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НАУЧНАЯ СТАТЬЯ УДК 582.24, 470.25, 470.24
РЕДКИЕ ВИДЫ МИКСОМИЦЕТОВ ПОЛИСТОВО-ЛОВАТСКОЙ БОЛОТНОЙ СИСТЕМЫ (ЕВРОПЕЙСКАЯ ЧАСТЬ РОССИИ)
Никита Иванович Борзов, Владимир Иванович Гмошинский
Московский государственный университет имени М.В. Ломоносова, биологический факультет
Автор, ответственный за переписку: Никита Иванович Борзов, borzovnikita@bk.ru
Аннотация. Исследование посвящено описанию редких видов миксомицетов: Craterium muscorum, Dianema harveyi, Didymium bahiense, D. flexuosum, Lampro-derma gulielmae, Valtocarpus trechisporus, найденных на территории Полистово-Ловатской болотной системы. Морфология плодовых тел описана с использованием световой и сканирующей электронной микроскопии, проиллюстрирована микрофотографиями спор и капиллиция. Приведено сравнение с морфологически близкими видами.
Ключевые слова: биоразнообразие, морфология, Полистовский государственный заповедник, Рдейский государственный заповедник, СЭМ
DOI: 10.55959/MSU0027-1403-BB-2024-129-3-43-56
Благодарности. Мы выражаем нашу благодарность сотрудникам межкафедральной лаборатории электронной микроскопии (ILEM) биологического факультета МГУ. Мы благодарны Надежде Ивановне Киреевой за помощь в подготовке иллюстраций к настоящему исследованию.
Финансирование. Полевая работа В. И. Гмошинского финансирована Полистов-ским заповедиком (проект ФОИВ-1-22_66_3), идентификация и изучение образцов проведено в рамках проекта Министерства науки и высшего образования Российской Федерации (№ 075-15-2021-1396).
Для цитирования: Борзов Н.И., Гмошинский В.И. Редкие виды миксомицетов Полистово-Ловатской болотной системы (европейская часть России) // Бюл. МОИП. Отд. биол. 2024. Т. 129. Вып. 3. С. 43-56.
ORIGINAL ARTICLE
RARE SPECIES OF MYXOMYCETES OF POLISTOVO-LOVATSKAYA BOG SYSTEM (EUROPEAN PART OF RUSSIA)
Nikita I. Borzov, Vladimir I. Gmoshinskiy
M.V. Lomonosov Moscow State University, Biological Faculty Corresponding author: Nikita I. Borzov, borzovnikita@bk.ru
Abstract. The study is devoted to the description of rare myxomycete species: Craterium muscorum, Dianema harveyi, Didymium bahiense, D. flexuosum, Lamproderma gulielmae, Valtocarpus trechisporus, found in the Polistovo-Lovatskaya bog system. The morphology of the fruiting bodies is described using light and electron scanning microscopy, and illustrated with microphotographs of spores and capillicium. Comparisons with morphologically similar species are also provided.
© Борзов Н.И., Гмошинский В.И., 2024
Keywords: biodiversity, morphology, Polistovsky Nature reserve, Rdeysky Nature Reserve, SEM
Acknowledgments. We express our gratitude to the staff of the Interdepartmental Electron Microscopy Laboratory (ILEM) of the Faculty of Biology, MSU. We are grateful to Nadezhda I. Kireeva for help in preparing the illustration on the current study.
Financial Support. The field work of V. I. Gmoshinskiy was financed by the Polistovsky Reserve (research project FOIV-1-22-66-3) and the identification and study of specimens was carried out as part of the state task of the Ministry of Science and Higher Education of the Russian Federation. (№ 075-15-2021-1396).
For citation: Borzov N.I., Gmoshinskiy V.I. Rare species of Myxomycetes of Polistovo-Lovatskaya bog system (European part of Russia) // Byul. MOIP. Otd. biol. 2024. T. 129. Vyp. 3. S. 43-56.
Myxomycetes (Myxogastrea) are a class of soil predatory amoeboid organisms that inhabit all ecosystems with plant debris (Stephenson, Schnittler, 2016) and including over 1100 species (Lado, 2005-2023). The main food of myxomycetes are soil bacteria, unicellular algae, fungal spores, and yeast. Thus, myxomycetes regulate the abundance of these organisms, primarily bacteria, as the most numerous representatives of the microbial community.
