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Myxomycetes of boreal forests
of the Laplandskiy State Nature
Biosphere Reserve
(Kola Peninsula, Russia)
Yuri K. Novozhilov
1
, Oleg N. Shchepin
1
,
Vladimir I. Gmoshinskiy
2
, Martin Schnittler
3
1
Komarov Botanical Institute of the Russian Academy
of Sciences, Laboratory of Systematics and
Geography of Fungi, Prof. Popov Street 2, 197376 St.
Petersburg, Russia
2
Lomonosov Moscow State University,
Faculty of Biology, Mycology and Algology Dept.,
119234 Moscow, Russia
3
Institute of Botany and Landscape Ecology,
Greifswald University,
Soldmannstr. 15, D-17489 Greifswald, Germany
* Corresponding author:
yurinovozhilov@gmail.com
Keywords: Amoebozoa, Arctic, Kola Peninsula,
Myxogastria, Russia, biodiversity, mountainous tundra,
northern taiga, slime molds, species inventory.
Article info:
Received: 01 July 2020
Accepted: 21 September 2020
Published online: 22 November 2020
Corresponding Editor: Nikki Heherson A. Dagamac
Abstract
Northern taiga forests and subalpine plant com-
munities of the Laplandskiy State Nature Biosphere
Reserve (Kola Peninsula, Russia) were surveyed for
myxomycetes. A total of 1675 specimens of myxo-
mycete fruit bodies (sporocarps) were registered,
among them 1584 records from eld collections and
92 obtained from 210 moist chamber cultures of
ground litter, bark of living trees, wood, and weath-
ered dung of moose and willow ptarmigan. Most of
125 taxa (124 morphospecies and one variety) rep-
resenting 34 genera were recorded only in the eld
(104 taxa from 32 genera), but some were exclusively
obtained from moist chamber cultures (8 taxa from
5 genera). All of the recorded species are new for the
Laplandskiy Reserve. Species numbers decreased
among the four studied forest associations along the
elevation and mositure gradient, and the Shannon
index showed a similar trend: spruce forest (PICa; 84
taxa, H’=3.8), spruce-peat moss forest (PICb; 70, 3.5),
dry spruce-pine forest (PIN; 62, 3.7), subalpine birch
forest (SB; 30, 2.7). e estimated completeness of
the survey according to the Chao1 estimator was
66%, indicating that most of the more common spe-
doi:10.29203/ka.2020.501
Karstenia, Volume 58 (2020), Issue 2, pages 292–315
www.karstenia.fi
ORIGINAL
RESEARCH
293
cies should have been recovered. e trend among
forest associations runs mainly parallel to diversity:
PICa and PICb 83%, PIN 47%, SB 57%. e myxo-
mycete assemblage of dry coniferous forests is the
most distinctive among the three forest types and
shows the highest number of indicator species. e
overall degree of specialization of myxomycetes is
higher for substrate type than for forest associa-
tions. Among substrate types, species diversity and
richness increase from litter over bark to wood.
Introduction
Taiga, or boreal forest, is the biggest biome in
Russia. In the European part of Russia, the south-
ern and middle subzones of this biome are rather
well surveyed for myxomycetes (Novozhilov 1980,
1999, Schnittler & Novozhilov 1996, Novozhilov &
Lebedev 2006, Barsukova et al. 2012, Erastova &
Novozhilov 2015, Gmoshinskiy & Matveev 2016,
Gmoshinskiy et al. 2019). However, the myxomycet-
es of the northern border of this biome, including
mountain subalpine birch forests, remained poorly
studied (Stephenson et al. 2000).
A number of studies are published for Finland
and Norway, where taiga forests show a similar
structure and species composition (Karsten 1882,
Hintikka 1919, Härkönen 1979, Johannesen 1984b,
Johannesen 1984a, Elvebakk et al. 1996, Härkönen
et al. 1999, Härkönen & Varis 2013, Johannesen &
Vetlesen 2020). In contrast, only four local species in-
ventories have been carried out in the Kola Peninsu-
la and adjacent territories (Schnittler & Novozhilov
1996, Novozhilov & Schnittler 1997, Novozhilov et al.
1998, Erastova et al. 2017).
To ll this gap, we have conducted a quantita-
tive survey that allowed to compile the rst anno-
tated checklist of myxomycete species of the Lap-
landskiy State Nature Biosphere Reserve (western
Kola Peninsula). An eort was made to record or
collect every myxomycete colony of sporocarps and
to obtain samples of dierent substrate types for
moist chamber cultures.
e primary objectives of this study were (1)
to obtain baseline data on myxomycete abundance
and biodiversity of the northern taiga forests, (2) to
estimate the degree of completeness that can be
achieved in this type of survey, and (3) to determine
to which extent myxomycete assemblages vary be-
tween dierent vegetation types and substrates
within the studied area.
Materials and methods
Study area
e Laplandskiy Reserve is located in the western
part of the Kola Peninsula about 120 km south of
Murmansk, between 67°10’–68°05’ N and 31°45’–
32°45’ E, covering the major portion of the Chu-
natundra and the Salnye Tundry Mountains (Fig. 1).
e reserve encompasses 2,784km2. Its territory be-
longs entirely to the northern taiga zone (Ogureeva
2010). e border of the orohemiarctic zone runs
70–90 km apart from the reserve’s northern border
(Ahti et al. 1968). e southern border is 120–130
km north of the Arctic Circle. e main mountain
range of the reserve, the Chunatundra, stretches
from west to east for 40 km. e southern foothill
of the massif is enclosed by an arched tectonic de-
pression lled with Chunozero lake, which reaches
up to 40 m depth. e northernmost and highest
peak, Ebruchorr, reaches 1115 m a.s.l. e climate
of the Laplandskiy Reserve is rather humid, but as
well continental, with long cold winters and short
cool summers (Dfc in Köppen climate classica-
tion), which is typical for the Russian taiga ecore-
gion (Kottek et al. 2006; Hogan 2012).
Average monthly temperatures vary from
–13.5 °C
in February to 13.5 °C in July. e average
annual precipitation is about 450 mm, with ca. 40%
falling as snow. e snow cover usually lasts from
mid-October until end of May. Its distribution and
the time of its melting in the mountains depend on
the relief and slope exposition. Precipitation is low-
est towards the end of the winter ( from February to
April, about 30 mm per month), the maximum is in
August and September (Koroleva 1999; Marshall et
al. 2016; Barkan 2020).
e mountain slopes are covered with northern
294
Fig. 1. Maps of the study area and images of typical habitats. a: schematic map of the study area, inset:
geographical position of the Laplandskiy State Nature Biosphere Reserve within the Kola Peninsula,
numbers and circles indicate the 44 sampled localities, b: subalpine birch forest (SB), c: spruce forest
(PICa), d: understory layer in spruce-peat moss forest (PICb), e: dry spruce-pine forest (PIN). Map
sources: Mapbox online service, localities indicated via QGIS.
295
taiga forest, featuring spruce, pine and birch as the
dominating tree species. e elevational timberline
is already reached at ca. 380 m a.s.l.; it decreases near
lakes but increases in sheltered valleys. e rounded
gneiss peaks with average heights of 500–800 m a.s.l.
are covered by arctic mountain tundra, consisting
of heathlands dominated by dwarf shrubs and li-
chens. Snow-bed plant communities are most com-
mon in the middle oroarctic zone, at about 600±80
m a.s.l., and are represented by moss cushions and
grass elds which occupy a fairly narrow area near
long-lasting snow patches (Koroleva 1999).
Localities and vegetation types:
e Laplandskiy State Nature Biosphere Reserve
was exhaustively surveyed during three expeditions
(three weeks in June 2015, three weeks in August
2015, and one week in July 2016).
is study was carried out along the slopes of
the Chunatundra mountain range near lake Chu-
nozero (Fig. 1a), focusing mainly on the northern tai-
ga forest (dominated by spruce) and the subalpine
birch forest (SB). e myxomycetes in the mountain
tundra were not specically studied because they
had been thoroughly investigated in the adjacent
Khibiny Mountains, situated only 45 km east from
the reserve (Schnittler & Novozhilov 1996, Erastova
et al. 2017).
e following brief description refers to veg-
etation types named and classied by Ahti et al.
(1968). Names of vascular plants are listed accord-
ing to Czerepanov (1995). Each studied locality was
assigned to one of the vegetation types. Numbers of
the studied localities are shown in Fig. 1a. Geographi-
cal data and vegetation types of all studied localities
are given below.
Northern taiga spruce forests
Spruce forests mainly with Picea obovata Ledeb. with
well-developed undergrowth of Prunus padus L. and
Sorbus aucuparia L. are most common in the reserve
between 150–250 m a.s.l. Some of these forests are
heavily damaged by res around the Salnye Tundry
mountains and greatly aected by airborne chemical
pollution and res in the surroundings of the Chu-
natundra mountains (Lukina & Chernen’kova 2008).
e most common type of soil is podzol, if the
soil is not waterlogged. Its upper layer is acidic due
to falling needles, favoring lichens and some moss-
es as pioneer plants, which are on favorable sites
replaced by dwarf shrubs like Vaccinium myrtillus
L. and V. vitis-idaea L. e moss layer is well devel-
oped: Pleurozium schreberi (Brid.) Mitt., Hylocomi-
um splendens (Hedw.) Schimp., and Dicranum spp.
dominate drier sites; in wet depressions Sphagnum
spp. prevails (Lukina & Chernen’kova 2008).
In pure spruce forests trees may reach 80 cm
in diameter and form a closed canopy, therefore
litter and logs are shaded and dry out slowly. ese
habitats are rich in coarse woody debris at dier-
ent stages of decomposition. ere are windfallen
trees (uprooted together with the surrounding soil)
and uneven-aged “regeneration gaps” in the birch
and spruce forest canopy (Lukina & Chernen’kova
2008). e fallen trees are mainly shaded and wet, so
decay proceeds faster than in the drier spruce forest
mixed with pine. e stock of the litter (mainly dead
mosses and needles) under tree crowns reaches 40
t/ha (Lukina & Chernen’kova 2008).
