Yuri N. Sundukov, PhD, Senior Researcher of the Laboratory of Entomology of the Zoology Department of the Federal Scientific Center of East Asia Terrestrial Biodiversity (690022, Russia, Vladivostok, Avenue of 100-letiya Vladivostoka 159); iD ORCID:; e-mail:
Kirill V. Makarov, PhD, Professor of the Zoology and Ecology Department of the Institute of Biology and Chemistry of the Moscow State Pedagogical University (129164, Russia, Moscow, Kibalchicha Street 6, build. 3); iD ORCID:; e-mail:

Reference to article

Sundukov Yu.N., Makarov K.V. 2021. The ground beetles of the tribus Trechini (Carabidae) on the Southern Kuril Islands. Nature Conservation Research 6(4): 15–51.

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This article is devoted to the inventory of ground beetles of the tribe Trechini on the southern Kuril Islands. Until the beginning of the XXI century, only one species of tribe was noted from the islands. As a result of our survey in 2008–2018 on the islands of Kunashir, Shikotan, Yuri, and Polonskogo, 764 specimens of ground beetles of the tribe Trechini were collected and studied, belonging to four species from two genera: Blemus discus (Kunashir Island), Trechus nakaguroi (Kunashir Island), T. dorsistriatus (Kunashir, Shikotan, Polonskogo and Yuri islands) and T. nigricornis (Kunashir and Yuri islands). To clarify their taxonomic status, type specimens of T. alexandrovi, T. dorsistriatus, T. nigricornis, Epaphius arsenjevi, and E. plutenkoi, as well as 475 specimens of these species from other parts of their ranges, were studied as comparative materials. Based on the study of this material, the following taxonomic changes were proposed: T. dorsistriatus laferi ssp. nov.; Blemus discus alexandrovi, stat. rest.; T. nigricornis, stat. resurr.; T. nakaguroi sachalinensis, stat. nov.; T. nigricornis arsenjevi, stat. nov.; Epaphius arsenjevi = Epaphius plutenkoi, syn. nov. The holotype of T. nigricornis was designated. Blemus discus alexandrovi has been recorded for the first time in the fauna of the Kuril Islands. The features of distribution and ecology of all species on the surveyed islands are discussed. Currently, B. discus is found only on the west coast of Kunashir Island, suggesting a recent colonisation of this island. Trechus nakaguroi inhabits only the mountainous dark coniferous forests of the northern Kunashir, which indicates a relict distribution of this species. Trechus dorsistriatus is the most abundant species of Trechini, inhabiting the most diverse biotopes on all the islands of the South Kurile. Trechus nigricornis is an inhabitant of moss mires in river floodplains. Unlike the specimens of this species from the mainland and Hokkaido, all specimens of T. nigricornis from the Southern Kuril Islands are wingless; we associate this with the impossibility of flying in the cold and foggy climate of the Kuril Islands. The proposed hypothesis on the formation of the Trechini fauna of the South Kurile is based on the geological youth of the Kuril Islands, which makes it necessary to take into account that by the time ground beetles colonised their ranges, most of the currently existing Trechini groups already existed, and their modern distribution was largely determined by climate and landscape changes in the Quaternary period. The compiled identification key includes the four species of the South Kuril fauna and Trechus ephippiatus, repeatedly erroneously indicated from Kunashir Island.


Coleoptera, fauna, fauna formation hypothesis, key, Russian Far East, systematic

Artice information

Received: 15.04.2021. Revised: 15.07.2021. Accepted: 23.07.2021.