The Polistovo-Lovatskaya bog system is one of the largest complexes of raised bogs in Europe and the largest system of sphagnum bogs in the Northwest Russia (Vagner, 2006). It occupies an area of 134 thousand hectares and comprises fifteen merged bogs. Most of the territory is open raised bogs, with some forested uplands (so called bog islands). The studied bog area is located in the southern taiga zone. The whole bog system is located in the center of the Priilmenskaya lowland, which is inclined in the north-northeastern direction and in the northern part changes to the Ilmen Lake basin. The system includes raised sphagnum bogs, birch-aspen forests with an admixture of oak and linden, as well as black-alder forests, spruce and pine forests (Korolkova et al., 2020). In the eastern part of the Polistovo-Lovatskaya system lies the Rdeysky Nature Reserve and the western part is occupied by the Polistovsky Nature Reserve, which were organized at the same time on May 25, 1994 (Korolkova et al., 2020).
In 2018, a study of the species diversity of myxomycetes of the Polistovo-Lovatskaya bog system was initiated. Prior to our research, this area was extremely poorly studied - only 45 species were known for the Pskov and Novgorod regions (Jacze-wski,1907; Lebedev, Sinetenkova, 2013). A total
of 60 perennial plots have been established in the bog system, mainly in the southern part of the bog system.
Previously, the first data on myxomycete diversity in the Polistovsky (Gmoshinskiy, Matveev, 2019) and Rdeysky Nature Reserves (Borzov et al., 2021) were published. This paper presents detailed descriptions of rare species recorded in the bog system during the study period.
Materials and methods
Myxomycetes fruiting bodies were studied on the territory of the Polistovo-Lovatskaya bog system in the field and with the use of the moist chamber (MC) method.
Field collection of material
Field collection of sporocarps was carried out according to the standard methodology (Martin, Alexopoulos, 1969). Substrate fragments with myxomycete fruiting bodies were glued to U-shaped paper bases using a crystal-clear glue and placed in matchboxes. In the stationary conditions, the boxes were opened and dried at room temperature without direct sunlight. Samples were frozen for 3-4 days in a refrigerator at -18 °C, redried, and put in zip lock bags before being placed in the herbarium.
Moist chamber method
Different plant substrates were used to set up moist chamber cultures. Samples of 10-20 g were placed in paper bags and dried at room temperature (Matveev et al., 2014). The moist chamber cultures were set up according to the standard technique (Stephenson, 1985). Petri dishes 9 cm in diameter were used, with filter paper and substrate fragments
placed at the bottom. The substrates were poured with distilled water. The pH was measured one day after the experiment start using an Aquilon pH-420 pH-meter. Chambers were checked once a week using an MBS-10 stereomicroscope. The observed specimens were collected and placed in a herbarium as those collected in the field. The duration of moist chamber experiment was three months.
Morphological study
The morphology of sporocarps was studied using an MBS-10 and a Micromed MC2 Zoom microscopes. Photographs were obtained with a Micromed 3 var. 3LED M and the E3CMOS06300
camera. A series of photographs from different focal planes were combined using Helicon Focus ver. 6.0.18. The sporocarps structures were measured using ToupView 3.7 and ImageJ ver. 1.52a. Photographs of spores and capillitium were also obtained using a Jeol JSM-6380 LA (Jeol, Tokyo, Japan) and a Quattro S (Thermo Fisher Scientific, Waltham, MA, USA) scanning electron microscopes in the Interdepartmental Laboratory of Electron Microscopy (ILEM) at the Faculty of Biology of Moscow State University. The specimens were prepared using Au-Pd alloy, without additional critical point drying.
Both classical and modern identification keys (Martin, Alexopoulos, 1969; Poulain et al, 2011;
Fig. 1. Craterium muscorum (MYX 12433). A, B - sporangia, C - columella, D, E - spores in light microscope and in SEM (Scanning Electron Microscope) respectively. Scale bars: A - 0.5 mm, B - 1 mm, C - 100 ^m, D - 20 ^m, E - 4 ^m
Gmoshinskiy et al, 2021) were used to identify morphospecies.
The specimens were deposited in the collection of the Department of Mycology and Algology, Faculty of Biology, Lomonosov Moscow State University (MYX).
