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Амурский зоологический журнал, 2021, т. XIII, № 2
Amurian Zoological Journal, 2021, vol. XIII, no. 2
www.azjournal.ru
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https://www.doi.org/10.33910/2686-9519-2021-13-2-167-182 http://zoobank.org/References/09A7051D-7885-4355-9218-7C98FF9BCC77
UDC 595.61
Review of the millipedes of the Sikhote-Alin State Nature Biosphere Reserve (Far East of Russia), with detection of the morphological variability of Diplomaragna terricolor
(Attems, 1899) (Diplopoda)
E. V. MikhaljovaH, M. E. Sergeev
Federal Scientific Center of the East Asia Terrestrial Biodiversity, Far Eastern Branch of the Russian Academy of Sciences, 159 Stoletiya Vladivostoka Ave., 690022, Vladivostok, Russia
Authors
Elena V. Mikhaljova
E-mail: Mikhaljova@biosoil.ru
SPIN: 4160-3223
Scopus Author ID: 7801402261
ResearcherlD: L-3304-2016
ORCID: 0000-0001-7132-8677
Maksim E. Sergeev
E-mail: eksgauster@inbox.ru
SPIN: 7313-0891
Scopus Author ID: 57207933239
ORCID: 0000-0001-9078-001X
Abstract. The millipede fauna of the Sikhote-Alin State Nature Biosphere Reserve (Far East, Russia) is reviewed, at present containing 17 recognizable species from 15 genera, 9 families and 5 orders. A brief historical account and new faunistic records as well as a key to all Diplopoda taxa presently known from the Sikhote-Alin State Nature Biosphere Reserve are provided. Morphological variability of Diplomaragna terricolor (Attems, 1899) is revealed and analyzed, including the material from other territories of the Primorsky Krai and Khabarovsky Krai. The species diversity of Diplopoda within the reserve is mapped.
Copyright: © The Authors (2021). Published by Herzen State Pedagogical
University of Russia. Open access under Keywords: diplopods, faunistics, variability, key, new faunistic records, CC BY-NC License 4.0. Russian Far East.
Обзор двупарноногих многоножек Сихотэ-Алинского природного биосферного заповедника (Дальний Восток России) с обнаружением морфологической изменчивости Diplomaragna terricolor (Attems, 1899) (Diplopoda)
Е. В. Михалеван, М. Е. Сергеев
Федеральный научный центр биоразнообразия наземной биоты Восточной Азии Дальневосточного отделения РАН, пр. Столетия Владивостока, д. 159, 690022, г. Владивосток, Россия
Сведения об авторах Михалева Елена Валентиновна E-mail: Mikhaljova@biosoil.ru SPIN-код: 4160-3223 Scopus Author ID: 7801402261 ResearcherlD: L-3304-2016 ORCID: 0000-0001-7132-8677 Сергеев Максим Евгеньевич E-mail: eksgauster@inbox.ru SPIN-код: 7313-0891 Scopus Author ID: 57207933239 ORCID: 0000-0001-9078-001X
Права: © Авторы (2021). Опубликовано Российским государственным педагогическим университетом им. А. И. Герцена. Открытый доступ на условиях лицензии CC BY-NC 4.0.
Аннотация. Представлен обзор фауны двупарноногих многоножек Сихотэ-Алинского государственного природного биосферного заповедника, которая к настоящему времени включает 17 видов из 15 родов, 9 семейств и 5 отрядов. Приведены краткая история исследований, новые фаунистические находки и ключ для определения всех таксонов Diplopoda, известных к настоящему времени с территории Сихотэ-Алинского заповедника. Выявлена и проанализирована с привлечением материала из других территорий Приморского и Хабаровского краев морфологическая изменчивость Diplomaragna Ьетсо1ог (Attems, 1899). Распространение видов в пределах заповедника нанесено на карту.
Ключевые слова: диплоподы, фаунистика, изменчивость, ключ, новые фаунистические находки, Дальний Восток России.
Introduction
The Sikhote-Alin State Nature Biosphere Reserve is the largest specially protected natural area in the south of the Russian Far East. It was founded in 1935. The reserve is located on the eastern and western slopes of the central Sikhote-Alin mountain range. The main part of the reserve territory is located within two administrative districts of the Primorsky Krai: Terneysky (eastern macroslope) and Kras-noarmeysky (western macroslope). A small portion of the south-west segment (natural landmark Abrek) belongs to the Dalnegorsky District. The Sikhote-Alin State Nature Biosphere Reserve extends from the coast of the Sea of Japan more than 90 kilometers inland. Currently, the area of its territory is more than 400.000 ha, including sea area.
Mountain slopes of different steepness occupy up to 80% of the reserve territory. The gentle and wide western slopes and short steep eastern ones distinguish the Sikhote-Alin relief. Medium-high mountains with elevations of 500-800 m a.s.l. are dominants in the reserve territory; the highest point is the Glukhomanka Mountain (1598 m a.s.l.). Leveled areas occupy little (about 20%) of the reserve territory; these are mainly river and stream valleys. The density of the reserve hy-drographic network is on average 0.7-0.9 km per 1 km2. The majority of the rivers are located on the eastern macroslope. Serebryanka, Dzhigitovka and Taiezhnaya are the largest rivers. The flow of these rivers is fast, there are many rocky rapids, and sometimes small waterfalls. The largest river of the western macroslope of the middle Sikhote-Alin is the Kolumbe, it belongs to the Amur River basin. It is a calmer river with extensive stretches of deep water and wide swampy valleys.
The monsoon climate is caused by the interaction of marine and continental air masses. The western and eastern macroslopes within the Sikhote-Alin Nature Biosphere Reserve differ substantially by the main climate parameters. The climate of the eastern macroslope is characterized by high humidity due to the constant influence of the Sea
of Japan and the Pacific Ocean. The climate of the isolated from marine influence western macroslope is more continental (Vasiliev et al. 1985; Pimenova 2016a; 2016b).
97% of the reserve's area is covered by forests. Pinus koraiensis, Picea ajanensis and Quercus mongolica are the dominant tree species of forest ecosystems. They occupy more than 70% of the forest vegetation of the reserve (Pimenova 2016c).
