ISSN 0013-8738, Entomological Review, 2014, Vol. 94, No. 6, pp. 846–879. © Pleiades Publishing, Inc., 2014. Original Russian Text © L.A. Akhmetova, А.V. Frolov, 2014, published in Entomologicheskoe Obozrenie, 2014, Vol. 93, No. 2, pp. 403–447.

A Review of the Scarab Tribe (Coleoptera, ) of the Fauna of Russia L. A. Akhmetova and А. V. Frolov Zoological Institute, Russian Academy of Sciences, St. Petersburg, 199034 Russia e-mail: [email protected] Received January 10, 2014

Abstract—An annotated list of 183 species of the dung-beetle tribe Aphodiini of the Russian fauna is given. The highest species diversity is characteristic of the southern mountainous regions of the country, the richest local fau- nas being known from the Lower Volga area. The greatest numbers of species are known from the steppe and broadleaved forest zones and from the nemoral types of altitudinal zonation of vegetation. The bulk of the fauna is formed by the species with wide ranges; 111 species occur in two or more landscape zones or altitudinal zonation types. Six groups of faunistically similar zoogeographic provinces are distinguished within the territory of Russia. DOI: 10.1134/S0013873814060074

Scarab of the tribe Aphodiini is a large, The regional faunistic lists, new distribution records world-wide distributed group of . The world of the aphodiines in Russia and adjacent territories, fauna comprises about 30 genera and over 2000 spe- new species descriptions, and diagnostic keys to some cies, including about 13 genera and 700 species in the groups and for some regions are available in the works Palaearctic Region (Dellacasa, 1988; Dellacasa, Della- of Medvedev (1965, 1976, 1979), Medvedev and Er- casa, 2006). Most aphodiines are coprophages feeding molenko (1969), Nikritin (1969а, 1969б), Medvedev on herbivore dung. In temperate regions including and Nikritin (1974), Berlov (1979, 1985, 1989), Niko- Russia, aphodiines dominate dung-beetle communities lajev (1987, 1998), Isajev (1995), Arzanov et al. both in terms of species richness and abundance. The (1996), Kabakov and Frolov (1996), Nikitsky et al. type genus of the aphodiines, Hellwig (sensu (1996), Novikov (1996, 1998), Kabakov (1998), lato), is the largest scarab beetle genus in Russia and Zinchenko (1999, 2003а, 2003б, 2011), Zinchenko one of the largest genera of insects. et al. (2002), Shokhin (2002, 2005, 2007), Gusakov (2004, 2006), Bezborodov and Berlov (2005), The earliest records of the aphodiines of the Rus- Makarov et al. (2009), Bezborodov (2009), Tarasov sian Empire are available in works of Faldermann (2008), Shabalin and Berlov (2009), Zinchenko and (1835a, 1835b, 1838), Gebler (1848), Ménétriés Kyzyl-ool (2010), Shabalin and Bezborodov (2012), (1832, 1849), and Motschulsky (1860). In addition to Zinchenko and Bezborodov (2013), as well as the au- the lists of known species collected in the Caucasus, thors of the present work (Frolov, 1995, 1996, 1997, Transcucasus, and West Siberia, they include descrip- 1999, 2001a, 2001b, 2001c, 2002, 2009; Akhmetova tions of new species. Reitter (1893) summarized the and Frolov, 2008a, b; 2009; Akhmetova, 2006, 2009; data about the Palaearctic scarab beetles. His work Frolov and Akhmetova, 2006, 2013). contains identification keys and numerous descriptions of new species largely from the Caucasus and Middle Despite the rather large number of publications Asia. Aphodiines of Southern Russia and Middle about the aphodiines of Russia, there has been no Asia have been actively studied by Russian coleo- work providing a comprehensive account of the taxo- pterologists in the late XIX and the early XX century nomic composition and geographical distribution of (D. Koshantschikow, 1891, 1894a, 1894b, 1894c, the species of our fauna. Available monographs 1894d; Semenov, 1898a, 1899, 1904, 1905; Semenov (Schmidt, 1922; Balthasar, 1964) are obsolete in terms and Medvedev, 1928; W. Koshantschikov, 1911а, of classification, nomenclature, and distribution data. 1911b, 1912, 1913, 1916). Olsoufieff (1918) summa- Furthermore, they are largely compilations of older rized the available data on the Caucasian copro- literature. Extensive materials from the collection of phagous species. the Zoological Institute of Russian Academy of

846 A REVIEW OF THE SCARAB BEETLE TRIBE 847

Fig. 1. Map of distribution of relative species richness of Russian Aphodiini fauna. Mean number of species per territory unit: 1, 1–10; 2, 11–20; 3, 21–30; 4, 31–40; 5, 41–50; 6, 51–60; 7, 61–70; 8, 71–80.

Sciences (ZIN), accumulated during many decades, tion coefficients. Average linkage was used as a clus- remained mostly unstudied. tering method (Fig. 7.)

MATERIALS AND METHODS Classification and nomenclature of the Aphodiini follow Medvedev and Nikritin (1970), Nikolaev This work is based on the rich collection of ZIN, in- (1979), and Kabakov and Frolov (1996): Aphodius is cluding vast material of more than 60 species collected treated sensu lato, including Heptaulacus Mulsant, by the authors in Lower Volga Region and the south- Sugrames Reitter and Mothon Semenov et Medvedev ern Far East. In addition, the materials from the fol- as subgenera. lowing institutions and private collections have been studied: Moscow Pedagogical State University, Mos- Genus Aphodius Hellwig, 1798 cow, Zoological museum, Lomonosov Moscow State University, Moscow, A.V. Ivanov’s collection, Ekate- Aphodius Illiger, 1798. rinburg, Soil biology institute FEB RAS, Vladivostok. Subgenus Acanthobodilus Dellacasa, 1983 The numbers of specimens of many studied species are approximate since not all the materials available in the 1. Aphodius (Acanthobodilus) immundus Creu- Zoological Institute, specifically those stored on cotton tzer, 1799. layers and in vials with alcohol, have been processed The species occurs throughout Europe, in Morocco, and integrated into the main collection. Egypt, the Caucasus, the Transcaucasus, Asia Minor, The map of distribution of relative species richness Syria, Kazakhstan, Middle and Central Asia. In Rus- in the territory of Russia (Fig. 1) was generated with sia, it occurs from western borders up to southern ArcGIS software. For special zoogeographic regioni- Yakutia. In the ZIN collection, there are 500 spms. zation, the scheme of the general zoogeographic sub- from the majority of the regions of the country. divisions of the Palaearctic Region (Emeljanov, 1974) was used. As the operational zoogeographic units, The species is common in the major part of its range provinces were used. In the cases when a province and prefers open biotopes. The beetles feed on horse, completely fits the studied area, its complete fauna cow and donkey dung; they are attracted to light and was included in the analysis. In the cases when only occur from April to September. a part of a province was situated in the territory of 2. Aphodius (Acanthobodilus) languidulus Russia, only whose the species occurring within this A. Schmidt, 1922. part were utilized. Provincial aphodiine faunas were compared by constructing similarity dendrograms The distribution range of this species includes based on pairwise calculations of Jaccard and correla- North-Eastern China, Korean Peninsula and Japan. In

