Myxomycetes of the Urals

МИКОЛОГИЯ И ФИТОПАТОЛОГИЯ Том 44 2010 Вып.4

УДК 582.241(234.850)

ФЕФЕЛОВ К.А. МИКСОМИЦЕТЫ УРАЛА

Myxomycetes (plasmodial slime molds, class Myxogasteromycetes) are common inha- bitants in terrestrial ecosystems. They prefer to develop on different decaying plant materials. The first records of myxomycetes species from the Ural Mountains reported by A. A. Yachevskii (1907), where common species, like Lycogala epidendrum, were reported from Perm region. Lateron, A. V. Sirko collected myxomycetes in the 1960—1970 s. Ho- wever, her collections were identified only partly and are not published. The first special studies on myxomycetes have been begun at the end of 20th century. In the Subarctic and Arctic areas of Russia 31 species from the Polar Ural (Novozhilov et al., 1998; Karatygin et al., 1999) are known. The data on myxomycetes of south parts of the Urals include records from Sverdlovsk region (Novozhilov, Fefelov, 2001; Mukhin et al., 2003; Fefelov, 2006; Plotnikov, Fefelov, 2008) and the South Ural (Fefelov, 2003). Besides, there are known investigations on myxomycetes of adjoining territories of the West Siberia: West-Siberian plain (Fefelov, 2002) and the Altay Mountains (Novozhilov, 1986; Barsukova, 2000; Novo- zhilov et al., in press). Our data are based on published records and collections obtained by author in numerous expeditions in the Urals. The field works were carried out during the field seasons of 1996—2007 in all vegetation types and typical habitats of studied region using transect method. Different substrates (decaying logs, litter, fungi, mosses, bark of living trees, dung of herbivorous mammals and birds) were studied to find mature myxomycete sporophores (fruit bodies). Additionally, bark from living trees, litter, dung of herbivorous animals, mosses were collected for moist chamber cultures. Altogether 1854 moist chamber cultures (Hдrkцnen, 1977; Novozhilov et al., 2000) were made in the Institute of Plants and Animals Ecology UrD RAS. A record is defined herein as one or more fruiting bodies of a species that developed from a moist chamber culture or found from one microhabitat (e. g. trunk, log etc.). Nomenclature used herein follows Lado (2001) for myxomycetes, and Gorchakov- skii (1975) for vascular plants and plant communities. Collections of the myxomycetes in- clud about 4000 specimens and deposited in the Institute of Plant and Animal Ecology RAS (SVER) and partly in the Komarov Botanical Institute RAS (LE), Russia. To compare the species communities in different geographical areas, an index of com- munity similarity (CC = Chekanovskii coefficient) was used as proposed by Zaitzev (1984). The formula CC = 2c/(a + b) used to calculate this index is based on the presence or absence of species; a, b — number of species in comparable lists of geographical areas; c — number of common species. Therefore, the value of CC ranges from 0 (the data sets share no common species) to 1 (all species are present in both data sets). As an indicator for species diversity, we used the mean number of species per genus (S/G). Occurrence of a particular species was estimated with a simple percentage scale.

