P r o t is t o lo g y
Protistology 1 (4), 161–178 (2000)
August, 2000
Biodiversity of plasmodial slime moulds (Myxogastria):
measurement and interpretation
Yuri K. Novozhilova, Martin Schnittlerb, InnaV.
Zemlianskaiac and Konstantin A. Fefelovd
a
V.L.Komarov Botanical Institute of the Russian Academy of Sciences, St.
Petersburg, Russia,
b
Fairmont State College, Fairmont, West Virginia, U.S.A.,
c
Volgograd Medical Academy, Department of Pharmacology and Botany,
Volgograd, Russia,
d
Ural State University, Department of Botany, Yekaterinburg, Russia
Summary
For myxomycetes the understanding of their diversity and of their ecological function remains
underdeveloped. Various problems in recording myxomycetes and analysis of their diversity
are discussed by the examples taken from tundra, boreal, and arid areas of Russia and
Kazakhstan. Recent advances in inventory of some regions of these areas are summarised. A
rapid technique of moist chamber cultures can be used to obtain quantitative estimates of
myxomycete species diversity and species abundance. Substrate sampling and species isolation
by the moist chamber technique are indispensable for myxomycete inventory, measurement of
species richness, and species abundance. General principles for the analysis of myxomycete
diversity are discussed.
Key words: slime moulds, Mycetozoa, Myxomycetes, biodiversity, ecology, distribution, habitats
Introduction
General patterns of community structure of terrestrial
macro-organisms (plants, animals, and macrofungi) are
well known. Some mathematics methods are used for their
studying, from which the most popular are the quantitative analysis of information diversity, species abundance,
and estimation of similarity of communities (Whittaker,
1960, 1977; Chernov, 1975; Vasilevich, 1969;
Pianka,1973; Pesenko, 1982; Magurran, 1988; Mukhin,
1993; Watling, 1994; Chernov and Matveeva, 1997). Similar research of micro-organisms remains very
underdeveloped and has been limited, to a large extent, by
methodical difficulties of their inventory (Chernov, 1997;
Dobrovol’skaia et al., 1999). Considerable success here
can be expected in the studies of separate groups that can
be easily recorded and identified. One of them is plasmodial myxomycetes or “slime moulds” (Myxogastria).
Myxomycetes (plasmodial slime moulds), are a group
of protista comprising about 1000 species (Mitchell, 2000).
Myxomycetes in ecological terms are predators of bacteria, holding maintenance between bacterial and fungal
© 2000 by Russia, Protistology.
decay (Madelin, 1984). The life cycle of myxomycetes
includes two trophic stages: uninucleate myxoflagellates
or amoebae, and a multi-nucleate plasmodium (Fig. 1).
The entire plasmodium turns almost all into fruit bodies,
called sporocarps (sporangia, aethalia, pseudoaethalia, or
plasmodiocarps). The plasmodium is very mobile, covered only by a simple membrane, live mostly inside the
substrate and can achieve macroscopic dimensions, clearly
visible in the field in only one group (Physarales). The
dispersal units are spores, which can be dispersed via air
or insects. Various dormant stages (microcysts, sclerotia,
and spores) may interrupt this cycle under harsh environmental conditions (Gray and Alexopoulos, 1968;
Blackwell et al., 1984).
Unique among living organisms is the combination
of single-cell stages living as true microorganisms (Feest,
1987) with sporocarps. Due to this feature the myxomycetes can be safely preserved as herbarium specimens.
This peculiarity permits to accumulate data on occurrence,
ecology and geography of myxomycetes into a comprehensive computer database, which will be electronically
accessible to the scientific community over the Internet.
162 · Yuri K. Novozhilov, Martin Schnittler, InnaV. Zemlianskaia and Konstantin A. Fefelov
× Fig. 1. Schematic drawing of the general life cycle of the
plasmodial slime moulds (myxomycetes). 1 – developing
sporocarps and sporogenesis, 2 – germinating spore,
3 – myxamoeba, 4 – microcysts, 5 – swarm cell, 6, 7 – zygote
formatting, 8 – plasmodium formatting and mature fan-shaped
plasmodium, 9 – sclerotium (macrocysts).
These observations serve as a basis for additional research
emphasising all forms of myxomycete diversity in the
world, including qualitative composition, community diversity, adaptive types, and functional (ecological) groups.
Up to the present time, about 3000 (or probably more,
due to under-representation of publications on biochemical aspects) papers focusing on myxomycetes inventory
have been published. About 1200 of these are local or regional species lists, and about 400 describe new taxa
(Schnittler and Mitchell, 2000).
Except in a very few arctic (Schinner, 1983; GØtzsche,
1984, 1989, 1990; Stephenson and Laursen, 1993, 1998;
Novozhilov et al., 1998a, 1998b, 1999; Stephenson et al.,
2000), temperate (Stephenson, 1988, 1989; NannengaBremekamp, 1991; Ing, 1994), boreal (Schnittler and
Novozhilov, 1996), desert (Blackwell and Gilbertson,
1980; Novozhilov and Goloubeva, 1986; Schnittler and
Novozhilov, 1999; Schnittler, 2000), mediterranean (Lado,
1993, 1994), tropical (Alexopoulos, 1970; Alexopoulos
and Saenz, 1975; Farr, 1976; Maimoni-Rodella and
Gottsberger, 1980; Eliasson, 1991) communities, evaluation of species diversity is scarce and inventory is widely
scattered. Russia, with its vast arctic, boreal territories, is
not an exception in this respect.
In this paper we want to determine methodical approaches in study of myxomycete biodiversity and to
discuss these methods on examples of diversity measurement in some local and regional myxomycete biotas of
Russia and Kazakhstan.
Evaluation of α-diversity
For myxomycetes typically α-diversity indices are
calculated using Shannon’s formula (Shannon and Weaver
1963; Stephenson, 1988, 1989; Novozhilov et al., 1999).
Species diversity (H′) = - ∑ Pi log Pi, where Pi is the relative abundance of a particular species (the proportion of
the total number of individuals represented by species i).
Maximum values for this diversity index are usually observed when there are many species with equal abundance.
Values decrease with both a reduction in the number of
species and an increase in abundance of a very few species. Species diversity (H′) is a function of both the number
of species present (species richness) and the evenness with
which individuals are distributed among these species (species equitability). Stephenson (1988) has calculated the
equitability component (J′) using the formula suggested
by Pielou (1975). J′ = H′/ H′max where H′max represents the
maximum possible diversity for the number of species (S)
present in the community (i.e., all of these species are
equally abundant) and is calculated as H′max = log S.
Commonly employed α-diversity indices require an
examination of at least three components: species richness (number of species), evenness (a gauge of how evenly
the individuals are distributed among the samples), and
abundance or the total number of organisms per sample
(Magurran, 1988; Vasilevich, 1992a; Bills, 1995; Miller,
1995).
Measurement of species richness and species
abundance
The estimation of myxomycete diversity has been limited by the difficulties in defining individuals.
Myxomycetes refer to the group of organisms for which it
is difficult to define a module unit in ecological research
requiring evaluation of species abundance. Until the publication of a method for the enumeration of myxomycetes
in soil (Feest and Madelin, 1985; Feest, 1987; Madelin,
1990), no data regarding the numbers of myxogastrid
propagules in any substrate had been reported. In their
papers, Feest and Madelin showed it was possible to obtain numerical data for soil using a unit (the
plasmodium-forming unit, PFU) analogous to the cloneforming unit of dictyostelids used by Cavender (1989).