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Russia, the species occurs in southern Khabarovsk tains of Middle Asia; it was imported to North Amer- Terr. and Primorskii Territory. It was also reported ica. In Russia, it is distributed throughout the Euro- from Amur Region (Berlov, 1989). In ZIN collection, pean Part and occurs in south of West Siberia. In the there are 20 spms. ZIN collection, there are 100 spms. from Leningrad, Yaroslavl, Nizhny Novgorod, Moscow, Kursk, Li- The species occurs in open biotopes, pastures, in petsk, Saratov, Rostov provinces, Krasnodar Terr., cow dung. The beetles fly in July and August. Adygea, Stavropol Terr., Daghestan, Kirovsk Prov., Subgenus Mulsant, 1842 Tatarstan and Bashkiria. A coprophagous species feeding on cow, horse, 3. Aphodius (Acrossus) bimaculatus (Laxmann, donkey and sheep dung. 1770). The species occurs in Central and Eastern Europe, 7. Aphodius (Acrossus) planicollis Reitter, 1890. North and Eastern Kazakhstan, Kyrgyzstan. In Russia, This species is endemic to the Caucasus. In Russia, it is known from a few localities in the European Part it is known from a few localities in the North Cauca- and West Siberia (up to Krasnoyarsk in the north-east) sus. In the ZIN collection, there are 10 spms. from (Kabakov and Frolov, 1996). In the ZIN collection, Krasnodar and Stavropol territories and Severnaya there are 200 spms. from Leningrad, Pskov, Smolensk, Osetia. Yaroslavl, Vladimir, Moscow, Saratov, Volgograd, Forest dweller, the beetles are active from May to Rostov provinces, Krasnodar Terr., Astrakhan, Sa- July. mara, Orenburg, Novosibirsk, Tomsk provinces, Altai Terr., and Khakassia. 8. Aphodius (Acrossus) rufipes (Linnaeus, 1758). The species is included in the Red Book of the Rus- The species occurs in most of Europe, except for sian Federation (Nikitsky, 2001). The beetles and lar- extreme north, in North Africa (Tunisia), the Tran- vae feed on horse dung (Frolov and Akhmetova, scaucasus, Kazakhstan, and Middle Asia; it was im- 2006). ported to North America. In Russia, it is widespread. In the ZIN collection, there are 400 spms. from major- 4. Aphodius (Acrossus) binaevulus Heyden, ity of regions of Russia. 1887. The species occurs in cow and horse dung and pre- The species is known from North Korea and North- fers forest biotopes. It is common in the major part of Eastern China. In Russia, it occurs in Amur Region its range, and is often attracted to light. and Primorskii Territory. In the ZIN collection, there are 60 spms. from Amur Prov., Khabarovsk and Pri- 9. Aphodius (Acrossus) superatratus Nomura morsky territories. et Nakane, 1951.

5. Aphodius (Acrossus) depressus (Kugelann, —A. arsenjevi Berlov, 1989. 1792). The species occurs in North Korea and Japan. In The species is widely distributed in Europe; it also Russia, it is known from a few localities in Amur Re- occurs in the Caucasus, Asia Minor, Kazakhstan, gion and Primorskii Territory. In the ZIN collection, Middle Asia, Mongolia, and China. In Russia, it oc- there are 8 spms. from Primorskii Territory. curs throughout the country. In the ZIN collection, Subgenus Aganocrossus Reitter, 1895 there are 300 spms. from the majority of regions. The species occurs in cow, horse and sheep dung, 10. Aphodius (Aganocrossus) urostigma Harold, less frequently in the dung of wild pigs, deer, roedeer, 1862. and hares. It prefers forest biotopes. The species is widely distributed outside Russia in South and South-Eastern Asia. In Russia, it is known 6. Aphodius (Acrossus) luridus (Fabricius, 1775). from a few findings from the southernmost Primorsky The species occurs in Europe, North Africa, the Terr. (Shabalin and Berlov, 2008). The beetles were Transcaucasus, Asia Minor, Kazakhstan, the moun- collected in cow dung in September.