340 We estimated the index of specificity for each territory (SI) to compare the species lists. This index was used only for comparison of two or several territories. The value of SI ranges from 0 (all species of the one territory are found on another) to 1 (no common species for this and other territories). The Urals (Ural Ridge, Ural Mountains) is north-south oriented ridge over 2000 km long, which extended across Russia from tundra of Arctic till steppe of Kazakhstan. The he- ight of some mountains is 1895 m in the north part and 1639 m in the south part of the ridge. The climate is heterogeneous and moderately continental, but with quite specific characters. The Ural Mts are a natural barrier on the way of air masses moving from west to east and therefore they deter the influence of the Atlantic Ocean. Climatic differentiation between the west and east slopes of the ridge is result of deformation of air current. Annual precipitation in the highest places on the north is over 800 mm and about 600 mm on the south. To the west and east this parameter is decrease till 300—400 mm. The driest regions are situa- ted in the eastern parts of study area. The west slopes adjoined with plains have maximum precipitation especially in summer period. Other climatic effect known as «rain shadow» is occurred over the Urals to the east lowlands. Precipitation on this territory has minimum value in all study area, even including hills of the western Urals (Kaigorodov, 1955; Chiki- shev, 1966; Kuvshinova, 1968). Vegetation of the region is rather various. Taiga occupies major portion of the territory and is represented by different plant communities. Dominate tree species include: fir, larch, pine, and spruce. In the South Ural, taiga turns into broad-leaved forests with elm, linden, maple, and oak. Steppe communities occur in the southern areas of the Urals. Tundra and forest-tundra cover large territories in the north (Gorchakovskii, 1975). In geography, the Ural Mts are divided into 5 geographical areas: Polar, Subpolar, North, Middle and the South Ural (Borisevich, 1968). Main study plots and geographical areas are shown on the figure below. Arctic territories of the Ural Mountains (AU) include the Polar (PU) and the Subpo- lar (SPU) Urals. This area begins on Barents Sea coast and stretches till 64°00S N, with maximum elevation in the Subpolar Ural (Narodnaya Mt., 1895 m). Annual average air temperature (AT) is –5—8 °С, AT of July (ATJ) is 5—15 °С, period without frost (PWF) — 67—91 days, annual precipitation (AP) — 400—600 mm, average temperature of soil (ATS) is –1.5 °С. Vegetation is represented by different types of tundra and forest-tundra communities with lichens, mosses, grasses and shrubs. Dominants are Duschekia fruticosa, Betula nana, B. tortuosa, Larix sibirica, L. sukaczewii, Picea obovata and numerous species of Salix. South- ward vegetation is formed by open coniferous forests (canopy coverage is less than 60 %) with spruce (Picea obovata) and birch (Betula pubescens) in the western regions. In the eastern regions dominant trees are larch (Larix sibirica) and pines (Pinus sibirica and P. sil- vestris). North Ural (NU: 64°00S—59°15S N) is characterized by high range of altitude. The highest point is Telpos-Iz Mt (1617 м). Main parameters of the climate are: AT is –1.5 °С, ATJ is 10—20 °С, PWF — 67—91 days, AP — 560—605 mm, ATS is –1.5 °С. The North Ural includes north and middle taiga vegetation subzones. Main tree species in forests of the north taiga is spruce, but along the river banks on the sand soil it is pine. Eastwards there are pine and spruce sometimes spruce-cedar pine forests. Aspen (Populus tremula) form rare riparian forests in the river valleys. In understory lichens and mosses prevail on soil. Westwards, spruce and pine forests prevail whereas eastwards — pine and fir-spruce-cedar pine forests. Secondary birch forests widely distributed in this area together with coniferous forests. In the middle taiga aspen grows out of river valley. Shrubs and tall herbs are well developed. In the highland, mountain tundra, as well as birch and larch open mountain forests dominate. Middle Ural (MU: 59°15S—55°54S N) is low part of the Ural Mts. Elevationis up to 1000 m. Climate: AT is — 0.8 °С, ATJ is — 15—20 °С, PWF — 84—115 days, AP — 460—620 mm, ATS is — 1.5 °С. Heve there are two vegetation zones: middle and south taiga. In the west, fir-spruce forests prevail over pine forests. In the east, there is an opposite situation. Species of birch (B. pu-

341 Studied areas and geographical division of the Ural Mountains. 1 — main study plots, 2 — border of geographical areas, 3 — border of the Ural Mountains. PU — Polar Ural, SPU — Subpolar Ural, NU — North Ural, MU — Middle Ural, SU — South Ural. bescens and, rarely, B. pendula) are presented in the coniferous forests as subdominants. Therefore, some areas are occupied by secondary forests. In the west, broad-leaved trees: linden (Tilia cordata), maple (Acer platanoides), elms (Ulmus laevis, U. scabra) are found in the spruce forests. Understory is good developed and has high species diversity. Vascular plants have particular importance in the overground coverage. Mosses coverage is sicker. South Ural (SU: 55°54S—51°00S N) is divided on two parts: north — mountainous and south — plateau. In this area mountain system is very wide (about 120 km) and represented by chains of mountains including the highest: Yremel Mt (1568 m) and Yamantau Mt (1639 m). Climate: AT is — 1.6 °С, ATJ is — 15—25 °С, PWF — 104—142 days, AP — 400— 570 mm, ATS is — 2.5 °С.

342 In this part of the Ural Mts vegetation is most various. There are four vegetation zones: taiga, broad-leaved forests, forest-steppe and steppe. Broad-leaved forest zone is characterized by mixed broad-leaved-coniferous forests and European broad-leaved forests with oak (Quercus robur), hazel (Corylus avellana) and spindle (Euonymus verrucosa) and others ha- bitants of the south taiga. Forest-steppe includes a scattered forest islands with aspen, birch and pine within the north steppe and meadow vegetation. Aspen, birch, cottonwood (Popu- lus nigra and P. alba), pine and willow (Salix alba) are found in the steppe only in wet places near water. Analogs of tundra and forest-tundra, opened birch and fir-spruce park forests are distributed in highlands of the South Ural.