Within these spruce forests, three associations
can be dierentiated according to the combina-
tion of dominant dwarf shrub species, lichens and
mosses. We surveyed myxomycetes in moderately
dry spruce forest (ground layer dominated by bry-
ophytes), moist spruce-peat moss forest (ground
layer dominated by Sphagnum spp.), and in very dry,
open spruce-pine forest (ground layer with lichens,
bryophytes, and dwarf Ericaceae shrubs).
SPRUCE FOREST (PICa) – localities 14–16, 20, 25,
34, 35, 36, 37, 38, 39, 44, 45 (Fig. 1c).
In this association, spruce is at its optimum:
the tallest trees can reach 50–80 cm diameter at
breast height (DBH). e understory layer is frag-
mented but sometimes well-developed, with trees
of Prunus padus L., Sorbus aucuparia L. and a rich
herb layer, dominated by dwarf shrubs and herbs,
such as Vaccinium spp., Empetrum nigrum L., Gera-
nium sylvaticum L., and Cornus suecica L. It is rich in
mosses of the genus Dicranum, and ferns in habitats
with sucient spring-water moisture.
Reserve station “Nizhn’aja Chuna”, ecological
route El’njun-2. LOC. 14: SE slope, 292±10 m a.s.l.,
67°39'11.2''N 32°37'20.1''E; LOC. 15: SE slope, 302±10
296
m a.s.l., 67°39'10.6''N 32°37' 25.8''E; LOC. 16: SE slope,
259±10 m a.s.l., 67°39'08.5''N 32°37'47.3''E; LOC. 20:
S slope, 111±10 m a.s.l., 67°39'04.3''N 32°39'03.3''E;
LOC. 25: S slope, 232±10 m a.s.l., 67°39'20.4''N
32°39'50.5''E; LOC. 44: S slope, 227±10 m a.s.l.,
67°39'41.4''N 32°37'47.7''E; LOC. 45: S slope, 167±10
m a.s.l., 67°39'01.6''N 32°39'09.0''E; brook “Ruchei
Kupletskogo”, southern shore of lake Chunozero.
LOC. 34: W slope, 169±10 m a.s.l., 67°40'55.6''N
32°22'01.3''E; LOC. 35: W slope, 162±10 m a.s.l.,
67°40'50.4''N 32°22'09.3''E; LOC. 36: SW slope, 176±10
m a.s.l., 67°40'18.5''N 32°26'11.5''E; near Seidapakhta
Mt., southern shore of lake Chunozero. LOC. 37: SW
slope, 134±10 m a.s.l., 67°40'13.7''N 32°25'43.9''E;
LOC. 38: SW slope, 174±10 m a.s.l., 67°40'18.7''N
32°26'00.0''E; LOC. 39: SW slope, 176±10 m a.s.l.,
67°40'18.5''N 32°26'11.5''E.
SPRUCE-PEAT MOSS FOREST (PICb) – localities
19, 26–31, 33, 40–43 (Fig. 1d). is spruce forest type
occurs at the foot of slopes and near rivulets where
the Sphagnum species constitute a nearly continu-
ous vegetation carpet.
Reserve station “Nizhn’aja Chuna”, ecological
route El’njun-2. LOC. 19: S slope, 368±10 m a.s.l.,
67°40'05.5''N 32°36'21.2''E; LOC. 26: W slope, 167±10
m a.s.l., 67°39'29.3''N 32°39'21.3''E; LOC. 27: W slope,
138±10 m a.s.l., 67°39'04.1''N 32°37'21.3''E; LOC. 28:
S slope, 145±10 m a.s.l., 67°39'02.6''N 32°37'24.4''E;
LOC. 29: S slope, 145±10 m a.s.l., 67°39'02.6''N
32°37'47.0''E; LOC. 30: S slope, 140±10 m a.s.l.,
67°39'15.9''N 32°37'50.7''E; near Seidapakhta Mt.,
southern shore of lake Chunozero. LOC. 31: SW slope,
176±10 m a.s.l., 67°40'18.5''N 32°26'11.5''E; brook
“Ruchei Kupletskogo”, near the waterfall. LOC. 33:
W slope, 191±10 m a.s.l., 67°41'01.7''N 32°21'52.5''E;
eastern shore of lake El’javr. LOC. 40: W slope,
153±10 m a.s.l., 67°39'45.5''N 32°39'59.6''E; LOC. 41:
W slope, 154±10 m a.s.l., 67°40'23.3''N 32°39'37.7''E;
LOC. 42: W slope, 164±10 m a.s.l., 67°40'23.5''N
32°39'37.9''E; LOC. 43: E slope, 222±10 m a.s.l.,
67°40'36.9''N 32°38'30.9''E.
DRY SPRUCE-PINE FOREST (PIN) – localities 21–
24 (Fig. 1e).
A dry, open forest with medium-sized, 30–40
cm DBH scattered trees of Picea obovata Ledeb. and
Pinus sylvestris L. accompanied by many saplings of
both species, together with Juniperus communis L.,
and Betula pubescens Ehrh. Due to the rather large
distances between trees (5–20 m) and their narrow,
pyramid-shaped form, the canopy is not closed.
is plant community is very rich in lichens (Clado-
nia mitis Sandst., C. rangiferina (L.) Weber ex Wigg.,
C. stellaris (Opiz) Pouzar & Vezda). Mosses (Polytri-
chum piliferum Hedwig, Hylocomium splendens
(Hedw.) Schimp., Dicranum fuscescens Turner, D.
drummondii Müll. Hal.) and liverworts (Barbilopho-
zia hatcheri (A. Evans) Loeske and Ptilidium ciliare
(L.) Hampe) occur on hillslopes and terraces with
good drainage (Neshataev & Neshataeva 2002). e
podzol soil is shallow and sandy. e water reten-
tion is low and the soil is moist only a short time af-
ter rainfalls. Many of the spruce and pine trees have
been blown down by wind, so numerous fallen trees
and branches lie on the soil. Spruce logs lie elevated
on their branches for a long time and therefore the
stems do not touch the ground directly. As a conse-
quence, the decay of the stemwood is delayed, and
the bark is much longer attached to the wood. Litter
and logs are mostly exposed to sunlight and there-
fore dry out quickly after rain.
Cordon “Volok”, southern shore of lake Saydala-
mina. LOC. 21: S slope, 104±10 m a.s.l., 67°39'56.7''N
32°22'07.9''E; LOC. 22: S slope, 104±10 m a.s.l.,
67°39'57.7''N 32°21'58.1''E; LOC. 23: S slope, 35±10
m a.s.l., 67°40'02.1''N 32°22'22.9''E. Reserve station
“Ni z h n’a j a Chuna”. LOC. 24: S slope, 185±10 m a.s.l.,
67°39'08.9''N 32°39'41.9''E.
SUBALPINE BIRCH FOREST (SB) – localities 1–13,
17–18 (Fig. 1b).
is type of vegetation (Hämet-Ahti 1963, Weh-
berg et al. 2005) is common between 350 and 500 m
a.s.l., mostly on S-exp. slopes sheltered from the pre-
vailing cold winds from north and north-east, look-
ing in some places like a forest tundra. It is a light
forest with crooked Betula czerepanovii N.I. Orlova
and a second species of rowan, Sorbus gorodkovii
Pojark., forming a low canopy. Creeping individuals
of spruce, aspen and rowan constitute a shrub layer
together with willows, juniper, dwarf birch (Betula
nana L.) and several species of Ericaceae (Empetrum
hermaphroditum Lange ex Hagerup, Vaccinium ulig-
inosum L., and Phyllodoce caerulea L. Bab.). Beside
the lichens Cladonia spp. and Cetraria spp. prevail-
297
ing at drier patches, the herb layer is represented
by several ferns and a rather diverse community
of owering plants. On dry convex slopes, lichens,
blueberries (Vaccinium myrtillus L.) or crowberries
(Empetrum nigrum L.) prevail, whereas at moister
sites cushions of Sphagnum spp. occur together
with blueberries or crowberries. Along streams,
birch is replaced by willow stands.
Reserve station “Nizhn’aja Chuna”, ecological
route El’njun-2. LOC. 1: SE slope, 317±10 m a.s.l.,
67°39'17.8''N 32°37'03.4''E; LOC. 2: SE slope, 338±10
m a.s.l., 67°39'20.9''N 32°36'57.2''E; LOC. 3: SE slope,
363±10 m a.s.l., 67°39'23.7''N 32°36'53.8''E; LOC. 4:
SE slope, 379±10 m a.s.l., 67°39'24.4''N 32°36'51.7''E;
LOC. 5: SE slope, 387±10 m a.s.l., 67°39'24.6''N
32°36'49.0''E; LOC.6: S slope, 404±10 m a.s.l.,
67°39'27.5''N 32°36'31.9''E; LOC. 7: N slope, 406±10
m a.s.l., 67°39'51.6''N 32°35'52.1''E; LOC. 8: S slope,
437±10 m a.s.l., 67°39'36.1''N 32°36'14.0''E; LOC. 9: S
slope, 402±10 m a.s.l., 67°39'27.8''N 32°36'29.0''E; L OC.
10: S slope, 522±10 m a.s.l., 67°40'06.0''N 32°34'14.7''E;
LOC. 11: E slope, 450±10 m a.s.l., 67°39'26.2''N
32°36'57.0''E; LOC. 12: E slope, 464±10 m a.s.l.,
67°39'30.5''N 32°37'02.2''E; LOC. 13: E slope, 442±10
m a.s.l., 67°39'31.5''N 32°37'11.5''E; LOC. 17: N slope,
428±10 m a.s.l., 67°39'37.6''N 32°36'10.4''E; LOC. 18:
SE slope, 362±10 m a.s.l., 67°39'24.7''N 32°36'57.9''E.