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Subgenus A. Schmidt, 1913 16. Aphodius (Agolius) montanus Erichson, 1848. 11. Aphodius (Agoliinus) amurensis Iablokov, The species is distributed in the mountains of Cen- 1972. tral and Eastern Europe, and it was recently found in the South Urals (Zinchenko, 2011). In the ZIN collec- The species is known from two localities in South- tion, there are no specimens of this species from Rus- ern Sikhote Alin: Shkotovo Distr., Pejshula Village, sia. and Lazo Nature Reserve, the pass to Uglovaja Bay (Yablokov-Khnzorian, 1972; Akhmetova, Frolov, Subgenus Agrilinus Mulsant et Rey, 1870 2009). In the ZIN collection, there are 10 spms. from the Lazo Nature Reserve. 17. Aphodius (Agrilinus) aleutus Eschscholtz, 1822. The species occurs in Japanese deer dung in mixed forests. —A. ursinus Motschulsky, 1845. The species was recorded by Berlov (1989) from 12. Aphodius (Agoliinus) guttatus Eschscholtz, Yakutia and Kamchatka as A . aleutus ursinus, and by 1823. Shabalin and Bezborodov (2012) from the northern The species is known from Canada (Alberta, British Kuril Islands. The range of the nominative subspecies Columbia, Newfoundland, Ontario, Quebec) and USA includes the western part of Canada (South-West (Alaska, Massachusetts) (Gordon, Skelley, 2007). It Alberta, South British Columbia) and the West USA was recently found in Kamchatka (Akhmetova, Frolov, (West Washington, Oregon, and the northern coast of 2009): Karaginsky Isl. (3 spms. in ZIN collection). California) (Gordon, Skelley, 2007). In the ZIN collec- tion, there are no specimens from Russia. 13. Aphodius (Agoliinus) piceus Gyllenhal, 1808. 18. Aphodius (Agrilinus) ater (De Geer, 1774). The species range is not yet well known. In the lit- erature, there are records of it from Central and South The species is distributed in Europe, North Africa, Europe (Dellacasa, Dellacasa, 2006), Ulyanovsk Prov. Kazakhstan, Middle Asia (except for deserts), and Mongolia. In Russia, it occurs throughout the country. (Isajev, 1995), Tuva, Irkutsk Prov., Buryatia and Za- In the ZIN collection, there are 100 spms. from the baykalskii Terr. (Berlov 1989). In the ZIN collection, majority of the regions of Russia. there are 500 spms. from Murmansk, Arkhangelsk, and Tyumen provinces. The species occurs in open biotopes and in forests, in cow and sheep dung. It was also found in gopher This coprophagous species inhabits forests and oc- (Nikolajev, 1987) and marmot (Zinchenko, 1999) curs in the dung of horses and wild ungulates from holes. March to July. 19. Aphodius (Agrilinus) breviusculus Motschul- 14. Aphodius (Agoliinus) satunini Olsoufiev, sky, 1866. 1918. The species is known from the Kuril islands (Itu- The species is known only from the original de- rup), as well as from Japan and the Korean Peninsula scription from the Sochi environs (Olsoufieff, 1918). (Berlov 1989). In the ZIN collection, there are no Its type depository is unknown. Shokhin (2007) sug- specimens of this species from Russia. gested that this name might be a synonym of A. (Agri- 20. Aphodius (Agrilinus) constans Duftschmid, linus) fasciatus (Olivier, 1789). 1805. Subgenus Agolius Mulsant et Rey, 1870 The species occurs in Central and South Europe, Ukraine, the Caucasus, the Transcaucasus, Asia Mi- 15. Aphodius (Agolius) falcispinis W. Ko- nor, and Turkmenistan. It was reported from Rostov shantschikov, 1912. Prov. of Russia (Shokhin, 2007). The species is known from East Kazakhstan, North- Western China, South Siberia, and Mongolia. In the 21. Aphodius (Agrilinus) convexus Erichson, ZIN collection, there are 10 spms. from Altai and Za- 1848. baykalskii Terr. —A. emerichi Reitter, 1892.

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According to Dellacasa and Dellacasa (2006), this try in the taiga zone; in Dzungarian Ala Tau and Tien- species is widely distributed in the Palaearctic Region; Shan it occurs in the coniferous forest altitudinal zone. however its range requires clarification. In the ZIN collection, there are 300 spms. from Kare- lia, Murmansk, and Arkhangelsk provinces, the Re- 22. Aphodius (Agrilinus) fasciatus (Olivier, 1789). public of Komi, Tyumen Prov., the Republic of Altai, —A. putridus (Herbst, 1789); Krasnoyarsk Terr., Tuva, Buryatia, Yakutia, Amur, Magadan and Kamchatka provinces and Khabarovsk —A. uliginosus Hardy, 1847. Terr. The species occurs in Northern, Central and Eastern Europe, Eastern Kazakhstan, Mongolia, and it was A coprophagous species occurring in ungulate dung. also imported to North America. In Russia, it occurs 27. Aphodius (Agrilinus) nemoralis Erichson, throughout the country except for steppe and desert 1848. zones. In the ZIN collection, there are 100 spms. from Karelia, Leningrad, Yaroslavl, Penza, Kirovsk, Chely- The species occurs in mixed and deciduous forests abinsk, Tyumen, Tomsk provinces, Altai and Kras- in Northern, Central and Eastern Europe. In Russia, it noyarsk Territories, Tuva, Irkutsk Prov., Buryatia, is known from the western border up to Volga Region. Yakutia, Amur and Sakhalin provinces, Khabarovsk In the ZIN collection, there are 30 spms. from Lenin- Ter. grad, Moscow, and Ulyanovsk provinces. The species mostly inhabits forest biotopes. It oc- A forest dweller feeding on the dung of wild ungu- curs in wild ungulate dung, as well as in cow and lates. The adults are active from April to June. horse dung. 28. Aphodius (Agrilinus) nikolajevi Berlov, 1989. 23. Aphodius (Agrilinus) hasegawai Nomura et Nakane, 1951. The species was described from South Primorskii The species is distributed in Japan. It was also re- Territory and also reported from Zabaykalskii Terr. ported from South Sakhalin and the Kunashir islands and Amur Prov. (Berlov 1989; Bezborodov and Ber- (Shabalin and Bezborodov, 2012). In the ZIN collec- lov, 2005). In the ZIN collection, there are no speci- tion, there are no specimens from Russia. mens of this species. 24. Aphodius (Agrilinus) inexpectatus Balthasar, 29. Aphodius (Agrilinus) pratensis Nomura et Na- 1935. kane, 1951. The species occurs in North Korea and Japan (Hon- Reported from Sakhalin (Berlov, 1989) and Primor- shu Island) (Stebnicka and Galante, 1991). In Russia, skii Territory (Gusakov, 2009). In the ZIN collection, it is known from Amur Region (Bezborodov and Ber- there are no specimens of this species from Russia. lov, 2005) and Primorskii Territory. In the ZIN collec- tion, there are 8 spms. from South Primorskii Terri- 30. Aphodius (Agrilinus) tenax Balthasar, 1932. tory. The species was described from Sino-Tibetan 25. Aphodius (Agrilinus) isajevi Kabakov, 1994. Mountains. It was reported from Vladivostok environs (Balthasar, 1964) and the lower Amur River (Berlov, The species is known from a few localities in the 1979). In the ZIN collection, there are no specimens of lower Volga and Don basins. In the ZIN collection, this species from Russia. there is a type series from Ulyanovsk Prov. The species is a specialist nidicolous occurring in 31. Aphodius (Agrilinus) uniformis Waterhouse, steppe biotopes in native colonies of marmots (Mar- 1875. mota bobac); it is not found in the reacclimatized The species occurs in south Primorsky Terr., as well marmot colonies (Isajev, 1995). as in Sakhalin and Kunashir islands. In the ZIN collec- 26. Aphodius (Agrilinus) lapponum Gyllenhal, tion, there are 100 spms. from Russia. 1806. The species occurs in open biotopes, pastures, The species is distributed from North Europe to in cow dung; it is common in the major part of its Kamchatka. In Russia, it occurs throughout the coun- range.

