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ISSN 1995-4255, Contemporary Problems of Ecology, 2023, Vol. 16, No. 4, pp. 403–425. © Pleiades Publishing, Ltd., 2023.
Russian Text © The Author(s), 2023, published in Sibirskii Ekologicheskii Zhurnal, 2023, No. 4, pp. 402–427.
Fungal Diversity of Native and Alien Woody Leguminous Plants
in the Middle Urals
A. G. Shiryaeva, *, I. V. Zmitrovicha, b, P. Zhaoc, S. A. Senatora, d, and T. S. Bulgakova, e
a Institute of Plant and Animal Ecology, Ural Branch, Russian Academy of Sciences, Ekaterinburg, 620144 Russia
b Komarov Botanical Institute, Russian Academy of Sciences, St. Petersburg, Russia
c Institute of Microbiology, Chinese Academy of Sciences, Beijing, China
d Tsitsin Main Botanical Garden, Russian Academy of Sciences, Moscow, Russia
e Subtropical Research Centre, Russian Academy of Sciences, Sochi, Russia
*e-mail: anton.g.shiryaev@gmail.com
Received February 17, 2023; revised February 28, 2023; accepted March 7, 2023
Abstract—The biodiversity of wood-inhabiting fungi on woody leguminous plants (WLPs) growing in the
Middle Urals (Russia) has been studied for the first time. From 2002 to 2022, in Sverdlovsk oblast as a model
region, 136 species of wood-inhabiting fungi were identified on WLPs: 127 species of Basidiomycota and
9 species of Ascomycota. Fungi develop on 12 out of 20 species of WLPs. The largest number of fungal species
was found on the alien Caragana arborescens (115 species/84.5% of the total number of species), while two
species were collected on Caragana decorticans, C. ussuriensis, and Laburnum alpinum each and one species
was collected on Genista florida. A total of 122 species of fungi were found on nine alien WLPs, which is
4.1 times more than on three native species. The largest number of substrate-specific fungal species can be
found to develop on C. arborescens (85/62.5%), four species on Chamaecytisus ruthenicus (2.9%), three spe-
cies on Maackia amurensis (2.2%), two on Genista tinctoria and Robinia pseudoacacia each (1.5%), and one
species on Caragana ussuriensis (0.7%). Nectria cinnabarina develops on the maximum number of substrates,
seven WLP species; Xylodon sambuci on six species; and Peniophora cinerea and Schizophyllum commune on
four species. In contrast, 71.3% of fungal species were found on one WLP species, and 27.2% of species are
characterized by a single finding. For the first time for Sverdlovsk oblast, 14 fungal species are indicated, of
which 86% were found in the parks of Ekaterinburg city and tree-lines along the roads, but only 14% were in
natural conditions. In order to reveal the latitudinal–zonal specificity for the distribution of species richness
of the WLP associated mycobiota, we use Aphyllophoroids as the largest group of fungi among all analyzed
(75% of species), and Caragana arborescens, or Siberian peashrub is the richest plant substrate. Changes in
the fungal diversity were studied along a meridional transect stretching for 800 km along 60° E, from the mid-
dle boreal subzone of Sverdlovsk oblast to the steppes of Chelyabinsk oblast (Russia) and Kostanay oblast
(Kazakhstan). In each of the five vegetation zones/subzones, as well as in Ekaterinburg city, six sites were
studied, the area of which varies from 0.9 to 6.8 ha. The aboveground phytomass of C. arborescens is maximal
in the forest steppe (8.9–11.7 t/ha), and minimal at the edges of the transect (2.4–5.8 t/ha). A positive cor-
relation was found between the aboveground plant phytomass and the species richness of mycobiota, while
there was no correlation with climatic parameters. Notable differences were found in Ekaterinburg city: the
Siberian peashrub phytomass was two times lower than in the forest steppe, but the species richness of myco-
biota was similar to the forest steppe. A similar result was obtained for the α diversity (average number of fun-
gal species at the sites and Shannon index) of mycobiota: an increase in the parameters from the middle
boreal subzone to the forest steppe and a decrease in the steppe. The Whittaker and Czekanowski–Sørensen
indices (β diversity) increase towards the steppe, which is due to a strong relationship with the mean annual
temperature and precipitation. A range of fungal species gravitating towards northern, southern, and urban-
ized conditions has been revealed. In the north of transect, local species of fungi predominate, while in the
south and in Ekaterinburg city, the role of biogeographically distant (alien) taxa is high. In this regard, the
species composition of mycobiota of Siberian peashrub is divided into two clusters, northern (boreal) and
southern (nemoral-steppe) ones, including Ekaterinburg city. To the south, the species richness of patho-
genic fungi increases, but this parameter does not correlate with the C. arborescens phytomass. In plantings of
invasive Siberian peashrub, the species richness of the poroid fungi is similar to that of the corticioid fungi at
the local and regional level, which differs significantly from natural conditions. A high level of pathogenic
fungi was also revealed compared to natural conditions. The results can be used to optimize the conception
of Greenway planning in Ekaterinburg city and help prevent a number of environmental problems arising
after the rapid implementation of the strategy for developing the city and the surrounding areas.
Keywords: Russia, Kazakhstan, anthropogenic impact, biogeography, ecology, phytopathology, invasion,
climate
DOI: 10.1134/S1995425523040091
404
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
SHIRYAEV et al.
Green spaces in the cities of the Urals and urban
forests perform important environmental functions:
they promote the formation of favorable living condi-
tions and are recreational areas (Rysin, L.P. and
Rysin, S.L., 2012). Many kilometers of artificial forest
belts along fields, highways, and railways contribute to
reducing wind speed and protecting roads from snow
drifts, and field protective forest belts contribute to
snow retention (in the winter) and protection from dry
winds (in the summer), preserving soil fertility and
increasing crop yields (Burnatsky, 1952; Shortt and
Vamosi, 2012). Urban plantings of woody plants, as
well as forest belts, are artificial communities, often
consisting of introduced (alien) plant species. Being
part of the ecological framework of many regions, they
are historically and genetically alien components of
the landscape, but are necessary for its ecological opti-
mization under conditions of intensive economic
activity (Safonov et al., 2013).
Native and alien woody legumeinous plants (here-
inafter referred to as WLPs), trees and shrubs of the
family Fabaceae (Leguminosae), occupy an important
place among the vast range of plants used in urban land-
scaping and the creation of forest belts in different
regions of the Urals. Representatives of this family of
plants play an important role in the lives of people: they
have long been cultivated as food and honey-bearing,
ornamental, and forage plants, and they are also a
source of valuable wood (Knyazev, 2014). Many repre-
sentatives of Fabaceae of the Ural flora are ornamental
plants that deserve wide introduction into the practice
of floriculture (Merker, 2005). Due to the symbiosis
with nitrogen-fixing bacteria, legumes are able to
actively enrich the soil with nitrogen available to plants
(Shortt and Vamosi, 2012); therefore, even rare legumi-
nous species can be an important element of the stabil-
ity of the ecosystem the and maintenance of biodiver-
sity in biocenoses (Lashchinskii and Revyakina, 1986).
Since the beginning of the 20th century, repeated
attempts have been made to introduce WLPs that are
exotic for the region to the Middle Urals, including
Sverdlovsk oblast, one of the most economically
developed regions of Russia (Govorukhin, 1937;
Merker, 2005). However, the harsh continental cli-
mate did not contribute to the successful acclimatiza-
tion of WLPs; many such attempts were unsuccessful:
the plants died during frosty winters (Merker, 2009;
Knyazev, 2014). Nevertheless, the Central Asian
Caragana arborescens Lam. proved to be adapted to
local climatic conditions distinguished by drought
resistance, winter hardiness, and salt resistance while
being in general undemanding in regards to soil fertil-
ity (Shortt and Vamosi, 2012). This predetermined its
widespread use in the creation of forest belts along
highways and railways and agricultural land in order to
reduce the wind speed and to retain snow (Merker,
2005). C. arborescens is also used in stabilizing the
slopes of ravines and recultivating man-made land-
scapes. The decorativeness of C. arborescens and its
resistance to gases and heavy metals, as well as the
ability to quickly restore foliage after damage, pro-
motes its active use in the landscaping of settlements
(Bukharina et al., 2007; Kopylova, 2017), especially
because C. arborescens does not suffer from dust, tol-
erates intensive cutting, and is a medicinal plant
(Bukharina et al., 2007; Shortt and Vamosi, 2012).
Due to global warming in recent decades (Report …,
2020), the climate in Sverdlovsk oblast becomes more
comfortable for WLPs introduced from more southern
regions. For this reason, the list of introduced and
invasive plant species is steadily expanding, although
most WLP species are undoubtedly still able to survive
in winter exclusively with the help of humans
(Knyazev, 2014). The increase in the diversity and
phytomass of WLPs alien to the region is a leading fac-
tor for the penetration of new species of fungi into the
Middle Urals, which can cause various diseases of the
plants under consideration (Shiryaev et al., 2022a).
Over 50 years (1970–2020), the phytomass of alien
woody plants in the parks of Ekaterinburg (the admin-
istrative center and the largest city of Sverdlovsk
oblast) increased many times: the introduced East
Asian woody species Juglans mandshurica Maxim. and
Vitis amurensis Rupr. increased 5–7 times; North
American Acer negundo L. and Fraxinus pennsylvanica
Marshall increased 4–8 times; and European Quercus
robur L. and Acer platanoides L. increased 3–4 times
(Sostoyanie …, 2019; Shiryaev et al., 2022a). During
this period, at least 60 new species of phytopathogenic
micromycetes and more than 20 species of macromy-
cetes appeared in Ekaterinburg on alien species of
woody plants (Shiryaev et al., 2010, 2021, 2022a,
2022b; Bulgakov and Shiryaev, 2022). An expansion of
the range of host plants was observed for a number of
phytopathogenic fungi compared to the natural range
(Shiryaev et al., 2021, 2022b). However, the mycobi-
ota of WLPs in Sverdlovsk oblast was not specially
studied, so only single findings of fungi collected from
local WLPs (Stepanova and Sirko, 1970; Stepanova,
1977) and on an alien Siberian peashrub (Stepanova,
1971; Shiryaev et al., 2010) were known. The timely
detection of new fungal phytopathogens that have
appeared in the region due to the mass and uncon-
trolled import of seedlings and plant diseases caused by
them will contribute to maintaining the biosafety of
natural and anthropogenic habitats of the Middle Urals.
This work presents the results of studies aimed at
identifying the diversity of wood-inhabiting fungi
developing on native and alien species of woody
legumes in Sverdlovsk oblast, as well as identifying the
factors determining the spatial distribution of the fun-
gal biodiversity on the model object, Caragana arbo-
rescens, as the most common WLP species along the
eastern macroslope of the Urals. The hypothesis about
the positive correlation between the number of species
of fungi (including phytopathogenic ones) and the
WLP biomass was proposed and tested.