Individual free-living plasmodium is physically discrete
and functionally independent, that is the reason for describing plasmodia as individuals. Feest and Madelin
(1985) used the presence or absence of plasmodia as an
unambiguous indicator of the presence of myxomycetes
P r o t is t o lo g y · 163
in the study sites. However, this method permits to estimate a total abundance of myxogastrids in the soil but is
time-consuming and almost useless for estimation of abundance of separate species.
The current species concept for myxomycetes is almost entirely a morphological one, based mostly on the
characters of the sporocarps (Lister, 1925; Gray and
Alexopoulos, 1968; Martin and Alexopoulos, 1969;
Rammeloo, 1974; Eliasson, 1977; Nannenga-Bremekamp,
1991; Keller and Eliasson, 1992; Gilert, 1996; Keller and
Braun, 1999; Mitchell, 2000; Novozhilov and Goodkov,
2000). In virtually, the presence or absence of a species in
diversity analysis is dependent on the appearance of the
sporocarps in the study site (transect, plot,) and substrate
samples. Sporocarps can be used as the module units for
estimation of species richness and species abundance because they are extractable, identifiable, and quantifiable.
An advantage of this approach is the easy way to collect specimens: the colonies of sporocarps can simply be
dried, glued with the substrate in small boxes like matchboxes and stored like other herbarium collection. Eliasson
(1981) and Stephenson (1988) considered a “collection”
as one or more (colony) sporocarps originated from a single
plasmodium. In these studies, the sporocarps were regarded
as separate collections if the intervening distance was at
least 30 cm.
For approximate estimation of species abundance and
species productivity, a simple scale was proposed by
Stephenson et al. (1993), based on the proportion of a species in the total number of records (collections of
sporocarps) and dividing between R – rare (<0.5%); O –
occasional (0.5–1.5%); C – common (1.5–3%); A – abundant (>3%).
Spore numbers probably is more accurate measure of
resource allocation and species productivity (Morton et
al., 1995). Recently, Schnittler (2000) proposed to measure the spore numbers for species cultivated in the moist
chamber cultures. For each species recorded, the number
of spores per sporocarp was estimated, using average sporocarp size, their shape (i.e. half-globose, globose,
cylindrical) and spore diameters. The dimension of these
values was confirmed by counting the spore numbers of a
few selected sporocarps with a counting chamber (as used
for determination of erythrocyte levels). However, this
method evidently is laborious for mass-analysis of species abundance. Moreover, our knowledge about
myxomycete space distribution still is scarce to understand
the resolving possibility of this method. It is obvious that
Stephenson’s scale and the calculation of the spore numbers give us only an approximate estimation of the species
abundance and permit to evaluate virtually only “frequency
of species” (occurrence).
About 1–3 % of the species form rather large (cmrange) clustered fructifications called aethalia or
pseudaethalia, the majority (60–80 %) of the species has
colonies of fragile, single sporocarps in the mm-range, the
remaining 20–40 % show minute fructifications less than
0.5 mm in size. The latter are usually not observed in the
field. The percentage of these size-classes varies widely
between climatic zones as well as between ecological
groups, with a tendency to have more minute forms in
harsh, especially dry, environments.
As an obvious consequence, in additional to the field
collection substrate sampling and the moist chamber technique has usually applied for evaluation of α-diversity and
the ecological analysis of myxomycete community structure (Härkönen, 1977; Stephenson, 1985, 1988;
Novozhilov, 1988; Schnittler, 2000).
Moist chamber technique for assessing species
richness and species abundance in substrata
(microhabitats)
Myxomycetes live mostly very hidden, but are present
in all habitats with decaying organic material, ranging from
tropical rainforests over deserts up to the arctic tundra.
Numerous research of myxomycete diversity in different
microhabitats (substrata) show that species composition
varies considerably on the certain types of substrata (e.g.
on the bark of trees). However, the set of species that dominate in a certain sample of substrate (microhabitat) is rather
limited. In every substrate sample usually 1–4 species comprise more than 30–50% of all group (“potential
dominants”). With the help of moist chamber technique
(Gilbert and Martin, 1933) these dominants are revealed
rather easily and definitely, which gives an opportunity to
carry out mass analysis and receive statistically true results. This simple method permits to evaluate α-diversity
and abundance as “frequency of species” and determinate
the “nucleus” of species diversity.
The respective moist chamber technique does not need
sterile conditions and is easy to carry out. It requires nothing more than Petri dishes, filter or even toilette paper and
a good dissecting microscope for checking. The cultures
run up to 2 months, for coprophilous forms sometimes
longer (up to 3–4 months). To reveal more ecological data
with less effort, the following slight modifications are proposed:
– pooling of bark scales (about 3 to 8 scales of 1–2 cm
size for one tree or shrub, only including the non-living layer, with sampling 5–10 trees from the same
habitat (equal tree species, height, and light exposition);
– measuring of the pH in the moist chamber after 2–5
days, ideally with a solid state probe (substrata often
show buffering capacity, therefore being stable for a
longer period);
– optionally, for statistical evaluations the number of
sporocarps in a moist chamber can be estimated.
For moist chambers, substrate pieces are densely
placed on filter paper in Petri dishes (63.6 cm2), with the
pieces touching but not overlapping each other and with
the outer side of bark uppermost. Cultures are watered
164 · Yuri K. Novozhilov, Martin Schnittler, InnaV. Zemlianskaia and Konstantin A. Fefelov
with distilled water adjusted to pH 7.0 and maintained up
to 2 months under diffuse daylight and at room temperature (22–23 °C). At five times (days 2, 6, 11, 21 and 40
after start) the chambers are checked with a high-magnification dissecting microscope. Mature fructifications are
boxed, and sporocarps of minute species are immediately
preserved in polyvinyl lactophenol or glycerol gelatine,
when calcareous structures are present in sporocarps.
It currently for all ecological analyses, one or more of
the following measures are applied: (i) number of records
per species, in this case the occurrence of one species in
one moist chamber constitutes a collection (record), (ii)
absolute abundance (number of sporocarps for one species in one moist chamber culture) and (iii) weighted
abundance, calculated by dividing the absolute number of
sporocarps recorded in a particular moist chamber through
the mean value from all moist chambers with this species
(Schnittler, 2000).
Consequently, the sum of all weighted abundances for
all cultures with a species is equal to the number of records
for this species.
time-consuming part, running up to 2 monthes and needing regularly checks from the 5th day on. About thirty
minutes per one moist chamber culture have to be invested,
considering also mounting of the often minute specimens
on permanent slides. Although not very costly in terms of
equipment, a well-done regional survey requires about 2–
4 months of pure working time. In special environments,
like arid areas, more time may be necessary due to the fact
that almost all species have to be detected by moist chambers.
The majority of the species having large sporocarps
(aethalium, plasmodiocarps) typically does not develop
in the moist chamber culture. For this reason this technique can be used as only an additional method for
evaluation of myxomycete diversity. The interpretation of
moist chamber detection has to be done cautiously, because spores of species regularly not occurring in the region
may be trapped and develop into fructifications. Colonies
including numerous sporocarps, developing in a short time,
point to a microcyst population (a dormant stage) occurring naturally.