Knowledge of the millipede fauna of the Sikhote-Alin State Nature Biosphere Reserve is still patchy and incomplete. The first data on diplopods of the territory derive from the paper by Mikhaljova (1990) in which one new species was described. Later she described two other new species (Mikhaljova 1997a). The review of the millipede fauna of Siberia and the Far East of Russia refers to 10 species recorded in the Sikhote-Alin Biosphere Reserve (Mikhaljova 1993). One of these species (Diplomaragna kuruma Mikhaljova, 1997) was transferred to Pacifiosoma (Mikhaljova 2000). Also, another additional species was mentioned from the reserve territory (Mikhaljova 1997b). In addition, the first ecological observations of general millipede abundance and of the species composition in oakwood ecosystems of the Sikhote-Alin Biosphere Reserve appeared as well (Gromyko 1990). Data on some millipedes are contained in the general survey of the reserve's invertebrates (Gromyko, Potikha 2006). The regional reviews by Mikhaljova (1998; 2004; 2017) include information on the distribution ranges of diplopod species occurring in the Sikhote-Alin Biosphere Reserve.
Material and methods
All material treated here is deposited in the collections of the Federal Scientific Center of the East Asia Terrestrial Biodiversity, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok, Russia (FSCB). Specimens are kept in 70-75% ethanol. In the process of studying the material, the gonopods and some other parts were dissected from the males and mounted in glycerin as temporary micro-preparations. Specimens were studied and il-
lustrated using standard stereomicroscopic (MBS-1) and drawing equipment (RA-6).
SEM micrographs were prepared at the Centre of Collective Use "Biotechnology and Gene Engineering" of the FSCB using a Merlin 62-15 scanning electron microscope. Mounts for SEM were made through air-drying after the transfer to acetone via 96% alcohol, mounting on stubs, and coating with carbon. After the examination, SEM material was removed from stubs and returned to alcohol.
The family-level classification adopted here is that of Shelley (2003). Publication and synonymy lists of each species include the literature references for the Sikhote-Alin State Nature Biosphere Reserve only.
All the natural landmarks in the Sikhote-Alin Nature Reserve are areas of its territory traditionally designated; they are tied to the reserve's cordons and include parts of the basins of the main rivers or large streams (Pimenova 2016d). The natural landmarks:
I, 2. Abrek: floodplain of the Upolnomochen-ny Stream (45°09'402"N, 136°46'721"E); same locality, floodplain of the Skrytaya River (45°06'290"N, 136°45'319"E);
3, 4, 5. Blagodatnoe: upper reaches of the Su-khoy Stream (44°58'57"N, 136°31'09"E); same locality, environs of the Blagodatnoe Lake (44°57'12"N, 136°32'48"E); Ozerny Stream (44°57'51"N, 136°29'46"E);
6. Kunaleika: floodplain of the Khanov Stream (44°53'825"N, 136°20'241"E);
7. Kuruma: floodplain of the Kuruma River (44°54'937"N, 136°12'752"E);
8. Nevidimka: floodplain of the Lianovaya River (44°55'575"N, 136°05'505"E);
9. Kabany: floodplain of the Kabany Stream (45°06'36"N, 135°51'59"E);
10. surroundings of the Glukhomanka Mountain: upper reaches of the Irtysh Stream (45°10'07"N, 135°47'48"E);
II. Sporny: upper part of the Serebryanka River, floodplain of the Sporny Stream (45°09'427"N, 135°54'154"E);
12. Zymoveyny: Zymoveyny Stream, floodplain of the Serebryanka River (45°08'317"N, 136°18'935"E);
13. Beloborodovsky: upper reaches of the Yas-naya River (45°16'594"N, 136°23'934"E);
14. Yasnaya: floodplain of the Zabolochen-naya River (45°19'619"N, 136°28'381"E);
15. Solontsovy: floodplain of the Za-bolochennaya River, Solontsovy Stream (45°18'877"N, 136°28'578"E);
16. Ust-Solontsovy: Solontsovy Stream (45°24'750"N, 136°30'542"E);
17. Sakhalinsky: floodplain of the Sakhalinsky Stream (45°28'231"N, 136°14'277"E);
18. Nechet: floodplain of the Krivoy Stream (45°29'974"N, 136°32'775"E).
Results and discussion
Order Polyxenida Verhoeff, 1934 Family Polyxenidaе Lucas, 1840 Genus Polyxenus Latreille, 1802/1803
Polyxenus sp.
Polyxenus sp. — Mikhaljova 1993: 34; 2004: 41; 2017: 35; Ganin 1997: 124; 2011: 341; Gro-myko, Potikha 2006: 232.
Material. Russia: 6 specimens (FSCB), Primorsky Krai, Sikhote-Alin State Nature Biosphere Reserve, Yasnaya, Quercus forest, 28.05.1985, leg. M. N. Gromyko.
Remarks. This material still requires a closer determination. At present, only the family Polyxenidae and the genus Polyxenus Latreille, 1802/1803 are known from the Asian part of Russia: Polyxenus sp. belonging to the Polyxenus shinoharai Ishii, 1983 group was recorded in Lazovsky District, Primorsky Krai (M. Short's [Deakin University, Australia] personal communication).
Above material is a second find of representatives of Polyxenida taxa in the territories lying between the Ural Mountains in the west and the Pacific Ocean in the east. These specimens most likely belong to the same genus and species group. The reason that we venture to determine these specimens as belonging to Polyxenus is geographical evidence.
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Yasnaya.
Oder Polyzoniida Cook, 1895 Family Polyzoniidae Newport, 1844 Genus Angarozonium Shelley, 1998
Angarozonium bonum (Mikhaljova, 1979)
Polyzonium bonum — Gromyko 1990: 63; Mikhaljova 1993: 4, Map 1; Ganin 1997: 123. Angarozonium bonum — Mikhaljova 1998: 15, Figs. 36-41, Map 2; Ganin 2011: 340, 344.
Material. Russia: 1$, 1? (FSCB), Primor-sky Krai, Sikhote-Alin State Nature Biosphere Reserve: Blagodatnoe, Quercus forest with Corylus, 24.07.1986; 1$, 2 ? (FSCB), Kabany, Pinus koraiensis forest with Betula costata, 13.08.1986; 2?, 2 juv. (FSCB), Zimoveyny, Pi-nus koraiensis forest with Larix, 24.08.1979, all leg. M. N. Gromyko.