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
FUNGAL DIVERSITY OF NATIVE AND ALIEN WOODY LEGUMINOUS PLANTS 405
2. MATERIALS AND METHODS
2.1. Study Area
Sverdlovsk oblast is the largest region of the Middle
Urals (Fig. 1). Ekaterinburg city (56°50′ N; 60°35′ E;
280 m above sea level) is the administrative center of
Sverdlovsk oblast, with a population of 1.6 million
people and an area of 1112 km2, located on the border
between Europe and Asia (Sostoyanie…, 2019). Over
the past 10 years, the average annual temperature in
Ekaterinburg has fluctuated between 3.1 and 5.3°C
(Fick and Hijmans, 2017; Federal Service…, 2021).
The climate is continental, with pronounced seasonal-
ity. The average monthly temperature in July is
19.4°C, with an absolute maximum of 39.6°C. The
average monthly temperature in January is –14.3°C
and the absolute minimum is –46.7°C. The average
annual precipitation in the period from 2010 to 2020
was 537 mm/year (Federal Service…, 2021).
In the north of Sverdlovsk oblast, in the middle
boreal subzone zone, the average annual temperature
is 0.6°C, while in the south of the region, in the forest-
steppe and hemiboreal subzone (mixed coniferous–
deciduous forests), it reaches 3.7°C (Federal Ser-
vice…, 2021). Marshes are widespread in the lowland
parts in the north of the region, and mountain tundra
and permafrost are widespread in the mountains. In
the hemiboreal forests, spruce, fir and pine forests
predominate, in which broad-leaved woody plants
(Acer platanoides L., Quercus robur L., Tilia cordata Mill.,
Ulmus laevis Pall. etc.), as well as Betula pendula Roth.,
Populus tremula L., Salix spp., Sorbus aucuparia L., etc.,
are present. Birch and aspen stands with undergrowths
of T. cordata, B. pendula, P. tremula, S. aucuparia,
Crataegus sanguinea Pall., etc., are widespread in the
forest-steppe zone (Merker, 2009).
2.2. Woody Plants of the Family Fabaceae
in Sverdlovsk Oblast
In this work we considered trees and shrubs (pha-
nerophytes according to the Raunkier system, includ-
ing nanophanerophytes) of woody plants of the family
Fabaceae (Raunkier, 1937), while hamephytes and
hemicryptophytes, for example, representatives of the
genera Astragalus L., Hedysarum L., Oxytropis DC.,
etc., are excluded.
Twenty WLP species currently grow in the natural
and anthropogenic conditions of the open ground in
Sverdlovsk oblast. Three species of them are native
plants: the Russian peashrub (Caragana frutex (L.)
K.Koch.), Russian broom (Chamaecytisus ruthenicus
Fig. 1. (a) Location of Sverdlovsk oblast. The red line is the border between Europe and Asia. (b) Latitudinal–zonal transect along
60° E, f rom th e mid dle ta iga of Sverd lovsk oblast to the steppes of C helya binsk ob last (Russia ) and Kosta nay obla st (Kazakh stan).
Kazakhstan
Kazakhstan
(a)
Perm
1
(b)
Yekaterinburg
Kurgan
Chelyabinsk
Ufa
Kostan ay
Orenburg Orsk
2
34
5
6
78
9
10 11 12
19
20
21
22
23
24
25
26
27
28
29
30 31
32
33
34
35
36
406
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
SHIRYAEV et al.
(Fisch. ex Vol.) Klask.), and dyer’s greenweed (Genista
tinctoria L.). These are low shrubs 0.3–1.5 m in height,
found in steppe or forest biotopes in open areas in pine
and deciduous forests and meadows, from the alpine
border of the forest and the middle boreal to the forest
steppe. Seventeen alien species were introduced from
other biogeographic regions (Fig. 2): four species have
natural habitats in Southern Europe: Genista florida L.,
Laburnum alpinum (Mill.) Bercht. & J.Presl, L., ana-
gyroides Medik. L., and watereri (G. Kirchn.) Dippel
(= L. alpinum × L. anagyroides); four species in Cen-
tral Asia (including mountains of southern Siberia):
Caragana arborescens Lam., C. decorticans Hemsl.,
C. spinosa (L.) Vahl ex Hornem., and C. turkestanica
Kom.; five species in East Asia: Caragana boisii
C.K. Schneid., C. microphylla Lam., C. ussuriensis
(Regel.) Pojark., Lespedeza bicolor Turcz., and Maackia
amurensis Rupr.; and four species in North America:
Amorpha fruticosa L., Gleditsia triacanthos L., Gymno-
cladus dioicus (L.) K. Kotch., and Robinia pseudoaca-
cia L. All introduced WLP species grow exclusively in
botanical gardens and parks of Ekaterinburg, with the
exception of C. arborescens, which is widespread in
Sverdlovsk oblast in parks and forest belts.
2.3. Latitudinal–Zonal (meridional) Transect
with Caragana arborescens
Caragana arborescens is a shrub reaching about 6 m
in height. Its natural range is Eastern Kazakhstan and
Kyrgyzstan, the mountains of Southern Siberia (Altai,
Sayan), and the northwest and northeast of China and
Mongolia (Shortt and Vamosi, 2012). The secondary
range of the Siberian peashrub covers almost the
entire territory of the forest zone of Russia (Knyazev,
2014). The acclimatization of the Siberian peashrub in
the Urals began in the first decade of the 20th century,
and mass plantings, some fragments of which have
survived to the present, were made in the 1950s
(Merker, 2009). C. arborescens is widespread in the
cities of the Urals and predominates in forest belts
along highways, railways, and agricultural fields from
the middle taiga to the steppe.
In the forest belts with Siberian peashrub, the set of
accompanying woody plants changes significantly
with latitude: Elaeagnus angustifolia L. and Prunus spi-
nosa L. were planted together in the steppe; Populus
balsamifera L. s. l., Betula pendula Roth, B. pubescens
Ehrh., Acer tataricum L., Syringa vulgaris L., S. josi-
kaea J. Jacq. ex Rchb., and Lonicera tatarica L. in the
forest steppe; Acer negundo L., Malus sylvestris Mill.,
P. balsamifera, and Syringa spp. in the hemiboreal sub-
zone; Pinus sylvestris L. and A. negundo in the south
boreal subzone; and Picea obovata Ledeb., P. s yl ve st ri s,
Salix caprea L., and Prunus padus L. in the middle
boreal subzone. Pure plantings of C. arborescens or
mixed with A. negundo, P. b a ls a mi fe r a , Lonicera tatar-
ica, and Syringa spp. are widespread in the central part
of Ekaterinburg. The names of plant species are given
according to the World Flora Online database (2023).
Over 20 years (2002–2022), the species richness of
wood-destroying fungi on C. arborescens was studied
on a latitudinal–zonal (meridional) transect 800 km
in length, stretching along the eastern macroslope of
the Ural-Novaya Zemlya mountain country (Fiziko-
geograficheskoe…, 1966) along 60° E (Fig. 1, Table 1).
The mycobiota of Siberian peashrub was studied at
36 sites, from the middle boreal subzone in Sverdlovsk
oblast (Severouralsk town surroundings, 60°09′ N) to
the steppe zone of Kazakhstan (Kostanay oblast) and
Chelyabinsk oblast, Russia (the settlement of Kartaly,
53°03′ N). Ekaterinburg city is located on the border
between the southern boreal and hemiboreal sub-
zones, i.e., in the middle of the studied latitudinal–
zonal transect.
In each of the five latitudinal divisions (natural and
climatic zones/subzones), as well as in Ekaterinburg,
six sites have been studied, the size of which varies
Fig. 2. Number of species of woody leguminous plants in Sverdlovsk oblast from different biogeographic regions. The total num-
ber of plant species and the number of species on which fungi were identified are given.
0 1 2 3 4 5 6
Total
North American
East Asian
Central Asian
European
Native With fungi
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
FUNGAL DIVERSITY OF NATIVE AND ALIEN WOODY LEGUMINOUS PLANTS 407
Table 1. Location of 36 sites studied in six latitudinal divisions located along the meridional transect
SО, Sverdlovsk oblast, Russia; ChО, Chelyabinsk oblast, Russia; KZ, Kostanay oblast, Kazakhstan.
Latitudinal
subdivision No. Name of the site Coordinates Area,
ha
Species of fungi,
individuals
Middle boreal 1 Surroundings of Severouralsk town, SO 60°09′ N, 60°00′ E2.9 8
2 Surroundings of Koziy village, SO 59°59′ N, 60°04′ E4 9
3 Surroundings of Karpinsk town, SO 59°46′ N, 60°03′ E3.3 10
4 Surroundings of Serov town, SO 59°37′ N, 60°38′ E3.8 10
5 Surroundings of Lobva village, SO 59°09′ N, 60°27′ E4.4 11
6 Surroundings of Verkhoturye town, SO 58°52′ N, 60°41′ E5.7 12
South boreal 7 Surroundings of Nizhny Tagil town, SO 57°58′ N, 59°51′ E3.9 9
8 Surroundings of Lipovka village, SO 57°27′ N, 61°10′ E3.6 10
9 Surroundings of Novouralsk town, SO 57°16′ N, 60°11 ′ E5.6 13
10 Surroundings of Kolpakovka village, SO 57°27′ N, 58°49′ E5.2 11
11 Surroundings of Losiny village, SO 57°08′ N, 61°04′ E3.6 10
12 Surroundings of Reftinskiy village, SO 57°05′ N, 61°40′ E 3.7 10
Ekaterinburg 13 av. Kosmonavtov, SO 56°55′ N, 60°36′ E1.8 10
14 Arboretum on st. Pervomaiskaya, SO 56°50′ N, 60°39′ E1.5 10
15 Mayakovsky Park of Culture and Recreation, SO 56°48′ N, 60°38′ E2.7 10
16 Park of 50 Years Anniversary of the Soviet Union, SO 56°49′ N, 60°37′ E3.2 11
17 Vigorov Garden of Medical Cultures, SO 56°49′ N, 60°39′ E3.2 12
18 Botanical Garden, Ural Branch, Russian Academy of
Sciences, SO
56°47′ N, 60°36′ E 4.8 15
Hemiboreal 19 Surroundings of Klenovskoe village, SO 56°47′ N, 58°36′ E2.2 10
20 Surroundings of Polovinka village, SO 56°32′ N, 59°13′ E3.7 11
21 Surroundings of Ledyanka village, SO 56°40′ N, 59°51′ E4 12
22 Surroundings of Verkhnyaya Sysert town, SO 56°26, 60°46′ E5.3 12
23 Surroundings of Verkhniy Ufalei town, ChO 56°01′ N, 60°17′ E4.5 13
24 Surroundings of Selyankino village, ChO 55°13′ N, 60°11 ′ E6 16
Forest-steppe 25 Surroundings of Krasnoufimsk town, SO 56°43′ N, 57°42′ E4.1 14
26 Surroundings of Bolshaya Tavra village, SO 56°09′ N, 58°00′ E3.6 15
27 Surroundings of Pokrovskoye village, SO 56°58′ N, 61°37′ E6.8 20
28 Surroundings of Maly Kuyash village, ChO 55°51′ N, 61°10′ E4.0 18
29 Surroundings of Muslyumovo village, ChO 55°35′ N, 61°36′ E3.9 17
30 Surroundings of Kukushka village, ChO 54°32′ N, 60°30′ E5.5 17
Steppe 31 Surroundings of Troitsk town, ChO 54°06′ N, 61°28′ E1.8 9
32 Surroundings of Karabalyk village, KZ 53°44′ N, 62°02′ E в 1.5 8
33 Surroundings of Slavenka village, KZ 53°19′ N, 62°02′ E2.3 9
34 Surroundings of Varna village, ChO 53°23′ N, 60°58′ E2.4 11
35 Surroundings of Parizh village, ChO 53°18′ N, 60°06′ E1.8 10
36 Surroundings of Kartaly town, ChO 53°03′ N, 60°38′ E0.9 6
408
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
SHIRYAEV et al.
from 0.9 to 6.8 ha. The work was carried out only
within the forest belts or urban plantings. There are
25 sites in Sverdlovsk oblast, 9 sites in Chelyabinsk
oblast, and 2 sites in Kostanay oblast (Fig. 1). Studies
at the sites were conducted from two to five times: in
Sverdlovsk oblast in 2005–2022, in Chelyabinsk oblast
in 2006–2012, in Kostanay oblast in 2002–2009.