Limitation of inventory studies and completeness of
evaluation of species richness
The examples above demonstrate how many factors
can influence on spatial distribution of myxomycetes in
the biotope. The spatial distribution of myxomycetes is
reflected in the species/sample number curve. At present
time the problem “number species/area” is studied insufficiently for myxomycetes and invites further
investigations.
Our experience shows that the completeness of the
evaluation of species richness depends on the number of
samples for moist chamber culture and from their distribution in the space. The sample plot 0.1 ha (1000 sq. m) is
typically sufficient. The bootstrap analysis shows a steep
increase in the mean cumulated number of recorded species over the first 50 samples putted in Petri dish. In
virtually the curve “species/sample” shows that 30 substrate samples (60 square cm each) are typically sufficient
to record all of the more common species.
According to the experience of the authors, a oneseason project gives a comparable and fairly complete
species list for an area, although annual shifts in the species abundance is known for many species as it was shown
below. With recording all colonies and describing the microhabitats, in a wooded area of temperate or boreal zone
an experienced collector can observe up to 50–70 developments during a field day (about 6 hours), occasionally
much more if mass fructifications occur. For example with
about 5 major vegetation types per area, and half a day for
each, four field surveys of 3 to 4 days each give a fairly
complete picture of the species inventory.
As in most protista, species determination is time-consuming and requires a good compound as well as a
dissecting microscope. The moist chambers are the most
Ecological groups of myxomycetes and distribution of
myxomycetes in biotope
The role of the biotope and microhabitat (substrate)
are considerable factors that influence the myxomycete
distribution. Literature and our data show that the following 4 substrate types are the most different from each other
as the myxomycetes environment and have different adaptive ecological groups of myxomycetes:
1. LIGNICOLOUS MYXOMYCETES (on coarse decaying wood
debris of all stages and sizes): the largest group, including 30–70 % of the total species number, with a
higher proportion in temperate and boreal zones; 70–
80 % of these slime moulds have macroscopical size
(mm-range), relatively easy to collect, moist chambers technique works not for all species; often sharply
defined sporulation peaks for certain species especially
in temperate zones, here mid-summer to late autumn
(Eliasson and Strid, 1976).
2. CORTICOLOUS MYXOMYCETES (inhabiting the bark of living trees and shrubs): 20–40, up to 80 % (in deserts)
of the total species number; almost all minute forms,
making moist chamber technique indispensable which
gives good and stable results in this group; can be
studied throughout the year (Gilbert and Martin, 1933;
Keller and Brooks, 1976, 1977; Härkönen, 1977,
1978; Schnittler and Novozhilov, 1999).
3. SOIL AND LITTER SPECIES (living in the upper soil layer
and fructificating on all kinds of herbaceous or other
small-sized plant refuse): 20–70 % of the species to
expect, mainly Physarales, with much higher proportion in the tropics; mostly of macroscopic size but
difficult to find in the field, for smaller forms moist
chamber techniques works reasonable; often pronounced sporulation peaks in temperate zone
P r o t is t o lo g y · 165
mid-summer to early autumn (Keller and Brooks,
1971; Härkönen, 1981)
4. COPROHILOUS MYXOMYCETES (on dung of herbivores
mammals and birds): a small group with only a few
specialised species typically in higher abundance in
arid areas, on dung with basic pH; moist chambers
running up to 4 months are necessary for detection;
seldom found in the field, but detectable throughout
the year by moist chamber method (Eliasson and
Lundqvist, 1979; Cox, 1981; Eliasson and Keller,
1999).
Some species are hardly related to concrete substrate
group. These species associated with rather specific habitats and their distribution strongly depends from
microclimate conditions in habitats.
BRYOPHILOUS OR MOSS-INHABITING MYXOMYCETES (associated with mosses, but more probably with slime algae, on
wood or rocks provided with trickling water in humid ravines). This group includes less than 5 %, mostly
macroscopic forms, temperate and boreal zones preferentially with sporulation peak in late autumn, and can be
collected in the field only, moist chamber techniques almost always fails to work. Typically small forms (e.g.
Barbeyella minutissima, Colloderma oculatum) have to
be collected with lenses or dissecting microscops in the
field.
NIVICOLOUS MYXOMYCETES (on plant refuse near the melting snow, preferentially in higher mountains with high
snow cover in winter). Almost all nivicolous species are
more or less conspicuous, and easy to collected for an experienced worker (Neubert et al., 1995). They occur very
locally but often with mass fructifications with sporulation peak near snow melting. For nivicolous myxomycetes
the relationship of the percentage of solid precipitation to
monthly averages of air temperature and humidity is a very
important factor (Kowalski, 1975).
The moist chamber technique gives insufficient results for briophilous and nivicolous species.
Estimation of β-diversity and niche breadth (NB)
It is known that one of the approach of β-diversity
study is the estimation of species composition along the
gradient of environment and comparison of species composition of different communities and habitats (Wittaker,
1960; MacArthur, 1965; Wilson, Mohler, 1983; Vasilevich,
1992b). Since the discovering of the right microhabitat is
the key for the stable detection of the species, this could
become the most important tool for enhancing our knowledge about biodiversity of myxomycetes. With the aim to
gather as much as possible data about microhabitat a
standardised, modular-build description system was used
(Schnittler et al., 1996).
For example estimation of niche breadth (NB) on the
basis of moist chamber data was carried out in the research
of myxomycetes of upland forests of south-western Virginia (Stephenson, 1988) and Kazakh deserts (Schnittler
2000). Values for NB were calculated using the formula
NB = 1/s ΣPij 2, where s is the number of states for the
environmental (microhabitat) parameter defining a niche
dimension, and Pij the proportion of species i associated
with state j divided by the total abundance of species i
across all states (Feinsinger et al., 1981). As abundance
measures, either records or total abundances were used.
In both cases, values range from 1/s (all individuals of
one species are associated with one resource state) to 1.0
(equal numbers of individuals are associated with each
resource state). In the same manner, niche overlap (NO)
was computed, using the symmetrical index NOik = ΣPij
Pik /sqrt(ΣPij 2) (ΣPik 2), with Pij and Pik as the proportions of
the ith resource state by the jth and kth species, respectively (Levins, 1968; Pianka 1973). Values for niche
overlap range from 0 to 1 too.
For analysis of myxomycete associations, the Cole
(1949) index of interspecific association and its standard
error was computed. It is based on a 2 x 2 contingency
table for presence and absence of a pair of species in one
moist chamber, ranging from –1 (the species never occur
together) to 1 (the species occur always together). With a
chi-square test the significance level of deviations between
observed association frequencies and those expected by
chance was determined.