Results
Craterium muscorum Ing
Material examined: MYX 12433. Russia, Pskov Region, Bezhanitskii District, Polistovsky State Nature Reserve, eco-trail "Plavnitskoe bog", Eloviy Island (N 57.104010, E 30.371710). Broadleaved forest dominated by Alnus glutinosa (L.) Gaertn and Betula pendula Roth. On deciduous wood, leg. Gmoshinskiy V.I., Matveev A.V., Borzov N.I., 30.08.2019, det. Gmoshinskiy V.I.
Description: sporocarps subglobose, oval, or obovate stalked sporangia, clustered, 1 mm high, 0.5-0.7 mm in diameter, purplish-brown, dull red (Fig. 1). The peridium is light brown to almost grey or pinkish in the upper part, dark reddish-brown, almost black in the lower part, dehiscing at the top and remaining as an irregular calyx at the base of sporotheca, occupying 1/2 to 3/4 of its diameter. The hypothallus is membranous, disc-shaped, light red, often with inclusions of lime. The stalk is rough, black, slightly wrinkled, equal to or slightly larger than diameter of sporotheca, filled with light lime granules visible on break. The columella is dark, cylindrical, 1/3 to 1/2 in diameter of sporo-theca, gradually thinning in the upper part. Capil-litium well-developed, badhamioid, represented by a network of tubes filled with whitish or slightly pinkish lime granules, abundantly branched and anastomosed. The spores are black in mass; purplish-brown in transmitted light, globose, with a light spot occupying about a half of diameter, (12.2) 12.9-16.1 (16.5) ^m in diameter (Mean = 14.28, SD = 0.92, n = 100), ornamented with large warts united in irregular lines and meshes. SEM shows that the ornamentation is represented by a large irregular net with 4-7 meshes per visible part of the spore, formed by ribs 0.3-0.6 ^m high topped by spherical outgrowths (cristate reticulate type by Rammeloo, 1975).
Notes: our specimen was represented by sporangia that were not fully formed yet. For the same reason, there are no sporangia with a ruptured peridium in the sample, but in the most formed ones, it is noticeable that the peridium is very thin and membranous in the upper part of the sporangium. Despite the changes in peridium colour, the process of spore morphogenesis is complete, and their
size and incomplete coarse-meshed ornamentation is consistent with the range given for this species in most publications (Ing, 1982; 1999, Lado et al., 2018).
The morphologically closest species is Dia-chea dictyospora (Rostaf.) J.M. Garcia-Martin, J.C. Zamora & Lado, which is distinguished by its small, obpyriform stalked sporangia (0.5-0.7 mm in diameter and up to 1 mm in height), the presense of lime in the stalk and peridium structures, a cylindrical and elongated columella (Zhao et al., 2018). Spores in C. dictyospora with a small meshed (broken) reticulation, warts low, many comfluents in rows, not forming a border (Nannenga-Breme-kamp, 2022). D. muscorum is very similar in appearance to D. obovata (Peck) J.M. GarciaMartin, J.C. Zamora & Lado, which is distinguished by its fine spinulose spore ornamentation (Poulain et al., 2011). Craterium retisporum G. Moreno, D.W. Mitch. & S.L. Stephenson also has spores with a warty-meshed ornamentation, but is characterized by smaller white or light brown conical sporangia (0.2-0.5 mm diameter), a flattened sporangial cap with a well-defined cleavage line (Moreno et al., 2009).
The correct identification of this sample was confirmed by comparing the nucleotide sequences of the genes 18S rDNA, EF1a and mt SSU with sample MM28650 collected in France (Novozhilov et al., 2023). This is the first find of this C. musco-rum for Russia.
Dianema harveyi Rex
Material studied: MYX 12466. Russia, Pskov Region, Bezhanitskii District, vicinity of Polistovsky Nature Reserve, Tereshikha tract (N 57.048006, E 30.334258). Broadleaf forest with Alnus glutinosa, Betula pendula, Picea abies (L.) H.Karst, Populus tremula L. On deciduous wood, leg. Gmoshinskiy V.I., Matveev A.V., Borzov N.I., 31.08.2019, det. Gmoshinskiy V.I.