Distribution. Russia: Far East (Primorsky Krai, southern part of Khabarovsky Krai).
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Blagodatnoe, Kabany, Yasnaya, Zi-moveyny.
Order Julida Brandt, 1833 Family Julidae Leach, 1814 Genus Pacifiiulus Mikhaljova, 1982
Pacifiiulus amurensis (Gerstfeldt, 1859)
Pacifiilus imbricatus (sic!) — Gromyko 1990: 66. Pacifiiulus imbricatus — Mikhaljova 1993: 12, Map 2; Ganin 1997: 123. Pacifiiulus amurensis — Ganin 2011: 340, 344.
Material. Russia: 1? (FSCB), Primorsky Krai, Sikhote-Alin State Nature Biosphere Reserve: Blagodatnoe, environs of Blagodatnoe Lake, Quercus forest, 44°57'14"N, 136°32'50"E, 13.05.2016; 2$, 1? (FSCB), Blagodatnoe, Sukhoy Stream (upper reaches), 600-700 m a.s.l., 44°58'57" N, 136°31'09"E, 9-10.07.2020; 1?, 1 juv., 1 fragment (FSCB), Sakhalinsky, Sakhalinsky Stream (upper reaches), 45°41'47"N, 136°37'13"E, 30.05.2020; 1? (FSCB), Kuruma, floodplain of Kuruma River, 44°91'52"N, 136°21'18"E, 6-9.06.2020; 1? (FSCB), Abrek, Upolnomochenny Stream, 45°15'89"N, 136°77'75"E, 29.06-1.07.2020; all leg. M. E. Sergeev.
Distribution. Russia: Far East (Primor-sky Krai, southern part of Khabarovsky Krai, Jewish Autonomous Oblast, Amur Oblast),
Siberia (Republic of Altai, southern part of Krasnoyarsky Krai, Republic of Khakassia, Republic of Tyva, Zabaikalsky Krai, Republic of Buryatia); Northeast China.
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Abrek, Blagodatnoe, Kunaleika, Kuruma, Nechet, Sakhalinsky, Solontsovy, Yasnaya, Zimoveyny.
Family Mongoliulidae Pocock, 1903 Genus Koiulus Enghoff, Jensen et Mikhaljova, 2017 Koiulus interruptus Enghoff, Jensen et Mikhaljova, 2017
Material. Russia: 1$, 1?, 1 juv. (FSCB), Primorsky Krai, Sikhote-Alin State Nature Biosphere Reserve, Zimoveyny, floodplain of the Serebryanka River, Zimoveyny Stream, 45°08'29"N, 136°18'33"E, 26.10.2015; leg. M. E. Sergeev.
Distribution. Russia: Far East (Primorsky Krai, southern part of Khabarovsky Krai).
Remarks. This species has hitherto been only known from the southern part of the Khabarovsky Krai (upper course of Ko River and environs of Zolotoy village). So, it is new to the fauna of the Sikhote-Alin State Nature Biosphere Reserve.
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Zimoveyny.
Genus Kopidoiulus Attems, 1909
Kopidoiulus continentalis Golovatch, 1979
Kopidoiulus continentalis — Mikhaljova 1997b: 146.
Material. Russia: (FSCB), Primorski Krai, Sikhote-Alin State Nature Biosphere Reserve: Zimoveyny, Pinus koraiensis forest with Larix, 24.08.1978; 1? (FSCB), Zimoveyny, Pinus koraiensis forest with Larix, 25.09.1979; 1 juv. (subadult male) (FSCB), Zimoveyny, watershed between the Zimoveiny and Sukhoy streams, Pinus koraiensis forest with Quercus, 20.05.1980; 1? (FSCB), Sporny Stream, Picea forest, 19.08.1981, all leg. M.N. Gromyko; 1$ (FSCB), Beloborodovsky, Yasnaya River (upper reaches), 45°29'39"N, 136°38'29"E, 18-20.05.2016. 2? (FSCB), Dalnegorsky Dis-
trict, near Glukhomanka Mts., Irtysh Stream (upper reaches), 45°17'0324"N, 135°77'78"E, 18.09.2015; all leg. M. E. Sergeev.
Distribution. Russia: Far East (Primor-sky Krai, southern part of Khabarovsky Krai); Northeast China.
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Beloborodovsky, Solontsovy, Sporny, Yasnaya, Zimoveyny + near Glukhomanka Mts., Dalnegorsky District.
Genus Skleroprotopus Attems, 1901
Skleroprotopus coreanus (Pocock, 1895)
Skleroprotopus coreanus — Mikhaljova 1993: 15; Ganin 1997: 123; 2011: 340, 344.
Material. Russia: 1$ (FSCB), Primorsky Krai, Sikhote-Alin State Nature Biosphere Reserve: Zimoveyny, Pinus koraiensis with Quercus forest, 28.08.1978; 1$ (FSCB), Zimoveyny, valley Pinus koraiensis forest, 13.09.1978, all leg. M. N. Gromyko; 2$, 1$, 1 juv. (FSCB), Belo-borodovsky, Yasnaya River (upper reaches), 45°29'39''N, 136°38'29"E, 18-20.05.2016; 1$ (FSCB), Sakhalinsky, Sakhalinsky Stream (upper reaches), 45°41'47"N, 136°37'13"E, 30.05.2020; 1$ (FSCB), Abrek, Skrytaya River floodplain, 45°09'82"N, 136°69'08"E, 3-5.07.2020, all leg. M. E. Sergeev.
Distribution. Russia: Far East (Primorsky Krai, southern part of Khabarovsky Krai, Jewish Autonomous Oblast, Amur Oblast); Korea.
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Abrek, Beloborodovsky, Blagodatnoe, Kuruma, Nevidimka, Sakhalinsky, Ust-Solontsovy, Yasnaya, Zimoveyny.
Order Chordeumatida Pocock, 1894
Family Diplomaragnidae Attems, 1907
Genus Orientyla Mikhaljova, 2000
Orientyla dahurica (Gerstfeldt, 1859)
Material. Russia: 1$ (FSCB), Primorsky Krai, Sikhote-Alin State Nature Biosphere Reserve, Sporny, Serebryanka River (upper reaches), environs of Bromleevskaya Izba, 45°19'40"N, 135°99'01"E, 31.07.-2.08.2020, leg. M. E. Sergeev.