Results and discussion

Two hundred and twenty-three species representing forty-four genera were identified from the 3947 specimens in the Ural Mts (see table). One hundred ninety eight species are presented from field collections and sixty six from moist chamber cultures. Twenty four species are obtained only with moist chamber cultures. Percent of field and laboratory specimens are presented in the table. For the first time 5 species of myxomycetes are found in Russia: Dictydiaethalium ferrugineum, Licea pygmaea, Diderma nigrum, Didymium comatum, and Stemonitis marjana. Almost 50 % of all specimens are represented by eighteen species and only six species — more than 3 % of total number of specimens (see table): Physarum album, Comatricha nigra, Trichia decipiens, T. varia, Lycogala epidendrum and Arcyria cinerea. According to Stephenson et al. (1993), the species is recognized as «abun- dant» if we should worked in the study plots. However, almost all collections are received in the field on transects. Therefore we estimate the occurrence of species and call them «frequent». All these species widely distributed in the study area. Additionally 12 species have occurrence of 1.39—2.66 %: Arcyria incarnata, Cribraria cancellata, Diderma radiatum, Ec- hinostelium minutum, Enerthenema papillatum, Licea minima, Metatrichia vesparia, Para- diacheopsis fimbriata, Stemonitis axifera, Trichia botrytis, Arcyria pomiformis and Trichia favoginea. All of these species are found in all geographical areas of the Urals excluding of the two last ones, which are not found in the Polar Ural. Probably, this is result of low study activity here, because these species are recorded regularly in other Arctic areas of the world (Novozhilov et al., 1998). Only 30 species are ubiquitous. Fourty seven species are found in three geographical areas. Sixty three — in two and 83 — in one correspondingly. Ten species have discontinuous range (Craterium leucocephalum, Dictidiaetalium plumbeum, Echi- nostelium elachiston, Licea kleistobolus, L. operculata, Lamproderma scintillans, Macbri- deola cornea, Perichaena chrysosperma, Paradiaceopsis solitaria and Stemonitis flavogenita). All of them are found in the South Ural. They appear in the Arctic again and never met in the Middle and North Ural. Species composition of myxomycete assemblages changes gradually by zonal gradient (in the row AU-NU-MU-SU the coefficient of similarity CC is 0.38—0.73—0.64 correspondingly). The Arctic myxomycete biota is more distinctive and specific (CC = 0.35—0.38). The greatest number of species belongs to the next genera: Physarum (34 species), Crib- raria (22), Arcyria (15), Licea (14), Trichia (13), Diderma (11), Comatricha (10), Didymi- um (10) and Stemonitis (10). Species/genus ratio for the entire study area is 5.1, and is comparable with values from other studied regions in temperate zone (Novozhilov, 1999). Fourty seven myxomycete species were registered in the Arctic part of the Urals (see table). Most frequent species are Arcyria cinerea, Echinostelium minutum, Licea parasitica, Physarum album and Trichia munda. Arcyria major, Diderma globosum, Lindbladia tubulina, Licea parasitica and Physarum oblatum were found only there. SI is 0.11. Six species belong to the genera Licea, Physarum and Trichia. Other genera have 1—3 species. Mean of S/G coefficient is 2.0. The specific character of myxomycete biota is an absence of Clastodermataceae which is probably general characteristic of the taxonomical structure of myxomycete biota in Arc- tic and Subarctic (Novozhilov et al., 1998).