Specimen collection and substrate sampling
For each forest type, an eort was made to examine
all types of microhabitats upon which sporocarps of
myxomycetes could potentially be found. Specimens
were identied to the lowest possible taxonomic lev-
el according to Martin and Alexopoulos (1969) and
various original descriptions from the literature
(Farr 1976, Poulain et al. 2011), using a morphospe-
cies concept. Myxomycete nomenclature follows
Lado (2005–2020). Common and easily recognizable
species were sometimes only recorded instead of
being collected, whereas rare and not easily recog-
nizable species were always preserved as herbarium
specimens. e genus Ceratiomyxa was included in
this study due to its ecological equivalence to the
true myxomycetes. We dened a record (colony) as
all sporocarps of one species that shared the same
substrate and clustered together (thus likely to have
developed from one plasmodium). For the purpose
of making determinations, sporocarps were pre-
served as permanent slides in polyvinyl-lactophenol
and/or glycerol gelatin, to distinguish between lime-
less and lime-containing structures.
Substrate samples were collected randomly
within predened plots of homogenous habitat
structure, covering about 500 m2 and placed within
two forest types (PICa: localities 15, 16; SB: local-
ities 1, 4). For all plots, geographical coordinates
were obtained with a hand-held GPS device and the
WGS 84 coordinate system. A total of 321 substrate
samples were collected for moist chamber cultures
(Database S1). All myxomycete substrates were
classied as following: “l” for forest ground litter, “b”
for bark of living trees, “w” for woody debris, “d” for
weathered dung of moose (Alces alces L.) and wil-
low ptarmigan (Lagopus lagopus L.). Moist chamber
cultures were set up in Petri dishes of 9 cm diam.,
with substrate pieces placed on a layer of lter pa-
per. After 24 h, the pH of the waterlogged substrates
was measured using a pH meter Hanna HI98128.
Cultures were incubated under ambient light and at
room temperature (20–24°C) for up to 90 days and
examined for the presence of myxomycetes on six
occasions (days 2–4, 6–8, 11–14, 20–22, 40–44 and
85–90) under high magnication with a dissecting
microscope. A ‘record’ was dened herein as one
or more fruiting bodies of a species that developed
from a moist chamber culture.
Data analysis
To estimate completeness of the survey, the quanti-
tative collection data (every fructication recorded)
was used to construct an individual-based species
accumulation curve with the program EstimateS
(Gotelli & Colwell 2011, Colwell 2014). Two diversity
indices were calculated: Shannon’s diversity index
H´ = –∑Pi ln Pi, where Pi is the relative abundance
(the proportion of the total number of individuals
or records represented by the ith species) of a par-
ticular species (Magurran 2004), and the inverse
Simpson’s dominance index D = 1/∑Pi2. e mean
number of species per genus (S/G) was used as an
indicator of overall taxonomic diversity.
Myxomycete assemblages from dierent veg-
etation communities and substrates were com-
298
pared by using the adjusted incidence-based
Chao-Sørensen index (Ccs). It was computed with
EstimateS and used for a cluster analysis by the
weighted pair-group method (WPGMA) with the
program Statistica 10. Graphs were created with
SigmaPlot 10.0. Species classied as rare (relative
frequency <0.5 % of all records) were excluded from
the abovementioned analysis (Chao et al. 2005,
2006). ere are two sources of a bias in the sam-
pling eort of the dierent plant associations stud-
ied: a) our eorts to collect sporocarps in the eld
were relatively similar in PICa (13 localities), PICb
(12), and SB (15) but were smaller in PIN (4) due to
the relative rarity of this association; b) the moist
chamber culture technique was used only for PICa
and SB (Table 1). erefore, only eld specimens were
used in the comparative analysis of species diversity
and species richness between the forest types, and
data were raried as described above.
To reveal the associations of the species with
dierent types of substrates and plant communi-
ties, an indicator species analysis was performed
(McCune & Meord 2006) using the ‘multipatt’
function from ‘indicspecies’ package in R with ‘In-
dVal.g’ as association function and 9999 permuta-
tions (De Cáceres & Legendre 2009).
All microscopic measurements and observa-
tions were made under a light microscope Zeiss
Axio Imager A1 with dierential interference con-
trast (DIC). Mean spore diameter was calculated
from 10 spore measurements from each collection.
Air-dried sporocarps were studied with a Zeiss
motorized stereo microscope Discovery V20, and
a JSM-6390 LA scanning electron microscope at
10–15 kV at the Core Facility Center of the Koma-
rov Botanical Institute of the Russian Academy of
Sciences, St. Petersburg. For the latter, specimens
were mounted on copper stubs using double-sided
sticky lm and sputter-coated with gold. Voucher
specimens were deposited in the collection of the
rst author in the mycological herbarium of the
Komarov Botanical Institute RAS, Laboratory of
Systematics and Geography of Fungi (LE).
Results and Discussion
The following annotated checklist was compiled
based on our quantitative survey. In the annotated
species list, each taxon name is followed by abbrevi-
ations for names of myxomycete species (mentioned
in Figs. 6, 7, Tables S1, S2, Supplementary Database S),
and an abundance estimation in brackets, including
the abundance class according to the ACOR scale
(Stephenson 1993), and records from eld / moist
chamber cultures. Next, the occurrence of a species
in the four different forest types and in the major sub-
strate types is listed. The abbreviations are explained
in the Materials and Methods. The next string
preceded by the abbreviation “Loc.” lists all locali-
ties where a species was found (see Figs. 1–2). Finally,
all or some (indicated by the string “...”) specimen
numbers are given referring to the herbarium LE.
Nivicolous species recorded in the adjacent Khibiny
Mountains (Novozhilov & Schnittler 1997, Erastova
et al. 2017) are labeled as «Khi».
Annotated species list
Arcyria anis Rostaf. [ARCa, O, 11/0] PICA: 4,
PICB: 2, PIN: 5; W: 11; LO C. 14, 21, 27, 41; LE306177…
All specimens displayed a typical capillitium
connected to the stalk apex and ornamented by
warts, spines, cogs, half-rings and rings and with a
number of ridges which may unite locally to form a
reticulum.
Arcyria cinerea (Bull.) Pers. [ARCcin, C, 9/27]
SB: 20, PICa: 12, PICb: 4; B: 13, L: 2, D: 9, W: 12; LO C.
1, 4, 14, 15, 16, 18, 30, 31, 35, 38, 42; LE306141…
All specimens found in the eld show the typ-
ical morphology with long grey sporothecae. Spec-
imens from moist chamber cultures with the bark
of living trees or dung may have yellowish ovoid
sporocarps and resemble A. pomiformis. e dif-
ferentiation between these species was thorough-
ly described in Farr (1962) and Härkönen (1977b).
Following these authors, we assigned to A. cinerea
the specimens with a capillitium that shows two or
three parts, including the part at base of sporotheca
with smooth thick tubes, becoming thinner and or-
namented by short spines in the middle part and by
long spines in the upper part.
Arcyria ferruginea Saut. [ARCfer, R, 4/0] PICa:
2, PICb: 2; W: 4; LOC. 25, 38, 42; LE306225…
e capillitium of our specimens has charac-
teristic features including a large brownish-orange
capillitium connected to the stalk apex and orna-
299
mented with warts and spirally arranged rings and
half-rings and usually reticulate ridges. Spores pale
ochraceous, large, 9–11 µm diam.
Arcyria helvetica (Meyl.) H. Neubert, Nowotny
& K. Baumann [ARChel, R, 2/0] PICa: 2; W: 2; LO C.
25, 39; LE306224, 306612.
Both specimens have the typical conspicuous
peridium persisting partly as a wine-red, slightly iri-
descent, funnel-shaped calyculus comprising about
one third of the sporotheca.
Arcyria incarnata (Pers. ex J.F. Gmel.) Pers.
[ARCinc, C, 40/1] PICa: 22, PICb: 4, PIN: 15; W:
41; LOC. 14, 15, 20, 21, 22, 23, 33, 38, 39, 40, 43, 44;
LE306035…
Arcyria minuta Buchet [ARCmin, R, 1/0] PIN:
1; W: 1; LOC. 21; LE306357.
Arcyria obvelata (Oeder) Onsberg [ARCobv, O,
24/0] PICa: 9, PICb: 12, PIN: 3; W: 24; LOC. 14, 20,
21, 22, 25, 27, 30, 33, 35, 38, 41, 42, 43, 44; LE306090…
Arcyria oerstedii Rostaf. [ARCoer, R, 7/0] PICa:
4, PICb: 1, PIN: 2; W: 7; LOC. 14, 21, 23, 25, 30, 34;
LE306360…
All specimens represent typical form, display-
ing numerous spines on the threads of the capilliti-
um which easily detaches from the calyculus.
Arcyria pomiformis (Leers) Rostaf. [ARCpom,
O, 23/2] PICa: 4, PICb: 2, PIN: 19; W: 25; LOC. 14, 15,
16, 21, 22, 23, 29, 33; LE306331… Fig. 2a.
is is one of the commonest lignicolous spe-
cies in the dry spruce-pine forest of the reserve,
forming minute sporocarps on pine and spruce,
from large logs to small branches, if the surface is
still solid and smooth with dense and relatively dry
wood. It often occurs on burned wood as well. Our
collections show the typical ovate to globose pale
ochraceous sporangia and the capillitium marked
with cogs, short spines or half rings.
Arcyria stipata (Schwein.) Lister [ARCsti, R,
1/0] PICb: 1, W: 1; LOC. 33; LE306556.
Only one specimen which shows the typical
capillitium with bulb-like thickenings and numerous
free ends. e tubules are 3–5 µm diam., bearing 3–4
spirals intermixed with spines, cogs, half-rings or oc-
casional rings and reticulations. Spores are covered
by small verrucae and large warts, 7–8 µm diam.
Badhamia foliicola Lister [BADfol, R, 3/0]
PICa: 3; W: 3; LOC. 20; LE 306072…
Barbeyella minutissima Meyl. [BARmin,
R, 8/0] PICa: 2, PICb: 6; W: 8; LOC. 20, 30, 34;
LE306496…, Fig. 2b.
is species is common on the lower side of
moss-covered, moderately to strongly decayed large
logs of spruce lying on the ground in spruce-peat
moss forest (Fig. 2). Distribution and ecology of this
species were described in detail (Schnittler et al.
2000, Stephenson et al. 2019).
Calomyxa metallica (Berk.) Nieuwl. [CALmet,
R, 3/0] PICa: 2, PICb: 1; W: 3; LOC. 14, 28; LE306147…
Ceratiomyxa fruticulosa (O.F.Mull.) T.Macbr.