The full text of the article

Ananina T.L. 2012. Ecological classification of ground beetles (Coleoptera, Carabidae) from Barguzinskiy ridge (on the basis of temperature, humidity and vegetative cover criteria preference). Proceedings of Samara Scientific Centre of RAS 14(1): 1868–1871. [In Russian]
Ananina T.L. 2014. About characteristic of carabid beetles fauna (Coleoptera, Carabidae) of the Eastern Baikal Area. Baikalskij Zoologičeskij Žurnal 2(15): 24–30. [In Russian]
Aukema B. 1991. Fecundity in relation to wing-morph of three closely related species of the melanocephalus group of the genus Calathus (Coleoptera: Carabidae). Oecologia 87(1): 118–126. DOI: 10.1007/BF00323789
Barkalov V.Yu. 2009. Flora of the Kuril Islands. Vladivostok: Dalnauka. 468 p. [In Russian]
Bazarova V.B. 2014. Spreading of broadleaved species in Amur River Basin in the Holocene. Botanica Pacifica 3(2): 47–54. DOI: 10.17581/bp.2014.03205
Belousov I.A. 2017. Tribe Trechini Bonelli, 1810. In: I. Löbl, D. Löbl (Eds.): Catalogue of Palaearctic Coleoptera. Vol. 1: Archostemata – Myxophaga – Adephaga. Revised and Updated Edition. Leiden; Boston: Brill. P. 357–455.
Belova V.A. 1985. Vegetation and climate of the Late Cenozoic of the south of Eastern Siberia. Novosibirsk: Nauka. 160 p. [In Russian]
Berg L.S. 1914. Les poissons du fleuve Toumen-oula (Corée), collectionés par A.I. Czerski. Annals of the Zoological Museum of the Imperial Academy of Sciences 19(4): 554–561. [In Russian]
Berlov E.Y., Berlov O.E. 1997. The ground beetles (Coleoptera, Carabidae) of Sakhalin Island. Vestnik IrGSHA 4: 52–56. [In Russian]
Bersenev I.I., Besverkhy V.L. 1991. Chapter 5. The Sea of Japan and the Pacific shelf of the Japanese Islands. In: M.N. Alekseev (Ed.): Atlas of paleogeographic maps «Eurasian shelves in the Mesozoic and Cenozoic». Vol. 1. Moscow: Geologigal Institute of AS USSR. P. 5(1–11). [In Russian]
Bousquet Y. 2012. Catalogue of Geadephaga (Coleoptera, Adephaga) of America, north of Mexico. ZooKeys 245(Spl. Iss.): 1–1722. DOI: 10.3897/zookeys.245.3416
Budilov P.V. 2013. The first data on the ground beetle population (Coleoptera, Carabidae) at the National Park «Anyuiskiy». In: E.A. Grigorieva (Ed.): Modern issues of regional development. Birobidzhan: ICARP FEB RAS. P. 126–128. [In Russian]
Budilov P.V. 2014. Finds of new species of ground beetles (Coleoptera, Carabidae) for the territory of the Jewish Autonomous Region. Regionalnye problemy 17(1): 40–42. [In Russian]
Budilov P.V. 2017. Ground beetles (Coleoptera, Carabidae) in the smalleaved forest of restricted area of the «Bastak» Nature Reserve, Jewish Autonomous Region. A.I. Kurentsov's Annual Memorial Meetings 28: 136–145. [In Russian]
Darlington P.J. 1943. Carabidae of mountains and islands: data on the evolution of isolated faunas, and on atrophy of wings. Ecological Monographs 13(1): 37–61. DOI: 10.2307/1943589
Dubatolov V.V., Kosterin O.E. 2000. Nemoral species of Lepidoptera (Insecta) in Siberia: a novel view on their history and the timing of their range disjunctions. Entomologica Fennica 11(3): 141–166. DOI: 10.33338/ef.84061
Fischhuber M. 1977. Die Arten der Gattung Epaphius Stephens mit Beschreibung einer neuen Species (Col., Carabidae). Koleopterologische Rundschau 53: 3–7.