Distribution. Russia: Siberia (eastern part of Zabaikalsky Krai, border between Zabaikal-
sky Krai and Amur Oblast), Far East (Primor-sky Krai, southern part of Khabarovsky Krai, Jewish Autonomous Oblast, Amur Oblast); North Korea.
Remarks. This is the first record of O. da-hurica in the Sikhote-Alin State Nature Biosphere Reserve.
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Sporny.
Genus Pacifiosoma Mikhaljova, 2000
Pacifiosoma kuruma (Mikhaljova, 1997)
Diplomaragna kuruma Mikhaljova 1997a: 125, Figs. 4-5.
Diplomaragna kuruma — Mikhaljova 1998: 36, Figs. 123-124, Map 8. Pacifiosoma kuruma — Mikhaljova 2000: 170; 2004: 156, Figs. 367-368, Map 20; 2017: 193, Figs 407-408, Map 26; Ganin 2011: 345.
Distribution. Russia: Far East (Primorsky Krai, Sikhote-Alin State Nature Biosphere Reserve).
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Kuruma.
Genus Diplomaragna Attems, 1907
Diplomaragna terricolor (Attems, 1899)
Figs 1-24
Diplomaragna terricolor — Mikhaljova 1993: 29; 1998: 27, Figs. 86-87, Map 6.
Material. Russia: 1$ (FSCB), Primorsky Krai: Ussuriysky Nature Reserve, mixed forest, litter, 25.09.1977; 2$, 1$ (FSCB), Vladivostok, Okeanskaya Station, forest, litter, 26.04.1980; 2 $ (FSCB), near Vladivostok, Popova Island, forest litter, 27.09.1980, all leg. E. V. Mikhaljova; 1$, 4$ (FSCB), Anuchinsky District, near 35 km NW of Arseniev, forest, August 1986, leg. V. D. Bakurov; 2$ (FSCB), Chuguevsky District, Verkhneussuriysky Research Station, ca. 43°50'N, 134°15'E, 31.07.-5.08.1998, leg. Yu. M. Marusik; 1$ (FSCB), Khasansky District, about 6 km NW of Zanadvorovka village, Gusevsky Mine, mixed forest on slope, litter, 18.08.2002, leg. E. V. Mikhaljova; 1$, 2$, fragments (FSCB), Kedrovaya Pad Nature Reserve, forest, pitfall traps, 13.06.2004, leg. V. N. Kuznetsov; 1$ (FSCB), Shkotovs-
9 10 11 12
Figs. 1-12. Diplomaragna terricolor (Attems, 1899). Variation in structure of the posterior angiocoxal process of posterior gonopod. 1 — from Anuchinsky District, Primorsky Krai; 2 — from Mt. Ko, Khabarovsky Krai; 3 — from Vladivostok, Okeanskaya Station, Primorsky Krai; (1st copy); 4 — from Vladivostok, Okeanskaya Station, Primorsky Krai; (2st copy); 5 — from Kedrovaya Pad Nature Reserve, Primorsky Krai; 6 — from Popova Island, Primorsky Krai; (1st copy); 7 — from Ussuriysky Nature Reserve, Primorsky Krai; 8 — from Lazovsky Nature Reserve, Primorsky Krai; 9 — from Sikhote-Alin Nature Biosphere Reserve (floodplain terrace, Yasnaya River), Primorsky Krai; 10 — from Chuguevsky District, Verkhneussuriysky Research Station, Primorsky Krai; 11 — from Popova Island, Primorsky Krai; (2st copy); 12 — from Sikhote-Alin Nature Biosphere Reserve, Blagodatnoe, Ozerny Stream, Primorsky Krai; a — apical outgrowth; b — finger-shaped process; m — mesal protrusion. Scale in mm (0.5)
Рис. 1-12. Diplomaragna terricolor (Attems, 1899). Вариации структуры заднего ангиококсального отростка заднего гонопода: 1 — из Анучинского района Приморского края; 2 — с горы Ко в Хабаровском крае; 3 — из Владивостока, станция Океанская (1-й экземпляр); 4 — из Владивостока, станция Океанская (2-й экземпляр); 5 — из заповедника Кедровая Падь в Приморском крае; 6 — с острова Попова в Приморском крае (1-й экземпляр); 7 — из Уссурийского заповедника в Приморском крае; 8 — из Лазовского заповедника в Приморском крае; 9 — из Сихотэ-Алинского заповедника (надпойменная терраса реки Ясная) в Приморском крае; 10 — из Чугуевского района (Верхнеуссурийский научно-исследовательский стационар) в Приморском крае; 11 — с острова Попова в Приморском крае (2-й экземпляр); 12 — из Сихотэ-Алинского заповедника (урочище Благодатное, ключ Озёрный) в Приморском крае; a — вершинный вырост; b — пальцевидный отросток; m — срединный выступ. Масштаб в мм (0.5)
ky District, environs of Anisimovka, Litovka Mt., about 100 m a.s.l., Pinus koraiensis forest, 11.09.2006, leg. A. A. Rodionov; 2$, 2? (FSCB), Sikhote-Alin Nature Biosphere Reserve, Blago-datnoe, Ozerny Stream, 29.09.2015, leg. M. E. Sergeev.