[CERfru, O, 14/0] PICa: 5, PICb: 5, PIN: 4; W: 14; LOC.
14, 20, 21, 23, 28, 30, 31; LE306294…
Clastoderma debaryanum A. Blytt [CLAdeb,
R, 6/0] PICa: 1, PICb: 5; W: 6; LOC. 30, 39, 41;
LE306496…
Collaria arcyrionema (Rostaf.) Nann.-Bremek.
ex Lado [COLanm, O, 9/0] PICa: 4, PICb: 5; W: 9;
LOC. 14, 20, 30, 43; LE 306094…
Colloderma oculatum (C. Lippert) G. Lister
[CODocu, A, 66/0] PICa: 28, PICb: 32, PIN: 6; W: 66;
LOC. 14, 21, 22, 23, 25, 29, 30, 35, 37, 38, 39, 40, 41, 43;
LE306499…, Fig. 2c.
All specimens were found on decorticated
wood with a slimy appearance, covered with algae
and liverworts mostly in shady habitats or at the
margins of bogs. All specimens possess small sporo-
carps (0.3–0.5 mm) with very iridescent, thin and
translucent peridia (Fig. 2c). A columella was not
found. e capillitium is almost colorless, arising
from the base, forming a large-meshed net, the tu-
bulae up to 1.5 µm in diameter, smooth. e same
minute, scattered fructications were frequently
observed in other regions of Fennoscandia, also on
large wet coniferous logs (Eliasson 1981, Schnittler
& Novozhilov 1996, Härkönen & Varis 2013).
is species occurred often with B. minutissima
and Lepidoderma tigrinum, the species that are able
to fruit on very thin slimy layers of liverworts, cov-
ered with a water lm.
Comatricha elegans (Racib.) G. Lister [COMe-
le, R, 7/0] PICa: 1, PICb: 1, PIN: 5; W: 7; LOC. 21, 34,
43; LE 306312…
Comatricha ellae Härk. [COMell, R, 4/0] PICa:
1, PIN: 3; W: 4; LOC. 21, 22, 38; LE 306353...
Comatricha laxa Rostaf. [COMlax, R, 3/0]
PICa: 1, PIN: 2; W: 3; LOC. 21, 22, 34; LE306303…
Comatricha nigra (Pers. ex J.F. Gmel.) J. Schröt.
300
[COMnig, C, 45/1] SB: 1, PICa: 10, PICb: 12, PIN: 23;
B: 1, W: 45; LOC. 1, 14, 19, 20, 21, 22, 23, 24, 29, 30, 33,
35, 37, 38, 40, 41, 42, 43; LE306327…, Fig. 2d.
is is one of the most common lignicolous
species in the reserve. It prefers large coarse woody
debris, occurring usually on dry wood in open
spruce and pine-lichen forest. is ecological niche
is as well occupied by C. elegans and C. ellae, but C.
nigra diers from both species by larger size (1.5–2
mm height), longer stalk and an undeveloped sur-
face capillitium net (Härkönen 1977a).
Cribraria argillacea (Pers. ex J.F. Gmel.) Pers.
[CRIarg, O, 13/0] PICa: 5, PICb: 5, PIN: 3; W: 13; LO C.
14, 20, 21, 22, 23, 28, 30, 35, 44; LE306398...
As typical for the genus, this species grows pref-
erably on strongly decayed wood of spruce.
Cribraria aurantiaca Schrad. [CRIaur, O,
10/1] SB: 1, PICa: 4, PICb: 3, PIN: 3; L: 1, W: 10; LOC.
1, 14, 21, 22, 33, 39, 41, 43; LE306444…
Cribraria cancellata (Batsch) Nann.-Bremek.
[CRIcan, C, 37/0] PICa: 11, PICb: 11, PIN: 15; W: 37;
LOC. 14, 21, 23, 30, 35, 36, 41, 43, 44; LE306302…
Cribraria intricata Schrad. [CRIint, R, 1/0]
PIN: 1; W: 1; LOC. 21; LE307691.
Cribraria languescens Rex [CRIlan, R, 1/0]
PICa: 1; W: 1; LOC. 14; LE306133.
Cribraria macrocarpa Schrad. [CRImac, R,
3/0] PICa: 1, PICb: 1, PIN: 1; W: 3; LOC. 14, 22, 43;
LE306236…, Fig. 2e.
Only three specimens but with the typical large
peridial cup with numerous dark ribs radiating up-
wards from the base of sporotheca, with many per-
forations in the upper part with dark nodes.
Cribraria microcarpa (Schrad.) Pers. [CRI-
mac, O, 9/6] SB: 1, PICa: 4, PICb: 10; L: 1, W: 14; LO C.
1, 14, 15, 27, 30, 37, 38; LE306168…
Cribraria oregana H.C. Gilbert [CRIore, C,
44/2] PICa: 10, PICb: 19, PIN: 17; W: 46; LOC. 14, 15,
21, 22, 23, 24, 30, 33, 35, 37, 42, 43; LE306103…
Cribraria persoonii Nann.-Bremek. [CRIper, R,
8/0] PICa: 2, PICb: 3, PIN: 3; W: 8; LOC. 14, 23, 33, 37,
43; LE306154…
Cribraria piriformis Schrad. [CRIpir, O, 9/0]
PICa: 2, PICb: 2, PIN: 5; W: 9; LOC. 14, 21, 22, 30, 33;
LE306181…
Cribraria purpurea Schrad. [CRIpur, R, 3/0]
PICa: 3; W: 3; LOC. 39; LE306609…, Fig. 2f.
Cribraria rufa (Roth) Rostaf. [CRIruf, O, 12/0]
PICa: 4, PICb: 3, PIN: 5; W: 12; LOC. 14, 21, 23, 25, 27,
33, 39, 43; LE306214…, Fig. 2g.
Our collections include bright orange-red
sporocarps up to 2 mm height. e peridial cup
is well-developed and comprises nearly half the
sporotheca, the peridial net is wide-meshed, con-
sisting of attened orange threads with small at
nodes which are often imperceptible, calcic gran-
ules are pale and up to 2 µm in diam.
Cribraria splendens (Schrad.) Pers. [CRIspl, O,
12/0] PICa: 8, PICb: 4; W: 12; LOC. 14, 33; LE306541…
Cribraria stellifera Nowotny & H. Neubert
[CRIste, R, 1/0] PICb: 1; W: 1; LOC. 29; LE306493.
is species, recorded as new for Russia, is rep-
resented by a small colony of long-stalked sporo-
carps. e peridial cup remains as a small basal disk
only, but stretches over the larger part of the sporo-
theca as a regular peridial net with thickened nodes,
which are connected with each other forming trian-
gular to quadrangular meshes, the nodes rounded
to elliptical, relatively large, densely adorned with
calcic granules, appearing star-like due to many
free-ending threads, which often repeatedly branch.
Cribraria tenella Schrad. [CRIten, R, 7/0]
PICa: 4, PICb: 1, PIN: 2; W: 7; LOC. 14, 21, 24, 25, 30;
LE306183…
Cribraria violacea Rex [CRIvio, R, 6/0] PICa: 6;
W: 6; LOC. 20; LE306078…
Cribraria vulgaris Schrad. [CRIvul, R, 6/0]
PICa: 4, PIN: 2; W: 6; LOC. 14, 21, 35; LE306156…
Dianema corticatum Lister [DNMcor, O,
12/0] PICb: 3, PIN: 9; W: 12; LOC. 21, 22, 27, 29, 33;
LE306240… Fig. 2h.
Occuring in small colonies of depressed sporo-
carps on medium-decayed coniferous wood with-
out bark. e yellowish, cartilaginous peridium is
very typical and allows to assign to this species even
remnants of fructications from the previous year.
Dianema harveyi Rex [DNMhar, R, 1/0] PICa:
1; W: 1; LOC. 14; LE307822.
Occurring in the same habitat as the previous
species, but much rarer.
Dianema repens G. Lister & Cran [DNMrep, R,
3/0] PIN: 3; W: 3; LOC. 21; LE307686.
Occurring in the same habitat as D. corticatum,
but probably rarer and/or less conspicuous due to
the dull greyish-brown spore mass in small sporo-
carps.
301
Fig. 2. Morphological traits of sporocarps of encountered myxomycete species as seen under the dissecting
microscope. a: Arcyria pomiformis (LE306331), b: Barbeyella minutissima (LE306496, DM),
c: Colloderma oculatum (LE306499), d: Comatricha nigra (LE306327), e: Cribraria macrocarpa (LE306234),
f: Cribraria purpurea (LE306614), g: Cribraria rufa (LE306214), h: Dianema corticatum (LE306337).
Scale bars: a–g = 200 μm, h = 500 μm.
302
KhiDiderma alpinum (Meyl.) Meyl. [DIDalp, R,
8/0] SB: 8; L: 8; LOC. 3, 4, 6, 7, 8; LE305754...
All specimens are recognizable by plasmodio-
carps growing densely together and sharing a con-
spicuous white hypothallus with a broad margin,
with a matt white outer peridium layer densely ad-
hering to the inner layer and the hypothallus.
Diderma montanum (Meyl.) Meyl. [DIDmon,
R, 1/0] PIN: 1; W: 1; LOC. 22; LE306423.
KhiDiderma niveum (Rostaf.) T. Macbr. [DIDniv,
C, 26/0] SB: 26; L: 26; LOC. 3, 4, 6, 8, 17; LE305733…
All specimens possess sporocarps with a dark
ferruginous columella.
Diderma radiatum (L.) Morgan [DIDrad, O,
16/0] PICa: 9, PICb: 6, PIN: 1; W: 16; LOC. 14, 21, 25,
30, 33, 38, 41, 43, 44; LE306220…
Diderma rugosum (Rex) T. Macbr. [DIDrug, R,
1/0] PICb: 1; W: 1; LOC. 30; LE307714.
Diderma umbilicatum Pers. [DIDumb, R, 2/0]
PICa: 1; PIN: 1; W: 2; LOC. 14, 21; LE306126, 306464.