The aboveground phytomass is the weight of dry
living (biomass) and dead (litter) plants that were pre-
served the anatomical structure of plants at the time of
accounting. The aboveground phytomass of Siberian
peashrub was examined in October 2022 after leaf fall.
The material from three plots measuring 10 × 10 m
from four sites in each latitudinal subdivision and
Ekaterinburg (72 plots in total) was studied. The cut
phytomass was dried to an absolutely dry weight at
105°C and weighed (Metody…, 2002).
In the analysis we used the De Martonne aridity
index (Bykov, 1983), which indicates the degree of
dryness (aridity) of the climate, calculated as the ratio
where R is the annual sum of precipitation (cm) and
tis the average annual temperature (°С).
2.4. Study of Mycobiota
The fruit bodies of fungi identified only on wood
(dry, windfall, living, and dead) were taken into
account. Fungal species forming fruit bodies on soil,
leaves, and litter were excluded from the study. When
determining the number of species on the site, the
average number of species, etc., the accounting unit of
the fungus, a specimen is understood as one species
inhabiting a discrete unit of the substrate (branch or
trunk) (Zmitrovich et al., 2018). The species richness
of mycobiota per unit area was determined as the aver-
age number of species per hectare (species/ha).
The following designations are given in the list of
fungal species (Table 2). The total number of samples
of fungi (the number of the site where the sample was
found: the number of samples and the substrate). For
example, 1(20:1Chr) means that one sample was
found in this latitudinal subdivision (one sample was
found on Chamaecytisus ruthenicus at site no. 20).
Species of fungi that are new to Sverdlovsk oblast are
marked with an asterisk (*). The names of fungal spe-
cies are given according to the Index Fungorum data-
base (2023).
2.5. Statistical Analysis
Ward’s method and the Pearson correlation coeffi-
cient were used to construct a dendrogram of the sim-
ilarity of the fungal species composition on different
WLP species, as well as individual latitudinal divisions
(climatic zones/subzones and Ekaterinburg). Spear-
man’s rank correlation coefficient (rs) was used to
(
)
+
/10,
Rt
establish a linear relationship between different param-
eters of microbiota, climate, and phytomass. The
Mann–Whitney U-test was used to evaluate the differ-
ences between two independent samples in order to
compare the levels of species richness between alien
and native species of WLPs.
The alpha diversity of mycobiota was measured as
the average number of fungal species at six sites within
one latitudinal subdivision, as well as by the value of the
Shannon index. The beta diversity was estimated as an
average of the Czekanowski–Sørensen index (Cs)
between six sites within one latitudinal subdivision, as
well as Whittaker’s Index (Metody…, 2002):
where S is the total number of species and α is the
average number of species per site.
An approach based on a sample generation algo-
rithm was used to estimate the expected number of
species on the transect (Colwell et al., 2012). This
approach is based on the construction of a rarefaction
curve using a special algorithm for random multiple
permutation of data within samples from the number
of detected samples. To calculate the expected number
of species in the general population from which the
sample was taken, an adjusted Chao1 index (index of
adjusted dataset) was used, which was calculated based
on the registration of the number of species repre-
sented by one sample.
An indirect method of assessing the completeness
of the identification of species richness is also applied,
the Turing coefficient (C), which is based on the ratio
of the number of singleton species (represented by a
single finding) to the total number of identified species
(Good, 1979),
where f1 is the number of singleton species and S is the
total number of identifies species. The potential num-
ber of species (T) can be calculated as
Statistical data processing was performed by means
of EstimateS 9.10 and MS Excel 2007 statistical soft-
ware packages.
3. RESULTS
3.1. Species Richness of Wood-Inhabiting Fungi
on Plants of the family Fabaceae in Sverdlovsk Oblast
Over 20 years of studies, 136 species of fungi have
been identified on WLPs: 127 species of Basidiomy-
cota and 9 species of Ascomycota (Table 2). Among
them, 14 species are reported for the first time for
Sverdlovsk oblast, 8 species of Basidiomycota: Fibulo-
myces mutabilis (Bres.) Jülich, Lachnella alboviolas-
cens (Alb. & Schwein.) Fr., Lentaria surculus (Berk.)
Corner, Phloeomana minutula (Sacc.) Redhead, Sang-
β= α−
/1,
wS
(
)
=− ×
1
1 / 100%,
CfS
=
/.
TSC
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
FUNGAL DIVERSITY OF NATIVE AND ALIEN WOODY LEGUMINOUS PLANTS 409
Table 2. Spatial distribution of the species composition of wood-destroying fungi on woody leguminous plants
Species Middle boreal South boreal Ekaterinburg Hemiboreal Forest steppe Steppe
BASIDIOMYCOTA
Aphyllophoroid fungi
Amphinema byssoides (Pers.)
J. Erikss
2(17:1Ma,
16:1Lb)
1(25:1Ca)
Antrodia albida (Fr.) Donk 1(17:1Ma)
Antrodiella faginea Vampola
& Pouzar
2(15:1Ca; : 1Ca)
A. leucoxantha (Bres.) Mietti-
nen & Niemelä
1(25:1Gt) 1(31:1Gt)
A. romellii (Donk) Niemelä 2(14:1Ca;
15: 1Ca)
A. serpula (P. Karst.) Spirin &
Niemelä
1(5:1Ca) 2(9:1Ca; 11:1Ca) 1(18:1Ca)
Athelia epiphylla Pers. 1(21:1Chr) 1(27:1Cf)
Baltazaria galactina (Fr.)
Leal-Dutra, Dentinger &
G.W. Griff.
2 (3:1Ca; 5:1Ca) 1(10:1Ca) 3(21:2Ca;
23:2Ca)
Bjerkandera adusta (Willd.)
P. Ka r st.
4(1:1Ca; 2: 1Ca;
4: 1Ca; 6: 1Ca)
3(7:1Ca; 11:1Ca;
12: 1Ca)
6(15:2Ca;
16:3Ca; 17:1Rp;)
2(19:1Ca;
24:1Ca)
4(25:2Ca;
29:1Ca; 30:1Ca)
Byssocorticium atrovirens (Fr.)
Bondartsev & Singer
2(25:1Ca;
27:1Ca)
Ceraceomyces microsporus
K.H. Larss.
1(18:1Ca)
Ceriporia viridans (Berk. &
Broome) Donk
2(15:1Ca;
18: 1Ca)
Cerioporus squamosus (Huds.)
Quél.
1(9:1Ca) 3(13:2Ca;
17: 1Ca)
2(19:1Ca;
24:1Ca)
2(26:1Ca;
30:1Ca)
2(32:1Ca;
34:1Ca)
C. varius (Pers.) Zmitr. &
Kovalenko
3(27:2Ca;
29:1Ca)
Cerrena unicolor (Bull.) Mur-
rill
3(7:1Ca; 9:1Ca;
12: 1Ca)
4(13:1Ca;
15:1Ca; 17:1Ma;
17:1Rp )
3(20:2Ca;
23:1Ca)
2(25:1Ca;
30:1Ca)
Chondrostereum purpureum
(Pers.) Pouzar
2(21:1Chr;
24:1Chr)
Clavulina cinerea (Bull.)
J. Schröt.
2(3:1Ca; 6:1Ca)
Cyanosporus caesius (Schrad.)
McGinty
2(18:2Ca) 1(22:1Ca) 2(28:1Ca;
30:1Ca)
1(34:1Ca)
Cylindrobasidium laeve (Pers.)
Chamuris
1(15:1Af) 1(28:1Cf) 2(33:1Ca;
35:1Ca)
Daedalea xantha (Fr.) A. Roy
& A.B. De
2(27:1Ca;
30:1Ca)
Erythricium laetum (P. Karst.)
J. Erikss. & Hjortstam
2(27:1Chr,
28:1Gt)
*Fibulomyces mutabilis (Bres.)
Jülich
2(22:1Ca,
1Chr)
Fomitoporia punctata
(P. Karst.) Murrill
2(21:1Ca;
23:1Ca)
1(25:1Ca) 2(32:2Ca)
F. r ob us ta (P. Karst.) Fiasson
& Niemelä
1(26:1Ca) 3(33:1Ca;
35:2Ca)
Fomitopsis pinicola (Sw.)
P. Ka r st.
2(7:1Ca; 12:1Ca) 1(27:1Ca) 2(33:1Ca)
410
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
SHIRYAEV et al.
Fuscoporia contigua (Pers.)
G. Cunn.
1(23:1Ca) 2(26:1Ca;
29:1Ca)
3(31:2Ca;
34:1Ca)
F. ferruginosa (Schrad.)
Murrill
1(26:1Ca) 2(32:1Ca;
35:1Ca)
Ganoderma applanatum
(Pers.) Pat.
1(18:1Ca) 1(31:1Ca)
Hapalopilus rutilans (Pers.)
Murrill
2(2:1Ca; 4:1Ca) 2(18:2Ca) 2(25:1Ca;
30:1Ca)
1(33:1Ca)
Hymennochaete cinnamomea
(Pers.) Bres.
1(22:1Ca) 1(28:1Ca) 1(35:1Ca)
Hyphoderma transiens (Bres.)
Parmasto
2(16:1Ca;
17:1Ma )
1(27:1Ca)
Hypochnicium bombycinum
(Sommerf.) J. Erikss.
1(1:1Ca) 1(9:1Ca)
Hypochnicium sp. nova ined. 1(25:1Ca)
Inonotus hispidus (Bull.)
P. K arst .
2(15:2Ca) 2(32:2Ca)
Irpex lacteus (Fr.) Fr. 4(2:1Ca; 5:2Ca;
6:1Ca)
7(7:1Ca; 8:1Ca;
9:3Ca; 11:2Ca)
12(13:2Ca;
14:3Ca; 15:4Ca;
16:3Ca )
8(19:2Ca;
20:1Ca;
21:3Ca ;
23:2Ca)
11 ( 25 : 4 C a ;
27:3Ca; 29:4Ca)
6(33:2Ca;
34:3Сa;
35:1Ca)
Junghuhnia nitida (Pers.)
Ryvarden
2(1:1Ca; 4:1Ca) 2(11:2Ca)
*Lachnella alboviolascens
(Alb. & Schwein.) Fr.