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Subgenus Alocoderus A. Schmidt, 1913 Subgenus Amidorus Mulsant et Rey, 1870 32. Aphodius (Alocoderus) digitalis D. Koshan- 36. Aphodius (Amidorus) alagoezi Olsoufiev, tschikow, 1894. 1918. In Russia, it is known from only 2 localities in the The species was described from Armenia (Aragats vicinity of Dosang Station in Astrakhan Prov. Mountain). In Russia, it is known from Elbrus (Tara- (Shokhin, 2007; Akhmetova and Frolov, 2008). sov, 2008).

33. Aphodius (Alocoderus) hydroсhaeris (Fabri- 37. Aphodius (Amidorus) obscurus (Fabricius, cius, 1798). 1792). —A. hydroсhoeris (Fabricius, 1798). The species occurs in Central and South Europe, the Carpathian Mountains, the Caucasus, and Asia Minor. The distribution range of this species includes Cen- In Russia, it is known from the North Caucasus. In the tral and South Europe, North Africa, Armenia, Azer- ZIN collection, there are 20 spms. from Krasnodar baijan, Middle East, and Kazakhstan. In Russia, it is Terr., Adygea, Karachay-Cherkessia, Kabardino Bal- mostly distributed in the steppe and desert zones of the kariya, and Daghestan. North Caucasus and the Volga Region. In the ZIN collection, there are 50 spms. from Voronezh, Rostov, The species occurs in cow dung in the mountains, Volgograd, Astrakhan provinces, Adygea and Da- up to 2000-3000 m a.s.l. in the Caucasus. ghestan. 38. Aphodius (Amidorus) thermicola Erichson, This coprophagous species mostly occurs in horse 1848. dung. The distribution range of this species includes Cen- 34. Aphodius (Alocoderus) rufus (Moll, 1782). tral, Southern and Eastern Europe, the Caucasus, the Transcaucasus, and Asia Minor. Tarasov (2008) re- —A. scybalarius (Fabricius, 1781). ported it from Krasnodar Terr., North Osetia and The species is distributed in Europe, the Transcau- Daghestan. In the ZIN collection, there are 6 spms. casus, and Western Kazakhstan. In Russia, it occurs from Krasnodar Terr. throughout the country up to Irkutsk Prov. in the east. Subgenus Ammoecius Mulsant, 1842 In the ZIN collection, there are 200 spms. from Kare- lia, Leningrad, Novgorod, Pskov, Tver, Kirovsk, Yaro- 39. Aphodius (Ammoecius) brevis Erichson, 1848. slavl provinces, Chuvashia, Moscow, Ryazan, Penza, The species is distributed in Central and Eastern Volgograd provinces, Adygea, Karachay-Cherkessia, Europe, the Caucasus and the Transcaucasus. In Rus- Daghestan, Ulyanovsk, Samara provinces, Bashkiria, sia, it occurs in the European Part and South Siberia Sverdlovsk, Chelyabinsk, Tomsk provinces, the Re- (up to Transbaikal in the east). In the ZIN collection, public of Altai, and Irkutsk Prov. there are 70 spms. from Leningrad, Saratov provinces, The species occurs from May to September in open Krasnodar Terr., Samara, Kurgan, Tyumen, Tomsk, biotopes, pastures, in cow and horse dung. Irkutsk provinces, the Republic of Altai. The species occurs in dry cow, horse and, rarely, elk 35. Aphodius (Alocoderus) sordidus (Fabricius, dung. 1775). The species is known from North and Central Subgenus Aparammoecius Petrovitz, 1958 Europe, North Kazakhstan, Mongolia, China, Korean 40. Aphodius (Aparammoecius) leisteri Medve- Peninsula, and Japan. In Russia, it occurs throughout dev, 1968. the country from Leningrad Prov. to Kunashir Island. In the ZIN collection, there are 300 spms. from the This species is endemic to the Caucasus. It was de- majority of the regions of the country. scribed from Abkhazia. In the ZIN collection, there are 2 spms. from North Osetia (Alagir Distr., Yu.E. Ko- The species occurs in cow dung in open biotopes. marov leg.).