Didymium diorme (Pers.) Gray [DDYdif, R,
0/2] SB: 2; D: 2; LOC. 1; LE306964…
Didymium melanospermum (Pers.) T. Macbr.
[DDYmel, C, 27/0] PICa: 5, PICb: 15, PIN: 7; L: 1, W:
26; LOC. 14, 21, 22, 27, 31, 33, 34, 35, 36, 41, 42, 43;
LE306228…
Didymium squamulosum (Alb. & Schwein.)
Fr. [DDYsqu, O, 15/0] PICa: 15; L: 15; LOC. 20;
LE306041…
Enerthenema papillatum (Pers.) Rostaf. [ENE-
pap, O, 23/0] PICa: 4, PICb: 3, PIN: 16; W: 23; LOC .
20, 21, 22, 23, 33, 43; LE306040…
is lignicolous species is often associated with
Arcyria pomiformis, Comatricha nigra and Licea
minima. It forms sporocarps on large spruce and
pine logs in dry open forest.
Fuligo leviderma H. Neubert, Nowotny & K.
Baumann [FULlev, R, 1/0] PIN: 1; W: 1; LOC. 23;
LE306810.
Fuligo muscorum Alb. & Schwein. [FUL-
mus, R, 3/0] PICa: 1, PICb: 2; L: 3; LOC. 27, 33, 36;
LE319311…, Fig. 3a, b.
is species forms large sporocarps on living
ferns, grasses and adjacent bryophytes, where bright
orange plasmodium moves up from ground litter.
Fuligo septica var. candida (Pers.) R.E. Fr.
[FULcan, R, 7/0] PICa: 2, PICb: 5; W: 4, L: 3; LOC. 14,
28; LE307766…
Fuligo septica (L.) F.H. Wigg. [FULsep, R, 3/0]
PICa: 2, PIN: 1; W: 3; LOC. 14, 23; LE319312…
Hemitrichia clavata (Pers.) Rostaf. [HEMcla,
R, 5/0] PICa: 3, PICb: 1, PIN: 1; W: 5; LOC. 14, 22, 28;
LE306144…
KhiLamproderma arcyrioides (Sommerf.)
Rostaf. [LAMarc, O, 10, 10/0] SB: 10; liv: 10; LOC. 4;
LE305736…
KhiLamproderma cacographicum Bozonnet,
Mar. Mey. & Poulain [LAMcac, R, 1/0] SB: 1; liv: 1;
LOC. 4; LE305744.
Lamproderma columbinum (Pers.) Rostaf.
[LAMcol, O, 15/0] PICa: 6, PICb: 9; W: 15; LOC. 14,
25, 30, 35, 38, 43; LE30650042…, Fig. 3c.
Khi
Lamproderma maculatum Kowalski [LAM-
mac, R, 6/0] SB: 6; L: 6; LOC. 5, 17; LE305776…Fig. 3d.
KhiLamproderma ovoideum Meyl. [LAMovo, O,
14/0] SB: 14; L: 14; LOC. 3, 4, 17; LE305758…
Lamproderma zonatum Mar. Mey. & Poulain
[LAMzon, R, 3/0] SB: 3; L: 3; LO C. 12, 17; LE305840…,
Fig. 3e.
It is the rst record of this species in Fen-
noskandia. e specimens have typical sessile, ellip-
soid sporocarps 0.5–1.5 mm wide, blue with bronze
shining zone (Fig. 3e); they are very similar to spec-
imens found in the Leningrad region (Erastova &
Novozhilov 2015). Columella reaching 1/2 to 2/3 of
the height of the sporocarp. Capillitium dense, with
ner threads at the periphery. Spores pale brown to
moderately dark (10.5–)11–13.5(–14) µm, warted or
nely spinulose with a rather dense, regularly dis-
tributed ornamentation.
Leocarpus fragilis (Dicks.) Rostaf. [LEOfra, C,
26/0] PICb: 12, PICa: 7, PIN: 7; L: 2, W: 24; LOC. 14,
21, 22, 27, 31, 33, 34, 36, 37, 43; LE 306192…
KhiLepidoderma carestianum (Rabenh.) Ros-
taf. [LEPcar, R, 4/0] SB: 4; L: 4; LOC. 3, 4; LE305766…
KhiLepidoderma chailletii Rostaf. [LEPcha, O,
16/0] SB: 16; L: 3, L: 13; LOC. 2, 3, 4, 5, 10, 11, 17, 18;
LE305731…
Lepidoderma tigrinum (Schrad.) Rostaf. [LEP-
tig, R, 3/0] PICa: 1, PICb: 1, PIN: 1; W: 3; LOC. 21, 30,
35; LE306452…
Licea chelonoides Nann.-Bremek. [LICche, R,
1/0] PIN: 1; W: 1; LOC. 21; LE319290.
Licea minima Fr. [LICmin, O, 14/4] SB: 1, PICa:
7, PICb: 2, PIN: 8; L: 1, W: 17; LOC. 4, 14, 15, 16, 21,
30, 35; LE307656…
303
Licea nannengae Pando & Lado [LICnan, R,
0/1] SB: 1; L: 1; LOC. 1; LE306974.
Licea pusilla Schrad. [LICpus, O, 19/2] SB: 1,
PICa: 5, PICb: 8, PIN: 7; L: 2, W: 19; LOC. 1, 15, 21, 23,
29, 37, 43; LE306307…
Licea pygmaea (Meyl.) Ing [LICpyg, C, 41/6]
PICa: 24, PICb: 16, PIN: 7; W: 47; LOC. 14, 15, 16, 20,
21, 22, 23, 24, 25, 27, 30, 37, 38, 41, 42, 43; LE306074…
Licea variabilis Schrad. [LICvar, A, 61/0] PICa:
26, PICb: 13, PIN: 22; W: 61; LOC. 14, 20, 21, 22, 23, 25,
29, 30, 33, 35, 37, 39, 41, 42, 43; LE306085…
Lindbladia tubulina Fr. [R, 1, 1/0] PICb: 1; W:
1; LOC. 30; LE307554.
Lycogala epidendrum (L.) Fr. [LYCepi, A, 58/0]
PICa: 12, PICb: 20, PIN: 26; W: 58; LOC. 14, 20, 21, 22,
23, 25, 27, 28, 30, 33, 34, 35, 41, 43; LE306031…
KhiMeriderma aggregatum ad int. (Meyl.) Mar.
Mey. & Poulain [MERagg, R, 1/0] SB: 1; L: 1; LOC. 3;
LE305771.
Metatrichia oriformis (Schwein.) Nann.-
Bremek. [METo, R, 1/0] PICa: 1; W: 1; LOC. 35;
LE307632.
Metatrichia vesparia (Batsch) Nann.-Bremek.
ex G. W. Martin & Alexop [METves, R, 1/0] PICa: 1;
W: 1; LOC. 14; LE306173.
Paradiacheopsis mbriata (G. Lister & Cran)
Hertel ex Nann.-Bremek. [PARm, O, 0/12] SB: 7,
PICa: 5; B: 9, W: 3; LOC. 1, 16, 18; LE306961…
Paradiacheopsis solitaria (Nann.-Bremek.)
Nann.-Bremek. [PARsol, R, 0/1] SB: 1; B: 1; LOC. 18;
LE319313.
Perichaena chrysosperma (Curr.) Lister [PER-
chr, R, 0/3] SB: 3; B: 3; LOC. 1, 4; LE307248…
Perichaena corticalis (Batsch) Rostaf. [PER-
cor, R, 0/1] SB: 1; B: 1; LOC. 4; LE306989.
Perichaena vermicularis (Schwein.) Rostaf.
[PERver, R, 2/1] SB: 1, PICa: 2; B: 1, W: 2; LOC. 1, 14;
LE307767...
KhiPhysarum albescens Ellis ex T. Macbr. [PHY-
alb, C, 26/0] SB: 26; L: 26; LOC. 5, 6, 7, 8, 9, 13, 17;
LE305803…
is was the most abundant nivicolous myxo-
mycete in subalpine birch forests of the reserve. All
collections show a morphology identical with that
of specimens found recently in the Khibiny Mts.
(Erastova et al. 2017) and are already recognizable
in the eld by their large, subglobose to obovoid
sporocarps on weak stalks formed by strands of
the hypothallus and the variegated color of lime at
the peridium surface ranging from yellow to pale
orange (limeless specimens appear dark blue and
iridescent).
Physarum album (Bull.) Chevall. [PHYalu, A,
177/0] PICa: 64, PICb: 93, PIN: 20; W: 177; LOC. 14,
19, 20, 21, 22, 23, 24, 25, 27, 28, 29, 30, 31, 33, 34, 35,
37, 38, 39, 40, 41, 42, 43; LE306060…
Physarum cinereum (Batsch) Pers. [PHYcin, R,
2/0] PICa: 2; L: 1, W: 1; LOC. 14, 20; LE306048…
Physarum compressum Alb. & Schwein. [PHY-
com, O, 0/15] SB: 9, PICa: 6; B: 5, D: 10; LOC. 1, 4, 16;
LE306955…, Fig. 3f.
is species is one of the most common species
in the reserve, appearing in moist chamber cultures
on droppings of willow ptarmigan and moose. is
species was frequently registered in dierent re-
gions on dung of birds which feed on seeds, buds or
other plant parts (Eliasson & Lundqvist 1979, Elias-
son 2013).
Physarum confertum T. Macbr. [PHYcon, R,
1/0] PICb: 1; L: 1; LOC. 28; LE306512.
Physarum globuliferum (Bull.) Pers. [PHYglo,
R, 1/0] PICa: 1; W: 1; LOC. 39; LE306611.
Physarum gyrosum Rostaf. [PHYgyr, R, 1/0]
PICa: 1; W: 1; LOC. 14; LE307844.
Physarum lateritium (Berk. & Ravenel) Mor-
gan [PHYlat, R, 1/0] PICb: 1; L: 1; LOC. 31. LE307701.
Physarum leucopus Link [PHYlcp, R, 1/0]
PICb: 1; W: 1; LOC. 43; LE306693.