343 Myxomycete distribution in geographical areas of the Ural Mountains

Distribution Species NS PS, % fc, % mc, % AU NU MU SU

ECHINOSTELIALES Clastodermataceae Barbeyella minutissima 21 0.53 100 0 — 10 10 1 Clastoderma debaryanum 26 0.66 100 0—224— Echinosteliaceae Echinostelium arboreum 2 0.05 0 100 — — — 2 E. brooksii 5 0.13 0 100 — 4 1 — E. elachiston 7 0.18 0 100—3—4 E. minutum 55 1.39 2 98 10 5 11 29 LICEALES Cribrariaceae Cribraria argillacea 19 0.48 1 0 — 8 9 2 C. atrofusca 12 0.30 100 0 — — 12 — C. aurantiaca 17 0.43 100 0—5111 C. cancellata 86 2.18 100 0 1 31 41 13 C. confusa 1 0.03 100 0—1—— C. costata 1 0.03 100 0—1—— C. exigua 5 0.13 100 0—5—— C. intricata 3 0.08 100 0 — — 3 — C. languescens 2 0.05 100 0—2—— C. macrocarpa 4 0.10 100 0 — 2 2 — C. microcarpa 5 0.13 100 0 — — 4 1 C. minutissima 3 0.08 0 100 — — — 3 C. mirabilis 5 0.13 100 0 — 1 4 — C. montana 1 0.03 100 0—1—— C. oregana 1 0.03 100 0 — — — 1 C. piriformis 4 0.10 100 0 — 1 2 1 C. purpurea 13 0.33 100 0—112— C. rufa 43 1.09 100 0 1 11 30 1 C. splendens 5 0.13 100 0 — 3 2 — C. tenella 31 0.79 400 0—5233 C. violacea 19 0.48 55 45—1126 C. vulgaris 31 0.79 100 0 — 10 20 1 Lindbladia tubulina 2 0.05 100 0 2 — — — Liceaceae Kelleromyxa fimicola 3 0.08 0 100 — — — 3 Licea belmontiana 6 0.15 60 40 3 2 1 — L. biforis 5 0.13 25 75 — — 3 2 L. castanea 1 0.03 0 100—1—— L. chelonoides 9 0.23 100 0 — 3 5 1 L. iridis 1 0.03 0 100 — — — 1 L. kleistobolus 24 0.61 0 100 1 — 11 12 L. minima 67 1.70 59 41 2 18 42 5 L. operculata 7 0.18 0 100 1 — — 6 L. parasitica 10 0.25 0 100 10 — — — L. pusilla 9 0.23 100 0 — 7 2 — L. pygmaea 3 0.08 0 100 — — 3 —

344 Continuation of the table

Distribution Species NS PS, % fc, % mc, % AU NU MU SU

L. tenera 1 0.03 100 0—1—— L. testudinacea 5 0.13 20 80 — 3 1 1 L. variabilis 19 0.48 100 0 1 2 16 Reticulariaceae Dictydiaethalium ferrugineum 1 0.03 100 0 — — 1 — D. plumbeum 2 0.05 100 0 1 — — 1 Lycogala epidendrum 125 3.17 100 0 6 42 65 12 L. exiguum 51 1.29 100 0 — 10 35 6 L. flavofuscum 3 0.08 100 0 — — 3 — Reticularia intermedia 7 0.18 100 0 — 3 4 — R. lycoperdon 7 0.18 100 0 — 2 5 — R. splendens 7 0.18 100 0 — 2 4 1 Tubulifera arachnoidea 34 0.86 100 0 — 11 20 3 T. casparyi 2 0.05 100 0—2—— T. dictyoderma 4 0.10 100 0 — 2 2 — PHYSARALES Didymiaceae Diderma brooksii 2 0.05 100 0—2—— D. floriforme 6 0.15 100 0 — 1 3 2 D. globosum 1 0.03 100 0 1 — — — D. lyallii 1 0.03 100 0—1—— D. montanum 4 0.10 100 0—4—— D. nigrum 1 0.03 100 0—1—— D. ochraceum 1 0.03 100 0—1—— D. radiatum 68 1.72 100 0 2 19 46 1 D. simplex 1 0.03 100 0—1—— D. trevelyanii 1 0.03 100 0 — — — 1 D. umbilicatum 1 0.03 100 0—1—— Didymium anellus 6 0.15 0 100 — — — 6 D. clavus 14 0.35 93 74442 D. comatum 6 0.15 0 100 — — — 6 D. difforme 51 1.29 12 88 — — 5 46 D. dubium 10 0.25 0 100 — — 4 6 D. iridis 8 0.20 13 87 — — 3 5 D. melanospermum 12 0.30 100 0255— D. minus 10 0.25 100 0 — 5 4 1 D. nigripes 19 0.48 31 69 — 2 7 10 D. squamulosum 15 0.38 50 50 — — 1 14 Lepidoderma tigrinum 10 0.25 100 0 — 10 — — Mucilago crustacea 7 0.18 100 0133 Physaraceae Badhamia macrocarpa 6 0.15 100 0 — — 5 1 B. nitens 1 0.03 100 0—1—— B. panicea 3 0.08 100 0 — 1 2 — B. papaveracea 1 0.03 100 0 — — 1 — B. utricularis 14 0.35 100 0 — 2 10 2 Craterium aureum 3 0.08 100 0 — — 3 — C. leucocephalum 10 0.25 100 0 4 — 4 2