3(16:3Ca)
Laetiporus sulphureus (Bull.)
Murrill
1(3:1Ca) 1(18:1Ca) 2(27:1Ca;
30:1Ca)
2(35:2Ca)
*Lentaria surculus (Berk.)
Corner
1(27:1Ca)
Leucogyrophana mollusca
(Fr.) Pouzar
2(18:2Ca) 2(21:1Ca;
24:1Ca)
1(26:1Ca)
Lyomyces crustosus (Pers.)
P. K arst .
1(25:1Ca) 2(32:1Ca,
1Cf)
L. erastii (Saaren. & Kotir.)
Hjortstam & Ryvarden
4( 15: 2C a; 18: 1 Gf;
15: 1Gt )
2(28:1Ca, 1Chr) 2(34:1Ca;
35:1Ca)
L. pruni (Lasch) Riebesehl &
Langer
2(26:1Ca;
29:1Ca)
2(33:2Ca)
Mutatoderma mutatum (Peck)
C.E. Gómez
2(25:1Ca;
30:1Ca)
Neofavolus alveolarius (DC.)
Sotome & T. Hatt.
2(2:1Ca; 6:1Ca) 2(8:2Ca) 5(15:3Ca;
16:2Ca)
7(19:2Ca;
20:1Ca;
21:2Ca ;
23:1Ca;
24:1Ca)
13(25:4Ca;
26:5Ca; 28:2Ca;
30:2Ca)
19(31:4Ca;
32:5Ca;
34:6Ca;
35:4Ca)
Odontia calcicola (Bourdot &
Galzin) Kõljalg
2(22:1Ca;
24:1Ca)
Oxyporus corticola (Fr.)
Ryvarden
2(18:2Ca) 2(26:2Ca) 1(31:1Ca)
O. latemarginatus (Durieu &
Mont.) Donk
1(21:1Ca) 2(33:1Ca;
35:1Ca)
Species Middle boreal South boreal Ekaterinburg Hemiboreal Forest steppe Steppe
Table 2. (Contd.)
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
FUNGAL DIVERSITY OF NATIVE AND ALIEN WOODY LEGUMINOUS PLANTS 411
Peniophora cinerea (Pers.)
Cooke
3(16:2Ca;
14:1Ma)
4(20:1Ca;
23:2Ca,
1Chr)
5(25:2Ca;
28:2Ca; 30:1Ca)
4(32:2Ca;
34:1Ca;
36:1Cf)
P. i n ca rn at a (Pers.) P. Karst. 3(3:1Ca; 5:1Ca;
6:1Ca)
2(9:2Ca)
P. l i mi ta ta (Chaillet ex Fr.)
Cooke
2(15:2Ca) 1(22:1Ca) 2(35:1Ca;
36:1Ca)
P. l yc ii (Pers.) Höhn. &
Litsch.
1(17:1Rp) 2(26:1Gt;
28:1Ca)
2(33:1Ca;
36:1Ca)
P. violaceolivida (Sommerf.)
Massee
3(20:1Ca;
23:2Chr)
1(28:1Ca)
Perenniporia sp. nova ined. 1(26:1Ca)
Phaeoclavulina flaccida (Fr.)
Giachini
1(4:1Ca) 1(12:1Ca)
Phanerochaete alnea (Fr.)
P. K arst .
1(20:1Chr)
Ph. laevis (Fr.) J. Erikss. &
Ryvarden
1(18:1Ca) 1(21:1Ca) 1(31:1Ca)
Phylloporia sp. nova ined. 1(26:1Ca)
Piloderma byssinum
(P. Karst.) Jülich
2(20:1Ca;
24:1Chr)
Podofomes mollis (Sommerf.)
Gorjón
1(3:1Ca) 2(10:2Ca) 1(28:1Ca)
P. stereoides (Fr.) Gorjón 2(31:1Ca;
35:1Ca)
Porostereum spadiceum (Pers.)
Hjortstam & Ryvarden
2(16:2Ca) 3(19:2Ca;
23:1Ca)
2(29:1Ca;
30:1Ca)
Pseudotomentella tristis
(P. Karst.) M.J. Larsen
2(26:1Ca;
29:1Ca)
Pterula subulata Fr. 2(9:2Ca)
Pycnoporellus fulgens (Fr.)
Donk
1(27:1Ca)
Raduliporus aneirinus (Som-
merf.) Spirin & Zmitr.
3(25:2Ca;
30:1Ca)
Radulomyces confluens (Fr.)
M.P. Christ.
2(4:1Ca; 6:1Ca) 3(8:2Ca; 10:1Ca)
R. rickii (Bres.) M.P. Christ. 4(13:1Ca;
15: 3Ca)
2(21:1Ca;
24:1Ca)
2(27:1Ca;
30:1Ca)
Rigidoporus sanguinolentus
(Alb. & Schwein.) Donk
2(22:1Ca;
24:1Ca)
1(28:1Ca)
*Sanghuangporus cf. baumii
(Pilát) L.W. Zhou &Y.C. Dai
1(27:1Ca)
Schizophyllum commune Fr. 3(1:1Ca; 5:1Ca;
6:1Ca)
4(7:2Ca; 9:2Ca) 20(13:4Ca;
14:5Ca; 15:4Ca;
16:4Ca, 1La;
17: 1М a, 1 L b )
5(19:1Ca;
20:2Ca;
22:1Ca;
24:1Ca)
5(25:2Ca;
27:1Ca; 29:2Ca)
6(31:2Ca;
34:2Ca;
35:2Ca)
Schizopora paradoxa
(Schrad.) Donk
4(14:2Ca;
16:1Cd, 1Ca)
4(26:2Ca;
29:1Ca; 30:1Ca)
3(32:2Ca;
35:1Ca)
Scopuloides hydnoides (Cooke
& Massee) Hjortstam &
Ryvarden
3(27:2Ca;
29:1Ca)
Species Middle boreal South boreal Ekaterinburg Hemiboreal Forest steppe Steppe
Table 2. (Contd.)
412
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
SHIRYAEV et al.
Sertulicium niveocremeum
(Höhn. & Litsch.) Spirin &
K.H. Larss.
2(22:1Ca,
1Ca)
1(26:1Ca) 2(32:2Ca)
Sistotrema muscicola (Pers.) S.
Lundell
2(2:1Ca; 6:1Ca) 3(10:3Ca)
*S. octosporum (J. Schröt. ex
Höhn. & Litsch.) Hallenb.
1(17:1Rp) 2(23:1Ca,1
Chr)
Skeletocutis nivea (Jungh.)
Jean Keller
1(12:1Ca) 3(18:3Ca) 2(26:1Ca;
30:1Ca)
S. bourdotii Saliba & A. David 2(16:2Ca) 3(20:2Ca;
24:1Ca)
2(25:1Ca;
30:1Ca)
2(33:1Ca;
35:1Ca)
S. fimbriatum (Pers.) J. Erikss. 18(1:3Ca; 3:2Ca;
4:4Ca; 5:4Ca;
6:5Ca)
9(7:1Ca; 8 :2Ca;
9:1Ca; 10:2Ca;
11:3 C a )
5(13:1Ca;
14:2Ca;
17: 2Ca)
4(19:1Ca;
21:1Ca ;
24:1Ca)
3(27:1Ca;
29:1Ca; 30:1Ca)
3(31:2Ca;
34:1Ca)
S. laeticolor
(Berk. & M.A. Curtis) Banker
3(13:2Ca;
18: 1Ca)
2(22:1Ca;
:1Ca)
S. ochraceum
(Pers. ex J.F. Gmel.) Gray
5(8:2Ca; 9:2Ca;
10:1Ca)
4(14:1Ca;
16:3Ca)
4(26:1Ca;
29:2Ca; 30:1Ca)
Stereum hirsutum (Willd.)
Pers.
2(2:1Ca; 5:1Ca) 3(7:3Ca) 3(13:2Ca;
17:1Rp )
S. subtomentosum Pouzar 2(16:2Ca) 1(28:1Ca)
Subulicystidium longisporum
(Pat.) Parmasto
1(18:1Ca) 3(20:2Ca;
24:1Chr)
Tomentella cinerascens
(P. Karst.) Höhn. & Litsch.
1(4:1Ca) 3(9:3Ca)
T. botr yo id es (Schwein.)
Bourdot & Galzin
2(23:1Ca;
24:1Chr)
2(25:1Ca;
30:1Ca)
1(32:1Ca)
T. bryophila (Pers.)
M.J. Larsen
3(8:1Ca; 11:2Ca) 1(26:1Ca)
T. f i brosa (Berk. & M.A. Cur-
tis) Kõljalg
1(21:1Ca) 1(31:1Ca)
T. stupo sa (Link) Stalpers 2(22:1Ca,1
Chr)
Trametes hirsuta (Wulfen)
Lloyd
9(13:3Ca;
14:2Ca; 16:4Ca)
3(20:1Ca;
23:2Ca)
5(31:2Ca;
34:2Ca;
36:1Ca)
T. ochra cea (Pers.) Gilb. &
Ryvarden
3(14:1Ca;
18: 2Ca)
4(19:2Ca;
22:2Ca)
4(26:1Ca;
28:2Ca; 30:1Ca)
3(33:2Ca;
35:1Ca)
T. ve rs i color (L.) Lloyd 3(1:1Ca; 5:1Ca;
6:1Ca)
3(7:1Ca; 10:2Ca) 2(17:1Ca, 1Ma)
Trechispora cohaerens
(Schwein.) Jülich & Stalper s
2(27:1Ca;
29:1Ca)
T. fa ri n acea (Pers.) Liberta 2(22:2Ca) 1(33:1Cf)
T. mollusca (Pers.) Liberta 3(2:1Ca; 3:1Ca;
6:1Ca)
2(9:2Ca) 2(14:1Ma;
16:1Ca)
Typ hul a micans (Pers.)
Berthier
2(4:1Ca; 5:3Ca) 2(12:2Ca) 2(17:1Rp, 1Lb) 3(19:2Ca;
23:1Ca)
3(27:1Ca;
28:1Ca)
2(31:1Ca;
36:1Ca)
T. spathulata (Corner)
Berthier
2(11:2Chr) 1(18:1Chr) 2(21:2Chr)
*Xylodon borealis (Kotir. &
Saaren.) Hjortstam &
Ryvarden
1(12:1Ca)
Species Middle boreal South boreal Ekaterinburg Hemiboreal Forest steppe Steppe
Table 2. (Contd.)