Physarum leucophaeum Fr. [PHYlph, C, 27/1]
SB: 1, PICa: 8, PICb: 10, PIN: 9; L: 1, W: 27; LOC. 4, 14,
20, 21, 22, 23, 31, 34, 41; LE306202…
Physarum notabile T. Macbr. [PHYnot, R, 2/0]
PICa: 2; W: 2; LOC. 14, 34; LE306263…
Physarum pusillum (Berk. & M. A. Curtis) G.
Lister [PHYpus, R, 0/1] SB: 1; B: 1; LOC. 4; LE306982.
Physarum rubiginosum Fr. [PHYrub, R, 1/0]
PICa: 1; W: 1; LOC. 14; LE306143.
Physarum viride (Bull.) Pers. [PHYvde, A, 73/0]
SB: 2, PICa: 29, PICb: 33, PIN: 9; L: 1, W: 72; LOC. 14,
19, 20, 21, 22, 27, 28, 30, 33, 34, 35, 37, 38, 39, 40, 41,
42, 43, 44; LE306038…
Physarum virescens Ditmar [PHYvir, R, 1, 1/0]
PIN: 1; W: 1; LOC. 21; LE307672.
Siphoptychium reticulatum Leontyev, Schnit-
tler et S.L. Stephenson [SIPret, R, 1, 1/0] PICb: 1; W:
1; LOC. 28; LE306515, Fig. 3g.
304
We found only one pseudoaethalium on strong-
ly decomposed wood of spruce, which is very similar
with the type description of S. reticulatum (Leontyev et
al. 2019). e key characters of the species are slightly
convex, polygonal sporothecal tips (0.3–)0.5–0.7(–1.0)
mm diam., with bright lines between sporothecal tips,
giving the pseudoaethalial surface a reticulate ap-
pearance. is taxon was so far most likely recorded
in Russia as Tubifera dictyoderma (Novozhilov et al.,
2017); detailed investigations have to be carried out to
ascertain if both species occur in Russia.
Stemonaria laxiretis Nann.-Bremek. & Y.
Yamam. [STAlax, R, 3/0] PICa: 3; W: 3; LOC. 20;
LE306034…
Our specimens t well the type description of
this species (Nannenga-Bremekamp & Yamamoto
1990) and are characterized by lax, rather slender
capillitial threads with some dark membranous ex-
pansions, constituting a lax internal net of one or two
meshes across the radius with many long free ends.
Stemonitis axifera (Bull.) T. Macbr. [STEaxi, C,
46/0] PICa: 16, PICb: 24, PIN: 6; W: 46; LOC. 14, 21,
23, 27, 30, 33, 34, 37, 38, 40, 41, 42, 43; LE306169…
Stemonitis capillitionodosaG. Moreno, D.W.
Mitch., C. Rojas & S.L. Stephenson [STEcap, R, 2/0]
PICb: 2; W: 2; LOC. 33; LE306548, 306833.
We found this rare species twice; both speci-
mens possess a typical columella with expansions at
the apex and large, densely spinulose spores (9–11
µm diam.). If observed under SEM, these spinules
appear as coral-like warts (Vlasenko et al. 2020).
Previously one sequence of this species (LE306548)
was submitted to GenBank under No. MH930789 but
with the incorrect determination Stemonitis laxila
(Shchepin et al. 2019). A phylogenetic analysis of the
18S rDNA sequences (SSU) revealed 100% similari-
ty with the studied specimen of S. capillitionodosa
(LE306833, GenBank No. MN610391). e SEM study
of these specimens conrmed that both specimens
belong to S. capillitionodosa, characterized by the
unique ornamentation of its spores in the form of
coral-shaped warts (Vlasenko et al. 2020).
Stemonitis fusca Roth [STEfus, R, 6/2] SB: 2,
PICa: 3, PICb: 2, PIN: 1; B: 2, W: 6; LOC. 1, 14, 21, 31,
33; LE306270…
Stemonitis lignicola Nann.-Bremek. [STElig, O,
11/0] PICb: 6, PIN: 5; W: 11; LOC. 21, 23, 28, 30, 31;
LE306501…
Stemonitis pallida Wingate [STEpal, R, 2/0]
PICa: 1, PIN: 1; W: 2; LOC. 21, 39; LE306297…
Stemonitis smithii T. Macbr. [STEsmi, R, 6/0]
PICa: 1, PICb: 3, PIN: 2; W: 6; LOC. 20, 21, 22, 30, 43;
LE306069…
Stemonitis splendens Rostaf. [STEspl, R, 2/0]
PIN: 2; W: 2; LOC. 21; LE306296…
Stemonitopsis aequalis (Peck) Y. Yamam.
[STEaeq, O, 17/0] PICa: 2, PICb: 9, PIN: 6; W: 17;
LOC. 21, 28, 29, 30, 33, 35, 37, 43; LE306319…
Stemonitopsis amoena (Nann.-Bremek.)
Nann.-Bremek. [STEamo, R, 4/0] PIN: 4; W: 4; L OC.
21, 22; LE306317…
Stemonitopsis gracilis (G. Lister) Nann.-
Bremek. [STEgra, R, 5/0] PICa: 1, PICb: 1, PIN: 3; W:
5; LOC. 21, 22, 27, 34; LE306236…
Stemonitopsis hyperopta (Meyl.) Nann.-
Bremek. [STEhyp, A, 53/0] PICa: 11, PICb: 23, PIN:
19; W: 53; LOC. 14, 21, 22, 23, 27, 28, 29, 30, 33, 34, 37,
39, 41, 42, 43; LE306182…
Stemonitopsis typhina (F.H. Wigg.) Nann.-
Bremek. [STEtyp, C, 26/0] PICa: 13, PICb: 10, PIN: 3;
W: 26; LOC. 14, 21, 22, 30, 31, 36, 41; LE306123…
Symphytocarpus accidus (Lister) Ing &
Nann.-Bremek. [SYMa, R, 1/0] PIN: 1; W: 1; L OC.
23; LE307636.
Trichia botrytis (J.F. Gmel.) Pers. [TRIbot, C,
42/0] PICa: 15, PICb: 23, PIN: 4; W: 42; LOC. 14, 20,
21, 28, 30, 33, 34, 37, 39, 41, 42, 43, 44; LE306071…
Trichia contorta (Ditmar) Rostaf. [TRIcon, R,
3/0] PICa: 1, PICb: 2; W: 3; LOC. 14, 31; LE306582…
Trichia decipiens (Pers.) T. Macbr. [TRIdec, C,
34/0] PICa: 21, PICb: 13, ; W: 34; LOC. 14, 20, 30, 33,
34, 35, 41, 42, 43; LE306033…
Trichia erecta Rex [TRIere, R, 4/0] PICa: 3,
PICb: 1, ; W: 4; LOC. 14, 31; LE306149…
Trichia favoginea (Batsch) Pers. [TRIfav, R,
5/0] PICa: 5; W: 5; LOC. 14, 25, 38, 44; LE306171…
Trichia lutescens (Lister) Lister [TRIlut, R, 1/0]
PIN: 1; W: 1; LOC. 22; LE306404.
Trichia persimilis P. Karst. [TRIper, R, 7/0]
PICa: 3, PICb: 1, PIN: 3; W: 7; LOC. 14, 20, 22, 23, 43;
LE306032…
Trichia scabra Rostaf. [TRIsca, R, 7/0] PICa: 1,
PICb: 4, PIN: 2; W: 7; LOC. 14, 22, 31; LE306258…
Trichia sordida Johannesen [TRIsor, R, 1/0]
SB: 1, L: 1; LOC. 3; LE305783, Fig. 3h.
In spite of intensive previous studies in the
305
Khibiny Mts. (Erastova et al. 2017), this nivicolous
species was not registered and appears to be a rare
species in the study area. Our specimen is a small
colony of sessile dull ochraceous sporangia with
dark lines and spots. e capillitium represents
smooth elaters with long gradually tapering tips.
Large, densely warted spores t well to the typical
description of the species (Johannesen 1984b).
Trichia subfusca Rex [TRIsub, R, 1/0] PICb: 1;
W: 1; LOC. 33; LE307623.
Trichia varia (Pers. ex J.F.Gmel.) Pers. [TRIvar,
C, 44/0] PICa: 19, PICb: 16, PIN: 9; W: 43, L: 1; LOC.
14, 20, 22, 23, 24, 26, 27, 28, 31, 33, 35, 37, 39, 41, 42,
44; LE306062…
Tubifera applanata (Leontyev & Fefelov) Le-
ontyev & Fefelov [TUBapp, R, 1/0] PICa: 1; W: 1;
LOC. 35; LE306573.
Our collection is represented by one pseudoae-
thalium with the typical sporothecal tips which are
isodiametric, roughly hexagonal, similar to each
other in size and occur in regular rows (Leontyev
et al. 2015).
Tubifera ferruginosa (Batsch) J.F. Gmel. [O, 13,
13/0] PICa: 4, PICb: 3, PIN: 6; W: 13; LOC. 14, 21, 23,
25, 30, 43, 44; LE306186…
Species diversity
e species list given above was compiled from
1675 records of myxomycetes. Of these, 1583 were
recorded in the eld, whereas the other 92 records
were obtained from 210 moist chamber cultures. In
total, 124 morphospecies and one variety belong-
ing to 34 genera were identied, all recorded as
new for the Laplandskiy Reserve. Most taxa were
collected in the eld (117); only 21 species were
recorded in moist chamber (with 13 taxa retrieved
by both methods). is is reected by the propor-
tions of species found exclusively in the eld (103
morphospecies and one variety of 32 genera, 83.2%)
or in moist chambers (8 species of 5 genera, 5.6%);
the remaining species (13 species of 7 genera, 8.8%)
were recorded with both methods. Non-fruiting
plasmodia and sclerotia were not included in these
numbers (114 records).
erefore, records from the eld and from the
moist chamber cultures largely complemented each
other (Fig. 4). Sixty two percent of all taxa (77) were
classied as rare for the whole study area (<0.5 %
of all records); most were represented by only one
(33) or two (7) records. Abundant species (>3 % of
Fig. 4. Frequency distribution of all 1675 determinable records of myxomycetes from 125 taxa. The black sections of
the bars indicate records collected in the field, whereas the grey sections indicate records from 210 moist chamber
cultures. Three thin vertical dotted lines show the delimitation of the ACOR classes.