Ganzei K.S., Ivanov A.N. 2012. Landscape diversity of the Kuril Islands. Geography and Natural Resources 33(2): 142–148. DOI: 10.1134/S1875372812020072
ICZN. 2004. International Code of Zoological Nomenclature. Fourth edition. Saint Petersburg. 221 p. [In Russian]
Igarashi Y., Zharov A.E. 2011. Climate and vegetation change during the late Pleistocene and early Holocene in Sakhalin and Hokkaido, northeast Asia. Quaternary International 237(1–2): 24–31. DOI: 10.1016/j.quaint.2011.01.005
Igarashi Y., Yamamoto M., Ikehara K. 2011. Climate and vegetation in Hokkaido, northern Japan, since the LGM: Pollen records from core GH02-1030 off Tokachi in the northwestern Pacific. Journal of Asian Earth Sciences 40(6): 1102–1110. DOI: 10.1016/j.jseaes.2010.08.001
Ikeda H., Kubota K., Cho Y.B., Liang H., Sota T. 2009. Different phylogeographic patterns in two Japanese Silpha species (Coleoptera: Silphidae) affected by climatic gradients and topography. Biological Journal of the Linnean Society 98(2): 452–467. DOI: 10.1111/j.1095-8312.2009.01283.x
Jäch M.A., Delgado J.A. 2014. Revision of the Palearctic species of the genus Ochthebius Leach XXIX. The Asian species of the O. vandykei group (Coleoptera: Hydraenidae). Koleopterologische Rundschau 84: 81–100. DOI: 10.1076/aqin.
Jeannel R. 1928. Monographie des Trechinae. Morphologie comparée et distribution géographique d'un groupe de coléoptères. (Troisième livraison). Les Trechini cavemicoles. L'Abeille, Journal d'Entomologie 35: 1–808.
Jeannel R. 1962. Les Trechini de l'extréme-orient. Revue Française d'Entomologie 29: 171–207.
Keleynikova S.I. 1976. Types of Coleoptera of V.I. Moschulsky in the collection of the Zoological Museum of Moscow State University. Proceedings of the Zoological Museum of the Moscow State University 15: 183–224. [In Russian]
Khobrakova L.Ts., Shilenkov V.G., Dudko R.Yu. 2014. The ground beetles (Coleoptera, Carabidae) of Buryatia. Ulan-Ude: Buryat Scientific Center of SB RAS Press. 380 p. [In Russian]
Kholin S.K., Storozhenko S.Yu., Sidorenko V.S., Lafer G.Sh., Tanabe S. 2005. The ground beetles (Coleoptera: Carabidae) assemblages on the Muravjev-Amursky Peninsula, Russian Far East. Far Eastern Entomologist 156: 1–11.
Kimoto Sh., Yasuda N. 1995. Carabidae in Hokkaido. Tokai University Press. 317 p.
Kosterin O.E. 2002. Western range limits and isolates of eastern odonate species in Siberia and their putative origins. Odonatologica 34(3): 219–242.
Kryzhanovskij O.L., Molodova L.P. 1973. Faunistical and ecological complexes of ground-beetles (Coleoptera, Carabidae) of South Sakhalin. Entomologicheskoye Obozrenie 52(1): 63–75. [In Russian]
Kryzhanovskij O.L., Okhotina M.V., Bromlei G.F., Lafer G.Sh. 1975. A review of the ground-beetles (Coleoptera, Carabidae) of the Kuril Islands. Proceedings of the Institute of Biology and Soil Science FEB AS USSR 28(131): 119–142. [In Russian]
Kryzhanovskij O.L., Belousov I.A., Kabak I.I., Kataev B.M., Makarov K.V., Shilenkov V.G. 1995. A checklist of the ground-beetles of Russia and adjacent lands (Insecta, Coleoptera, Carabidae). Sofia–Moscow: Pensoft Publishers. 271 p.
Kuberskaya O.V. 2017. Biotopic distribution of ground beetles (Coleoptera, Carabidae) in the Komsomolsky State Nature Reserve. In: E.Ya. Frisman (Ed.): XII Far Eastern Conference of Nature Conservation Problems (Birobidzhan, 10–13 October 2017). Birobidzhan: ICARP FEB RAS. P. 96–100. [In Russian]
Kwon Y.J., Lee S. 1986. Check list of superfamily Caraboidea from Korea. Insecta Koreana 6(1): 1–56.