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
FUNGAL DIVERSITY OF NATIVE AND ALIEN WOODY LEGUMINOUS PLANTS 413
X. brevisetus (P. Karst.) Hjort-
stam & Ryvarden
2(27:1Ca;
30:1Ca)
X. flaviporus (Berk. &
M.A. Curtis ex Cooke) Rie-
besehl & Langer
3(3:1Ca; 4:2Ca) 3(8:1Ca; 10:1Ca;
12: 1Ca)
2(18:1Cu, 1Af) 4(20:1Ca;
22:3Ca)
2(26:1Ca;
28:1Ca)
2(36:1Ca;
35:1Ca)
X. nesporii (Bres.) Hjortstam
& Ryvarden
3(27:2Ca;
29:1Ca)
X. sambuci (Pers.) Ţura,
Zmitr., Wasser & Spirin
12( 1:1 Ca; 2:1Ca;
3:3Ca; 4:2Ca;
5:3Ca; 6:2Ca)
14(7:2Ca; 8:1Gt;
9:3Ca; 10:3Ca;
11:2Ca; 12:3Ca)
15(13:3Ca;
14:1Rp, 2Ca;
16:1Cu, 3Ca;
18:1Af, 4Ca)
16(19:3Ca,
1Gt;
20:4Ca;
22:6Ca;
23:2Ca)
12(25:3Ca, :1Cf;
26:4Ca; 27:3Ca;
28:2Ca)
15(31:3Ca
, 1Cf;
33:3Ca;
34:4Ca;
35:4Ca)
Agaricoid fungi
Coprinopsis atramentaria
(Bull.) Redhead, Vilgalys &
Moncalvo
1(18:1Cu)
Crepidotus calolepis (Fr.)
P. K arst .
1(18:1Ca)
C. caspari Velen. 1(18:1Ca)
C. subverrucisporus Pilát 1(18:1Ca)
Flammulina velutipes (Curtis)
Singer
1(18:1Ca)
Galerina marginata (Batsch)
Kühner
1(18:1Ca)
Gymnopilus junonius (Fr.)
P.D . Or ton
1(18:1Ca)
Hemimycena sp. 1(18:1Ca)
Hypholoma lateritium
(Schaeff.) P. Kumm.
1(27:1Ca)
Megacollybia platyphylla
(Pers.) Kotl. & Pouzar
1(18:1Ca)
Mycena abramsii (Murrill)
Murrill
1(18:1Ca)
M. galericulata (Scop.) Gray 1(14:1Ma)
Pholiota limonella (Peck)
Sacc.
1(18:1Ca)
*Phloeomana minutula
(Sacc.) Redhead
1(18:1Ca)
Pleurotus pulmonarius (Fr.)
Quél.
1(5:1Ca) 2(8:1Ca; 12:1Ca) 5(13:1Ca;
15: 2Ca; 18:2Ca)
9(20:3Ca;
22:4Ca;
24:2Ca)
11 ( 25 : 3 C a ;
26:3Ca; 27:2Ca;
28:2Ca; 30:1Ca)
4(31:2Ca;
32:1Ca;
35:1Ca)
Pluteus salicinus (Pers.)
P. Kum m.
1(18:1Ca)
*Simocybe haustellaris (Fr.)
Watling
1(22:1Ca)
S. sumptuosa (P.D. Orton)
Singer
1(18:1Ca)
Gasteroid fungi and heterobasidiomycetes
Apioperdon pyriforme
(Schaeff.) Vizzini
1(10:1Ca) 1(19:1Ca)
Crucibulum laeve (Huds.)
Kambly
1(2:1Ca)
Species Middle boreal South boreal Ekaterinburg Hemiboreal Forest steppe Steppe
Table 2. (Contd.)
414
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
SHIRYAEV et al.
huangporus сf. baumii (Pilát) L.W. Zhou & Y.C. Dai,
Simocybe haustellaris (Fr.) Watling, Sistotrema octospo-
rum (J. Schröt. ex Höhn. & Litsch.) Hallenb., and
Xylodon borealis (Kotir. & Saaren.) Hjortstam & Ryvar-
den and 6 species of Ascomycota: Camarosporidiella
mackenziei Wanas., Bulgakov & K.D. Hyde, C. rob-
iniicola (Wijayaw., Camporesi & K.D. Hyde) Wijayaw.,
Wanas. & K.D. Hyde, Diaporthe caraganae Jacz.,
D. oncostoma (Duby) Fuckel, Keissleriella genistae
(G.Winter) E. Müll., and Stromatonectria caraganae
(Höhn.) Jaklitsch & Voglmayr. Nine of them are sapro-
bic macromycetes, and seven species are phytopatho-
genic macro- and micromycetes.
Half of all species new to Sverdlovsk oblast were
identified in botanical gardens and parks of Ekaterin-
burg, and five species (36%) were identified in forest
belts along highways and agricultural lands. Only two
species (12.5%) were collected in the natural commu-
nities: Keissleriella genistae on Genista tinctoria and
Camarosporidiella caraganicola on Caragana frutex. It
should be mentioned that three identified species of
basidiomycetes, Hypochnicium sp., Perreniporia sp.,
and Phylloporia sp., are probably new to science.
In Sverdlovsk oblast, wood-inhabiting fungi were
detected on 12 WLP species: Amorpha fruticosa, Cara-
gana arborescens, C. decorticans, C. frutex, C. ussurien-
sis, Chamaecytisus ruthenicus, Genista florida, G. tinc-
toria, Laburnum alpinum, Lespedeza bicolor, Maackia
amurensis, and Robinia pseudoacacia (Fig. 3).
Abbreviations: Af, Amorpha fruticosa; Ca, Caragana arborescens; C, C. decorticans; Cf, C. frutex; Cu, C. ussuriensis; Chr, Chamaecytisus
ruthenicus; Gf, Genista florida; Gt, G. tinctoria; La, Laburnum alpinum; Lb, Lespedeza bicolor; Ma, Maackia amurensis; and Rp, Robinia
pseudoacacia.
Cyathus striatus Willd. 1(8:1Ca) 1(18:1Ma)
Dacrymyces lacrymalis (Pers.)
Nees
1(11:1Ca)
Exidia cartilaginea S. Lundell
& Neuhoff
1(17:1Ma)
E. nigricans (With.)
P. R obe rts
2(18:1La;
17:1Rp )
1(26:1Ca) 1(31:1Ca)
Tremella mesenterica
(Schaeff.) Pers.
1(21:1Ca)
ASCOMYCOTA
*Camarosporidiella macken-
ziei Wanas., Bulgakov &
K.D. Hyde
1(17:1Ca) 1(25:1Ca)
*Camarosporidiella robinii-
cola (Wijayaw., Camporesi &
K.D. Hyde) Wijayaw.,
Wanas. & K.D. Hyde
1(17:1Rp)
Camarosporidiella caragani-
cola (Phukhams., Bulgakov &
K.D. Hyde) Phukhams.,
Wanas. & K.D. Hyde
1(25:1Chr)
*Diaporthe caraganae Jacz. 1(17:1Ca) 1(25:1Ca)
*Diaporthe oncostoma (Duby)
Fuckel
1(17:1Rp)
*Keissleriella сf.genistae
(G. Winter) E. Müll.
1(18:1Gt)
Nectria cinnabarina (Tode)
Fr.
2(1:2Ca) 3(8:1Ca; 10:1Ca;
11:1 C a )
6(13:1Ca;
14:1Chr, 1Rp;
17:1Ma; 18:1La,
1Af)
3(19:1Ca;
20:2Ca)
2(25:1Ca;
27:1Cf)
1(35:1Ca)
*Stromatonectria caraganae
(Höhn.) Jaklitsch & Vogl-
mayr
1(15:1Ca)
Xylaria hypoxylon (L.) Grev. 1(20:1Ca)
Species Middle boreal South boreal Ekaterinburg Hemiboreal Forest steppe Steppe
Table 2. (Contd.)
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
FUNGAL DIVERSITY OF NATIVE AND ALIEN WOODY LEGUMINOUS PLANTS 415
Fungi were detected on all species of native WLPs;
on 60% of East Asian; and on 50% of plant species of
European, North American, and Central Asian origin
(Fig. 2).
Most species of wood-inhabiting fungi (115) were
found on the Central Asian species Caragana arbo-
rescens (Fig. 3), which constitutes 84.5% of the total
number of fungal species. This is followed by native
Chamaecytisus ruthenicus (17) and the East Asian
Maackia amurensis (12). No more than ten species of
fungi were collected on the other plants. The range of
the species richness of the mycobiota of alien WLPs
varies from 115 to 1 species, while that of native ones
from 17 to 5 species. Consequently, there is no statisti-
cal difference between the species richness of native
and alien plants: U = 7, z = –1.19, p = 0.26 (Monte
Carlo: p = 0.24). Nevertheless, 122 species of fungi
were identified on 9 alien WLP species, which is four
times more than on 3 native WLP species. At the same
time, the greatest number of fungal species is found on
the alien species C. arborescens, which is more than
4 times greater compared to the richest native sub-
strate (Ch. ruthenicus). This indicates that the intro-
duced plants are a valuable food resource for native
fungi.
Ninety-five out of 136 fungal species (69.8%) were
identified on only one of the WLP species (substrate-
specific). At the same time, 85 species were collected
only on C. arborescens, which amounts to 62.5% of the
total number of fungal species (Fig. 3). Then Chamae-
cytisus ruthenicus follows, on which four specific spe-
cies (2.9%) were collected, three on Maackia amuren-
sis (2.2%), two on Genista tinctoria and Robinia pseu-
doacacia (1.5%) each, and one on Caragana
ussuriensis (0.7%). Single findings were recorded on
27.2% of species (Antrodia albida, Ceraceomyces
microsporus, Fibulomyces mutabilis, Lentaria surculus,
Phanerochaete alnea, Pycnoporellus fulgens, Sanguang-
porus cf. baumii etc.). On the contrary, the fungus Nec-
tria cinnabarina develops on the maximum number of
substrates (on seven WLP species), Xylodon sambuci
on six species, and Peniophora cinerea and Schizophil-
lum commune on four species.
Due to the high level of species richness and the
number of specific species of fungi on Caragana arbo-
rescens, this substrate forms a separate cluster when
compared with other WLP species (Fig. 4). The other
species are grouped into two large clusters. The first
cluster combines native plant species (Caragana frutex,
Chamaecytisus ruthenicus, and Genista tinctoria) and
the largest alien substrate in terms of the aboveground
phytomass (Maackia amurensis), and the second clus-
ter is formed exclusively by alien species with low abo-
veground phytomass (Amorpha fruticosa, Caragana
ussuriensis, Laburnum alpinum, Lespedeza bicolor, and
Robinia pseudoacacia).
Saprobic fungi dominate (112 species/82.3%) in the
general list of species (Table 2), while the number of plant
pathogens is noticeably smaller (26 species/19.1%).
Among the species characterized by obligate or facul-
Fig. 3. Number of fungal species on woody plants species of the family Fabaceae. Native plant species are marked with an asterisk
(*). The total number of fungal species and the number of substrate-specific fungi for each plant species are given.
100
80
140
120
60
40
20
0
Total
115
Number of species
Caragana arborescens
Specific
85
17
43333
12 766 5222 11
*Chamaecytisus ruthenicus
Maackia amurensis
*Caragana frutex
Caragana decorticans
*Genista tinctoria
Genista florida
Caragana ussuriensis
Lespedeza bicolor
Amorpha fruticosa
Robinia pseudoacacia
Laburnum alpinum
416
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
SHIRYAEV et al.
tative pathogenic activity, 18 are species of Basidiomy-
cota (Cerrena unicolor, Chondrostereum purpureum,
Fomitoporia punctata, F. robusta, Fuscoporia contigua,
F. ferruginosa, Ganoderma applanatum, Inonotus hispi-
dus, Irpex lacteus, Laetiporus sulphureus, Oxyporus cor-
ticola, Sanghuangporus cf. baumi, Schizophyllum com-
mune, Stereum hirsutum, S. subtomentosum, Trametes
hirsuta, T. ochracea, and Typhu la mi can s) and 8 are spe-
cies of Ascomycota (Camarosporidiella mackenziei,
C. robiniicola, C. caraganicola, Diaporthe caraganae,
D. oncostoma, Keissleriella genistae, Nectria cinnaba-
rina, and Stromatonectria caraganae).