Inherent problems in estimating species diversity:
seasonality and features of microhabitats
Numerous studies of myxomycetes have shown that
there are often dramatic shifts in the appearance of sporocarps from the fall to the spring in boreal and temperate
areas. The inventory of nivicolous species is a good example of this shift. Until recently, nivicolous species were
not found in Russia. Intensive studies in the Khibine mountains in spring (Novozhilov and Schnittler, 1997) showed
that this ecological group is abundant in this area. This
region provides good conditions for cryophilous-nivicole
litter myxomycetes. This ecological group can be separated into two subgroups, based on phenology and habitat
requirements. The first subgroup contains the ‘true’
nivicolous species: Diacheopsis effusa, Diderma niveum,
the Lepidoderma, and Lamproderma species (with the
exception of Lamproderma sauteri). According to
Schinner (1981, 1983), their habitat requirements can be
characterized as follows: open ground, a more or less thick
layer of herbaceous plant refuse, high snow cover in winter (may be for providing a dormant period, or simply for
protection from hard frosts), and an exposition, providing
enough water from the melting snow to keep the substrate
wet over 2–3 weeks, relatively high daily temperatures
(for plasmodium growth), alternating with lower night
temperatures (possibly for inducing fructification). These
habitats occur only very locally in the Khibine mountains,
because conditions like open ground, heavy snow pack in
winter and plants providing much herbaceous biomass are
somewhat contradictory. Only a short time after snow
166 · Yuri K. Novozhilov, Martin Schnittler, InnaV. Zemlianskaia and Konstantin A. Fefelov
melting the ground is open and able to warm up during
the day. Then the plants shoot very quickly and form a
dense cover shadowing the ground. Therefore, the developing window for these species is small, in the Khibine
Mountains probably the second half of June. We were already slightly too late and did not found fresh sporocarps
of the above-mentioned species. From the slopes around
the Kirovsk Botanical Garden obviously the best place
was the avalanche gutter described above, wet enough for
tall perennials and with high snow cover hindering tree
growth.
Surprisingly, also in the plains of the valley grounds
species of this group were found, especially on places with
high umbellifers. The reason may be frequent changes of
weather in the area. Probably the man-made meadows can
serve as a secondary facultative habitat.
The second group of species is more cryophilous,
growing predominantly in summer under cool and wet
conditions on litter, especially in shady woodland.
Physarum cinereum, Didymium deplanatum and D. dubium, perhaps Trichia alpina and Lamproderma sauteri,
can be placed here. Their ecological requirements may be
summarised as: shady ground, high moisture over a longer
period (about two months) a more or less thick layer of
herbaceous plant refuse.
Such habitats are common in the valleys of the Khibine
mountains, also indicated by the abundance of Physarum
cinereum. Hollow, collapsed previous year’s stems of
Cicerbita alpina or Cirsium heterophyllum form the most
important substrate. The microclimate is moist and cold;
here we found fresh sporocarps during the investigation
time (July), especially of Physarum cinereum and
Lamproderma sauteri.
Somewhat different are the microhabitats of two species. Trichia alpina was found predominantly on shaded
leafy litter (especially rowan) between small boulders.
Another exception was Lamproderma sauteri, preferring
moss layers on rocks and boulders with running water. An
extreme example was a place in the alpine tundra, a stony
depression on a NO-exposed slope. On the ground floor
there was a two meter pack of big granite stones; their
sides were covered with mosses, mainly the arctic-alpine
liverwort Gymnomitrium concinnatum (Lightf.) Corda.
Under the stones runs water from a nearby snowfield.
Lamproderma sauteri seems to be the most cryophilous
species, which does not need higher temperature in any
phase of its development. Also the other growth places
were very shady, often almost hidden, e.g. moss-covered
rocks under tree roots.
A shift in species composition and diversity can also
occur from year to year. For example the effects of seasonality was the occurrence of Colloderma oculatum in
moss communities on rocks on Srednii island of Keret
Archipelago in the White Sea (Schnittler and Novozhilov,
1996). There are frequent vertical steps in the granite rocks
on the island, which separate the damp woodlands from
the dry pine-lichen community. If water trickles over for a
long period of time, a thin cover of liverworts and bluegreen algae is formed, especially under big cushions of
mosses. During the summer 1993, these moss and liverworts layers provided a very good microhabitat for
Colloderma oculatum and Lepidoderma tigrinum, especially in eastern exposure on very thin (less than 0.5 cm),
slimy layers of liverworts, covered with a water film. This
microhabitat is found at 1–3 m height on rocks that are
provided with trickling water. The huge colonies, especially of Colloderma oculatum, suggest that moss layers
are a normal microhabitat. The communities are nevertheless unstable, and in the exceptionally warm summer
of 1994, when there was no trickling water, only dry scraps
of dead liverworts were found on these rocks.
Other example of a shift in species composition in
inventory studies is Barbeyella minutissima. The minute
myxomycete Barbeyella minutissima, described by
Meylan (1914) from the Swiss Jura Mountains, was long
thought to be exceedingly rare. Stephenson and Studlar
(1985) considered Barbeyella as strongly bryophilous. The
pattern of occurrence during the year shows that in southern regions Barbeyella fruits in the winter, whereas in more
northern regions the fructification peak occurs in September to early October (eastern North America) or
mid-October (Germany). Observation in the Northern
Ammergauer Alps (Schnittler and Novozhilov, 1998;
Schnittler et al., 2000) provide evidence that Barbeyella
can develop at temperatures between 0 and 10 °C. These
collections were made after a first period of frost in the
year, followed by a couple of warmer autumn days. Only
the most cool and shady parts of the narrow ravines investigated in this study harboured Barbeyella. Observations
and collections made in a narrow and cool valley system
appear to add yet another aspect to the ecology of this
myxomycete – the association with unicellular algae,
which form a slime layer providing continuous moisture
as well as a microenvironment suitable for microbial
growth. In five of seven collections of Barbeyella, algae
were clearly visible, forming a thin, slimy layer on the
wood surface.
General principles for analyses of myxomycete
diversity
1) Size and comparability of the area. To provide data
comparable with those on other regions, study areas
should be of minimal size, but include all major vegetation types of a region. In general, the simplest way
to representative data might be to choose only a certain number of areas limited in size, e.g. in National
Parks, reservations or near biological station of Universities which can be surveyed for a period of years
within other activities and by local people.
2) Thoroughness of investigation. To provide the full
species inventory, a systematic survey of all suitable
microhabitats should be carried out, together with
P r o t is t o lo g y · 167
extensive substrate sampling. Substrate sampling and
detection by the moist chamber method is indispensable. Checklists of myxomycetes should be considered
incomplete unless the list includes a systematic examination of diversity by moist chamber indirect
isolation method.
3) Repeated survey. To ensure the recording of all phenological groups, more than one field survey in a year
is recommended. Four surveys are optimal (spring,
midsummer, early and late autumn).
4) Ecological niches. To reveal the microhabitat requirements of the species, as much as possible data about
the microhabitat should be gathered besides collecting of specimens of substrata for moist chamber
cultures (see above).
As shown for the commoner species, primarily substrate features, with pH, texture of bark, and probably water
retention as the most important factors determinate ecological niches.
The practical examples of estimation and measurement of local and regional species diversity
The foregoing methodical approaches and general
principles of inventory studies of myxomycetes were used
in study of myxomycete biota of Russia and in some
neighbouring countries.
According to our database, 326 species are reported
from this area. The main biomes studied are: tundra and
forest-tundra (Yamal peninsula, Taimyr peninsula), taiga
or boreal coniferous forests (Tver’ province, Leningrad
province, Karelia, Kola Peninsula, Altai Krai, and Ural
Mountains), deciduous temperate forests (Northern
Caucase, Krasnodarskii Krai, and Vladivostok province),
mediterranean xerophilic forests “shibliak” (Crimea peninsula), forest steppe and steppe (Voronezh and Volgograd
provinces), and desert (Volgograd, Astrakhan’ provinces,
Kalmykia, and Mangyshlak peninsula in Kazakhstan).