Lafer G.Sh. 1977. Materials on the study of ground beetles (Coleoptera, Carabidae) of the dark-coniferous taiga zone of Sikhote-Alin' in the Primorsky Krai, USSR. Proceedings of the Institute of Biology and Soil Science FEB AS USSR 44(147): 5–34. [In Russian]
Lafer G.Sh. 1984. Ecological and faunal review of ground beetles (Coleoptera, Carabidae) of Sakhalin and Moneron. 1. Tribes Trachypachini – Bembidiini. In: Fauna and ecology of invertebrates of the Far East (pests and entomophages). Vladivostok: Dalnauka. P. 43–66. [In Russian]
Lafer G.Sh. 1989. 4. Family Carabidae – The Ground-beetles. In: P.A. Lehr (Ed.): Key to the insects of the USSR Far East. Vol. 3. Coleoptera. Part 1. Leningrad: Nauka. P. 71–222. [In Russian]
Lafer G.Sh. 1990. New information on the fauna of ground beetles (Coleoptera, Carabidae) of Iturup Island (Kuril Islands). In: V.A. Mazyuk (Ed.): Fauna and ecology of ground beetles. Kishinev. P. 41–43. [In Russian]
Lafer G.Sh. 2002. Ground beetles (Coleoptera, Carabidae) of southern oceanic islands of the Greate Kuril Ridge. Euroasian Entomological Journal 1(1): 47–66. [In Russian]
Lafer G.Sh. 2005. A check-list of ground beetles (Coleoptera: Cicindelidae, Carabidae) of the Muravjev-Amursky Peninsula, Primorskii Krai. Far Eastern Entomologist 151: 1–8.
Lafer G.Sh. 2006. Ground-beetles (Coleoptera: Cicindelidae, Carabidae) of Moneron Island. In: S.Yu. Storozhenko (Ed.): Flora and fauna of Moneron Island. Vladivostok: Dalnauka. P. 218–227. [In Russian]
Lindroth C.H. 1961. The ground beetles (Carabidae excl. Cicindelinae) of Canada and Alaska. Part 2. Opuscula Entomologica 20: 1–200.
Lindroth C.H. 1979. 2.41. The theory of glacial refugia. In: T.L. Erwin (Ed.): Carabid beetles: Their evolution, natural history, and classification. The Hague: Dr. W. Junk Publ. P. 385–394.
Lutshnik V. 1915. Description d'une nouvelle espèce du sous-genre Lasiotrechus Gаnglb. (Coleoptera, Carabldae). Russian Entomological Review 15(1): 74. [In Russian]
Molodova L.P. 1973. Fauna of soil invertebrates of South Sakhalin. In: M.S. Gilyarov (Ed.): Ecology of soil invertebrates. Moscow: Nauka. P. 60–73. [In Russian]
Molodova L.P. 1974. Zoogeographic composition and altitude distribution of soil invertebrates in South Sakhalin. Zoologicheskii Zhurnal 53(8): 1178–1186. [In Russian]
Moraveč P., Wrase D.W. 1995. Beitrag zur Systematik und Fauna der Trechodini und Trechini des Russischen Fernen Ostens mit der Beschreibung von zwei neuen Arten (Col., Carabidae). Linzer Biologische Beiträge 27 (1): 367–395.
Moraveč P., Wrase D.W. 1997. Zweiter Beitrag zur Systematik und Fauna der Trechodini und Trechini des Russischen Fernen Ostens mit der Beschreibung von zwei neuen Trechus-Arten (Coleoptera, Carabidae). Linzer Biologische Beiträge 29(2): 1057–1067.