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Subgenus Aphodaulacus W. Koshantschikov, 1911 46. Aphodius (Aphodius) fimetarius (Linnaeus, 1758). 41. Aphodius (Aphodaulacus) kizeritskyi Frolov, 2002. The species is widely distributed in Europe, North Africa, Kazakhstan, Middle and Central Asia, im- The species was described from Turkmenistan. It is ported to North America and Australia. In Russia, it known from a few localities in Karakum and Caspian occurs throughout the country up to Eastern Siberia in Lowland deserts. In the ZIN collection, there are 150 the east; in the ZIN collection, there are 300 spms. spms. from Astrakhan Prov. (Dosang environs). from the majority of the regions of the country. The species occurs in horse dung. In Russia, imagoes are active from April to October 42. Aphodius (Aphodaulacus) koltzei Reitter, occurring in open biotopes, pastures, in cow, sheep 1892. and horse dung. —A. medvedevi Nikritin, 1969; 47. Aphodius (Aphodius) foetens (Fabricius, —A. kurenzovi Nikritin, 1969. 1787). The species is known from Primorskii Territory, —A. aestivalis Stephens, 1839. Amur Region and North-Eastern China. In the ZIN The species is widely distributed in Europe, also collection, there are 11 spms. from Primorskii Terri- occurs in the Caucasus, the Transcaucasus, Asia Mi- tory. nor, and North Kazakhstan. In Russia, it is known from the European part and Siberia (up to Yakutia in 43. Aphodius (Aphodaulacus) nigrotessellatus the east). In the ZIN collection, there are 100 spms. Motschulsky, 1866. from the majority of the regions of the country. The distribution range of this species includes In Russia, the beetles occur in open biotopes, North-Eastern China and Japan. In Russia, it is known pastures, in cow and horse dung, from May to Octo- from Amur Region and Primorskii Territory. In the ber. ZIN collection, there are 72 spms. from Zabaykalskii, Khabarovsk and Primorsky territories. 48. Aphodius (Aphodius) swaneticus Reitter, 1892. Subgenus Aphodiellus A. Schmidt, 1913 The species is endemic to the Caucasus. In the ZIN 44. Aphodius (Aphodiellus) impunctatus Water- collection, there are 9 spms. from Krasnodar Terr. and house, 1875. Adygea. The species occurs in Japan, the Korean Peninsula, in North-Eastern China (from Beijing to Harbin). In Subgenus Biralus Mulsant et Rey, 1870 Russia, it is distributed in South Primorskii Territory. 49. Aphodius (Biralus) menetriesi Ménétriés, In the ZIN collection, there are 17 spms. 1849. The species occurs in pastures, in cow dung. A few The species is distributed in South-Eastern Europe, specimens were collected in a forest under rotten the Transcaucasus, South Kazakhstan, West and Mid- mushrooms. dle Asia. In the ZIN collection, there are 10 spms. from Volgograd and Astrakhan provinces. Subgenus Aphodius Hellwig, 1798 45. Aphodius (Aphodius) conjugatus (Panzer, The species occurs in open biotopes, in cow dung; it 1795). reaches middle altitudes in the mountain regions. The species occurs in Central and South Europe, the 50. Aphodius (Biralus) satellitus (Herbst, 1789). Caucasus, the Transcaucasus, and Asia Minor. In Rus- The species occurs in Central and South Europe, sia, it is known from the North Caucasus. In the ZIN North Africa, the Transcaucasus, Middle Asia. In Rus- collection, there are 15 spms. from Rostov Prov., sia, it is mainly distributed in North the Caucasus and Krasnodar Terr. and Daghestan. the Volga Region. In the ZIN collection, there are A coprophagous species occurring in cow dung. 40 spms. from Voronezh, Saratov, Volgograd, Rostov

ENTOMOLOGICAL REVIEW Vol. 94 No. 6 2014 A REVIEW OF THE SCARAB BEETLE TRIBE 853 provinces, Krasnodar and Stavropol territories, Astra- Astrakhan provinces, Krasnodar and Stavropol territo- khan and Samara provinces, and Daghestan. ries, Daghestan, Samara, Orenburg and Kurgan prov- inces. The species occurs in cow and horse dung, mostly in open biotopes. The species occurs in open biotopes, in cow dung. In Russia, the beetles are active from April to Subgenus Bodilus Mulsant et Rey, 1870 October.

51. Aphodius (Bodilus) gregarius Harold, 1871. 55. Aphodius (Bodilus) punctipennis Erichson, The species is distributed in South-Eastern Europe, 1848. Kazakhstan, Middle Asia, Mongolia and North-West The species occurs in Central, South and Eastern China. In Russia, it occurs in the steppe zone and the Europe, Egypt, the Transcaucasus, Iran, Afghanistan, forest-steppe subzone of the broadleaf forest zone South Kazakhstan and Middle Asia. In Russia, it is from Rostov Prov. up to southern West Siberia. In the distributed in the steppe zone and the forest-steppe ZIN collection, there are 60 spms. from Volgograd, subzone of the broadleaf forest zone from the western Astrakhan, Orenburg provinces and Khakassia. border to the Volga Region. In the ZIN collection, In the territory of Russia, the beetles occur in cow there are 90 spms. from Ryazan, Kursk, Belgorod, and horse dung from May to August. Penza, Volgograd, Rostov, Astrakhan, Samara prov- inces, Stavropol Terr., Daghestan and Bashkiria. 52. Aphodius (Bodilus) ictericus (Laicharting, 1781). 56. Aphodius (Bodilus) sordescens Harold, 1869. —A. nitidulus (Fabricius, 1792). The species occurs in North-Eastern Kazakhstan, Mongolia, and North China. In Russia, it is known The distribution range of this species includes the from the steppe zone of South Siberia, from the Trans- whole of Europe (except for the extreme north), North Urals to Transbaikal. In the ZIN collection, there are Africa, the Transcaucasus, Asia Minor, Iran and North 100 spms. from Chelyabinsk, Tomsk, and Kemerovo Kazakhstan. In Russia, it is widely distributed in the provinces, Altai and Krasnoyarsk territories, Khakas- European Part. In the ZIN collection, there are 200 sia, Tuva, Buryatia, Zabaykalskii Terr. spms. from Leningrad, Novgorod, Tver, Moscow, Ryazan, Kursk, Lipetsk, Saratov, Volgograd, Rostov Subgenus Calamosternus Motschulsky, 1859 provinces, Krasnodar Terr., Adygea, Astrakhan, Ki- rovsk, Samara provinces, Chechnia and Daghestan. 57. Aphodius (Calamosternus) granarius (Linna- eus, 1767). The species occurs in open biotopes, pastures in The species is distributed throughout Europe (except cow and horse dung from spring to late autumn. for the extreme north), West Asia, Kazakhstan, and Middle Asia; imported to North America. In Russia, it 53. Aphodius (Bodilus) longeciliatus Reitter, occurs from the western border up to Transbaikal Re- 1887. gion. In the ZIN collection, there are 200 spms. from The species occurs throughout Mongolia and in Kaliningrad, Leningrad, Tver, Nizhny Novgorod, Rya- China (Qinghai, Inner Mongolia provinces). It was zan, Belgorod, Voronezh, Volgograd provinces, Kras- reported from Buryatia (Berlov, 1989). nodar Terr., Kalmykia, Astrakhan Prov., Daghestan, Kirovsk and Samara provinces, Bashkiria, Tyumen 54. Aphodius (Bodilus) lugens Creutzer, 1799. Prov. and Zabaykalskii Terr. The species occurs in Central, South and Eastern The species is common in all the parts of its range. Europe, North Africa, the Transcaucasus, Asia Minor, It occurs in open biotopes in the dung of different Iran, Afghanistan, Kazakhstan and Middle Asia. In , in carrion, and in marmot holes. Russia, it is distributed in the steppe zone and the for- est-steppe subzone of the deciduous forest zone from 58. Aphodius (Calamosternus) sublimbatus Mo- the western border to the Trans-Urals. In the ZIN col- tschulsky, 1860. lection, there are 100 spms. from Tver, Kursk, Bel- The distribution range of this species includes gorod, Voronezh, Penza, Saratov, Volgograd, Rostov, North-Eastern China, the Korean Peninsula, Japan and