306
Fig. 3. Morphological traits of sporocarps of encountered myxomycete species. a, b: Fuligo muscorum (LE319311),
a: plasmodium on living grasses, b: immature sporocarps on living grasses, C: Lamproderma columbinum (LE306500),
sporocarp as seen under dissecting microscope (DM), d: Lamproderma maculatum (LE305776, DM),
e: Lamproderma zonatum (LE305840), f: Physarum compressum (LE306955, DM), g: Siphoptychium reticulatum
(LE306515, DM), h
: Trichia sordida (LE305783, DM). Scale bars: a, b = 3 cm, c = 200 μm, d, e, f, h = 500 μm, g = 1 cm.
307
all records) included Colloderma oculatum, Licea
variabilis, Lycogala epidendrum, Physarum album,
Ph. viride, and Stemonitopsis hyperopta. e follow-
ing species were classied as common (1.5–3 % of
all records): Arcyria cinerea, A. incarnata, Comatri-
cha nigra, Cribraria cancellata, C. oregana, Diderma
niveum, Didymium melanospermum, Leocarpus fra-
gilis, Licea pygmaea, Physarum albescens, Ph. leuco-
phaeum, Stemonitis axifera, Stemonitopsis typhoides,
Trichia botrytis, T. decipiens, and T. vari a. A few spe-
cies, including Arcyria cinerea, Cribraria microcar-
pa, Licea pygmaea, Paradiacheopsis mbriata, and
Physarum compressum, were observed most often in
moist chamber cultures (Fig. 4).
Eciency and completeness of the survey
For the study as a whole, values of Chao1 estimator
(Fig. 5a, Table 1) suggest that in spite of 28 days of eld
work in the reserve spread over June, July and Au-
gust of dierent years, our eort was not sucient
to recover all species (125 taxa from 1675 records,
completeness 66%, all gures given according to the
nal gure of the Chao1 estimator). However, a part
of this relative incompleteness is due to the dierent
species assemblages recovered by the two dierent
survey methods used ( eld collecting and moist
chambers). Figures for completeness increase if we
look at these two methods separately ( eld collect-
ing: 117 taxa from 1584 records, 70%; moist cham-
bers: 21 taxa from 92 records, 78%).
If we look at the species assemblages recovered
from the eld (ca. 95% of all records), all four forest
types seem to be exhaustively sampled (fc: PICa: 82
taxa from 546 records, completeness according to
the Chao1 estimator 84%; PICb: 70 taxa from 540
records, 84%; PIN: 62 taxa from 381 records, 83%;
SB: 13 taxa from 116 records, 87%, Fig. 5a, Table 1). e
raried data (at level of 116 records) show a similar
trend: PICa 46 taxa, 47%, PICb 41 taxa, 67%, PIN 43
taxa, 73% and SB 13 taxa. However, since we have
no data obtained from moist chamber cultures for
PICb and PIN and we were not able to visit PIN and
SB throughout the eld season, the species richness
and diversity were likely to be underestimated in
these vegetation types. We will address these sourc-
es of error within future investigations.
Seen in total, the main substrate types were rath-
er exhaustively sampled (Fig. 5b), but the complete-
ness varies greatly for dierent types of substrates,
depending on the collection method. Most exhaus-
tively sampled was coarse woody debris (w): 102 taxa,
1466 records, 88% completeness, Table 2. For this sub-
strate type, the moist chamber culture method did
not reveal additional information: all of the 8 species
(25 records) recorded were as well observed in the
eld. In contrast, this method is indispensable for
corticolous (10 taxa, 37 records) and coprophilous
(3 taxa, 21 records) myxomycetes, where all taxa were
exclusively observed in moist chamber.
Myxomycete diversity within vegetation types
of the reserve
At least when judged by fructication propensity,
myxomycetes are to some degree specialized on
vegetation ( forest association) and substrate type.
Despite the high number of 125 taxa recorded, only
16 were found to be widely distributed (present in
10 or more of the 44 studied localities, Database S1).
For eld records, both species richness and diversi-
ty increased signicantly from subalpine birch for-
est (SB: 13 species, H´= 2.1, D = 6.9) over open dry
spruce-pine forest (PIN: 62, H´= 3.7, D = 30.0) to the
two associations of spruce forest (PICb: 70 species,
H´= 3.5, D = 19.1; PICa: 82 species, H´= 3.8, D = 28;
Fig. 5a, Table 1).
Most of the abundant myxomycete species oc-
curred in two or three forest types of the reserve, al-
though preferences were visible in some cases. Only
in the subalpine birch forest (SB) we could reveal
nivicolous myxomycetes. Common species found in
SB and conned exclusively to litter and stems of liv-
ing dwarf shrubs were Diderma niveum (15% of 170
records of all found species), Physarum albescens
(14%), Lepidoderma chailletii (9%), and Lamproder-
ma ovoideum (8%). All these nivicolous species are
widely distributed in subalpine communities and
mountain alpine tundra (Erastova et al. 2017) and
appear to be abundant in mountain regions (Roniki-
er & Ronikier 2009).
None of the very common myxomycete spe-
cies (abundance according to the ACOR scale >3 %
of all records) was recorded exclusively in only one
forest type. e most common lignicolous species
in the reserve is Physarum album (11% of 1675 re-
308
Fig. 5. Individual-based species accumulation curves (thick lines) and the Chao 1 estimator (thin jagged lines) of
expected morphospecies richness; a: for the whole studied area and for the four forest types (only field specimens
considered; PICa – spruce forest, PICb – spruce-peat moss forest, PIN – dry spruce-pine forest, SB – subalpine
birch forest); b: for the three studied substrate types (b – bark, w – wood, l – ground litter).
Table 1. Statistical data for myxomycetes of the Laplandskiy Reserve obtained using field collections (fc) and moist
chamber cultures (mc), grouped according to the four major vegetation associations: subalpine birch forest (SB),
spruce forest (PICa), spruce-peat moss forest (PICb), and dry spruce-pine forest (PIN).
1 Only cultures that yielded sporocarps were counted as positive. BET - Betula czerepanovii, PIC – Picea obovata,
PIN – Pinus sylvestris. 2 The Chao1 estimator for forest types were calculated for the field collections only, whereas
for the whole area both sets of data (field collections and moist chamber cultures) were used. ND = no data.
PIN PICa PICb SB Whole area
Elevation range (m a.s.l.) 35–190 110–300 120–370 340–520 35–520
Main canopy tree species PIN PIC PIC BET PIC
Sampled localities 4 13 12 15 44
Records (fc/mc) 381/ND 546/38 540/ND 116/54 1583/92
Species (fc/mc) 62(62/ND) 84(82/10) 70/ND 30(13/17) 125(117/21)
Chao1 ± SD275 ± 9 98 ± 9 84 ± 9 15 ± 3 191 ± 33
Genera (fc/mc) 27/ND 34(30/5) 7/ND 14(14/8) 34(33/9)
Species/Genera 2.7 2.8 2.6 2.1 3.7
Shannon H´ (fc) 3.7 3.9 (3.8) 3.5 2.8 (2.2) 4.1
Simpson’s D (fc) 30 30 (28) 19 12 (6.7) 35
Number of mc ND 90 ND 120 210
% mc positive 1 ND 48 ND 60 34
309
cords). e most specialized lignicolous species in
pure spruce forests are Lamproderma columbinum
(spruce wood, PICa: 6, PICb: 9 records), Colloderma
oculatum (spruce wood, PICa: 28, PICb: 32 records)
and Barbeyella minutissima (PICa: 2, PICb: 6 re-
cords). e most common species of the dry spruce-
pine forest (PIN) are Lycogala epidendrum (45% of
58 records of this species), Comatricha nigra (50% of
46), Licea variabilis (36% of 61), Arcyria pomiformis
(76% of 25), Enerthenema papillatum (70% of 23),
Cribraria oregana (37% of 46), and Stemonitopsis hy-
peropta (36% of 53).
e overall diversity and the degree of special-
ization in species show opposite trends. A com-
parison based on adjusted incidence-based Chao-
Sørensen similarity (Ccs) revealed the myxomycete
assemblages of the two forest associations poorest
in species to be the most distinctive (SB: 30 spe-
cies, mean Ccs = 0.12±0.03; PIN 62 species, mean Ccs
= 0.59±0.02; Table 3). e main reason is likely the
general rarity of acidophilic and bryophilic myxo-
mycete species in these two communities. Many
species require for development the thick mats of
mosses often covering large logs of spruce and do
not fruit on the rather dry wood prevailing in SB and
PIN. In addition, nivicolous myxomycetes are a dis-
tinct component of the subalpine forests (SB).
A view from the opposite side presents the anal-
ysis of the indicator values of species (Fig. 6, Table S1).
Here, the two rather moist forest associations (PICa
and PICb) have the lowest number of indicator spe-
cies, the dry spruce-pine forest (PIN) the highest.
Probably, most species can fruit in the rather dense
and moist spruce forest, whereas only specialized
species can cope with the overall lower moisture
supply in the two other habitats (PIN and SB) where
the ground layer retains less water due to the rath-
er open canopy and the lower moss biomass. In the
Fig. 6. Schematic view of the forest associations and their typical myxomycete species revealed by an indicator
value analysis, sorted according to significance of indicator values (p value <0.05 *, <0.01**, <0.001***). Three types
of indicator values were calculated and stacked in the graphical visualization: the probability that the surveyed site
belongs to the target site group given the fact that the species has been found (component A, green bars), the
probability of finding the species in sites belonging to the site group (component B, ochraceous bars), and IndVal.g
influenced by both components (black bars). Abbreviations for myxomycete species and forest associations are
indicated in the annotated species list and in Material and Methods, correspondingly.