Moraveč P., Uéno S.I., Belousov I.A. 2003. Tribe Trechini Bonelli, 1810. In: I. Löbl, A. Smetana (Eds.): Catalogue of Palaearctic Coleoptera. Vol. 1: Archostemata – Myxophaga – Adephaga. Stenstrup: Apollo Books. P. 288–346.
Mori M. 2014. Trechinae of Japan. Kiberihamushi 37 (1): 26–34.
Motschulsky V. 1844. Insectes de la Sibérie rapportés d'un voyage fait en 1839 et 1840. Mémoires présentés a l'Académie Impériale des Sciences de St.-Pétersbourg par divers savans et lus dans ses assemblées 5: 1–274.
Movchan Yu.V., Petrusenko A.A. 1973. On studying the nutrition of Salvelinus malma Walb. from the Kuril Islands. Proceedings of the Zoological Museum of the AS UkrSSR 35: 46–50. [In Ukrainian]
Pankratov A.A. 2011. Ground beetles (Coleoptera, Carabidae) of wetlands in the south of Lake Baikal. Baikalskij Zoologičeskij Žurnal 1(6): 10–13. [In Russian]
Park J.K., Park J., Choi I.J., Choi E.Y. 2014. A catalog of the beetles (Coleoptera) recorded from the Korean indigenous species survey of the National Institute of Biological Resources (2011–2013). Entomological Research Bulletin 30(2): 100–103.
Pietsch T.W., Bogatov V.V., Storozhenko S.Yu., Lelej A.S., Barkalov V.Yu., Takahashi H., Joneson S.L., Kholin S.K., Glew K.A., Harpel J.A., Krestov P.V., Makarchenko E.A., Minakawa N., Ôhara M., Bennett D.J., Anderson T.R., Crawford R.L., Prozorova L.A., Kuwahara Y., Shedko S.V., Yabe M., Woods P.J., Stevenson D.E. 2012. Biodiversity and biogeography of Sakhalin Island. In: S.Yu. Storozhenko (Ed.): Flora and fauna of North-West Pacific islands. Vladivostok: Dalnauka. P. 11–78.
Razjigaeva N.G., Ganzey L.A., Arslanov Kh.A., Grebennikova T.A., Belyanina N.I., Mokhova L.M. 2011. Paleoenvironments of Kuril Islands in Late Pleistocene–Holocene: Climatic changes and volcanic eruption effects. Quaternary International 237(1–2): 4–14. DOI: 10.1016/j.quaint.2011.01.008
Razzhigaeva N.G., Ganzei L.A., Grebennikova T.A., Belyanina N.I., Mokhova L.M., Arslanov Kh.A., Chernov S.B. 2014. Landscape evolution of Kuril Islands in the Holocene. Izvestiya Rossiiskoi Akademii Nauk. Seriya Geograficheskaya 3: 43–50. DOI: 10.15356/0373-2444-2014-3-43-50 [In Russian]
Rogatnykh D.Yu. 2007. New records of ground beetles (Coleoptera, Carabidae) from Amur Region. Euroasian Entomological Journal 6(4): 493–495. [In Russian]
Rogatnykh D.Yu. 2009. Characteristics of the landscape-biotopic distribution of the population and life forms of ground beetles (Coleoptera: Carabidae) in the south of the Amur Region. Amurian Zoological Journal 1(1): 6–16. [In Russian]
Rogatnykh D.Yu., Yakubovich V.S., Kurenshchikov D.K. 2013. The characteristic of seasonal dynamics of life-form spectrum of ground beetles (Coleoptera, Carabidae) in secondary forest of Bolshehehtsirsky reserve, Khabarovskiy Krai, Russia. Euroasian Entomological Journal 12(3): 271–277. [In Russian]
Sasakawa K. 2007. Taxonomic studies on the Bembidion («Cillenus») complex (Coleoptera: Carabidae): A revision of the subgeneric taxonomy and description of a new species from Japan. Zootaxa 1575(1): 35–45. DOI: 10.11646/ZOOTAXA.1575.1.2
Shilenkov V.G. 1979. New information on the fauna of ground beetles (Coleoptera, Carabidae) of the Southern Baikal region. In: Beetles of the Far East and Eastern Siberia (new data on fauna and systematics). Vladivostok. P. 36–57. [In Russian]
Shilenkov V.G. 1982. Some notes on South Siberian Trechini (Coleoptera, Carabidae) with descriptions of three new species. Annales Historico-Naturales Musei Nationalis Hungarici 74: 85–91.