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Taiwan Island. In Russia, it is distributed in Primorskii 63. Aphodius (Chilothorax) comma Reitter, 1892. Territory and Amur Region. In the ZIN collection, The species occurs in North Kazakhstan, eastern there are 100 spms. from the Jewish Autonomous Middle Asia, and Mongolia. In Russia, it is known Province, Khabarovsk, and Primorsky territories. from South Siberia and the Amur Region. In the ZIN The species occurs in open biotopes, in cow and collection, there are 300 spms. from Tyumen, Novosi- horse dung. This is a common, locally abundant spe- birsk, Tomsk, and Kemerovo provinces, Altai and cies. Krasnoyarsk territories, the Republic of Altai, Khakas- sia, Tuva, Irkutsk Prov., Buryatia, Zabaykalskii Terr., 59. Aphodius (Calamosternus) trucidatus Harold, the Amur Prov., Yakutia, and Khabarovsk Terr. 1863. 64. Aphodius (Chilothorax) conspurcatus (Lin- —A. suturifer Reitter, 1892. naeus, 1758). The species occurs in South Europe, North Africa, The species is distributed in North and Central Transcaucasus, Middle East (Syria, Lebanon, Israel, Europe; it locally occurs in South Europe (Dellacasa, Iran, Iraq), Kazakhstan, Middle Asia, and Mongolia. Dellacasa, 2006). In the ZIN collection, there are 30 In Russia, the species is known from the Ciscaucasus spms. from Leningrad, Novgorod, and Yaroslavl prov- and Lower Volga Region (Arzanov et al., 1996; inces. Shokhin, 2007). In the ZIN collection, there are 2 spms. from Astrakhan Prov. (Lake Baskunchak and 65. Aphodius (Chilothorax) distinctus (Müller, Dosang). 1776). —A. inquinatus Herbst, 1783. Subgenus Carinaulus Tesar, 1945 The distribution range of this species includes the 60. Aphodius (Carinaulus) costatellus A. Schmidt, whole of Europe (except for the extreme north), North 1916. Africa, the Transcaucasus, Asia Minor, Kazakhstan, The species is known from several findings in the Middle Asia, Mongolia. It was imported to North Amur Region and Primorskii Territory. In the ZIN America. In Russia, it occurs throughout the country in collection, there are 8 spms. from the Amur Prov. and the European part, reaching Baikal region in the east. Primorskii Territory. In the ZIN collection, there are 300 spms. from the majority of the regions of the country. Subgenus Chilothorax Motschulsky, 1859 The species occurs in cow and horse dung, in mar- —Volinus Mulsant, 1870. mot holes, and also is attracted to light. In the territory of Russia the beetles are active from March to Octo- 61. Aphodius (Chilothorax) badenkoi Nikolajev, ber. It is a common, locally abundant species. 1987. The species is distributed in deserts of Middle Asia, 66. Aphodius (Chilothorax) exilimanus Kabakov, and the Northern Caspian Region. In Russia, it is 1994. known from single specimen from Astrakhan Prov. The species is known from a type series from Uly- (Dosang environs). anovsk Prov. (15 spms. in the ZIN collection). It is a specialist nidicolous species. All the specimens were 62. Aphodius (Chilothorax) clathratus Reitter, collected in the holes of Marmota bobac. 1892. The species is known from the Caucasus, the Tran- 67. Aphodius (Chilothorax) grafi Reitter, 1901. scaucasus, Turkey, Iran, and Middle Asia (Nikolajev, —A. kryzhanovskii Nikritin, 1969; 1987). Its distribution range needs clarification be- —А. schutovae Nikritin, 1969. cause some records of A. clathratus may refer to the very similar A. melanostictus. In Russia, it was found This species is widely distributed in Mongolia. In in Dosang environs (Astrakhan Prov.), in horse dung, Russia, it occurs in South Siberia from the Altai in fixed sands. Mountains to Transbaikal Region. In the ZIN collec-

ENTOMOLOGICAL REVIEW Vol. 94 No. 6 2014 A REVIEW OF THE SCARAB BEETLE TRIBE 855 tion, there are 100 spms. from the Republic of Altai, of Russia, the beetles are active from March to Octo- Khakassia, and Buryatia. ber. The species is common throughout its range.

68. Aphodius (Chilothorax) hahni Reitter, 1907. 74. Aphodius (Chilothorax) mongolaltaicus Niko- lajev, 1984. The species is known from a few localities in North- West Kazakhstan. In the ZIN collection, there is The species is known from East Kazakhstan and 1 spm. from Volgograd Prov. Mongolia. In Russia, it was found in South Altai. In the ZIN collection, there are 3 spms. from Chuya 69. Aphodius (Chilothorax) ivanovi Lebedev, Steppe. 1912. The species is distributed in the Middle Volga and 75. Aphodius (Chilothorax) paykulli (Bedel, Don basins (Medvedev, 1965). In the ZIN collection, 1798). there are 27 spms. from Penza and Ulyanovsk prov- —A. tessulatus Duftschmid, 1805. inces. The species occurs in the greatest part of Europe, The species occurs on sandy riversides. the Caucasus, the Transcaucasus and Asia Minor. In Russia, it is known from a few findings in North- 70. Aphodius (Chilothorax) jacobsoni W. Ko- Western Region and the North Caucasus. In the ZIN shantschikov, 1911. collection, there are 3 spms. from Kaliningrad and The species is distributed in Eastern Kazakhstan Leningrad provinces. and Mongolia. In Russia, it is known from South Sibe- ria. In the ZIN collection, there are 18 spms. from 76. Aphodius (Chilothorax) planus D. Koshan- Altai and Khakassia. tschikow, 1894.