345 Continuation of the table

Distribution Species NS PS, % fc, % mc, % AU NU MU SU

Craterium minutum 4 0.10 100 0 — 1 3 — Fuligo laevis 5 0.13 100 0 — — 4 1 F. leviderma 6 0.15 100 0 — — 5 1 F. luteonitens 1 0.03 100 0 — — — 1 F. muscorum 1 0.03 100 0 — — 1 — F. septica 43 1.09 100 0 2 8 30 3 Leocarpus fragilis 26 0.66 86 14 3 16 7 Physarum album 233 5.90 98 2 9 57 152 15 P. auriscalpium 2 0.05 100 0 — — 2 — P. bethelii 3 0.08 67 33 — — 2 1 P. bitectum 1 0.03 100 0 — — — 1 P. bivalve 6 0.15 67 33 2 1 1 2 P. cinereum 17 0.43 71 29 — 2 7 8 P. citrinum 4 0.10 100 0 — — 4 — P. confertum 1 0.03 100 0 — — 1 — P. conglomeratum 3 0.08 100 0 — 2 1 — P. compressum 6 0.15 33 67 — — 6 — P. contextum 2 0.05 100 0—2—— P. decipiens 4 0.10 0 100 — — 4 — P. diderma 5 0.13 100 0 — 2 2 1 P. didermoides 6 0.15 67 33 — 1 5 — P. dubium 1 0.03 100 0 — — 1 — P. famintzinii 2 0.05 100 0—2—— P. flavicomum 17 0.43 100 0 — 6 8 3 P. globuliferum 19 0.48 95 5 1 4 10 4 P. gyrosum 2 0.05 0 100 — — — 2 P. leucopus 36 0.91 100 0—9234 P. leucophaeum 18 0.46 100 0 1 5 10 2 P. notabile 35 0.89 70 30—81512 P. nudum 1 0.03 100 0 — — 1 — P. oblatum 2 0.05 100 0 2 — — — P. psittacinum 16 0.41 100 0—2122 P. pulcherripes 12 0.30 100 0—210— P. pusillum 1 0.03 100 0 — — — 1 P. rubiginosum 1 0.03 100 0—1—— P. serpula 4 0.10 75 25 — — — 4 P. straminipes 14 0.35 100 0 — — 12 2 P. sulphureum 2 0.05 100 0 — — 2 — P. vernum 3 0.08 33 67 — 1 2 — P. virescens 1 0.03 100 0 — — — 1 P. viride 43 1.09 100 0 2 19 20 2 Willkommlangea reticulata 3 0.08 100 0 — — 3 — STEMONITALES Stemonitaceae Collaria arcyrionema 34 0.86 100 0—6226 C. lurida 2 0.05 100 0 — 2 — Colloderma oculatum 7 0.18 100 0—7—— Comatricha alta 2 0.05 100 0 — 1 1 — C. elegans 15 0.38 87 13—411— C. ellae 6 0.15 100 0 1 1 3 1

346 Continuation of the table

Distribution Species NS PS, % fc, % mc, % AU NU MU SU

C. filamentosa 1 0.03 100 0 — — — 1 C. laxa 37 0.94 73 27 1 4 25 7 C. nigra 224 5.68 90 10 6 52 149 17 C. pulchella 41 1.04 97 3 — 23 15 3 C. rigidireta 2 0.05 50 50 — 1 1 — C. tenerrima 4 0.10 100 0 — 1 1 2 C. suksdorfii 1 0.03 100 0 — — 1 — Diachea leucopodia 1 0.03 100 0 — — — 1 Diacheopsis metallica 7 0.18 75 25 2 5 Enerthenema papillatum 97 2.46 75 25 4 25 61 7 Lachnobolus atrus 4 0.10 100 0 — — 4 — Lamproderma arcyrioides 1 0.03 100 0—1—— L. columbinum 21 0.53 100 0 — 20 1 — L. laxum 1 0.03 100 0 — — 1 — L. scintillans 19 0.48 5 95—1—18 Macbrideola cornea 16 0.41 100 0 2 — — 14 Paradiacheopsis cribrata 1 0.03 100 0 — — 1 — P. fimbriata 63 1.60 7 93 6 2 35 20 P. microcarpa 2 0.05 0 100 — — 2 — P. solitaria 5 0.13 0 100—4— 1 Stemonaria irregularis 16 0.41 100 0 — 7 8 1 S. longa 1 0.03 100 0—1—— Stemonitis axifera 77 1.95 98 2 3 31 31 12 S. flavogenita 3 0.08 100 0 — 2 1 S. fusca 45 1.14 100 0 1 11 32 1 S. herbatica 4 0.10 100 0 — 2 2 — S. lignicola 1 0.03 100 0 — — — 1 S. marjana 1 0.03 100 0 — — 1 — S. pallida 25 0.63 100 0 — 11 13 1 S. smithii 12 0.30 100 0 — 4 7 1 S. splendens 2 0.05 100 0—1— 1 S. virginiensis 3 0.08 100 0 — 2 1 — Stemonitopsis aequalis 13 0.33 100 0—112— S. amoena 2 0.05 100 0 — — — 2 S. hyperopta 12 0.30 100 0 — 5 7 — S. microspora 1 0.03 100 0—1—— S. subcaespitosa 1 0.03 0 100 — — — 1 S. typhina 44 1.11 100 0 1 11 21 11 Symphytocarpus flaccidus 3 0.08 100 0 — — — 3 S. herbaticus 1 0.03 100 0 — — — 1 S. impexus 1 0.03 100 0 — — — 1 TRICHIALES Dianemataceae Calomyxa metallica 11 0.28 100 0 2 2 6 1 Trichiaceae Arcyria affinis 5 0.13 100 0 — — — 5 A. minuta 4 0.10 100 0 — 1 2 1 A. cinerea 125 3.17 42 58 14 31 59 21 A. denudata 14 0.35 100 0 — 1 5 8