Description: sporophores are sessile sporangia or flattened plasmodiocarps, pinkish-brown, ochra-ceous, iridescent, 0.5-2 mm diameter, about 0.3 mm high, scattered along the substrate (Fig. 2). The peridium is thin, membranous, single, smooth or slightly rough, light yellow to almost transparent in transmitted light, without deposits of granular material; the inner surface is covered with small warts. Dehiscence of the peridium is irregular. Hypothallus is inconspicuous. Stalk and columella absent. Capillitium attached to both the base and inner peridium, consisting of thin, slightly twisted filaments that branch only at the ends. Us-
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Fig. 2. Dianema harveyi (MYX 12466). A - group of sporangia on the wood. B - spores in transmitted light. C, E, F - warted capillitium in SEM. D - warted spore in SEM. Scale bars: A - 1 mm, B - 20 ^m, C - 10 ^m, D-F- 2 ^m
ing SEM, it can be seen that the capillitium filaments are covered with warts on one side of the filament, while the other half is smooth. Spores dirty yellow in mass; light yellow in transmitted light, globose, with uniformly thickened sheath, (9.0)9.5-11.4 (12.2) (mean = 10.61, SD = 0.68, n = 30), uniformly ornamented with small warts. SEM shows that cylindrical, almost not expanded in upper part warts (baculae by Rammeloo, 1974) reach 0.4 pm in height and some of them may merge with each other to form short lines.
Notes: The most characteristic feature of Diane-ma harveyi is the presence of only slightly flattened sporangia, not clustered spores, ornamented with small warts and not nivicolous habitat (Poulain et al., 2011). D. harveyi var. verrucatum Y.Yamam. is distinguished by the presence of well-defined warts on the surface of the peridium and spores (Yamamo-
to, 2006). In contrast to most descriptions (Martin, Alexopoulos, 1969; Ing, 1999), the sporocarps in our specimens were rather scattered on the substrate surface, did not form dense clusters and did not form plasmodiocarps. However, the other characters were fully consistent with the descriptions (Rex, 1891). Among the species observed in non-nival conditions, Dianema corticatum Lister and D. andersonii Morgan in Macbride are the closest. The former is distinguished by the presence of somewhat larger spores (ellipsoidal, 10-15x8-10 pm) clustered together and the brown colouration of the peridium (Poulain, 2011; Kuhnt, 2014). D. andersonii is characterized by globose sporangia on a narrowed base and larger spores (10-12 pm diameter) ornamented by more sparsely arranged, smaller warts (Moreno et al., 2011).
D. harveyi is quite rare in Russia. It is probably sometimes overlooked during the fieldwork.
It has been registered only in the Altai Republic (Vlasenko et al., 2016), Murmansk (Novozhilov et al., 2020) and Moscow Region (Gmoshinskiy, Kireeva, 2022).
Didymium bahiense Gottsb.
Material studied: MYX 14130. Russia, Pskov Region, Bezhanitsky District, Polistovsky State Nature Reserve, Slepetnoe island (N 57.084458, E 30.513806). Mixed forest dominated by Alnus glutinosa and Betula pendula, Pinus sylvestris L. On the dead bark of deciduous tree, leg. Gmoshinskiy V. I., Matveev A. V., Borzov N. I., 03.09.2019, det. Gmoshinskiy V. I.
Description: sporophores are sporangia on long stalks, globose or somewhat flattened, deep umbil-icate below, 0.2-0.7 mm in diameter forming colonies of tightly adpressed sporangia, but without any loss of individuality, white (Fig. 3). The perid-ium is thin, membranous, practically colourless in
transmitted light, except for the basal part, where it is lightly ochre, evenly coloured, without any dark spots, uniformly and densely covered by numerous star-shaped lime crystals, which are similar in size to the diameter of spores. The peridium ruptures at the top into irregular petal-like lobes. The hy-pothallus is thin, membranous, blackish-brown, individual or common for several adjacent sporangia. The stalk is long, 0.7-0.9 mm long, longitudinally-wrinkled, limeless, almost black at base, to yellowish-brown or straw-brown in upper part. The pseudocolumella is saddle-shaped, flattened, white, calcareous, with rough surface. The capillitium is abundantly developed, composed of light brown, almost colourless filaments, poorly branched and slightly twisted, sometimes with nodose endings, attached to the pseudocolumella and peridium, resulting in laminae of peridium remaining attached to capillitium filaments after dehiscences. Spores dark brown in mass; purple-brown
Fig. 3. Didymium bahiense (MYX 14130). A-C - sporangia with saddle flattened pseudocolumella, light microscopy. D - sporangium in SEM. E, F - spores in SEM and in light microscope respectively. H - spores with capillitium in SEM. I - Peridium with lime crystals. Scale bars: A - 1 mm, B, C - 0,5 mm, D - 400 ^m, E - 2 ^m, F, H - 20 ^m.