71. Aphodius (Chilothorax) kerzhneri Nikolajev, —A. transvolgensis Semenov, 1898. 1984. The species is known from South Ukraine, North The species is distributed in Eastern Kazakhstan and Central Kazakhstan and Turkmenistan. In Russia, and Mongolia. In Russia, it is known from South Sibe- the species occurs in the steppe zone from the Ciscau- ria. In the ZIN collection, there are 22 spms. from casus to the south of West Siberia. In the ZIN collec- Altai. tion, there are 16 spms. from Saratov and Volgograd provinces, Krasnodar Terr., Orenburg and Novosibirsk The species occurs in gray marmot holes (Nikola- provinces. jev, Puntsagdulam, 1984; Zinchenko, 1999). The species occurs in gopher and marmot holes 72. Aphodius (Chilothorax) logunovi Zinchenko, (Isajev, 1995; Zinchenko, 1999). 2003. 77. Aphodius (Chilothorax) plustschewskii D. Ko- The species is known only from the type series from shantschikow, 1894. South-East Altai (3 spms., one of which is housed in the ZIN collection) (Zinchenko, 2003a). The species is distributed in the deserts of the North Caspian Region. In the ZIN collection, there are 100 73. Aphodius (Chilothorax) melanostictus spms. from Astrakhan Prov. W. Schmidt, 1840. A coprophagous species, feeding on horse dung. The species occurs in Central, South and Eastern Beetles are active in late fall. Europe, North Africa, West Asia, Kazakhstan, and Middle Asia. In Russia, it is distributed from western 78. Aphodius (Chilothorax) subpolitus Motschul- border to Transbaikal Region, mostly in the steppe sky, 1860. zone and the forest-steppe subzone of the deciduous The species is known only from the original de- forest zone. In the ZIN collection, there are 300 spms. scription from Siberia (without a more precise local- The species occurs in open biotopes, in the dung of ity) (Motschulsky, 1860). The type depository is un- domestic and in marmot holes. On the territory known.

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79. Aphodius (Chilothorax) sticticus (Panzer, 84. Aphodius (Colobopterus) indagator Manner- 1798). heim, 1849. —A. equestris (Panzer, 1798). The distribution range of this species includes The species occurs throughout Europe (except for Mongolia and North-Eastern China. In Russia, it is the extreme north), in the Trans-Caucasus, Asia Mi- known from Altai Mountains, south of East Siberia, nor, and Northern Iran. In Russia, it is widely distrib- Amur Region and Primorskii Territory. In the ZIN uted in the European part up to the South Urals Moun- collection, there are 45 spms. from Republic of Altai, tains. In the ZIN collection, there are 100 spms. from Irkutsk Prov., Buryatia, Zabaykalskii Terr., Amur Yaroslavl Prov., Chuvashia, Moscow, Ryazan, Kursk, Prov. and Primorsky Terr Belgorod, Penza, Samara provinces, Krasnodar Terr., 85. Aphodius (Colobopterus) notabilipennis Pet- and Bashkiria. rovitz, 1972. A forest dweller, feeding mostly on cow and horse The species is known from the Amur Region and dung. Primorskii Territory. In the ZIN collection, there are 100 spms. from Zabaykalskii Terr., the Jewish 80. Aphodius (Chilothorax) tanhensis Frolov, Autonomous Province, Khabarovsk and Primorsky 2001. territories. The species is known from Mongolia and the Altai Mountains. In the ZIN collection, there are 5 spms. 86. Aphodius (Colobopterus) propraetor Baltha- from Chuya steppe. sar, 1932. The distribution range of this species includes 81. Aphodius (Chilothorax) variicolor D. Koshan- North-Eastern China and the Korean Peninsula. In tschikow, 1894. Russia, it is widely distributed in southern Far East. In The species is distributed in the Caspian lowland the ZIN collection, there are 200 spms. from the Amur deserts. In the ZIN collection, there are 28 spms. from Prov., the Jewish Autonomous Province, Khabarovsk, Astrakhan Prov. and Primorsky territories, Sakhalin Prov. (Sakhalin and Kunashir islands). The beetles feed on horse dung; the larvae occur in barchan sands and apparently feed on the roots of A common, locally abundant coprophagous species cheat grass. occurring from April to August in cow and horse dung. Subgenus Colobopterus Mulsant, 1842 87. Aphodius (Colobopterus) quadratus Reiche, 82. Aphodius (Colobopterus) brignolii Carpaneto, 1847. 1973. —A. haroldianus Balthasar, 1932. The species is known from the North Caucasus, the Transcaucasus and North-Eastern Turkey. In the ZIN The species was described from Japan. In the ZIN collection, there are 11 spms. from Krasnodar Terr., collection, there are 6 spms. from Kunashir Island. Adygea, Karachay-Cherkessia and North Osetia. Subgenus Coprimorphus Mulsant, 1842 83. Aphodius (Colobopterus) erraticus (Linnaeus, 88. Aphodius (Coprimorphus) scrutator (Herbst, 1758). 1789). The species occurs in Europe (except for the ex- The species is distributed in Central and South treme north), West, Middle and Central Asia; it was Europe, Azores, the Caucasus, the Transcaucasus, imported to North America. In Russia, it is distributed South-West Asia. In Russia, it is known from the throughout the country. In the ZIN collection, there are northern foothills of the Caucasus. In the ZIN collec- 1000 spms. from the majority of the regions of Russia. tion, there are 10 specimens from Krasnodar, Stavro- pol territories and Daghestan. This eurybiont species is abundant throughout its range occurring mostly in open biotopes, in the dung The beetles mostly occur in open biotopes, in cow of domestic and wild animals. dung.