Some study areas where the most comprehensive and intensive of ecological studies of Myxomycetes have been
carried out are shown on the Fig. 2. and summarised in
tables 1, 2.
In the myxomycete biotas of different natural zones
of Russia there are different sets of main ecological groups.
For example in extreme conditions of high-latitude and
arid regions not only the reduction of species numbers takes
place but also the simplification and elimination of some
groups (for example lignicolous K-strategs with large
phaneroplasmodium of Physarales, (see table 1).
Corticolous myxomycetes adapted to the life in the bark
of trees and bushes reflect the climate conditions to a
greater extent than the habitants of litter, dung, and decayed wood. In spite of their wide distribution, certain
groups of myxomycetes have areas of mass reproduction.
The structure of desert corticolous myxomycetes is the
simplest one. The Echinostelium, Licea, and Perichaena
dominate here. In the steppe and forest steppe, there dominate Echinostelium minutum, Macbrideola cornea. In
temperate deciduous forests, there dominate Macbrideola
cornea, Cribraria violacea, Physarum decipiens, and Licea
operculata. Arcyria pomiformis, Macbrideola cornea, M.
synsporus, Echinostelium colliculosum, and Licea
kleistobolus dominate in Crimea “shibliak” communities.
Considerable differences between southern taiga and northern taiga are not observed. E. minutum, Paradiacheopsis
fimbriata, Licea parasitica, and L. minima dominate here.
Fig. 2. Location map showing the main collecting areas for plasmodial slime moulds in Russia and Kazakhstan. A dotted line
indicates the northern limit of boreal forests according to Word Atlas “Resources and Environment” (1998). White circles
indicate the collecting areas in tundra and forest tundra biomes, half-black ones – boreal forest (taiga) biomes, black ones –
desert and steppe biomes.
168 · Yuri K. Novozhilov, Martin Schnittler, InnaV. Zemlianskaia and Konstantin A. Fefelov
Table 1. Occurrence of myxomycetes in tundra, boreal, and desert zones of Russia and Kazakhstan.
(Explain remarks: the number before a slash indicates the total of all specimens collected in the field, whereas the number
after a slash indicates the total of all specimens obtained from moist chambers; abbreviations used for the study areas are:
KM = Khibine Mountains, PU = Polar Ural, PP = Plateau Putorana, TP = Taimyr Peninsula, CH = Chukchi Peninsula,
RK = Russian northern Karelia, LE = Leningrad province, SV = Sverdlovsk province, VO = Volgograd province,
KZ = Mangyshlak Peninsula; study areas are situated in: tundra zone and forest tundra, (KM, PU, PP, TP, CH); boreal zone,
taiga (RK, LE, SV); steppe and cold desert zones, intrazonal vegetation in riparian and wash woodlands (VO); central and
southern desert, desert vegetation on the eastern shore of the Caspean Sea in the Mangyshlak Peninsula (KZ); widely
distributed species (recorded from more than 4 study areas) are listed in bold; species with records from nivicolous situations are indicated with an asterisk).
Myxomycetes
Amaurochaeta atra
Arcyodes incarnata
Arcyodes incarnata
Arcyria cinerea
Arcyria denudata
Arcyria ferruginea
Arcyria helvetica
Arcyria incarnata
Arcyria insignis
Arcyria magna
Arcyria minuta
Arcyria obvelata
Arcyria oerstedtii
Arcyria pomiformis
Arcyria stipata
Badhamia affinis
Badhamia capsulifera
Badhamia panicea
Badhamia foliicola
Badhamia macrocarpa
Badhamia obovata
Badhamia populina
Badhamia utricularis
Brefeldia maxima
Calomyxa metallica
Ceratiomyxa fruticulosa
Clastoderma debaryanum
Colloderma oculatum
Comatricha cf. rigidereta
Comatricha dictyospora
Comatricha elegans
Comatricha irregularis
Comatricha laxa
Comatricha longa
Comatricha nigra
Comatricha pulchella
Comatricha tenerrima
Comatricha typhoides
Craterium aureum
Craterium leucocephalum
Craterium minutum
Cribraria argillacea
KM PU
1/–
1/5
PP
TP
CH
RK
LE
3/–
4/–
SV
2/–
VO
KZ
–/1 1/1
–/13 1/19 –/23 2/12 –/9
–/1 –/1
1/–
1/–
1/1
1/5
1/9
2/7
2/–
58/7 12/2 21/–
8/– 1/– 10/–
1/– 6/–
1/–
11/1 5/– 33/1 23/–
1/– 1/– 2/–
1/–
–/1
–/1
1/1
1/–
–/1
–/2
1/1
3/–
5/–
5/–
3/–
1/–
1/–
4/–
1/–
1/–
3/–
1/–
1/–
1/–
9/–
9/–
1/–
47/6 9/–
5/–
3/–
1/–
3/–
1/–
1/–
1/–
5/–
1/–
3/–
2/1
–/3
–/1
2/3
–/1
–/1
2/3
4/7
2/–
–/1
1/–
–/3
3/–
3/–
1/–
2/–
3/– 10/– 34/–
2/– 1/– 7/–
13/– 1/–
1/–
1/–
9/– 1/– 7/–
8/–
1/1 9/– 2/– 13/–
2/–
1/35 2/10 12/2 9/– 60/1
–/1
4/– 7/–
1/– 1/–
2/1 4/– 8/– 8/–
1/–
–/1
6/– 5/– 1/–
3/–
7/– 5/– 6/–
9/–
1/–
15/–
4/– –/6
2/–
1/–
11/–
2/–
P r o t is t o lo g y · 169
Table 1. Continuation
Cribraria aurantiaca
Cribraria cf. atrofusca
Cribraria intricata
Cribraria languescens
Cribraria macrocarpa
Cribraria microcarpa
Cribraria minutissima
Cribraria piriformis
Cribraria piriformis
Cribraria purpurea
Cribraria rufa
Cribraria splendens
Cribraria tenella
Cribraria violacea
Cribraria vulgaris
Diachea leucopodia
Diachea splendens
Diacheopsis effusa *
Diacheopsis sp. A
Diacheopsis sp. B
Dianema corticatum
Dictydiaethalium plumbeum
Dictydium cancellatum
Diderma asteroides
Diderma cf. simplex
Diderma deplanatum
Diderma floriforme
Diderma globosum
Diderma hemisphaericum
Diderma montanum
Diderma niveum *
Diderma radiatum
Diderma sauteri
Diderma sp. A
Diderma spumarioides
Diderma trevelyani
Didymium anellus agg.