In the natural conditions of Sverdlovsk oblast
(excluding Ekaterinburg), the total number of fungal
species on WLPs increases linearly in the southern
direction (rs = 0.98, p = 0.02), from the minimum val-
ues in the middle and southern boreal forests (27 and
35 species, respectively) to 69 species in the forest
steppe (Fig. 5). However, the microbiota of Ekaterin-
burg city, located on the border between the southern
boreal and hemiboreal, turns out to be the richest
(72 species) and is comparable to the forest steppe in
terms of the number of species. Nevertheless, taking
into account the urban mycobiota, the linear correla-
tion remains statistically significant (rs = 0.77, p = 0.05).
To study the relationship between the species rich-
ness of the microbiota and biotic and abiotic environ-
mental factors, aphyllophoroid fungi were selected as
the largest group of those developing on WLPs (75% of
the total number of species, Table 2). The Siberian
peashrub was chosen as a model substrate: the largest
number of fungi (84.5%) was collected on it and it is
the most common WLP species in the study region.
Consequently, Siberian pearshrub and aphyllopho-
roid fungi can be considered model groups of plants
and fungi to identify ecological and geographical pat-
terns of the formation of the diversity of the WLP-
associated mycobiota.
3.2. Relationship between the Aboveground Phytomass
of Caragana arborescens and the Diversity
of Aphyllophoroid Fungi along the Meridional Transect
The aboveground phytomass of C. arborescens in
the forest belts of the middle boreal subzone averages
4.6 t/ha (Fig. 6). It increases to 7.2 t/ha towards the
southern boreal subzone, and the maximum level
(10.4 t/ha) was recorded for the forest steppe. The bio-
mass decreases to the level of the middle boreal forests
on the southern edge of the transect, in the steppe. The
average phytomass of Siberian peashrub (4.6 t/ha) in
the urban f lora of Ekaterinburg corresponds to the min-
imum values at the northern and southern edges of the
transect, which is 2.3 times lower than this value in the
forest-steppe zone.
Ninety-three species of aphyllophoroid fungi were
collected on C. arborescens along the studied transect;
60 species were identified in Sverdlovsk oblast, 51 in
Chelyabinsk oblast, and 17 in Kostanay oblast
(Kazakhstan) (Table 2). The minimum number of
species was found on Siberian peashrub in the north of
the transect, in the middle and southern boreal sub-
zones—24 and 30 species, respectively (Table 3).
Forty-seven species were collected in Ekaterinburg,
and the maximum number of species were collected
in the forest steppe (61). The northern mycobiotes
(middle and southern boreal forests) are 1.8–2.6 times
poorer than the southern ones.
Fig. 4. Similarity of the species composition of fungi on ten species of the richest woody leguminous plants (on which three or
more species of fungi were identified).
0 0.4 0.8 1.4 1.60.2 0.6 1.0 1.2
Caragana arborescens
Linkage distance
Maackia amurensis
Chamaecytisus ruthenicus
Caragana frutex
Genista tinctoria
Caragana ussuriensis
Lespedeza bicolor
Amorpha fruticosa
Robinia pseudoacacia
Laburnum alpinum
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
FUNGAL DIVERSITY OF NATIVE AND ALIEN WOODY LEGUMINOUS PLANTS 417
According to the maximum average value of the
Chao1 index calculated for the species accumulation
curve (Fig. 7), the number of species of aphyllopho-
roid fungi on the transect is 101.6 ± 4.1 species, with
93 species collected by us (92% of the potential species
composition). Consequently, the identification of
diversity was sufficient to detect all the most frequent
fungal species along the transect. At the same time,
8 out of 93 species of aphyllophoroid fungi on the
studied transect were found only once (singletons);
therefore, the Turing coefficient is 0.91, or 91%, which
indirectly confirms the assumed species richness of
102 species.
The comparison of lists of fungal species for each
natural and climatic zone/subzone and Ekaterinburg
city along the transect shows that the northern
(boreal) communities of aphyllophoroid fungi on
Siberian peashrub (hereinafter AFSP) differ signifi-
cantly (U= 0, z = –1.94, p = 0.05) when compared to
the southern (steppe) and hemiboreal communities in
Ekaterinburg. The northern mycobiotes form a sepa-
rate cluster in relation to the southern and urban
mycobiota of Ekaterinburg (Fig. 8).
Six species of aphyllophoroid fungi (4.4% of the
total number of AFSP species) develop along the
entire spectrum of natural zonal conditions of the
transect (Table 2). Nevertheless, some of them have
increased abundance in the northern boreal part of the
transect (Steccherinum fimbriatum, Typ hula m i can s )
(Fig. 9) and others, on the contrary, in the southern
steppe part (Neofavolus alveolarius). Some species are
more frequent in the anthropogenic conditions of
Ekaterinburg (Schizophyllum commune), and Irpex
lacteus and Xylodon sambuci are evenly distributed
throughout the latitudinal spectrum. Of the other
groups of fungi, Nectria cinnabarina and Pleurotus pul-
monarius have the widest possible zonal distribution
spectrum.
4. DISCUSSION
Wood-destroying fungi were detected on 12 WLP
species, which is 60% of the total number of species of
woody plants of the family Fabaceae in natural and
anthropogenic conditions of Sverdlovsk oblast. This is
explained by the harshness of the climatic and edaphic
conditions of region for such thermophilic plants.
Despite the long growing time in Ekaterinburg, many
WLP species do not tolerate winter low temperatures,
and the upper part of the plants protruding above the
snow surface freezes annually. For example, this con-
cerns two species of plants, Gleditsia triacanthos and
Gymnocladus dioicus. The studied plants of these spe-
cies in Ekaterinburg are more than 10 years old, but in
2022 their height did not exceed 30 cm; i.e., they actu-
ally exist here as shrubs. Fungi were either not identi-
fied or no more than three species of fungi (Caragana
ussuriensis, Genista florida, or Laburnum alpinum,
Fig. 5. Total number of species of wood-inhabiting fungi on woody leguminous plants in different parts of Sverdlovsk oblast.
70
80
60
50
40
30
20
Middle boreal
27
Number of species
35
72
48
69
Southern boreal Ekaterinburg Hemiboreal Forest steppe
Fig. 6. Aboveground phytomass of Caragana arborescens
along the meridional transect (t/ha). The average level and
95% conf idence interval are given.
12
10
8
6
4
2
Middle
Phytomass, t/ha
Southern Ekaterinburg Hemi-
boreal
Forest Steppe
steppeborealboreal
418
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
SHIRYAEV et al.
Lespedeza bicolor) were detected on other species
(Laburnum anagyroides, L. × watereri, Caragana boi-
sii, C. microphylla, C. spinosa, and C. turkestanica),
represented by single low shrubs (less than 1.5 m in
height), which are regularly pruned in gardens and
parks. The findings of fungi on plants of the genera
Amorpha, Genista, Laburnum, Lespedeza, Maackia,
and Robinia in Sverdlovsk oblast are reported for the
first time.
Thirteen species of fungi new to Sverdlovsk oblast
were identified only on alien WLP species: Cama-
rosporidiella caraganicola, C. mackenziei, C. robinii-
cola, Diaporthe caraganae, D. oncostoma, Fibulomyces
mutabilis, Lachnella alboviolascens, Lentaria surculus,
Phloeomana minutula, Sanghuangporus cf. baumii,
Simocybe haustellaris, Stromatonectria caraganae, and
Xylodon borealis. The following native fungal species
were collected only on alien WLP species: Amphinema
byssoides, Cyathus striatus, Exidia nigricans, Tomentella
botryoides, Typhula micans, Xylodon flaviporus, etc.,
including Antrodiella romellii, Byssocorticium atrovirens,
Pseudotomentella tristis, etc., only on C. arborescens and
Antrodia albida, Exidia cartilaginea, and Mycena
galericulata only on M. amurensis.
Twenty-five species of wood-destroying fungi were
identified on all three native WLP species (C. frutex,
Ch. ruthenicus, and G. tinctoria) in Sverdlovsk oblast,
7 of which were found only on these substrates: Antro-
diella leucoxantha, Athelia epiphylla, Erythricium lae-
tum, Chondrostereum purpureum, and Typhula spathu-
lata, including 2 species which are new for region:
Stromatonectria caraganae and Keissleriella genistae. In
general, six species of pathogens develop on native
WLP species (Camasporidiella caraganicola, Chon-
drostereum purpureum, Keissleriella genistae, Nectria
cinnabarina, Stereum rugosum, and Stromatonectria
caraganae); the other species are saprobic and mycor-
rhiza-forming.
Sixteen fungal species are common for native and
alien WLPs: Cylindrobasidium laeve, Lyomyces crusto-
sus, Nectria cinnabarina, Peniophora cinerea, Pilo-
derma byssinum, Stereum hirsutum, etc. All these spe-
cies are native and widespread in the natural condi-
tions of Sverdlovsk oblast. All saprobic species of fungi
on WLPs are typical wood-inhabiting of native decid-
uous trees.
Pathogens identified for first time on alien plants in
region are of interest. Thus, Sanghuangporus cf. baumii
was collected on Siberian peashrub; it has a central
range in East Asia, where it develops on living and dry-
ing trunks of woody plants of the genera Syringa, Loni-
cea, Viburnum, etc. (Bondartseva and Parmasto, 1986).
This species of fungi was first identified on plants of the
family Fabaceae. The pathogenic micromecetes Dia-
Table 3. Diversity of meridional complexes of aphyllophoroid fungi developing on Caragana arborescens
The arithmetic mean and (±) standard error are given.