Material re-examined. Russia: 1^, 1?, 1 juv. (FSCB), Primorsky Krai: Churuevsky Dis-
trict, Verkhneussuriysky Research Station, Picea forest, litter, 2.09.1975, leg. E. V. Mikhaljova (specimens published by Mikhaljova 1997a);
(FSCB), Vladivostok, Okeanskaya Station, forest, litter, 8.05.1977, leg. E. V. Mikhaljova (specimen published by Mikhaljova 1997a);
(FSCB), Ussuriysky Nature Reserve, Picea forest, 20.05.1977, leg. E. V. Mikhaljova (spec-
imen published by Mikhaljova 1997a); 1$, 2 juv. (FSCB), Ussuriysky Nature Reserve, Abies holophylla forest, 26.10.1977, leg. R. Fe-dorova (specimens published by Mikhaljova 1997a); 1$, 1? (FSCB), near Vladivostok, Pop ova Island, Quercus, Tilia, Betula, Acer, etc., forest, litter, 9.10.1979, leg. E. V. Mikhaljova (specimens published by Mikhaljova 1993); 2$, 1? (FSCB), near Vladivostok, Pop-ova Island, Quercus, Tilia, Betula, Acer, etc., forest, litter, 7.09.1980, leg. E. V. Mikhaljova (specimens published by Mikhaljova 1993); 1$, 3 juv. (FSCB), Sikhote-Alin Nature Biosphere Reserve, floodplain terrace, Yasnaya River, valley Betula forest, 5.06.1984, leg. M. N. Gromyko (specimens published by Mikhaljova 1993); 1$, 1 juv. (FSCB), Gamo-va Peninsula, environs of Telyakovsky Cape,
leaved forest, near stream, litter, 17.06.2003, leg. E. V. Mikhaljova (specimens published by Mikhaljova and Marusik 2004); 1$, 1? (FSCB), Lazovsky Nature Reserve, Amerika Cordon, 15.05.2007, leg. S. A. Shabalin (specimens published by Mikhaljova 2009); 1$, 7 ? (IBSS), Khabarovsky Krai: Sikhote-Alin, Mt. Ko, 1000 m, 6.07.2007, leg. P. V. Budilov (specimens published by Mikhaljova 2016)
Distribution. Russia: Far East (Primorsky Krai, southern part of Khabarovsky Krai).
Remarks. This species was originally described from the environs of Vladivostok, Russia (Attems 1899). Later, based on topotype material, a re-description of this species was provided (Shear 1990).
In the course of our study of males of D. terricolor from the Blagodatnoe of the Sikho-
Figs. 13-16. Diplomaragna terricolor (Attems, 1899). Posterior angiocoxal processes of posterior gonopods. 13 — from Mt. Ko, Khabarovsky Krai; 14 — from Vladivostok, Okeanskaya Station, Primorsky Krai; 15 — from Lazovsky Nature Reserve, Primorsky Krai; 16 — from Sikhote-Alin Nature Biosphere Reserve, Blagodatnoe, Ozerny Stream, Primorsky Krai; a — apical outgrowth; b — finger-shaped process; m — mesal protrusion. Scales: 100 ^m
Рис. 13-16. Diplomaragna terricolor (Attems, 1899). Задние ангиококсальные отростки задних гоноподов: 13 — с горы Ко в Хабаровском крае; 14 — из Владивостока, станция Океанская; 15 — из Лазовского заповедника в Приморском крае; 16 — из Сихотэ-Алинского заповедника (урочище Благодатное, ключ Озёрный) в Приморском крае; a — вершинный вырост; b — пальцевидный отросток; m — срединный выступ. Масштабы: 100 ^m_
Figs. 17-20. Diplomaragna terricolor (Attems, 1899). Gonopods, caudal view. 17 — from Mt. Ko, Khabarovsky Krai; 18 — from Vladivostok, Okeanskaya Station, Primorsky Krai; 19 — from Lazovsky Nature Reserve, Primorsky Krai; 20 — from Sikhote-Alin Nature Biosphere Reserve, Blagodatnoe, Ozerny Stream, Primorsky Krai; a — apical outgrowth; b — finger-shaped process; m — mesal protrusion. Scales: 100 ^m
Рис. 17-20. Diplomaragna terricolor (Attems, 1899). Гоноподы, вид сзади: 17 — с горы Ко в Хабаровском крае; 18 — из Владивостока, станция Океанская; 19 — из Лазовского заповедника в Приморском крае; 20 — из Сихотэ-Алинского заповедника (урочище Благодатное, ключ Озёрный) в Приморском крае; a — вершинный вырост; b — пальцевидный отросток; m — срединный выступ. Масштабы: 100 |im
te-Alin Reserve, the unusual structure of their gonopods has been revealed (see Figs 16, 20). The gonopods differs in the configuration of the posterior angiocoxal processes with the very high apical oval outgrowths (a) (vs not high apical oval outgrowth according to the original description and re-description by Shear) and with broad mesal protrusions (m) carrying finger-shaped processes (b) directed to body dorsum (vs finger-shaped processes directed strictly forward according to the redescription by Shear).
Additional investigation of males of this species from different territories of the Pri-morsky and Khabarovsky Krai allowed discovering of the specific variations in the go-nopod structure of D. terricolor. The basic diagnostic characters of D. terricolor are the
configuration of the posterior gonopod's posterior angiocoxal process with arms and outgrowths of different size, and the architecture of the colpocoxite distal part. The above material appears to show certain variation exactly in the configuration of the posterior angiocoxal process. Thus, the apical oval outgrowth (a) of posterior angiocoxal process ranges from very low to very high (Figs. 1-16). Also, the finger-shaped process (b) is straight (Figs. 11, 12, 16) or more (Figs. 1-3, 5-6, 8-9, 13) or less (Figs. 4, 7, 10, 14, 15) strongly curved forward; its length varies too.
In addition, the variation of the coxal process of the male leg 11 (p) is revealed. Thus, its size ranges from small to large (Figs. 2124). An increase in the size of this coxal process correlates with an increase in the size of
the apical oval outgrowth (a) (see Figs. 13 and 21; 16 and 24). Moreover, we can distinguish two distinct extreme forms of the gonopods + male coxae 11. The first form (males from the Anuchinsky District of the Primorsky Krai and from the environs of Ko Mountain, the Khabarovsky Krai): a very low apical oval outgrowth (a) of the posterior angiocoxal process, a strongly curved forward process b, a small coxal process of leg 11 (Figs. 1, 2, 13, 17, 21). The second form (males from the Blagodatnoe, Sikhote-Alin Nature Reserve,): a very high apical oval outgrowth (a) of the posterior angiocoxal process, straight (directed to dorsum) process b, large coxal process of leg 11 (Figs. 12, 16, 20, 24). The differences between these extreme forms are linked by a complete series of transitions. At
that, the gonopods of a male from the flood-plain terrace of the Yasnaya River (Sikhote-Alin Reserve) (Fig. 9) belong to an intermediate transitional form in contrast to the second extreme gonopod form of the males from Blagodatnoe (Sikhote-Alin Reserve). It should be noted that the second gonopod form and forms close to one are more common for males of the eastern part of the Primorsky Krai. While the first gonopod form and forms close to one were mainly recorded in males from the central and southern parts of the Primorsky Krai as well as in the male from the south of Khabarovsky Krai. However, both forms can occur together. For example, males of two opposite forms (Figs. 6, 11) were found together in the same locality of the Popova Island, south of Primorsky Krai.