I - 10 ^m
Fig. 4. Didymiumflexuosum (MYX 22537). A-C, E - plasmodiocarps, arrows on B and C show columella. D, J - spores with vesicular spore-like bodies in transmitted light. Scale bars: A - 1 mm, B, C, E - 0,5 mm, F - 10 ^m, G - 30 ^m, H - 50 ^m, J - 100 ^m
in transmitted light, globose, with uniformly thickened threads, 8.9-10.8 цт diameter (Mean = 9.72, SD = 0.48, n = 48), ornamented with small warts forming 3-4 groups per visible part of spore. Under SEM, spores with flattened tops sometimes united in short, curved lines (pilate type by Rammeloo, 1974).
Notes: This species is rather difficult to identify and it belongs to the Didymium iridis-complex. Its main distinguishing feature is the saddle-shaped flattened white pseudocolumella (Stephenson, 2021). Many authoritative authors have suggested
that D. bahiense is only a form of D. iridis with an irregularly developed columella (Clark, Mires, 1999; Clark, Haskins, 2018). Others, in contrast, recognise the division of D. iridis-complex into: D. verrucisporum A. L. Welden, D. canariense Ing, and D. crassicolle Ing. D. verrucosporum is characterized by a white club-shaped columella (Martin, Alexopoulos, 1969), D. canariense Ing has a thin, disc-shaped columella at the base of the sporangium, yellow in colour (Ing, 2000). D. crassi-colle is characterized by the complete absence of a columella (Ing, 1982). D. megalosporum Berk.
& M. A. Curtis in Berkeley, is characterized by a yellowish-brown flattened columella (Stephenson, 2021) and also by a compressed-spherical or almost disc-shaped sporangia (Martin, Alexopoulos, 1969; Clark, Haskins, 2018). In addition, D. ni-gripes (Link) Fr is similar in external morphology, distinguished by its darkly coloured club-shaped columella and dark brown to almost black sporangia stems (Martin, Alexopoulos, 1969).
In Russia, this species has been recorded in the European part: in the Astrakhan Region on bark and leaf litter (Novozhilov et al., 2003), in Moscow, on live Chamaerops humilis L. bark in the greenhouses of the Aptekarsky Ogorod Botanical Garden of MSU (Matveev, Gmoshinskiy, 2022), and in the Far East (Novozhilov et al., 2017). However, it is likely to be quite often confused with closely related species from the D. iridis-complex. A thorough revision of the collections will therefore provide more information on its distribution.
Didymium flexuosum Yamash.
Material studied: MYX 22537. Russia, Pskov Region, Bezhanitsky District, Polistovsky State Nature Reserve, eco-trail "Plavnitskoe bog", Eloviy Island. Spruce (N 57.101996, E 30.387909). Forest with Picea abies, Betula pendula, Populus tremula. On the bark of living Picea abies (in moist chambers). pH 4.39, leg. Borzov N.I., 04.05.2022, det. Gmoshinskiy V.I.
Description: sporophores are short, laterally compressed plasmodiocarps, slightly curved, almost not branching, scattered over substrate, white or dark brown when lacking lime, 0.2-0.4 mm in diameter (Fig. 4). The hypothallus is thin, membranous, almost transparent, covered with small amount of lime granules. The peridium is membranous, practically transparent, covered with few angular lime crystals. The columella is thin, resembling a thin septum, running along the entire plas-modiocarp, slightly widened and attached to the base and very thin in the upper part, not attached to the inner surface of the peridium. The capillitium is represented by brown filaments branching from the columella and base of the sporangium, 1-2 pm diameter, dichotomously branching and interspersed with globular vesicles 15-25 pm diameter, which are colored and ornamented similarly to the spores. The spores are black in mass; light purplish-brown in transmitted light, globose, with uniformly thickened sheath, (9.8) 9.9-12.6 (14.3) pm diameter (Mean = 11.62, SD = 0.85; n = 100), ornamented with spines and irregular meshing. SEM shows that
the surface of the spore is covered with individual cells formed by merging warts, resulting in a network with non-resembling arched bridges. The rest of the spore is covered with irregular slightly elongated warts (by Rammeloo, 1975). The plasmodium is a white phaneroplasmodium.