Didymium annulisporum
Didymium clavus
Didymium crustaceum
Didymium difforme
Didymium dubium *
Didymium iridis
Didymium melanospermum
Didymium minus
Didymium nigripes
Didymium squamulosum
Echinostelium arboreum
Echinostelium brooksii
Echinostelium colliculosum
Echinostelium minutum
Enerthema papillatum
1/–
7/–
6/–
5/–
3/–
2/–
2/–
2/–
3/–
3/–
–/1
–/4
8/–
2/–
3/–
2/–
1/–
–/4
–/1
1/–
1/–
3/–
2/–
4/–
1/–
4/–
2/–
1/–
1/–
5/–
21/–
3/–
1/– 1/–
3/– 1/–
1/–
1/–
1/–
1/–
1/–
1/–
8/–
1/–
2/–
1/–
10/– 23/– 12/–
1/–
2/–
1/–
2/–
2/–
1/–
11/– 3/–
3/–
2/–
2/–
6/– 2/–
1/–
1/–
1/–
1/–
1/–
1/–
6/–
3/1
4/1
1/1
–/1
6/–
4/–
2/–
2/–
3/–
5/–
6/–
–/10 –/15 –/45 –/21 –/6
–/4 –/1 1/4 1/1 9/2
–/4
7/–
4/1
–/2
–/1
–/2
–/1
–/2
–/1
–/5
1/–
2/–
1/–
3/–
3/–
4/–
2/–
–/1
3/–
1/–
–/1
1/1
8/–
–/24
–/1
2/–
1/–
7/–
4/–
–/12
1/–
1/–
4/–
3/–
3/–
2/–
4/–
1/–
21/– 8/3
–/16
–/7
1/22
39/– 1/–
–/45
–/3
170 · Yuri K. Novozhilov, Martin Schnittler, InnaV. Zemlianskaia and Konstantin A. Fefelov
Table 1. Continuation
Enteridium intermedium
Enteridium lycoperdon
Enteridium oliviaceum
Enteridium splendens
var. juranum
Fuligo cinerea
Fuligo leviderma
Fuligo septica
Hemitrichia abietina
Hemitrichia clavata
Hemitrichia intorta
Hemitrichia karstenii
Hemitrichia serpula
Lamproderma arcyrioides *
Lamproderma arcyrionema
Lamproderma carestiae *
Lamproderma columbinum
Lamproderma fuscatum *
Lamproderma qulielmae
Lamproderma sauteri *
Lamproderma scintillans
Leocarpus fragilis
Lepidoderma aggregatum *
Lepidoderma carestianum *
Lepidoderma granuliferum *
Lepidoderma tigrinum
Licea biforis
Licea castanea
Licea cf. belmontiana
Licea denudescens
Licea kleistobolus
Licea marginata
Licea minima
Licea operculata
Licea parasitica
Licea pusilla
Licea sp. A
Licea sp. B
Licea testudinacea
Licea variabilis
Lindbladia tubulina
Lycogala epidendrum
Lycogala exiguum
Lycogala flavofuscum
Macbrideola cornea
Macbrideola oblonga
Metatrichia floriformis
Metatrichia vesparium
Mucilago crustacea
Oligonema flavidum
Oligonema fulvum
Oligonema schweinitzii
1/–
1/–
3/–
1/–
1/–
2/–
3/–
1/–
1/–
1/–
2/–
1/–
1/–
1/–
3/–
3/–
12/– 23/–
1/–
7/– 12/– 1/–
1/–
1/–
4/– 1/–
3/–
4/– 7/– 6/–
7/–
5/–
1/1
2/–
1/–
5/–
1/–
2/–
–/2
1/–
1/–
12/–
1/–
3/–
2/–
10/–
3/–
2/–
–/1
3/2
2/–
3/–
2/–
7/–
2/–
6/–
1/–
4/–
2/–
3/–
1/–
1/–
–/3
2/–
–/4
–/1
1/–
–/6
–/1
–/6
–/1
–/3 –/4
–/9
–/10
–/19 –/3
–/1
–/3
–/2
1/–
13/1 –/5
–/2
–/1 –/3
–/1
–/2
–/21
–/4
6/8
–/1
–/1
–/5
–/15
–/1
1/–
2/–
–/2
2/–
2/–
4/–
–/3
6/–
2/–
4/–
–/2
–/1
2/–
–/4
2/–
11/– 17/– 12/–
1/–
3/–
1/–
1/– –/1
–/25
1/–
1/–
13/–
1/–
6/–
7/–
1/–
1/–
21/– 5/–
11/–
6/–
1/–
P r o t is t o lo g y · 171
Table 1. Continuation
Paradiacheopsis cribrata
Paradiacheopsis fimbriata
Paradiacheopsis solitaria
Perichaena chrysosperma
Perichaena corticalis
Perichaena depressa
Perichaena liceoides
Perichaena minor
Perichaena taimyriensis
Perichaena vermicularis
Perichena quadrata
Physarum alpinum *
Physarum auriscalpium
Physarum bitectum
Physarum bivalve
Physarum cf. carneum
Physarum cf. confertum
Physarum conglomeratum
Physarum leucopus
Physarum cf. nudum
Physarum cinereum
Physarum citrinum
Physarum compressum
Physarum contextum
Physarum decipiens
Physarum didermoides
Physarum famintzinii
Physarum flavicomum
Physarum globuliferum
Physarum leucophaeum
Physarum notabile
Physarum nutans
Physarum oblatum
Physarum pezizoideum
Physarum psitacinum
Physarum pulcherrimum
Physarum pulcherripes
Physarum rubiginosum
Physarum serpula
Physarum sp. A
Physarum straminipes
Physarum sulphureum
Physarum tenerum
Physarum vernum
Physarum virescens
Physarum viride
Protophysarum phloiogenum
Prototrichia metallica
Stemonitis axifera
Stemonitis fusca
Stemonitis herbatica
Stemonitis hyperopta
–/2
–/20 –/3
–/6
–/2
–/1
–/5
–/2
–/3
–/3
–/1
–/5
1/2
1/3
–/2
–/3
–/1
–/2
3/–
1/–
–/2
1/–
–/2
–/1
–/2
–/4
–/6
–/2
3/36
–/1
6/–
1/–
3/–
1/–
–/15
–/3
–/1
–/1
1/–
1/–
3/–
–/11 2/–
1/–
–/4
4/–
1/–
1/–
1/–
3/–
5/–
1/–
3/–
3/–
/1
7/–
1/– 2/–
10/–
–/42
2/–
–/6
1/–
–/1
1/–
16/–
–/4
2/1
2/–
2/–
–/1
1/–
9/–
–/4
1/–
2/–
4/3
–/1
2/3
–/2
–/1
2/2
–/2
3/6
–/2
7/–
2/4
5/–
3/–
1/–
1/–
1/–
1/–
9/–
1/–
12/–
1/–
1/–
–/1
–/1
–/1
1/–
5/–
4/– 8/–
3/– 1/– 2/82
70/– 10/–
1/–
6/–
2/–
1/–
8/–
1/–
1/–
1/–
–/2
6/–
1/–
2/–
1/–
–/1
2/–
3/–
1/–
1/–
1/– 4/–
14/1 7/–
–/4
1/–
2/1
1/–
1/–
9/–
4/–
7/–
1/–
3/–
12/– 3/–
–/3
–/1
1/–
3/–
1/–
2/–
5/1
4/2
3/–
1/–
16/– 16/–
27/– 12/–
1/–
4/– 3/–
172 · Yuri K. Novozhilov, Martin Schnittler, InnaV. Zemlianskaia and Konstantin A. Fefelov
Table 1. Continuation
Stemonitis nigrescens
Stemonitis pallida
Stemonitis smithii
Stemonitis sp.