Parameters Middle
boreal
Southern
boreal Ekaterinburg Hemi-
boreal
Forest-
steppe Steppe
Number of sites 6 6 6 6 6 6
Area of sites, ha 24.1 25.6 17.2 25.7 33.9 10.7
Average area of sites, ha 4.1 ± 0.9 4.3 ± 0.8 2.9 ± 1.1 4.3 ± 1.2 5.6 ± 1.8 1.8 ± 0.5
Aboveground phytomass, t/ha 4.7 ± 0.8 7.1 ± 1.2 4.6 ± 1.0 8.8 ± 1.2 10.5 ± 1.4 4.5 ± 0.7
Average annual temperature, °С 1.2 1.9 3.5 3.3 3.7 4.1
Average annual precipitation, mm 627 588 565 532 475 353
Aridity index 63.9 60.7 60 56.6 51.2 39.4
Species of fungi 24 30 46 41 60 38
Samples 77 92 156 117 147 115
Average number of species 10.1 ± 1.2 10.5 ± 1.5 11.4 ± 2.6 12.3 ± 1.3 16.8 ± 2.1 8.8 ± 2.0
Average species diversity, species/ha 2.5 ± 0.3 2.5 ± 0.3 4.4 ± 1.4 3.1 ± 0.8 3.0 ± 0.3 5.2 ± 0.9
Shannon index, Н 2.56 2.94 3.45 3.38 3.75 3.24
Whittaker index, βw 0.68 0.91 1.98 1.19 1.55 2.41
Czekanowski–Sørensen index, Cs 0.69 ± 0.07 0.65 ± 0.07 0.34 ± 0.06 0.51 ± 0.09 0.39 ± 0.08 0.30 ± 0.0 4
Number of pathogenic species 3 4 10 6 8 9
Average species diversity of pathogens,
species/ha
0.12 0.19 0.52 0.28 0.31 0.84
Species of saprotrophs/pathogens 4.75 4.3 2.9 4 3.5 2.25
Corticioid/Poroid species 0.91 1.17 0.73 1.29 0.90 0.65
Corticioid/Poroid samples 3.36 2.89 1.92 2.44 1.73 1.07
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
FUNGAL DIVERSITY OF NATIVE AND ALIEN WOODY LEGUMINOUS PLANTS 419
porthe caraganae and Camarosporidiella mackenziei
were collected for the first time on dying branches of
C. arborescens. Camarosporidiella mackenziei was
already reported for Chelyabinsk oblast (Stepanova
and Sirko, 1970) on a similar substrate (Table 4),
which may indicate the expansion of its secondary
range to the north along the Urals. Camarosporidiella
robiniicola and Diaporthe oncostoma, typical pathogens
of Robinia within its natural distribution and second-
ary range, were collected on dying branches of Robinia
pseudoacacia for the first time in region (Michalopou-
los–Skarmoutsos, 1999; Vajna, 2002; Pem et al.,
2021).
Fruitbodies of Lentaria surculus were collected on
the decomposed deadwood of C. arborescens. This is
the second finding of this tropical fungus in the Urals.
Fig. 7. Results of bootstrap analysis used to assess the completeness of identification of species of aphyllophoroid fungi on Cara-
gana arborescens depending on the number of accounting units. The thin line is the average value of the Chao1 index (the expected
number of species) as the number of accounting units increases; the thick line is the individual-based rarefaction curve depending
on the number of accounting units.
100
120
80
60
40
20
0 200 400 600 800 1000 1200
Number of species
Number of accounting units
Fig. 8. Similarity of species composition of meridional complexes of aphyllophoroid fungi developing on Caragana arborescens.
0 0.4 0.8 1.4 1.6 1. 80.2 0.6 1.0 1.2
Linkage distance
Middle boreal
Southern boreal
Ekaterinburg
Hemiboreal
Forest steppe
Steppe
420
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
SHIRYAEV et al.
Previously it was collected in the vicinity of Perm city
in ruderal communities on dead stems of the invasive
Heraculum sosnowskyi Manden. (Shiryaev, 2014).
Consequently, all findings of this species of fungi in
the Urals were made on invasive plants: on the ligni-
fied stems of large herbaceous plants (Heraculum),
which persist for a relatively long time, as well as on a
shrub (Caragana) with heavily destroyed wood. Prob-
ably, on the verge of these two states (lignified stems
and heavily decomposed deciduous wood), there is
the possibility of penetration into the hemiboreal sub-
zone of this tropical fungus.
All 12 species of WLPs develop within Ekaterinburg
city. In the city, 72 species of fungi (52.9% of the total
number of fungal species) are collected, which is the
largest number of all latitudinal divisions represented
on the meridional transect (Fig. 5). Twenty-six species
of fungi were found exclusively in Ekaterinburg, of
which most were collected only on introduced plants:
Antrodia albida, Antrodiella faginea, Ceraceomyces
microsporus, Ceriporia viridans, Gymnopilus junonius,
Megacollybia platyphylla, and Pluteus salicinus, and
there are pathogens among them, Camarosporidiella
robiniicola, Diaporthe oncostoma, and Keissleriella
genistae. All three species of pathogens are new for
Sverdlovsk oblast. Thus, the mycobiota of Ekaterin-
burg turns out to be the richest according to the num-
ber of phytopathogenic fungi species per WLPs (ten
species), This is due to the fact that all introduced
WLPs known in the region with which substrate-spe-
cific pathogens are associated (Camarosporidiella
mackenziei, C. robiniicola, Diaporthe caraganae,
D. oncostoma, and Keissleriella genistae), as well as
some usually saprobic species of fungi that exhibit
pathogenic properties in urban conditions (Cerrena
unicolor, Irpex lacteus, Trametes ochracea, and Typhula
micans), develop in the parks and gardens of the city.
It should be noted that fungi forming mycorrhiza
Fig. 9. Distribution of the number of accounting units (samples) of fungi on Caragana arborescens along the meridional transect:
(a) the northern species Steccherinum fimbriatum, (b) southern species Neofavolus alveolarius, (c) anthropophilic species Schizo-
phyllum commune, and (d) eurybiont species Irpex lacteus.
(a)
Middle boreal
20
15
10
5
0
Southern boreal
Ekaterinburg
Hemiboreal
Forest steppe
Steppe
(b)
Middle boreal
20
15
10
5
0
Southern boreal
Ekaterinburg
Hemiboreal
Forest steppe
Steppe
(c)
Middle boreal
20
15
10
5
0
Southern boreal
Ekaterinburg
Hemiboreal
Forest steppe
Steppe
(d)
Middle boreal
16
12
8
4
0
Southern boreal
Ekaterinburg
Hemiboreal
Forest steppe
Steppe
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
FUNGAL DIVERSITY OF NATIVE AND ALIEN WOODY LEGUMINOUS PLANTS 421
(genera Clavulina, Odontia, Pseudotomentella, and
Tomentella) were not detected on WLPs in Ekaterin-
burg, in contrast to the natural conditions of the
region.
Diversity of aphyllophoroid fungi on Caragana arbo-
rescens along the meridional transect In the Middle
Urals, 115 species of wood-destroying fungi were iden-
tified (93 of which are aphyllophoroid) on Siberian
peashrub. This is four times more than the number of
fungal species known in its natural range, for example,
Altai-Sayan mountain country, where 27 species were
previously found on Siberian peashrub. Such a large
difference is probably explained by the fact that this
species was not the object of targeted mycological
studies in Southern Siberia. The data on the occur-
rence of aphyllophoroid fungi on Caragana arbo-
rescens in the Altai and Salair Ridge are presented in
the following publications (Shvartsman 1964;
Vlasenko, V.A. and Vlasenko, A.V., 2015) and data
from the Sayans are presented in (Beglyanova et al.,
1978; Kotiranta et al., 2017); agaricoid fungi were col-
lected in the Sayans and Altai (Perova and Gorbun-
ova, 2001; Kutafieva and Kosheleva, 2005; Gorbun-
ova, 2015). Pathogenic micromycetes were studied in
Altai and Salair, as well as in the cities of Novosibirsk,
Barnaul, and Krasnoyarsk (Rastitelnor.., 2014; Tomo-
shevich, 2015).
The introduction of the Siberian peashrub along
the eastern macroslope of the Urals, in the areas of the
transect, began in the first decades of the 20th century
(Protasov, 1927; Govorukhin, 1937), but by the pres-
ent, the century-old plantings have not been pre-
served. The currently existing plants were planted
mainly in the 1950s and 1960s, so the age of some
shrubs can reach 60–70 years. However, the Siberian
peashrub has almost disappeared in the forest belts of
the steppe Trans-Urals (Merker, 2009); small planting
areas remain only near settlements. Therefore, the age
of plantings does not affect the latitudinal dynamics of
the species richness of fungi.
The species diversity of phytopathogens is probably
indirectly affected by the age of plants. In the boreal
areas in the north of the transect, due to the large
masses of snow, the beginning of the fall of the aging
trunks of Siberian peashrub occurs at the age of 14–
20 years, whereas in the steppe and forest steppe, it
occurs at the age of 22–28 years. Therefore, the aver-
age age of living parts of plants in the north is lower
than in the south. Consequently, in the north, wood-
inhabiting pathogens do not have enough time to
develop on living wood, and in the south, a quarter of
a century is enough for the development of a rich
pathogenic mycobiota.
In view of the fact that the Siberian peashrub plant-
ings are man-made, their development depends
(mostly) on human activity, and not on abiotic factors
(Table 4). Almost none of the analyzed parameters of
mycobiota are related to the Siberian peashrub phyto-
mass, with the exception of the total and average num-
ber of species of fungi on the plots.
Significantly more mycobiota parameters are due
to abiotic environmental factors. At the northern
edge of the transect, the α diversity (the average
number of species at the site) is extremely low and
increases in the southern direction, reaching a maxi-
mum in the forest steppe (from 10.1 to 16.8), but then
declines sharply to a minimum level (8.8) in the
steppe. A similar trend was observed for the Shannon
index. The index values increase from 2.56 to 3.75
from the middle boreal subzone to the forest-steppe
Table 4. Correlation between the mycobiota parameters and abiotic environmental factors and aboveground phytomass of
Caragana arborescens
* p < 0.05, ** p < 0.01, and *** p < 0.005. The level below 0.05 is not significant.
For transects with Ekaterinburg/without Ekaterinburg.
Parameters Aboveground
phytomass
Average annual
temperature
Average annual
precipitation Aridity index
Number of fungal species 0.61/0.79* 0.78*/0.75 –0.43/–0.51 –0.38/–0.47
Average number of fungal species 0.85*/0.93** 0.29/0.31 –0.01/0.01 0.03/0.05
Species diversity, species/ha –0.48/–0.37 0.76*/0.74 –0.77*/–0.93** –0.72/–0.94*
Shannon index, Н 0.64/0.81 0.79*/0.77 –0.44/–0.51 –0.38/–0.47
Whittaker index, βw 0.25/–0.10 0.88**/0.88** –0.82*/–0.99*** –0.79*/–0.99***
Czekanowski–Sørensen index, Cs –0.09/–0.27 –0.97***/–0.97*** 0.80*/0.9 4** 0.75/0.92*
Number of pathogenic species –0.01/0.23 0.94***/0.97*** –0.73/–0.96** –0.68/–0.94**
Species diversity of pathogens, species/ha –0.39/–0.33 0.78/0.81* –0.84*/–0.95** –0.83*/–0.96**
Number of species of saprotrophs/pathogens 0.26/0.13 –0.87**/–0.86* 0.83*/0.99*** 0.81*/0.99***
Number of species of corticioid/poroid fungi 0.57/0.49 –0.38/–0.33 0.48/0.63 0.47/0.66
Number of samples of corticioid/poroid fungi –0.02/–0.10 –0.94***/–0.95** 0.91**/0.99*** 0.98**/0.99**
422
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
SHIRYAEV et al.
and decrease in the steppe, but not as sharply (3.24)
as the average number of species at the site.
Spatial changes in β diversity are opposite those of
α diversity. For example, the Whittaker index is mini-
mal in the north of the transect (0.68) and reaches a
maximum in the steppe (2.41), as well as in Ekaterin-
burg (1.98). The lowest value of the average similarity
between sites (the Czekanowski–Sørensen index) was
found in the steppe and Ekaterinburg (0.34 and 0.37),
while the maximum value was found in the middle and
southern boreal subzones (0.65 and 0.69, respec-
tively).