Shilenkov V.G., Anistshenko A.V. 1999. About new findings of carabids (Coleoptera, Carabidae) in the Baikal Siberia. Proceedings of the Faculty of Biology and Soil Science of the Irkutsk State University 1: 15–19. [In Russian]
Shilenkov V.G., Pankratov A.A. 2011. Features of the vertical distribution of ground beetles (Coleoptera, Carabidae) on the northern macroslope of the Khamar-Daban ridge. Bulletin of Irkutsk State University. Series «Biology. Ecology» 4(4): 146–148. [In Russian]
Sota T., Hayashi M. 2007. Comparative historical biogeography of Plateumaris leaf beetles (Coleoptera: Chrysomelidae) in Japan: interplay between fossil and molecular data. Journal of Biogeography 34(6): 977–993. DOI: 10.1111/j.1365-2699.2006.01672.x
Suenson E. 1957. Trechinae from the Far East with description of new species collected by E. Suenson. Entomologiske Meddelelser 28: 84–96.
Sundukov Yu.N. 2001. New data on the carabid fauna (Coleoptera, Carabidae) of the Russian Far East. Entomological Review 81(6): 729–732.
Sundukov Yu.N. 2003. Ground beetle fauna (Coleoptera, Caraboidea) of Sikhote-Alinsky Nature Reservation. Euroasian Entomological Journal 2(2): 109–115. [In Russian]
Sundukov Yu.N. 2009. Family Carabidae – Ground beetles. In: S.Yu. Storozhenko (Ed.): Insects of Lazovsky State Nature Reserve. Vladivostok: Dalnauka. P. 88–109. [In Russian]
Sundukov Yu.N. 2013. An annotated catalogue of the ground beetles (Coleoptera: Caraboidea) of Sikhote-Alin. Vladivostok: Dalnauka. 271 p. [In Russian]
Sundukov Yu.N. 2017. The ground beetles (Coleoptera, Carabidae) of the Yuri Island, Southern Kuriles. A.I. Kurentsov's Annual Memorial Meetings 28: 101–110. [In Russian]
Sundukov Yu.N. 2019. The main stages in the formation of the ground beetle fauna (Coleoptera, Carabidae) of the Sikhote-Alin, with endemics taken as an example. 1. Characteristics of taxa. Entomological Review 99(8): 1128–1144. DOI: 10.1134/S0013873819080086
Sundukov Yu.N. 2019. The ground beetles (Coleoptera, Carabidae) of the Polonskogo Island, Southern Kuriles. A.I. Kurentsov's Annual Memorial Meetings 30: 140–152. DOI: 10.25221/kurentzov.30.12 [In Russian]
Sundukov Yu.N. 2020. The ground beetles (Coleoptera, Carabidae) of the Sakhalin Botanical Garden, Yuzhno-Sakhalinsk. A.I. Kurentsov's Annual Memorial Meetings 31: 131–140. DOI: 10.25221/kurentzov.31.11 [In Russian]
Sundukov Yu.N., Makarov K.V. 2013. The ground beetles (Coleoptera, Carabidae) of Shikotan Island, Kuril Islands, Russia. Euroasian Entomological Journal 12(4): 339–348. [In Russian]
Sundukov Yu.N., Makarov K.V. 2019. Ground beetles (Coleoptera: Carabidae) of the Russian Far East: Additions and corrections to the Catalogue of Palaearctic Coleoptera, Volume 1 (2017). Invertebrate Zoology 16(3): 283–304. DOI: 10.15298/invertzool.16.3.07
Sundukov Yu.N., Makarov K.V. 2019. The Dokuchaev Mountain Ridge as the main faunal refugium of Kunashir Island. A.I. Kurentsov's Annual Memorial Meetings 30: 63–79. DOI: 10.25221/kurentzov.30.5 [In Russian]
Taira A. 1990. Formation of Japanese Archipelago. Tokyo: Iwanami-shinsho. 72 p. [In Japanese]
Tojo K., Sekiné K., Takenaka M., Isaka Y., Komaki Sh., Suzuki T., Schoville S.D. 2017. Species diversity of insects in Japan: Their origins and diversification processes. Entomological Science 20(1): 357–381. DOI: 10.1111/ens.12261
Uéno S.I. 1954. A list of adephagous beetles from the Ozegahara Moor, with the descriptions of new species. Scientific Researches of the Ozegahara Moor. Tokyo: Japan Society for the Promotion of Science. P. 718–726.