Stemonitis splendens
Stemonitis virginiensis
Stemonitopsis microspora
Stemonitopsis subcaespitosa
Symphytocarpus confluens
Symphytocarpus flaccidus
Trichia alpina *
Trichia botrytis
Trichia contorta
Trichia decipiens
Trichia erecta
Trichia favoginea
Trichia floriformis
Trichia lutescens
Trichia munda
Trichia scabra
Trichia subfusca
Trichia varia
Tubifera ferruginosa
Tubifera cf. microsperma
1/–
1/–
1/–
1/–
2/–
4/–
1/–
6/–
1/–
–/1
–/1
–/1
2/–
1/–
19/–
–/1
1/–
–/2
18/1
6/– –/2
2/–
1/–
1/–
1/–
–/2
–/10 –/1
–/2
1/1
1/–
2/–
–/2
–/5
–/1
–/5
2/–
2/1
1/–
4/–
6/–
1/–
2/–
1/–
4/– 4/– 19/–
4/– 5/– 6/– 1/–
16/– 10/– 20/– 1/–
1/–
12/– 1/–
2/–
7/–
3/–
1/–
1/–
In general, the myxomycete biota of the tundra zone of
the Polar Ural, the Taimyr Peninsula, and the Chukchi
Peninsula can be considered as impoverished biota of the
northern taiga subzone. In extreme conditions (tundra, and
desert ecosystems) the amount of species decreases and
some species disappear. At the same time occurrence of
Echinostelium minutum and Licea minima is not only lower
but in some habitats in tundra (on bark of Dushekia
fruticosa) and desert (on bark of Artemisia) is rather high.
Thus, probably in this case the inverse dependence between population density and species richness in extremal
conditions is obvious.
Due to the poor knowledge about myxomycete distribution we should be careful in making conclusions about
their geography. The majority of species in Russia have
polyzonal areas but some species show high abundance
and mass distribution in separate climatic zones (tables 1,
2). There are arct-alpine species (numerous nivicolous
species e.g. Lamproderma, Diderma, etc.), arct-boreal (e.g.
T. munda), boreal (Trichia varia), and desert-steppe species (Echinostelium colliculosum). The overall high degree
of similarity between the biotas of the 10 study areas (expressed as coefficient of community indices in table 3)
certainly suggests that most species of myxomycetes have
high dispersal capabilities. Exceptional was western
Kazakhstan, with extremely severe arid climatic conditions. The extreme fluctuations in air humidity favour
species with a short development time or those able to
3/1
3/–
1/–
5/–
1/–
7/– 2/–
1/–
9/– 14/– 1/–
10/– 19/– 3/–
2/–
survive repeated desiccation during development in desert
areas (Schnittler and Novozhilov, 1999).
In the tundra, myxomycetes are represented mainly
by multizonal and even cosmopolitan species. Many boreal species are widely distributed within the northern taiga
and can be found also in forest-tundra and tundra vegetation (table 1). However, only Echinostelium minutum (2
collections), Didymium dubium (2), Craterium
leucocephalum (1), Licea minima (1), and L. testudinacea
(1) were recorded for typical tundra.
In general, the species richness of myxomycetes decreases northwards. In the Taimyr Peninsula (Fig. 3) results
of moist chamber cultures obtained for all substrate types
are demonstrated this trend (table 4). Due to the patchy
nature of the vegetation, collections from one geographical locality may be assigned to more than one subzone.
The Shannon diversity index for the taiga (H’ = 1.31) is
slightly higher than the value for the forest-tundra (H’ =
1.28), whereas the value for the tundra is much lower (H’
= 0.99). However, this pattern differs among particular
ecological groups. For example, the mean value of the
number of wood-inhabiting species per moist chamber
culture decreases from 3.54 in the taiga to 1.66 and 1.13
in the forest-tundra and tundra, respectively. This correlates with a decrease in species richness and diversity.
Corticolous myxomycetes exhibit similar patterns. In contrast, litter-inhabiting myxomycetes exhibit a higher
diversity in forest-tundra (H’ = 1.01) and tundra (H’ =
P r o t is t o lo g y · 173
Table 2. Summary data for specimens of myxomycetes collected in the field and from moist chambers
(abbreviated as ‘MC’) for the ten study areas of Russia and Kazakhstan.
(Explain remarks: since more than one myxomycete species may occur in a given moist chamber, the number of specimens
obtained from a set of moist chambers for a single study area is usually higher than the total number of positive moist
chambers. Exceptions may occur in study areas where many moist chambers were prepared with litter, because these moist
chambers often have high proportions of plasmodia that cannot be induced to fruit. These non-fruiting plasmodia remain
unidentified and are not considered in the numbers of specimens from moist chambers. Moist chamber data could not be
reconstructed for Karelia, Leningrad, Volgograd, and Sverdlovsk provinces, and are therefore not included in the totals
given in the last column. ND – no data; abbreviations for study areas are the same as those used in Table 1).
Study area
KM PU
PP
TP
CH RK
LE
SV
VO
KZ
Total
Wood: MC positive 3
MC prepared 7
% positive
43
9
11
82
10
14
71
56
83
67
14
22
64
ND
ND
ND
ND
ND
92
137
67
Bark: MC positive
MC prepared
% positive
9
71
13
32
37
86
17
19
89
37
61
61
27
32
84
ND
ND
ND
ND
72
81
89
194
301
64
Litter: MC positive
MC prepared
% positive
5
32
16
13
19
68
3
14
21
20
49
41
19
31
61
ND
ND
ND
ND
29
35
83
89
180
50
Dung: MC positive
MC prepared
% positive
1
13
8
4
6
67
1
5
20
6
25
24
9
12
75
ND
ND
ND
ND
16
30
53
37
91
41
Total: MC positive
MC prepared
% positive
18 58
123 73
15 80
31
52
60
119 69
218 97
55 71
ND
ND
ND
ND
ND
ND
ND
ND
ND
ND
ND
ND
117
146
80
412
709
59
Specimens collected
in the field
Specimens from mc
Number of species
Number of genera
Species/genus ratio
101 22
27
14
68
341
493
821
352
10
2249
18
40
20
2.0
76
29
15
1.9
249
48
22
2.1
111
48
23
2.0
56
96
33
2.9
35
148
40
3.7
82
105
32
3.3
ND
77
25
3.0
323
27
12
2.3
1051
222
42
5.3
101
35
19
1.8
0.90) than in the taiga (H ‘= 0.58). Myxomycetes cultured
from dung in the taiga subzone occurred too sporadically
to indicate any distribution trends.
However, zonal limits of myxomycete distribution are
relative. Presumably, differences among myxomycete assemblages in taiga, forest-tundra, and tundra are more the
result of differences in the abundance of shared species
than actual differences in species composition.
Obviously, the main factors for the decrease in the
number of myxomycete species in desert and arctic regions are unfavourable hydro-temperature conditions and
the reduced range and extent of available microhabitats
(α-diversity). For example as shown for the common species in the desert regions of Kazakhstan, ecological niches
are determinated primarily by substratum features, with
pH, texture and probably water retention as the most im-
portant factors (Schnittler, 2000). The CCA (canonical
correspondence analysis), mean values for niche states
restricted to the microhabitat (pH, substratum type) were
lower than for climatic parameters (light, wind). This may
express difficulties in estimating the latter parameters for
a very small space, but more probably it reflects the generally higher importance of microhabitat features in
comparison to habitat-describing parameters for myxomycetes. Regarding the generally low utilization of space
(also in the most productive moist chambers not all substratum pieces had myxomycete colonies), a low niche
overlap in already one dimension seems to allow the coexistence of two species.