347 Continuation of the table

Distribution Species NS PS, % fc, % mc, % AU NU MU SU

Arcyria ferruginea 18 0.46 100 0 — 4 14 — A. helvetica 6 0.15 100 0 — 1 4 1 A. incarnata 63 1.60 96 4 2 18 38 5 A. insignis 9 0.23 100 0 — 1 7 1 A. major 1 0.03 100 0 1 — — — A. minuta 1 0.03 100 0 — — 1 — A. obvelata 35 0.89 100 0 — 10 18 7 A. occidentalis 2 0.05 100 0 — — 2 — A. oerstedii 8 0.20 100 0 — 4 3 1 A. pomiformis 105 2.66 87 13 — 24 73 8 A. stipata 12 0.30 100 0 — — 10 2 A. versicolor 2 0.05 100 0 — — 1 1 Arcyodes incarnata 4 0.10 75 25 — — 2 2 Hyporhamma abietina 4 0.10 100 0 — — 3 1 H. calyculata 3 0.08 100 0 — — — 3 H. clavata 49 1.24 100 0 — 11 30 8 H. karstenii 9 0.23 0 100 — — 1 8 H. leiotricha 5 0.13 25 75 — 1 4 — H. minor 7 0.18 0 100 — — 4 3 H. serpula 10 0.25 100 0 — — 2 8 Metatrichia floriformis 6 0.15 100 0 — 2 4 — M. rosea 4 0.10 100 0 — — 4 — M. vesparia 84 2.13 98 2 1 11 46 26 Oligonema fulvum 1 0.03 100 0 — — 1 — Perichaena chrysosperma 18 0.46 0 100 2 — — 16 P. corticalis 17 0.43 88 12 — 2 11 4 P. depressa 4 0.10 75 25 — 3 1 — P. pedata 1 0.03 100 0 — — 1 — P. syncarpon 1 0.03 0 100 — — 1 — P. vermicularis 3 0.08 0 100 — — 2 1 Prototrichia metallica 5 0.13 100 0 — 2 3 — Trichia alpine 2 0.05 100 0 — 1 1 — T. botrytis 84 2.13 87 13 3 24 38 19 T. contorta 33 0.84 100 0 — 8 20 5 T. decipiens 167 4.23 100 0 4 57 82 24 T. erecta 10 0.25 100 0 — 3 7 — T. favoginea 89 2.25 100 0 — 19 56 14 T. flavicoma 8 0.20 100 0 — 4 1 3 T. lutescens 13 0.33 100 0 2 7 2 2 T. munda 14 0.35 0 100 10 4 — — T. scabra 51 1.29 100 0 2 22 21 6 T. subfusca 33 0.84 100 0 — 27 6 — T. varia 151 3.83 100 0 3 56 60 32 T. verrucosa 7 0.18 100 0 — 4 3 — Total specimens 3947 100 — — 146 1049 2068 684 Total genera 44———24343631 Total species 223———47139160124 Note.NS— number of specimens; PS — percent of specimens in the collection; percent of total number of specimens collected in field (fc) and moist chamber culture (mc), correspondingly; AU — Arctic areas of the Urals: Polar and Subpolar (see Materials and methods), NU — North Ural, MU — Middle, SU — South Urals (number of specimens is in cells).