Notes: The most characteristic features of this species are the laterally flattened, curved plas-modicarps, with a thin, elongate columella running along the entire plasmodiocarp, and the presence of spore-like bodies in the capillitium structures and the characteristic irregular, warty-mesh ornamentation of the spores.
Different sources contain various photographs of spores of this species, obtained using scanning electron microscopy. The ornamentation of the spores of our specimens is fully consistent with that given by Neubert et al. (1995, p. 341), but not Sala-zar-Márquez et al. (2013, fig. 5, 6), where they are ornamented by large individual warts with expanded apex, which only occasionally merge into small groups to form a network-like pattern. However, the appearance of plasmodiocarps and the structure of the columella appear similar to our specimens.
This is rare species in Russia, previous findings were in Volgograd region, Moscow region and Tver region (Novozhilov et al., 2003; Gmoshinskiy, 2013; Matveev et al., 2018). Our samples are fully correspond to the original description (Martin, Brooks, 1938). From the bark of living Amelanchi-er canadensis (L.) Medik. collected in the botanical garden of Tver State University, a morphologically similar plasmodiocarp (MYX 6711) was obtained by the moist chamber method, which had all typical characters, except for the presence of spore-like bodies in the capillitium (Matveev et al., 2018). Having examined both specimens, we concluded that they undoubtedly belong to the same mor-phospecies. It is possible that in some cases sporelike bodies do not develop in capillitium structures. A similar structure of capillitium devoid of sporelike bodies is presented in pl. 118 in Emoto, 1977.
Lamproderma gulielmae Meyl.
Material studied: MYX 12338. Russia, Pskov Region, Bezhanitsky District, Polistovsky State Nature Reserve, southern part of the eco-trail "Plavnitskoe bog" (N 57.098263, E 30.372753). A small area of deciduous forest with dominant Betu-la pendula, Populus tremula. On a deciduous wood and leaf litter. leg. Gmoshinskiy V.I., Matveev A.V., Borzov N.I., 30.08.2019, det. Gmoshinskiy V.I. MYX 14099. Russia, Pskov Region, Bezhan-
Fig. 5. Lamprodermagulielmae (MYX 12338). A, C - sporangia. B - spores and capillitium in transmitted light. D — sporangium with peridium plates, SEM. E-G spores with capillitium in SEM. Scale bars: A - 1mm, B - 20, C - 0,5 mm, D — 300 ^m, E - 2 ^m, F - 4 ^m,
G - 20 ^m
itskii District, Polistovsky State Nature Reserve, Slepetnoe island (N 57.083305, E 30.512137). Secondary dead fir forest. Mixed forest with Bet-ula pendula, Picea abies, Populus tremula. On coniferous wood, leg. Gmoshinskiy V. I., Matveev A. V., Borzov N. I., 03.09.2019. det. Gmoshinskiy V.I Description: sporophores are long stalked sporangia, 1-2 mm in total height, the sporotheca is globose
or slightly elongated, 0.3-0.5 mm in diameter, gray-blue (Fig. 5 A, C). The peridium is membranous, shiny, bluish-gray, covered with large dark speckles, practically colourless in transmitted light and brown in places corresponding to speckles. Dehis-cence occurs into individual plates corresponding to the dark spots. The peridium is smooth, with dense and dark areas with capillitium endings attached to
them, and thinner between them. Along these thin areas its destruction occurs, which is clearly visible with SEM. The hypothallus is reddish-brown, membranous, shared with the adjacent sporangia. The stalk is long, 2-3 times larger than the diameter of the sporotheca, thin, smooth, black. The columella is an extension of stalk, black, that does not usually reach the middle of sporangium, abruptly terminates and transforms into a network of capillitium. The capillitium separates from the columella along its entire length, filaments thin, about 1-2 pm diameter, with few spindle-shaped thickenings up to 3 pm diameter, branching and anastomosing, with long loose ends, light brown, almost colorless at periphery. Spores dark brown in mass; light purplish brown in transmitted light, globose with light spot, (11.4) 11.8-15.3 (15.7) (mean = 13.86, SD = 0.92, n = 98), ornamented by individual warts evenly scattered on spore surface. Using SEM, it can be seen that the spores are covered with large pila on expanded bases, with only slightly developed caput, scattered over the spore surface and very rarely aggregated in small groups (pilate type by Rammeloo, 1975). Secondary ornamentation, represented by scattered globose warts, is visible between the warts.
