Inisheva, L.I., 2002. The Structure
of the Micromycete Complexes of Oligotrophic Peat Deposits in the Southern Taiga Subzone of
West Siberia. Microbiology 71, 575–581.
Grum-Grzhimaylo, O.A., Bilanenko, E.N., 2012. The micromycete complexes of bogs at
the Kandalaksha bay of the White Sea. Mycologia and Phytopatologia, 46, 297-305. (in Russian)
Hansen, Lise, Khudsen, Henning (Eds.), 1997. Nordic Macromycetes. Vol 3.
Heterobasidioid, Aphyllophoroid and Gastromycetoid basidiomycetes. Copenhagen: Nordsvamp.
Hansen, Lise, Khudsen, Henning (Eds.), 2000. Nordic Macromycetes. Vol 1. Ascomycetes.
Copenhagen: Nordsvamp.
Hansen, L., Knudsen, H., Dissing, H., 1992. Nordic Macromycetes. Vol 2. Polyporales,
Boletales, Agaricales, Russulales. Copenhagen: Nordsvamp.
Heilmann-Clausen, J., Verbeken, A., Vesterholt, J., 1998. The genus Lactarius. Danish
Mycological Society, Denmark.
Ivanov, K.E., 1981. Water movement in Mirelands. Academic Press, London.
Kalamees, K., 1982. The composition and seasonal dynamics of fungal cover on peat soils,
in: Mazing V.V., (Ed.), Peatland Ecosystems: Researches into the Plant Cover of Estonian Bogs
Ad Their Productivity. Valgus, Tallinn, pp. 12–29.
Kalamees, K., Raitviir, A., 1982. A list of higher fungi of Estonian peatlands, in: Mazing
V.V., (Ed.), Peatland Ecosystems: Researches into the Plant Cover of Estonian Bogs Ad Their
Productivity. Valgus, Tallinn, pp. 30–33.
Lange, M., 1948. The Agarics of Maglemose. Dansk Botanisk Arkiv Udgivet af Dansk
Botanisk Forening. Copenhagen 13, 1–141.
Liss, O.L., Abramova, L.I., Avetov, N.A., et al, 2001. Wetland systems of West Siberia and
their importance for nature conservation. Grif i K., Tula. (in Russian)
Lodge, D.J., 2004. Terrestrial and Lignicolous Macrofungi, in: Mueller, G. et al., (Eds.),
Biodiversity of Fungi. Inventory and Monitoring Methods. Elsevier Academic Press,
Amsterdam; Boston, pp. 127–172.
Moore, D., British Mycological Society, 2001. Fungal conservation: issues and solutions.
Cambridge University Press, Cambridge.
Mueller, G., Schmit, J., Leacock, P., Buyck, B., Cifuentes, J., Desjardin, D., Halling, R.,
Hjortstam, K., Iturriaga, T., Larsson, K.-H., Lodge, D., May, T., Minter, D., Rajchenberg, M.,
Redhead, S., Ryvarden, L., Trappe, J., Watling, R., Wu, Q., 2007. Global diversity and
distribution of macrofungi. Biodiversity and Conservation 16, 37–48.
Mueller, G.M., Bills, G.F., Foster, M.S., 2004. Biodiversity of fungi. Inventory and
monitoring methods. Elsevier Academic Press, Amsterdam; Boston.
Niskanen, T., Kytovuori, I., Bendiksen, E., 2008. Cortinarius (Pers.) Gray, in: Knudsen, H.
& Vesterholt, J. (eds.): Funga Nordica. pp. 661–777.
Noordeloos, M.E., 1992. Entoloma s.l. Giovanna Biella, Saronno.
Peregon, A., Maksyutov, S., Kosykh, N.P., Mironycheva-Tokareva, N.P., 2008. Map-based
inventory of wetland biomass and net primary production in western Siberia. J. Geophys. Res.
113, G01007.
Polyakova, A.V., Chernov, I.Y., Panikov, N.S., 2001. Yeast Diversity in Hydromorphic
Soils with Reference to a Grass–Sphagnum Wetland in Western Siberia and a Hummocky
Tundra Region at Cape Barrow (Alaska). Microbiology 70, 617–623.
Roberts, C., Ceska, O., Kroeger, P., Kendrick, B., 2004. Macrofungi from six habitats over
five years in Clayoquot Sound, Vancouver Island. Canadian Journal of Botany 82, 1518–1538.
Romanova, E.A., Bybina, R.T., Golitsyna, E.F., et al., 1977. Wetland typology map of west
Siberian lowland, scale 1:2,500,000. GUGK, Leningrad. (in Russian)
Rossman, A.Y., Tulloss, R.E., O’Dell, T.E., Thorn, R.G., 1998. Protocols for an All Taxa
Biodiversity Inventory of Fungi in a Costa Rican Conservation Area. Parkway Publishers, Inc.
Salo, K., 1993. The composition and structure of macrofungus communities in boreal
upland type forests and peatlands in North Karelia, Finland. Karstenia 33, 61–99.
Smith, A.H., Singer, R., 1964. A monograph on the genus Galerina Earle, 2nd ed.
University of Michigan Library, Ann Arbor, Michigan.
Sotek Z., Stasinska M., 2010. Diversity of macromycetes and the transformation of
vegetation in the Atlantic raised bog “Stramniczka”. Water-Environment-Rural Areas 10, 257–
270.
Stasinska, M., Sotek, Z., Gamrat, R., Prajs, B., 2005. Vascular plants and macromycetes of
the peat-bog in the Głowacz reserve (NW Poland). In: Wołejko, L.;Jasnowska, J., (Eds.): The
future of Polish mires, pp.183–188.
Thormann, M.N., 2006. The Role of Fungi in Boreal Peatlands, in: Wieder, R.K. et al.,
(Eds.), Boreal Peatland Ecosystems, Ecological Studies. Springer Berlin Heidelberg, pp. 101–
123.
Thormann, M.N., Rice, A.V., 2007. Fungi from peatlands. Fungal diversity 24, 241–299.
Vesterholt, J., 2005. The genus Hebeloma. Danish Mycological Society, Denmark.
Vasin, A.M., (Ed.)., 2003. Red Book of KHMAO (animals, plants, fungi). Pakrus,
Ekaterinburg. (in Russian)
Vitt, D.H., 2006. Functional Characteristics and Indicators of Boreal Peatlands, in: Wieder,
R.K. et al., (Eds.), Boreal Peatland Ecosystems, Ecological Studies. Springer Berlin Heidelberg,
pp. 9–24.
Download