Regardless of the method of assessment, β diversity
increases by 2.2–3.7 times in the southern direction.
Some sites in the steppe zone do not have any com-
mon species, which confirms the mosaic character of
the transformation of fungal communities and indi-
cates an increase in the spatial isolation of local popu-
lations. Perhaps, due to the fact that WLPs are south-
ern plants, the main spectrum of fungal species associ-
ated with them also has relatively southern features, in
contrast to the boreal mycobiota. The β diversity of the
urban mycobiota of Ekaterinburg is also 2–3 times
higher compared to natural conditions. The maximum
numbers of fungal species and α diversity (the average
number of species at the sites and the Shannon index)
were found in the ecotone zone, the forest steppe, but
this rich complex is destroyed further south in the
steppe under the effect of harsh arid conditions and a
sharp decrease in the aboveground phytomass of
C. arborescens. This, in turn, is reflected in the maxi-
mum density of fungal species along the entire transect
(2.1 times higher when compared with northern
mycobiotes). Perhaps the increase in β diversity in the
pollution gradient is due to the uneven elimination of
species in space. This is due to the fact that, with an
increase in the pessimality of conditions in the steppe,
instead of the elimination of fungal species, a decrease
in the number of local populations is observed. Even at
the lowest level of phytomass of C. arborescens, some
individuals remain in fragments of habitats where, for
various reasons, conditions may remain relatively
favorable.
A negative correlation was found between the spe-
cies diversity and the average annual precipitation
(r= –0.77 to –0.93, p = 0.01). A correlation was also
found with the average annual temperature and the
Shannon index (r = 0.79, p = 0.04).
For β diversity, a strong negative correlation
between the average annual temperature and the Cze-
kanowski–Sørensen index (r = –0.97, p = 0.01), as
well as a positive correlation with the Whittaker index
(r = 0.88, p = 0.01), was established. An inverse rela-
tionship was revealed with the level of precipitation: a
strong negative correlation with the Whittaker index
(r= –0.82 to –0.99, p = 0.001) and a positive one with
the Czekanowski–Sørensen index (r = 0.80 to 0.94,
p= 0.01).
The data obtained indicate that the number of phy-
topathogenic AFSP species is not related to phytomass
(r = –0.01, p = 0.99), but is strongly positively cor-
related with temperature (r = 0.94, p = 0.004) and
negatively with aridization (r = –0.94, p = 0.01). From
this we can conclude that, due to the ongoing increase
in temperatures and climate aridization in the region
(Report…, 2020), the number of pathogenic AFSP
species will increase.
The number of pathogenic species and the average
species diversity of pathogenic AFSP species increases
with decreasing latitude, reaching a maximum in the
steppe, as well as in Ekaterinburg (Table 3). The ratio
of saprobic and pathogenic AFSP species decreases in
the southern direction (and in Ekaterinburg): in the
north of the transect, in the middle boreal forests, the
number of saprobic species is 4.75 times higher than
the number of pathogenic ones, while in the steppe
this parameter decreases twice. The results are inter-
esting because the minimum phytomass of C. arbo-
rescens was recorded in the steppe and Ekaterinburg.
At the same time, a similar low level of phytomass of
the Siberian peashrub was found in the north of the
transect, although both parameters of mycobiota are
opposite in the harsh conditions of the middle boreal
subzone. It should be noted that the maximum num-
ber of pathogenic species was also identified on the
southern border of the forest zone and in anthropo-
genic conditions, for example, in Novosibirsk city,
where the ratio of saprobic and pathogenic species of
aphyllophoroid fungi on introduced woody plants is
one (Vlasenko, V.A. and Vlasenko, A.V., 2018). A sim-
ilar result was obtained for phytopathogenic micromy-
cetes: saprobics fall out in arid conditions of the city
and parasites remain attached to a moist living sub-
strate (Tomoshevich, 2015).
In the general list of AFSP, the number of poroid
and corticoid fungal species turned out to be similar
(1.0), although corticioid fungi predominate twice in
the natural forests of the Urals (Stepanova, 1971;
Shiryaev et al., 2010, 2012). Moreover, when the study
area decreases in some latitudinal–zonal subdivisions
of the transects (steppe, forest steppe, and Ekaterin-
burg), poroid species prevail in number (Table 4).
Nevertheless, the number of corticioid samples is
higher compared to poroid ones (for example, in the
middle boreal subzone by more than three times), but
this parameter decreases in the southern direction. It
should be noted that the dominance or a similar num-
ber of poroid fungi species compared to corticioid ones
was also found for mycobiotes of other anthropogeni-
cally altered territories, the cities of St. Petersburg, Tyu-
men, and Krasnoyarsk (Arefiev and Kazantseva, 2016;
Zmitrovich et al., 2018; Kryuchkova, 2022). Undoubt-
edly, this phenomenon requires further studies.
Some native species of fungi expand their natural
range due to the development of forest belts with Sibe-
rian peashrub. Two boreal species preferring conifer-
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
FUNGAL DIVERSITY OF NATIVE AND ALIEN WOODY LEGUMINOUS PLANTS 423
ous woody plants, Pycnoporellus fulgens and Daedalea
xantha, were detected only in C. arborescens plantings
in the forest steppe of the Trans-Urals. Both species
are extremely rare on deciduous substrates, and repre-
sentatives of the family Fabaceae were not indicated as
known host plants (Bondartseva, 1998; Ryvarden and
Melo, 2014). Perhaps such substrate specificity of
these species is explained by the severity of the conti-
nental semiarid climate of the forest steppe of the
Trans-Urals and regular grassroots fires in forest belts
of C. arborescens. Antrodiella serpula, Baltazaria galac-
tina, and Xylodon flaviporus, which were previously
known in the forest steppe, hemiboreal and southern
boreal, penetrate in the opposite direction, north of
their natural range (Shiryaev et al., 2010, 2021), and
were first collected in plantings of the Siberian
peashrub in the middle boreal subzone.
Fomitiporia robusta, included in the Red Data Book
of Chelyabinsk oblast (2017) with the status III (rare
species (NT means near-threatened)) and in Sverd-
lovsk oblast (Shiryaev et al., 2010) with the status Crit-
ically Endangered (CR C2a(i); D1), was detected in
steppe and forest-steppe zones and in forest belts with
C. arborescens. Consequently, forest belts with the
Siberian peashrub can be considered territories where
rare and endangered species of fungi are preserved.
Few species of fungi were found in urban parks,
where plants are annually subjected to sanitary prun-
ing (the removal of broken and dead branches) com-
pared to old abandoned plantings. Therefore, we can
recommend trimming broken branches, taking care of
plantings, and removing branches with frost cracks,
because this is the first and easiest way for the penetra-
tion of facultative and obligate pathogens. As the study
showed, many new southern (characteristic of mixed
and broad-leaved forests of Europe) species of fungi
most actively destroying the wood of living plants and
leading to the death of WLPs in forest belts and urban
plantings were identif ied in the southern part of the
transect, from the steppe to Ekaterinburg. The lack of
care for plantings leads to a rapid accumulation of
dead and weakened branches, which, in turn, pro-
motes the development of pathogens and accelerates
the fallout of trees in forest belts. In this regard, it is
still necessary to optimize the methods of pruning tree
shoots in parks of the cities of the Middle Urals. The
thickening of inner city deciduous plantings should
also be avoided.
CONCLUSIONS
Woody leguminous plants (WLPs) are a southern
group of plants in relation to zonal dendrocomplexes
of the Middle Urals and neighboring territories. This is
probably why the maximum values of the species rich-
ness of the wood–inhabiting mycobiota associated
with WLPs were recorded in the southern part of the
transect, in the steppe and forest steppe. Also, the
urban mycobiota of Ekaterinburg is characterized by
one of the highest values of biodiversity, which once
again confirms the increased level of diversity of dif-
ferent groups of living organisms, including fungi, in
anthropogenically altered territories. Botanical gar-
dens and parks of Ekaterinburg, as well as forest belts,
are places of concentration and migration channels of
alien phytopathogenic mycobiota.
In Sverdlovsk oblast, 95% of pathogenic species of
macromycetes detected on WLPs are native species.
Only Sanghuangporus cf. baumii can be considered an
alien species; the center of their range is located in East
Asia. A lower value was found for microscopic fungi,
where 50% of fungal species are specific to alien species
of WLPs and, apparently, were originally introduced
together with their host plants during introduction.
The maximum species richness of phytopatho-
genic fungi on Siberian peashrub was recorded in the
south of the transect, as well as in Ekaterinburg, which
is explained both by climatic conditions and by the
high frequency of mechanical damage to plants. The
common native saprobic species develop on the
wounds of damaged plants, which act as primary
wound pathogens. It many respects, due to such fac-
ultative parasites, the number of fungal species with
pronounced phytopathogenic activity found on alien
woody plants in urban conditions significantly
exceeds the same parameter for the natural habitats
of region.
The hypothesis about the correlation between the
species richness of fungi and the aboveground phyto-
mass of the host plant was partially confirmed (only
for wood-inhabiting basidiomycetes). The peak of the
phytomass of Siberian peashrub was recorded in the
forest steppe along with the maximum of the total
number of species of aphyllophoroid fungi and their α
diversity; however, the β diversity of associated fungi
turned out to be maximal in the steppe, where the phy-
tomass of the considered plant was minimal. We
should also mention the urban mycobiota of Ekaterin-
burg, the parameters of which are different from natu-
ral climatic complexes: one of the lowest levels of phy-
tomass of C. arborescens was detected in the city with
one of the highest levels of α and β diversity of myco-
biota. This study once again confirms the specificity of
the processes of formation of the taxonomic structure
of the mycobiota of urban ecosystems. The tested
hypothesis is not confirmed for wood-inhabiting
ascomycetes due to the fact that, for microscopic
pathogens, the leading factor of species richness is not
phytomass, but the number of plant species.
It should be noted that, despite large-scale studies,
the species composition of some groups of fungi on
WLPs is not fully studied, although, undoubtedly, the
total number of collected species is high and the
entire spectrum of WLPs developing in natural and
anthropogenic conditions of the Middle Urals is
maximally revealed. Further studies are required,
424
CONTEMPORARY PROBLEMS OF ECOLOGY Vol. 16 No. 4 2023
SHIRYAEV et al.
especially of micromycetes, as well as of xylotrophic
agaricoid fungi.
ACKNOWLEDGMENTS
We are grateful to I.V. Petrova, director of the Botanical
Garden of the Ural Branch, Russian Academy of Sciences;
V.V. Valdaiskikh, director of the Botanical Garden at the
Ural Federal University; and P.A. Martyushov, director of
the Vigorov Garden of Medical Cultures for assistance. We
also thank O.S. Shiryaeva for identifying agricoid fungi.
FUNDING
The study was supported by the Russian Science Foun-
dation, project no. 22-26-00228.
COMPLIANCE WITH ETHICAL STANDARDS
Conf lict of interests. The authors declare that they have
no conflict of interest.
Statement on the welfare of animals. All applicable inter-
national, national, and/or institutional guidelines for the
care and use of animals were followed
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