Notes: this species can be distinguished from other species of the genus Lamproderma with a mottled peridium due to the sporangia on very long, dark stalks and spores ornamented with small wart-hogs. Among the non-nival species characterised by the mottled peridium, L. puncticulatum Hark. and L. griseum K.S. Thind & T.N. Lakh. The former is distinguished by its shorter sporangia stems, which usually do not exceed its diameter (Poulain et al., 2011), and by the fact that sporangia are most often formed on the surface of mosses and liverworts (Schnittler et al., 2010), while in L. guliel-mae the stems are usually several times the length of the sporotheca diameter and sporangia are usually formed on leaf litter and rotten wood, which may sometimes be covered by moss. L. griseum is distinguished by its smaller spores (11-12.5 pm) and the absence of bluish tones in peridium coloration (Thind, Lakhanpal, 1968).
Compared to the data of Lizarraga et al. (2005, figs. 7 and 8), the spores of our specimens are ornamented with more irregularly scattered warts on the surface, which sometimes merge, and the tops of the warts are more extended. In addition, secondary ornamentation, formed by small warts, is stronger on the spore surface. In contrast, compared to the spore ornamentation given in Yatsiuk et al. (2017), the warts in our samples were more evenly and densely arranged.
Some authors indicate that L. guelmae forms fruiting bodies only in mountains (Martin, Alexo-poulos, 1969), while others directly indicate that this species belongs to the nival category (Kowalski, 1970, Ing, 1999). However, in contrast, modern publications indicate that this species is not nival (Poulain et al., 2011; Yatsiuk et al., 2017). In this situation, we are dealing either with morphologically similar species with similar sporophore morphology or with a species with broad ecological plasticity.
In Russia, Lamproderma gulielmae was previously found only in the north-west of the European part: in the Republic of Karelia and the Tver region bordering the study area (Bortnikov et al., 2020).
Valtocarpus trechisporus (Berk. ex Torrend) Gmoshinskiy, Prikhodko, Bortnikov, Shchepin & Novozh.
Material studied: MYX 17497. Russia, Pskov Region, Bezhanitsky District, Polistovsky State Reserve, Osinovy Island. Aspen, central part (N 57.09948, E 30.58180). Alnus glutinosa, Betula pendula, most of the logs are under water. On moss, leg. Gmoshinskiy V.I., Kireeva N.I., 12.07.2021. det. Gmoshinskiy V.I. MYX 18061. Russia, Pskov Region, Bezhanitsky District, Polistovsky State Nature Reserve. Ecotrope "Plavnitskoye bog", at the edge of the Spruce Island (N 57.104007, E 30.371716). Alnus glutinosa, Betula pendula. On Sphagnum spp., leg. Gmoshinskiy V.I., Kireeva N.I., 19.07.2021. det. Gmoshinskiy V.I.
Description: sporophores are presented by large, irregularly shaped pseudoethalia, consisting of deformed sporangia densely pressed to one another, united in a single mass at base and relatively free at the top, 3-7 mm high, up to 10 cm wide, dark brown (Fig. 6). The peridium is completely destroyed during the sporangium maturation. The hypothallus inconspicuous, thin, membranous. The sporangia stalks are virtually not expressed. The columella is opaque in transmitted light, black, smooth, usually curved, does not reach apex of sporangium, may branch. The capillitium is separated from the colu-mella along its entire length; represented by coarse threads, branches off from the column throughout its entire length, each thread branch several times, become thinner, and form loose ends at the periphery of the sporangium. Surface capillitium net is absent. Spores dark brown in mass; purple-brown in transmitted light, globose or elliptic, fine-meshed (9.5) 9.7-12.2 (12.5) pm (mean = 11.09, SD = 0.74, n = 107). SEM shows that the spore ornamentation is of the simple reticulate type (Rammeloo, 1974);
Fig. 6. Valtocarpus trechisporus (A-B - MYX18061; C-F - MYX 18560). A, B - pseudoethalia in nature. C - spores and capillitium in transmitted light. D, E - capillitium in SEM. F - reticulated spore in SEM. Scale bars: C - 20 цт, D, E - l0 цт, F - 3 цт
net presented by continuous vertical ridges, without perforations, up to 0.5 p,m. The plasmodium is white, large, turning reddish brown before sporo-phore formation and then black.