310
species, 1465 records), only 8 species of 25 records
were obtained in moist chamber cultures. Of the 102
species inhabiting wood, 86 were found exclusively
on this substrate. However, in terms of ACOR class-
es, only ten species, Physarum album (177 records),
Ph. viride (72), Colloderma oculatum (66), Licea var-
iabilis (61), Lycogala epidendrum (58), Stemonitopsis
hyperopta (53 records), Licea pygmaea (47), Cribrar-
ia oregana (46), Stemonitis axifera (46), and Comatri-
cha nigra (45) were abundant (>3 % of all records) in
the reserve, whereas 83 lignicolous species were rare
(< 0.5 % of 1465 records).
Litter was the second most diverse substrate
type, but only 17 of the 29 species occurred exclu-
sively on this substrate type. irteen species found
in litter were rare. Common species found only on
dierent types of ground litter including mosses
and dwarf shrubs were Diderma niveum (26 re-
cords), Physarum albescens (24 records), Lepidoder-
ma chailletii (16 records, all three nivicolous), and
Didymium squamulosum (15 records). Fuligo musco-
rum is the only myxomycete species that appears to
be restricted to wet mossy ground litter in the re-
serve. We collected in the eld only three specimens
with mature fruit bodies (Figs. 3a, b), but other four
records of plasmodia and immature sporocarps
analysis, these specialized species appear with rath-
er high indicator values (PIN: lignicolous species
growing on rather dry wood, SB: nivicolous species
developing in soil under long-lasting snowpatches).
Substrate-species relationships
e overall degree of specialization of myxomycet-
es is higher for substrate type than for forest asso-
ciations. With respect to the major substrate types
studied, diversity and species richness increased
from dung (3 species, H´= 0.9, D = 2.4, Sp/G = 1.0)
and bark (10 species, H´= 1.8, D = 4.7, Sp/G = 1.7)
over litter (29 species, H´= 2.7, D = 10.5, Sp/G =
3.2) to wood (103 species, H´= 3.9, D = 28.7, Sp/G =
3.2). is pattern basically remains the same when
comparing the individual-based species accumu-
lation curves (rareed values for species richness
at the level of 21 records: dung 3, bark 8, litter 14,
and wood 16, see Fig. 5b). e similarity between the
myxomycete assemblages associated with dierent
substrates in the reserve was very low (mean Ccs =
0.15 ± 0.09, Table 3).
Wood-inhabiting (lignicolous) myxomycetes
represented the largest myxomycete assemblage in
the reserve and were found mostly in the eld (102
Table 2. Statistical data for myxomycetes from the dierent substrate types. Abbreviations as in Table 1.
Bark Litter Wood Dung Total
Records (fc/mc) 0/37 143/9 1441/25 0/21 1584/92
Species (fc/mc) 0/10 22/7 101/8 0/3 117/21
Chao1 ± SD 15 ± 6 52 ± 16 141 ± 21 ND 191 ± 33
Genera 6 9 32 3 34
Species/Genera 1.7 2 3.2 1 3.7
Shannon H´ 1.8 2.7 3.9 ND 4.1
Simpson’s D 4.7 10.5 28.7 ND 35.0
Number of mc 90 50 30 40 210
% mc positive 1 32 12 53 50 34
Species per mc (mean±SD) 0.4 ± 0.6 0.2 ± 0.6 0.8 ± 1.0 0.5 ± 0.5 0.4 ± 0.7
1 Only cultures that yielded sporocarps were counted as positive.
311
were damaged by animals (apparently lemmings).
Bark was the least diverse substrate type, with
only 4 species recorded exclusively on this substrate.
e most abundant corticolous species were Arcyria
cinerea (36% of 36 records), Perichaena chrysosper-
ma (56% of 43 records), Paradiacheopsis mbriata
(25% of 12 records), and Physarum compressum
(14% of 15 records). While 51% of all bark moist
chamber cultures were positive (displayed evidence
of either plasmodia or sporocarps), only 32% yielded
sporocarps (Table 2).
In the reserve, three species of myxomycetes
have been recorded from weathered dung: Arcyria
cinerea, Didymium diorme, and Physarum compres-
sum. However, only the last species seems to be spe-
cialized on this substrate to some degree.
ese relationships are as well reected by the
indicator species analysis (Fig 7): all substrates have
several exclusive species with high indicator val-
ues. Wood shows the highest number of indicator
species. In contrast, only a single species, Arcyria
cinerea, has an indicator value for both wood and
dung (Table S2, not shown in Fig. 7).
A dissimilarity comparison of the four studied
forest types shows that the myxomycete assemblag-
es of the Laplandskiy Reserve fall into three groups:
one cluster formed by the two associations of mossy
spruce forests (PICa and PICb) and two further clus-
ters for each dry spruce-pine (PIN) and subalpine
birch forest (SB, Fig. 8).
Both myxomycete assemblages from mossy
spruce forests show a high degree of overlap. Dif-
ferences among assemblages in PICa and PICb are
more the result of dierences in the abundance of
shared species than actual dierences in species
composition. Exceptions are Colloderma oculatum
and Barbeyella minutissima. Both species seem to
require constantly high air moisture, are limited to
the spruce-peat moss forest (PICb) and are often
associated with Physarum viride, Lepidoderma tigri-
num, and Lamproderma columbinum (Schnittler et al.
2000, Stephenson et al. 2019). ese species form a
distinct ecological guild on large wet coarse conifer-
ous woody debris covered by mosses and liverworts
in boreal forests (Schnittler & Novozhilov 1998).
Among nivicolous species, only Trichia sordida
was found the rst time for the Kola peninsula, other
species of this guild were found in previous studies
in the adjacent Khibiny Mts. (Erastova et al. 2017).
Summarizing, the Laplandskiy Reserve har-
bours a myxomycete biota with relatively high spe-
cies richness and diversity (125 taxa, H´= 4.1, D =
35.1), reecting the rather high diversity of vegeta-
tion in the reserve. Cribraria is the most diverse ge-
nus, all 18 species were found on decayed coniferous
wood. Virtually all species known from temperate
and boreal zones are fairly common in the reserve,
like Physarum album, Ph. viride or Lycogala epiden-
drum. e proportions of species among myxomy-
Fig. 7. Indicator species for the four main substrate
types: decayed wood, forest floor litter, dung, and bark
of living trees. For explanation of bars, significance values
and abbreviations of myxomycete names see Fig. 6.
312
Fig. 8. Dendrogram of dissimilarity, based on the
Chao-Søerensen similarity index (Table 3), of the
relative occurrence of the species found in the forest
types of the reserve (abbreviations of the forest types as
explained in Material and Methods).
Table 3. Similarity of myxomycete assemblages between forest types (SB, PICa, PICb, and PIN) and substrate
types (b, d, w, l – see text for explanations). The table shows the adjusted incidence-based Chao-Sørensen similarity
index (Ccs, upper right) and the number of species shared (lower left) for 125 myxomycete species. The last two rows
indicate the mean figures of Ccs for a study site (Mean) and its standard deviation (SD).
Main forest types Substrates
SB PICa PICb PIN B D W L
SB - 0.18 0.12 0.06 b - 0.63 0.09 0.03
PICa 12 - 0.92 0.83 d 2 - 0.02 0.03
PICb 9 58 - 0.90 w 5 1 - 0.14
PIN 6 49 46 - l 1 1 12 -
Mean 0.12 0.65 0.64 0.60 0.25 0.23 0.08 0.06
SD 0.03 0.23 0.26 0.27 0.33 0.35 0.06 0.07
cete families revealed in the reserve are typical for
myxomycete biota of the taiga forest (Novozhilov
1980, 1999, Schnittler & Novozhilov 1996). If seen
according to the traditional system, the four most
common families recovered in the eld are Cribrar-
iaceae (19 species, 15%, of all 125 taxa), Physarace-
ae (20 species, 16%), Trichiaceae (28 species, 22%,),
and Stemonitidaceae (29 species, 23%), making up
for 76% of all taxa. ese proportions dier between
studied vegetation types. Both associations of pure
spruce forest were relatively rich in Stemonitidace-
ae (PICa: 17 species, 20% of all 84 taxa; PICb:, 15
species, 21% of all 70 taxa), Trichiaceae (PICa: 20
species, 24%; PICb: 16 species, 23%), Cribrariaceae
(PICa: 15 species, 18%; PICb: 14 species, 20%), and
Physaraceae (PICa: 13 species, 15%; PICb: 8 spe-
cies, 11%), whereas in the subalpine birch forest
the proportion of members from the Cribrariaceae
decreased (3 species, 10% of total 30 taxa). e taxo-
nomic structure of myxomycete biota in the reserve
corroborates results from other studies in boreal
forests of Fennoscandia (Härkönen & Varis 2013,
Johannesen & Vetlesen 2020). e main genera in
the reserve are Cribraria (18 species), Physarum
(15), Trichia (11), and Arcyria (10).
Interestingly, some genera common in the tem-
perate zone, especially in deciduous forests, like
Arcyodes, Brefeldia, Craterium, Diachea, Dictydiae-
thalium, Echinostelium, Mucilago, Prototrichia, and
Reticularia, have not been revealed in the reserve.
Further studies using the analysis of environmental
DNA would show if they are present there as amoe-
bal populations that rarely or never fruit or they are
completely absent in the northern boreal forests of
the reserve.
313
Acknowledgements
The authors are grateful to the administration of the Lapland-
skiy
State Nature Biosphere Reserve, Natalia Berlina and
Valeriy Barkan, for their assistance in the organization of the
field work. We gratefully acknowledge the technical support
(SEM) provided by Ludmila A. Kartzeva (The Core Facility
Center “Cell and Molecular Technologies in Plant Science”
at the Komarov Botanical Institute RAS, St. Petersburg,
Russia) and help from Elizaveta Shchepina and Daria Erastova
during the field work. This work was supported by Russian
Foundation for Basic Research (project 18-04-01232 A),
the state task of BIN RAS ‘Biodiversity, ecology, structural
and functional features of fungi and fungus-like protists’
(AAAA-A19-119020890079-6) for O. Shchepin and
Y. Novozhilov. The work with herbarium collections was
supported by the grant of Moscow State University for leading
scientific schools “Depository of the Living Systems” and the
state task (AAAA-A16-116021660084-1) for V. Gmoshinskiy.
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