348 One hundred thirty nine species are found in the North Ural. Approximately half of the specimens belongs to 13 frequent and common species in this area (Arcyria cinerea, A. pomiformis, Comatricha nigra, C. pulchella, Cribraria cancellata, Enerthenema papillatum, Lycogala epidendrum, Physarum album, Stemonitis axifera, Trichia botrytis, T. decipiens, T. subfusca and T. varia). Twenty four species are registered only in NU (Badhamia nitens, Cribraria confusa, C. costata, C. exigua, C. languescens, C. montana, Colloderma oculatum, Diderma brooksii, D. lyallii, D. montanum, D. nigrum, D. ochraceum, D. simplex, D. umbilicatum, Licea castanea, L. tenera, Lamproderma arcyrioides, Lepidoderma tigrinum, Physarum contextum, P. famintzinii, P. rubiginosum, Stemonaria longa, Stemonitop- sis microspora and Tubulifera casparyi). The genera Cribraria and Diderma include the most part of species found. SI is 0.17. Species-rich genera are Arcyria (10 species), Cribra- ria (17), Diderma (9), Licea (8), Physarum (18) and Trichia (13). S/G ratio is 4.1. The most diverse myxomycete biota is revealed in the Middle Ural (160 species). Fifte- en of them are frequent and common (50 % of specimens): Arcyria cinerea, A. incarnata, A. pomiformis, Comatricha nigra, Cribraria cancellata, Diderma radiatum, Enerthenema papillatum, Licea minima, Lycogala epidendrum, Metatrichia vesparia, Physarum album, Trichia botrytis, T. decipiens, T. favoginea and T. varia. Thirty species are registered only in MU: Arcyria minuta, A. occidentalis, Badhamia papaveracea, Cribraria atrofusca, C. intricata, Collaria lurida, Comatricha suksdorfii, Craterium aureum, Dictydiaethalium ferrugineum, Fuligo muscorum, Lycogala flavofuscum, Lamproderma laxum, Licea pygmaea, Lachnobolus atrus, Metatrichia rosea, Oligonema fulvum, Paradiacheopsis cribrata, P. microcarpa, Perichaena pedata, P. syncarpon, Physarum auriscalpium, P. citrinum, P. compressum, P. confertum, P. decipiens, P. dubium, P. nudum, P. sulphureum, Stemo- nitis marjana and Willkommlangea reticulata. SI is 0.18. In contrast to NU, Physarum includes the most part of myxomycete species (24). Other dominant genera are Arcyria (14 species), Comatricha (9), Cribraria (15), Didymium (8), Licea (9) and Trichia (12). S/G ratio is 4.4. In the South Ural one hundred twenty four species are found. Half of the specimens belongs to 13 species: Arcyria cinerea, Comatricha nigra, Didymium difforme, D. squamulosum, Echinostelium minutum, Lamproderma scintillans, Metatrichia vesparia, Paradiache- opsis fimbriata, Perichaena chrysosperma, Physarum album, Trichia botrytis, T. decipiens and T. varia. Twenty four species are revealed only in this area: Arcyria affinis, Comatricha filamentosa, Cribraria minutissima, Cribraria oregana, Diachea leucopodia, Diderma trevelyanii, Didymium anellus, D. comatum, Echinostelium arboretum, Fuligo luteonitens, He- mitrichia calyculata, Kelleromyxa fimicola, Licea iridis, Physarum bitectum, Physarum gyrosum, Physarum pusillum, Physarum serpula, Physarum virescens, Stemonitis lignicola, Stemonitopsis amoena, S. subcaespitosa, Symphytocarpus flaccidus, S. herbaticus and S. impexus. Like in the Middle Ural, Physarum species prevail among them, and it is inte- resting that all species of Symphytocarpus are found there. SI is 0.19. The dominants are Ar- cyria (12), Cribraria (11), Licea (7), Didymium (9), Physarum (18), Stemonitis (7) and Tric- hia (8). S/G ratio for this areas is 4.0. The structure of myxomycete assemblage varies in the different geographical regions from north to south. Number of species depends on field work intensity and specimen collecting technique used in the studies. It explains high number of species found in the Middle Ural in comparison with other regions. It is not obvious for the number of genera and S/G ratio. In the Arctic part of the Urals low species diversity as well as SI are not depended only on intensity of specimen collecting, but also from the ecological factors. SI increases from north to south. The average S/G ratio in the Arctic part of regions of the World varies from 1.6 to 2.8 (Novozhilov et al., 1998). Probably, it is explained by diversity of the plant communities and natural conditions in different areas of the Ural Mts. Compositions of the frequent species and rich genera are quite distinctive for each geographical area. Only Arcyria cinerea and Physarum album can be recognized as ubiquitous whereas other frequent species are unique for each area. Some taxa demonstrate quite specific distribution, e. g. Trichia botrytis-«morphotypes». These forms have similar morpholo- gical characters, but their distribution is different in the Ural Mts. T. munda is frequent in