Figs. 21-24. Diplomaragna terricolor (Attems, 1899). Coxae of male leg pair 11, front view. 21 — from Mt. Ko, Khabarovsky Krai; 22 — from Vladivostok, Okeanskaya Station, Primorsky Krai; 23 — from Lazovsky Nature Reserve, Primorsky Krai; 24 — from Sikhote-Alin Nature Biosphere Reserve, Blagodatnoe, Ozerny Stream, Primorsky Krai; p — coxal process. Scales: 100 ^m
Рис. 21-24. Diplomaragna terricolor (Attems, 1899). Коксы 11-й пары ног самца, вид спереди: 21 — с горы Ко в Хабаровском крае; 22 — из Владивостока, станция Океанская; 23 — из Лазовского заповедника в Приморском крае; 24 — из Сихотэ-Алинского заповедника (урочище Благодатное, ключ Озёрный) в Приморском крае; р — коксальный отросток. Масштабы: 100 ^m
The species is rather variable. However, the present material being unrepresentative, we have preferred not to make conclusion on the nature of variability. An investigation of this phenomenon is a very considerable project in itself, which can only be realized in future with further accumulation of abundant, representative material and using molecular genetic methods too. Now, it can only be assumed that the morphological variability of this species is due to the different ecological conditions of the eastern, southern and central territories of the Primorsky Krai and the Khabarovsky Krai.
Also, it is necessary to add to the description of D. terricolor the presence of a process on the male coxa 11 since this character was omitted in the re-description of D. terricolor (see Shear 1990).
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Blagodatnoe, Yasnaya.
Diplomaragna zimoveinaya Mikhaljova, 1997
Diplomaragna zimoveinaya Mikhaljova 1997a: 123, Figs. 1-3.
Diplomaragna zimoveinaya — Mikhaljova 1998: 28, Figs. 88-90, Map 6; 2000: 175; 2004: 140, Figs. 330-332, Map 18; 2017: 169, Figs. 363-365, Map 24; Ganin 2011: 340.
Material. Russia: 1$ (FSCB), Primorsky Krai, Sikhote-Alin State Nature Biosphere Reserve, Zimoveyny, Pinus koraiensis with Larix forest, 24.08.1978, leg. M. N. Gromyko.
Distribution. Russia: Far East (Primorsky Krai, Sikhote-Alin Nature Biosphere Reserve).
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Kunaleika, Solontsovy, Zimoveyny.
Family Caseyidae Verhoeff, 1909
Genus Underwoodia Cook & Collins, 1895
Underwoodia kurtschevae Golovatch, 1980
Underwoodia kurtschevae — Mikhaljova 1993: 17; Gromyko 1990: 63; Shelley 1993: 175; Ganin 1997: 123; 2011: 340, 344.
Distribution. Russia: Far East (Primorsky Krai, southern part of Khabarovsky Krai including Bolshoy Shantar Island, Jewish Au-
tonomous Oblast, Amur Oblast, Kamchatka Peninsula, Sakhalin Island, Moneron Island, Kuriles: Zeleny, Shikotan, Kunashir, Iturup, Urup, Chirpoi, Ketoi); North Korea.
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Blagodatnoe, Kabany, Kunaleika, Kuruma, Nechet, Nevidimka, Solontsovy, Sporny, Yasnaya, Zimoveyny.
Order Polydesmida Pocock, 1887 Family Xystodesmidae Cook, 1895 Genus Levizonus Attems, 1898
Levizonus distinctus Mikhaljova, 1990
Levizonus distinctus Mikhaljova 1990: 134, fig. 1. Levizonus distinctus — Mikhaljova 1993: 33; 1998: 53, Figs. 200-204, Map 12; 2004: 245, Figs. 612-616, Map 32; 2017: 295, Figs. 660663, Map 43; Ganin 2011: 341. Levizonus distinatus (sic!) — Ganin 1997: 124; Gromyko, Potikha 2006: 232.
Material. Russia: 1$, 1 juv. (FSCB), Primorsky Krai, Sikhote-Alin State Nature Biosphere Reserve: Zimoveyny, Pinus koraiensis with Rhododendron, 22.08.1978; 1$, 1? (FSCB), Ust-Solontsovy, burnt Pinus koraiensis with Quercus forest (5 year old), 4.06.1983; 1$, 2?, 12 juv., 2 fragments (FSCB), Yasnaya, valley Betula forest, floodplain terrace, 24.06.1984, all leg. M.N. Gromyko; 1$, 1? (FSCB), Abrek, floodplain of Skritaya River, 45°09'82"N, 136°69'08"E, 3-5.07.2020; 1$ (FSCB), Sporny, Serebryanka River (upper reaches), environs of Bromleevskaya Izba, 45°19'40"N, 135°99'01"E, 31.07.-2.08.2020, all leg. M.E. Sergeev.
Distribution. Russia: Far East (Primorsky Krai, Sikhote-Alin State Nature Biosphere Reserve).
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Abrek, Nechet, Ust-Solontsovy, So-lontsovy, Sporny, Yasnaya, Zimoveyny.
Levizonus malewitschi Lokschina et Golovatch, 1977
Levizonus malewitschi — Gromyko 1990: 63; Mikhaljova 1993: 33; Ganin 1997: 124; 2011: 341.
Distribution. Russia: Far East (Primorsky Krai).
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Blagodatnoe.
Family Paradoxosomatidae Daday, 1889 Genus Sichotanus Attems, 1914
Sichotanus eurygaster (Attems, 1898)
Sichotanus eurygaster — Gromyko 1990: 66; Mikhaljova 1993: 30; Ganin 1997: 124; 2011: 341.