Notes: This is a rather distinctive species, forming extensive sporophores in waterlogged forests on the margins of sphagnum bogs. Its most important differences are the presence of fine spores and the dark brown coloration of the sporangia. The closest species, Symphytocarpus amaurochaetoides Nann.-Bremek. has black sporangia and smaller spores (7.5-8.5 (9) p,m in diameter), which are also ornamented by a fine-meshed net of warts with connected by expanded tops. S. cristatus Nann.-Bremek. is characterized by the presence of spores ornamented with warts, which are collected in lines but do not form regular net. Another closely related species, Valtocarpus megaloplegmus Gmoshinskiy, Prikhodko, Bortnikov, Shchepin & Novozh. possesses coarse-meshed spores, with 10-12 cells
per visible part and a brown coloured spore mass. This species was recorded in the Rdeisky Reserve in 2020 (Borzov et al., 2021).
In Russia, this species has been previously recorded only in the Republic of Karelia (Novozhilov, 2005) and Tatarstan (Zemlyanskaya, Novozhilov, 2022).
Conclusion
Long-term studies in the Polistovo-Lovatskaya bog system have revealed several rare myxomycetes species new to Russia. But some of them are not rare in the traditional sense of the word. It is only when abiotic and biotic factors are combined that this species forms sporophores.
In this case, these species are formed everywhere. Valtocarpus trechisporus is the most striking example. Mass development of sporophores of this species was observed not only in the Polis-tovsky Reserve, but also on a considerable territory
of the European part of Russia. We also obtained specimens from the Central Forest State Biosphere Reserve, the Moscow (Zvenigorod Biological Station and Sima swamp) and Vladimir Regions (Su-dogodsky district). In all cases, the finds were made in July-August 2020, in similar biocenoses (mixed forest with birch, spruce, aspen and, sometimes, alder, with dense moss cover on the soil formed by species of the genera Sphagnum and Polytrichum). At the same time, sporophores were so abundant and large that their projective coverage on some test plots in the Polistovsky Reserve reached 3-5%. Thus, with such a massive development, this species simply cannot be missed during work in the field conditions. Consequently, some combination of environmental factors was observed in 2020 that led to such mass sporophore formation. However, such a phenomenon is not observed every year. This species may be a good illustration of the concept of latent diversity, according to which it is ar-
gued that the real number of myxomycete species present in an ecosystem is much higher than can be established by traditional: collection of material in the field or the moist chamber method (Shchepin et al, 2019). It appears that many extremely numerous species under natural conditions never, or very rarely, form sporophores and, as a result, cannot be accounted for by traditional methods of species diversity research.
Some species have phenological characteristics of sporophore formation (Gmoshinskiy, Matveev, 2016). Major peaks of sporulation can be defined for individual species. Thus, they can be observed only in a certain period of time.
Therefore, in order to carry out comprehensive studies of species diversity and to determine the degree of "rarity" of species, multi-year field studies in a variety of habitats are required, which should be carried out during the snow-free period.
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Информация об авторах
Никита Иванович Борзов - лаборант кафедры микологии и альгологии биологического факультета Московского государственного университета имени М.В. Ломоносова (borzovnikita@bk.ru);
Владимир Иванович Гмошинский - ст. препод. кафедры микологии и альгологии биологического факультета Московского государственного университета имени М.В. Ломоносова, канд. биол. наук (rubisco@list.ru).
Information about the authors
Borzov Nikita Ivanovich, laboratory assistant, department of the mycology and algology, Biological Faculty, Lomonosov Moscow State University.
Gmoshinskiy Vladimir Ivanovich, PhD, senior lecturer, department of the mycology and algology, Biological Faculty, Lomonosov Moscow State University.
Вклад авторов
Все авторы сделали эквивалентный вклад в подготовку публикации.
Contribution of the authors
the authors contributed equally to this article.
Конфликт интересов
Авторы заявляют об отсутствии конфликта интересов. Conflict of interests
The authors declare no conflicts of interests.
Статья поступила в редакцию 11.07.2023; одобрена после рецензирования 12.11.2023; принята к публикации 29.01.2024.
The article was submitted 11.07.2023; approved after reviewing 12.11.2023; accepted for publication 29.01.2024.