349 the Arctic. T. subfusca is more frequent species than T. botrytis in the North Ural, an opposite situation is registered in the Middle Ural where T. botrytis is dominant among the lignico- lous species in the South Ural. Licea, Physarum and Trichia are the richest genera in all study areas as well as Cribra- ria. The late genus is rare in the Arctic. Occurrence of Cribraria species decreases from north to south. The same pattern of distribution is established for Trichia, whereas Didymi- um is the richest genus in the South Ural. Some genera have specific geographical distribution, e. g. Diderma, Comatricha and Stemonitis associate with North, Middle and the South Urals, correspondingly. Absence of geographical data for some taxa can be only an artifact due to high underestimation in study areas. Every regional myxomycete biota has its own peculiarities in spite of gradually changes in the myxomycete assemblages by zonal gradient.

I acknowledge Dr. Yu. K. Novozhilov and Dr. V. A Mukhin for the discussion and help with identifying some specimens. Appreciation is extended to Dr. A. G. Shiryaev, Dr. I. V. Stavishenko and especially B. S. Plotnikov for collecting specimens of the myxomycetes.

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350 Novozhilov Yu.K.,Fefelov K.A.An annotated cheklist of the Myxomycetes of Sverd- lovsk region, West Siberian lowland, Russia / Mycology and phytopathology. 2001. Vol. 35, N 4. P. 41—52. Novozhilov Yu.K.,Schnittler M.,Stephenson S.The Myxomycetes of Russian Su- barctic and Arctic Areas / Mycology and phytopathology. 1998. Vol. 32, N 1. P. 18—29. Novozhilov Yu.K.,Schnittler M.,Vlasenko A.V.,Fefelov K.A.Myxomycete diversity of the Altay Mountains (southwestern Siberia, Russia) / Mycotaxon. 2010 (in press). Novozhilov Yu.K.,Schnittler M.,Zemlianskaia I.V.,Fefelov K.A.Biodiversi- ty of plasmodial slime moulds (Myxogastria): measurement and interpretation / Protistology. 2000. Vol. 1, N 4. P. 161—178. Plotnikov B.S.,Fefelov K.A.Myxomycetes under the pollution gradient of a copper plant in southern taiga, the Middle Ural / Mycology and phytopathology. 2008. Vol. 43, N 1. P. 33—44 (in Russ.). Stephenson S.L., Kalyanasundaram I., Lakhanpal T.N. A comparative biogeog- raphical study of myxomycetes in the mid-Appalachians of eastern North America and two regions of India / J. of Biogeography. 1993. Vol. 20. P. 645—657. Yachevskii A.A. Mycological flora of the European and Asian Russia. Slime Molds. M., 1907. Vol. 2. 410 p. (in Russ.). Zaitzev G.N.Mathematical statistics in the experimental botany. M.: Nauka, 1984. P. 339 (in Russ.).

Institute of Plants and Animals Ecology UrD RAS Received 4 VIII 2009 Yekaterinburg [email protected]

РЕЗЮМЕ

На территории Уральской горной страны выявлено 223 вида из 44 родов миксомицетов. Приведены сравнительные данные о миксомицетах физико-географических областей региона. Показано, что состав региональных биот миксомицетов изменяется с севера на юг постепенно. Это проявляется как на видовом, так и на родовом уровне. Географические области Урала от- личаются друг от друга по составу доминирующих видов, ведущих родов и по значениям ин- дексов специфичности. Ключевые слова: слизевики, биоразнообразие, география, Уральские горы.

SUMMARY

Two hundred and twenty-three species representing forty-four genera were found in the Ural Mts. Data on myxomycete assemblages from different geographical areas of the studied region are presented. The composition of species and genus components of myxomycete biota changes from North to South gradually. However, every geographical area is characterized by its own composition of frequent species and dominant genera, and index of specificity. Key words: slime molds, biodiversity, geography, Ural Mts.

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