Material. Russia: 1? (FSCB), Primorsky Krai, Sikhote-Alin State Nature Biosphere Reserve: Yasnaya, floodplain of Zabolochennaya River, 2.07.2015; 12$ (FSCB), Yasnaya, flood-plain of Zabolochennaya River, forest, pitfall traps, 44°53'25"N, 136°20'18"E, 3-5.07.2015; 1$ (FSCB), Beloborodovsky, Yasnaya River (upper reaches), 45°29'39"N, 136°38'29"E, 18-20.05.2016; 1? (FSCB), Sporny Stream (upper reaches), 29.06.2017; 1juv. (FSCB), Sporny, Serebryanka River (upper reaches), environs of Bromleevskaya Izba, 45°19'40"N, 135°99'01"E, 31.07.-2.08.2020; 1$ (FSCB), Sakhlinsky, Sakhalinsky Stream (upper reaches), 45°41'47"N, 136°37'13"E, 30.05.2020; 2$, 1? (FSCB), Kuruma, floodplain of Kuruma River, 44°91'52"N, 136°21'18"E, 6-9.06.2020; 1$ (FSCB), Abrek, floodplain of Skrytaya River, 45°09'82"N, 136°69'08"E, 3-5.07.2020; 1? (FSCB), Blagodatnoe, Sukhoy Stream (upper reaches), 600-700 m a.s.l., 44°58'57"N, 136°31'09"E, 9-10.07.2020; all leg. M. E. Sergeev.
Distribution. Russia: Far East (Primorsky Krai, southern part of Khabarovsky Krai, Jewish Autonomous Oblast); North and South Korea; Northeast China.
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Abrek, Beloborodovsky, Blago-datnoe, Kuruma, Sakhalinsky, Solontsovy, Sporny, Yasnaya.
Genus Cawjeekelia Golovatch, 1980
Cawjeekelia koreana (Golovatch, 1980)
Cawjeekelia koreana — Mikhaljova 1993: 30; Ganin 1997: 124; 2011: 341.
Material. Russia: 1$ (FSCB), Primorsky Krai, Sikhote-Alin State Nature Biosphere Reserve: Kuruma, 44°91'52"N, 136°21'18"E, 7.05.2017; 1$, 1? (FSCB), Blagodatnoe,
Sukhoy Stream (upper reaches), 600-700 m a.s.l., 44°58'57"N, 136°31'09"E, 9-10.07.2020, all leg. M. E. Sergeev.
Distribution. Russia: Far East (Primorsky Krai, Amur Oblast); North Korea.
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Blagodatnoe, Kuruma.
Family Polydesmidae Leach, 1815 Genus Epanerchodus Attems, 1901
Epanerchoduspolymorphus Mikhaljova et Golovatch, 1981
Material. Russia: 4$ (morpha simplifica-ta), 2?, 1 juv. (FSCB), Primorsky Krai, Sikho-te-Alin State Nature Biosphere Reserve: Kuru-ma, floodplain of Kuruma River, 44°91'52"N, 136°21'18"E, 6-9.06.2020, leg. M. E. Sergeev.
Distribution. Russia: Far East (Primorsky Krai); North Korea.
Remarks. This is the first record of Epanerchodus polymorphus in the Sikhote-Alin State Nature Biosphere Reserve.
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Kuruma.
Genus Uniramidesmus Golovatch, 1979
Uniramidesmus dentatus Mikhaljova, 1979
Uniramidesmus dentatus—Mikhaljova 1993: 32; Ganin 1997: 124; 2011: 341.
Distribution. Russia: Far East (Primor-sky Krai, southern part of Khabarovsky Krai, ?Amur Oblast).
Occurrences in the natural landmarks of the Sikhote-Alin State Nature Biosphere Reserve: Kuruma.
Conclusion
At present, 17 species from 15 genera, 9 families and 5 orders of Diplopoda are known from the Sikhote-Alin State Nature Biosphere Reserve. This is 36.9% of the total number of millipede species inhabiting the Pimorsky Krai. Three species are new to the reserve fauna. Rather this pattern expresses the degree of diplopod fauna study.
Generally, the reserve fauna of Diplopoda is relatively original. At the species level, the endemism amounts to at least 18.2% of the
total number of millipede species occurring there. Three species (Levizonus distinctus, Diplomaragna zimoveinaya, Pacifiosoma ku-ruma) are known from the Sikhote-Alin State Nature Biosphere Reserve only. Most of the species that populate the Sikhote-Alin State Nature Biosphere Reserve can be attributed to the Manchurian distribution pattern, which includes the Primorsky Krai, the southern part of the Khabarovsky Krai, the Amur Oblast, the Jewish Autonomous Oblast, Northeast China, and Korea. Only three species (Orientyla dahurica as well as parthe-nogenetic Pacifiiulus amurensis and Under-woodia kurtschevae) show wider distributions; both O. dahurica and P amurensis are also known from Siberia while U. kurtschevae also from the Kamchatka Peninsula.
Underwoodia is a genus showing a trans-Beringian distribution pattern (Golovatch 1980). Orientyla and Pacifiiulus are endemic to the Asian part of Russia.
The order Polydesmida is definitely dominant in the diplopod fauna of the reserve. It is represented by three families, five genera and six species. Families Paradoxosomatidae and Polydesmidae include the species being quite common in the southern part of the Russian Far East and the adjacent territories. However, family Xystodesmidae is represented by both Levizonus distinctus, endemic to the reserve, and by Levizonus malewitschi known from the central-eastern part of the Primorsky Krai only. The orders Polyzoniida and Polyxenida are the smallest; they include one species each.
The distribution of Diplopoda in the Sikhote-Alin State Nature Biosphere Reserve (Map) shows four of the species (Sichotanus eurygaster, Underwoodia kurtschevae, Sklero-protopus coreanus, Pacifiiulus amurensis) as being common and occurring at 8-10 natural landmarks of the reserve. Three species (Levizonus distinctus, Kopidoiulus continen-talis, Angarozonium bonum) are recorded in 4-7 natural landmarks. The remaining species tend to be more local in distribution. Of course, our map is provisional, reflecting the state of the art and the distribution of collecting efforts.
Millipedes occur in most of the biotopes, only excluding the steppe sea terrace within the natural landmark of Blagodatnoe (Gro-myko, Potikha 2006). Millipede abundance is a variable characteristic. Thus, it ranges from 0.2 to 4.1 ind./m2 in different dry Quercus forests, and from 23.3 to 57.1 ind./m2 in different humid Quercus forests (Gromyko 1990). Levizonus distinctus is the most numerous species; its maximum numbers (more 30 ind./m2) were registered in the valley with Pinus ko-raiensis forests (Gromyko, Poltikha 2006).