The majority of species found in the boreal zone belongs to the group of xylophils and occurs in association
with wood and bark debris (Eliasson, 1981; Schnittler and
174 · Yuri K. Novozhilov, Martin Schnittler, InnaV. Zemlianskaia and Konstantin A. Fefelov
Table 3. Pairwise comparisons of myxomycete biotas among the 10 study areas.
(Explain remarks: both coefficient of community indices (upper right) and numbers of species shared in common (lower left) are
given; abbreviations for study areas are the same as those in Table 1.)
KM
KM
PU
PP
TP
CH
RK
LE
SV
VO
KZ
11
14
15
18
27
31
23
23
4
PU
PP
TP
CH
RK
LE
SV
VO
KZ
0,29
0,41
0,44
0,34
0,58
0,55
0,41
0,67
0,44
0,63
0,40
0,38
0,34
0,43
0,46
0,33
0,38
0,26
0,35
0,28
0,62
0,32
0,36
0,27
0,42
0,38
0,58
0,57
0,39
0,29
0,36
0,30
0,40
0,47
0,56
0,49
0,12
0,06
0,11
0,21
0,16
0,10
0,16
0,14
0,12
15
24
28
25
29
25
16
2
21
17
21
23
18
19
3
30
31
34
32
19
8
33
41
29
25
6
76
58
41
8
72
63
14
45
9
6
Fig. 3. Map of the Taimyr Peninsula showing the location of the ten study sites (black rectangles). A dotted line shows the northern
boundary of the light larch taiga, whereas solid black lines indicate the boundaries of the tundra subzones: I- northern taiga;
II – forest-tundra; III – southern tundra; IV – typical tundra; V – arctic tundra; VI – polar desert (according to Chernov and
Matveyeva, 1997).
P r o t is t o lo g y · 175
Table 4. Results obtained from moist chamber cultures prepared with substratum samples
collected in the vegetation subzones of the Taimyr Peninsula.
Vegetation
Subzones
Number
of moist
chamber
cultures
Positive moist
chamber
cultures
(% of total)
Number
of
collections
Number
of
species
Average yield
(species per
moist chamber)
Mean ± SE
Shannon
diversity
index (H´)
I Taiga
Wood (w)
Litter (l)
Bark (b)
Dung (d)
II Forest tundra
Wood (w)
Bark (b)
Litter (l)
Dung (d)
III-IV Tundra
Wood (w)
Bark (b)
Litter (l)
Dung (d)
55
24
7
21
3
110
34
34
25
17
105
39
25
31
10
45 (82)
21 (88)
3 (43)
18 (86)
1 (33)
54 (49)
21 (62)
19 (56)
8 (32)
4 (24)
51 (49)
22 (56)
14 (56)
12 (39)
2 (20)
122
79
5
37
1
119
54
39
19
7
90
45
26
16
3
32
28
4
16
1
36
22
15
13
5
22
15
8
9
2
2.19 ± 0.26
3.54 ± 0.48
0.71 ± 0.40
1.76 ± 0.23
0.33 ± 0.33
1.08 ± 0.12
1.66 ± 0.27
1.15 ± 0.21
0.76 ± 0.28
0.41 ± 0.15
0.84 ± 0.10
1.13 ± 0.21
1.04 ± 0.20
0.52 ± 0.14
0.30 ± 0.21
1.31
1.26
0.58
1.06
0
1.28
1.24
0.94
1.01
0.67
0.99
0.98
0.76
0.90
0.28
All subzones
270
145 (54)
331
48
1.22 ± 0.09
1.32
Wood (w)
Bark (b)
Litter (l)
Dung (d)
97
80
63
30
64 (66)
51 (64)
23 (37)
7 (23)
178
102
40
11
37
21
16
8
1.82 ± 0.19
1.27 ± 0.13
0.63 ± 0.13
0.32 ± 0.11
1.29
1.03
1.05
0.88
Novozhilov, 1996). The xylophilic species diversity in
forest-tundra and tundra considerably decreases compared
to the boreal (taiga) zone, but the abundance of some species remains rather high (Novozhilov et al., 1998a, 1998b).
For example, towards the tundra zone in the Taimyr Peninsula xylophilic species may penetrate into woodless
territories inhabiting the tiny branchlets of shrubs in bush
“islands” communities. Small dead and dying twigs of
shrubs (Salix spp. and Duschekia) with exfoliating bark
represent a rather suitable substrate for some species. When
very moist, such bark easily becomes exfoliated, forming
numerous small gaps or “shelters” between wood and bark.
These “shelters” play a role of natural moist chambers,
where the plasmodium can survive under unfavorable conditions.
diversity in extreme conditions) as plants and animals. Our
research and literature data show that some species have
rather limited distribution connected with certain climatic
zones and habitats. Nevertheless, these patterns are seen
only at the scale of whole natural climatic zones and large
ecosystems (biomes) characterised by similar vegetation,
fauna, and climate when different groups of habitats are
studied. At the present level of knowledge, further comparative investigations in other regions of Russia,
especially steppe and desert biomes, are necessary to explore the whole richness of biotic diversity of myxomycetes
and reveal their world-wide distribution patterns. Using
of standardised sampling procedures and the introduction
of a systematic program for surveying small representative areas (“concrete biota”) throughout the large territory
of Russia would rise our knowledge on myxomycete
biodiversity to a new level with reasonable efforts.
Conclusion
In summary it may be said that myxomycetes with
their vagility, relatively short life cycle and high ecological adaptation demonstrate the same patterns of distribution
(constant structure of communities within certain biomes,
reduction of adaptive complexes and decrease of species
Acknowledgements
The work of the first author was supported in part by
grants (N 96–04–48209, N 98–04–48120, N 98–07–
176 · Yuri K. Novozhilov, Martin Schnittler, InnaV. Zemlianskaia and Konstantin A. Fefelov
90346) from the Russian Foundation for Basic Research
[RFBR].
We acknowledge logistical support provided by Dr.
D. Bolsheianov of the Arctic and Antarctic Research Institute, St. Petersburg, Russia. We are grateful to the
organisation ‘Biopractica’ (Dr. A. Balachonov, Dr. A.
Markov, Dr. M. Molitvin) for providing us the opportunity to do field work at the biostation of St. Petersburg
University. We are grateful to the staff of the Polar-Alpine
Botanical Garden for offering us the possibility for fieldwork around Kirovsk. Appreciation is extended to Dr. I.
Yu. Kirtsideli for collecting substrate samples for moist
chambers in some areas of the Taimyr Peninsula. For confirming determinations or loan of authentic or type
material, we are indebted to Mrs. M. Meyer, H. Keller, A.
Garcia, C. Lado, D.W. Mitchell, G. Moreno, D. Wrigley
de Basanta, J. Rammeloo, U. Eliasson, S.L. Stephenson,
and Y. Yamamoto. We also wish to express our thanks to
L.A. Karzeva, St. Petersburg, for technical assistance during the SEM-investigations.
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Address for correspondence: Yuri K. Novozhilov. V.L.Komarov Botanical Institute of the Russian Academy of Sciences, Prof. Popova Street, 2, 197376 St. Petersburg, Russia. E-mail: mixus@YN1091.spb.edu
The manuscript is presented by A.A.Dobrovolskij and A.V.Goodkov