Uéno Sh.I. 1960. Alpine trechids from Hokkaido, Japan. I. The Japanese representative of the subgenus Trechus. Memoirs of the College of Science, University of Kyoto (series B) 27: 133–137.
Uéno Sh.I. 1982. Origin and dispersal of the Trechina in East Asia (Coleoptera: Carabidae). Entomologia Generalis 8(1): 71–78.
Uéno Sh.I. 1992. Two new Epaphius (Coleoptera, Trechinae) from the Kushiro Moor, Northeast Japan. Elytra 20: 1–10.
Uéno Sh.I., Lafer G.Sh. 1994. Two relatives of Trechus nakaguroi (Coleoptera, Trechinae) with notes on the Trechus fauna of northeast Asia. Bulletin of the National Science Museum, Tokyo (series A) 20: 111–126.
Vertyankin A.V. 2014. Preliminary data on the ground beetles (Coleoptera, Carabidae) inhabits some forest biotopes in the southern part of Sakhalin Island. A.I. Kurentsov's Annual Memorial Meetings 25: 163–175. [In Russian]
Vertyankin A.V. 2015. The population of ground beetles (Coleoptera, Carabidae) in the valley of Lyutoga River, South Sakhalin. A.I. Kurentsov's Annual Memorial Meetings 26: 137–143. [In Russian]
Vertyankin A.V., Shabalin S.A. 2013. Provisional data on the population and domination of ground beetles (Coleoptera, Carabidae) in the meadows of Southern Sakhalin Island. A.I. Kurentsov's Annual Memorial Meetings 24: 179–188. [In Russian]
Watanabe T. 1989. The Coleoptera of Miyagi Prefeacture, Japan. Tokyo: Japanese Society of Coleopterology. 365 p.
Yahiro K., Yano K. 1997. Ground beetles (Coleoptera, Caraboidea) caught by a light trap during ten years. Esakia 37: 57–69.
Yoshimatsu S.I., Ito N., Nakatani Y., Yoshitake H. 2018. A list of ground beetles (Insecta: Coleoptera: Caraboidea) in Dr. Kazuo Tanaka collection preserved in the Insect Museum of Institute for Agro-Environmental Sciences, NARO. Bulletin of National Institute for Agro-Environmental Sciences 39: 15–191.
Yoshitake K., Kurihara T., Yoshimatsu Sh., Nakatani T., Yasuda K. 2011. A list of carabid specimens (Insecta: Coleoptera) collected by the late Dr. Akinobu Habu preserved in the Insect Museum of the National Institute for Agro-Environmental Sciences. Bulletin of National Institute for Agro-Environmental Sciences 28: 1–327.
Zerche L. 2004. Revision der Gattung Aegialites Mannerheim (Coleoptera: Salpingidae: Aegialitinae). Stuttgarter Beiträge zur Naturkunde, Serie A (Biologie) 666: 1–116.