Key to Diplopoda orders, families, genera and species occurring in the Sikhote-Alin State
Nature Biosphere Reserve
1(2) Cuticle soft, not impregnated with cal-cites. Head, most of pleura and telson with bundles of hollow serrate setae (=
trichomes) ..............................
order Polyxenida, family Polyxenidae, genus Polyxenus, Polyxenus sp. 2(1) Cuticle hard, impregnated with calcites. Complex trichomes absent, body bare or
with simple setae...................... 3
3(4) Head very small, elongated anteriorly into a rostrum. Body strongly flattened dorso-ventrally, without paraterga and macro-
chaetae...............order Polyzoniida,
family Polyzoniidae, genus Angarozonium, A. bonum.
4(3) Head larger, more or less ovoid, devoid of a rostrum. Body more or less cylindrical,
with paraterga and 3+3 macrochaetae____
....................................... 5
5(14) Telson with a pair of spinnerets
................... order Chordeumatida
6(13) Each metatergite with paraterga
...............family Diplomaragnidae
7(8) Colpocoxites of posterior gonopods flattened on sides, not sagittally. Lateral sheath
process present..........................
Genus Orientyla, O. dahurica 8(7) Colpocoxites of posterior gonopods different, if flattened, then frontocaudally. Lateral sheath process absent..............9
9(12) Gonopod anterior angiocoxal processes present.............genus Diplomaragna
Map. Species diversity of Diplopoda within the Sikhote-Alin State Nature Biosphere Reserve. The numbers of the natural landmarks (1-18) are explained in the "Material and Methods section"
Карта. Видовое разнообразие двупарноногих многоножек в Сихотэ-Алинском заповеднике. Номера урочищ (1-18) поясняются в разделе «Материал и методы»
10(11) Posterior angiocoxal process of 11(10) Posterior angiocoxal process of poste-
posterior gonopod like a caudally con- rior gonopod large, with broad mesal den-vex plate with an undulate edge, devoid of teeth. Body length 12-12.5 mm ticle protrusions. Body length 2°-22 mm ..........................D. zimoveinaya .............................D. terricolor
12(9) Gonopod anterior angiocoxal processes
absent.....genus Pacifiosoma, P. kuruma
13(6) Each metatergite without paraterga
......................... family Caseyi-
dae, genus Underwoodia, U. kurtschevae
14(5) Telson without a pair of spinnerets____
...................................... 15
15(26) Metatergites with paraterga. Eyes absent. Adult body with 20 segments, including
telson.................order Polydesmida
16(19) Paraterga well-developed, serrate at lateral margin, without peritremata. Body relatively slender; metaterga relatively flat, with three transverse rows ofbosses. Gonopod
coxites fused medially....................
.....................family Polydesmidae
17(18) Paraterga broad. Body relatively large
(adults >15 mm long) .................
.....genus Epanerchodus, E. polymorphus
18(17) Paraterga narrow. Body relatively small
(adults <15 mm long) .................
.......genus Uniramidesmus, U. dentatus
19(16) Paraterga relatively poorly-developed, with peritremata, non-serrate at lateral margin. Body stout, metaterga strongly convex, arched, without rows of bosses. Gonopods free from each other basally ...
...................................... 20
20(23) Adults poorly pigmented: whitish to yellowish with or without brown dorsal pattern. Gonopods simple, without
conspicuous solenomere branch............
____family Xystodesmidae, genus Levizonus
21(22) Apex of gonopod telopodite like two plates placed perpendicular to each other; one of the plates serrate at outer margin
...........................L. malewitschi
22(21) Apex of gonopod telopodite different. Solenomere with a large horn basally
.............................L. distinctus
23 (20) Adults strongly pigmented: brown to dark brown with yellow peritremata. Gonopods complex, with an evident
solenomere branch.......................
...............family Paradoxosomatidae
24(25) Peritremata of midbody segments poorly demarcated ventrally. Gonopod solenomere flagelliform....................
...........genus Sichotanus, S. eurygaster
25(24) Peritremata of midbody segments evidently demarcated ventrally. Gonopod
solenomere different ....................
............genus Cawjeekelia, C. koreana
26(15) Metatergites without paraterga, body subcylindrical Eyes present. Adult body
with more than 30 segments...............
..............................order Julida
27(28) Surface of metazonites completely striate.............................family Julidae, genus Pacifiiulus, P. amurensis 28(27) Surface of metazonites clearly striate
only below ozopore level ...............
....................family Mongoliulidae
29(30) Male leg 7 with very long coxal process
covering with spinose setae, large .........
.........genus Skleroprotopus, S. coreanus
30(29) Male leg 7 without very long coxal process covering with spinose setae, small ... 31 31(32) Posterior gonopod divided from the basis into two equally long branches
.............genus Koiulus, K. interruptus
32(31) Posterior gonopod not divided from the basis into two equally long branches .......genus Kopidoiulus, K. continentalis
Acknowledgements
We are grateful to Dr. M. N. Gromyko (Sik-hote-Alin State Nature Biosphere Reserve) who provided additional material for the present study. We are also thankful to Mr. G. M. Shaul-sky (Terney, Russia), Mr. G. A. Nacharkin and Mrs. E.A. Govorova (both Moscow, Russia) for their assistance in the material collecting. Our special thanks are extended to Mr. V. M. Kaza-rin (Vladivostok, Russia) for the help in preparation of scanning electron micrographs.
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For citation: Mikhaljova, E. V., Sergeev, M. E. (2021) Review of the millipedes of the Sikhote-Alin State Nature Biosphere Reserve (Far East of Russia), with detection of the morphological variability of Diplomaragna terricolor (Attems, 1899) (Diplopoda). Amurian Zoological Journal, vol. XIII, no. 2, pp. 167-182. https://www.doi. org/10.33910/2686-9519-2021-13-2-167-182
Received 21 January 2021; reviewed 16 February 2021; accepted 3 March 2021.
Для цитирования: Михалева, Е. В., Сергеев, М. Е. (2021) Обзор двупарноногих многоножек Сихотэ-Алин-ского природного биосферного заповедника (Дальний Восток России) с обнаружением морфологической изменчивости Diplomaragna terricolor (Attems, 1899) (Diplopoda). Амурский зоологический журнал, т. XIII, № 2, с. 167-182. https://www.doi.org/10.33910/2686-9519-2021-13-2-167-182 Получена 21 января 2021; прошла рецензирование 16 февраля 2021; принята 3 марта 2021.