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Subgenus Coptochiroides Balthasar, 1938 A coprophagous species occurring mostly in cow dung, in open biotopes, from April to September. 89. Aphodius (Coptochiroides) subcostatus Kolbe, 1886 94. Aphodius (Esymus) pusillus (Herbst, 1789). The distribution range of this species includes The distribution range of this species includes al- North-Eastern China, Korean Peninsula and Japan. In most whole Europe (except for extreme north), the Russia, it occurs in southern Primorskii Territory; In Caucasus, the Transcaucasus, Kazakhstan, Middle the ZIN collection, there are 200 spms. Asia, Asia Minor, Mongolia, North China, Korean Peninsula, and Japan. In Russia, it is distributed Subgenus Erytus Mulsant et Rey, 1870 throughout the country. In the ZIN collection, there are 90. Aphodius (Erytus) aequalis A. Schmidt, 1907. 300 spms. from the majority of the regions. The species occurs in South Europe, North Africa, It is a common, locally abundant species feeding on the Caucasus, Transcaucasus, Turkey, Iran, Syria, cow, horse, and sheep dung. Kazakhstan, Middle and Central Asia. In the ZIN col- Subgenus Eudolus Mulsant et Rey, 1870 lection, there are 30 specimens from Volgograd, Astrakhan provinces and Stavropol Terr. Russia. 95. Aphodius (Eudolus) quadriguttatus (Herbst, 1783). The beetles of this species mostly occur in cow dung in late spring. In the Lower Volga Terr. The bee- The species is distributed in Europe, North Africa, tles are active at nights. West Asia, Kazakhstan, and Middle Asia. In Russia, it mostly occurs in the Vorga Region and the Ciscauca- 91. Aphodius (Erytus) klugi A. Schmidt, 1910. sus. In the ZIN collection, there are 100 spms. from The distribution range of this species includes Saratov, Volgograd, Astrakhan, Samara provinces and Southern Europe, North Africa, South-Western Asia, Krasnodar Terr. Kazakhstan, Middle Southern Asia (Hindustan Penin- A coprophagous species occurring in the dung of sula and Himalaya). It was reported from Rostov, Vol- different animals. It prefers arid biotopes with sandy gograd, Astrakhan provinces (Arzanov et al., 1996) and sandy clay soils. and Kalmykia (Shokhin, 2007). Subgenus Euorodalus Dellacasa, 1983 92. Aphodius (Erytus) pruinosus Reitter, 1892. 96. Aphodius (Euorodalus) coenosus (Panzer, The distribution range of this species includes North 1798). Africa, South-Western Asia, Southern Kazakhstan, The distribution range of this species includes al- and Middle Asia. It was reported from Kalmykia (Ar- most the whole of Europe (except for the extreme zanov et al., 1996) and Astrakhan Prov. (Shokhin, north), the Transcaucasus, Asia Minor, West Kazakh- 2007). stan, and Tajikistan. In the ZIN collection, there are 40 spms. from Leningrad, Nizhny Novgorod, Vo- Subgenus Esymus Mulsant et Rey, 1870 ronezh, Saratov, Volgograd and Astrakhan provinces. 93. Aphodius (Esymus) merdarius (Fabricius, A coprophagous species occurring in the dung of 1775). cows, horses, and wild ungulates. The species occurs in different biotopes: in Eastern Europe, in forests; The species is distributed in Europe (Dellacasa, Isajev (1995) recorded it from steppes; we collected it Dellacasa, 2006), North Africa, the Caucasus, on sandy soils on the Akhtuba riverside in Dosang the Transcaucasus, Asia Minor, Iran, Kazakhstan, environs (Astrakhan Prov.). and Middle Asia. In Russia, the species occurs from the European part to the South Urals Mountains. In Subgenus Eupleurus Mulsant, 1842 the ZIN collection, there are 100 spms. from Lenin- grad, Novgorod, Tver, Vologda, Yaroslavl, Nizhny 97. Aphodius (Eupleurus) antiquus Faldermann, Novgorod, Moscow, Ryazan, Lipetsk, Volgograd, 1835. Samara provinces, Krasnodar Terr., Daghestan and The species is known from South Siberia and Mon- Bashkiria. golia. In the ZIN collection, there are 50 spms. from

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Altai and Krasnoyarsk territories, the Republic of Al- In Russia, the species is mostly distributed in the tai, Khakassia, Buryatia, Irkutsk Prov., Zabaykalskii steppe zone to the Cis-Urals Region. In the ZIN collec- Terr., Yakutia, Amur Prov., and the Jewish Autono- tion, there are 30 spms. from Samara and Astrakhan mous Province. provinces. 98. Aphodius (Eupleurus) subterraneus (Lin- In the Volga Region, the beetles occur in dry horse naeus, 1758). dung. The distribution range of this species includes al- 102. Aphodius (Heptaulacus) villosus Gyllenhal, most the whole of Europe (except for the extreme 1806. north), North Africa, the Caucasus, the Transcaucasus, Kazakhstan, Asia Minor and Middle Asia, Afghani- The species range includes Ukraine (the Carpathian stan, Mongolia, China, North the Korean Peninsula; it Mountains), North and Central Europe. The species is was imported in North America. In Russia, it is dis- known from Nizhny Novgorod, Volgograd, Rostov tributed throughout the country. In the ZIN collection, provinces (Kabakov and Frolov, 1996) and Karachay- there are 300 spms. from the majority of the regions. Cherkessia (Shokhin, 2007). In the ZIN collection, there are 9 spms. from Leningrad and Ulyanovsk prov- A common, locally abundant species occurring in inces, North Osetia, Tomsk Prov. And the Republic of open biotopes, pastures, in cow and horse dung. Altai.

Subgenus Heptaulacus Mulsant, 1842 Subgenus Mulsant et Rey, 1870 99. Aphodius (Heptaulacus) carinatus (Germar, 103. Aphodius (Labarrus) lividus (Olivier, 1789). 1824). The species occurs almost throughout Europe, ex- The species occurs in Central and South-Eastern cept for the extreme north, in North Africa, the Cauca- Europe, the Caucasus, Turkey, North Iran, Kazakh- sus, Asia Minor, Kazakhstan, and Middle Asia. In the stan, Mongolia, and North China. In Russia, it is dis- ZIN collection, there are 100 spms. from Belgorod, tributed in the steppe zone and the forest-steppe sub- Volgograd, Rostov, Astrakhan provinces, Krasnodar zone of the deciduous forest zone from the Ciscauca- Terr., Kalmykia, and Daghestan. sus to the Amur Region. In the ZIN collection, there are 23 spms. from North Osetia, Daghestan, Kurgan A coprophagous species occurring in horse and cow Prov., Altai and Krasnoyarsk territories, Irkutsk and dung. the Amur provinces. Subgenus Limarus Mulsant et Rey, 1870 100. Aphodius (Heptaulacus) sus (Herbst, 1783). 104. Aphodius (Limarus) maculatus Sturm, 1800. The distribution range of this species includes al- The species occurs in Central and South Europe, most the whole of Europe, North Africa, the Caucasus, Ukraine (the Carpathian Mountains), the Transcauca- the Transcaucasus, Asia Minor, North Iran, West Ka- sus, and Asia Minor. In the ZIN collection, there are zakhstan, and Turkmenistan. In the ZIN collection, specimens from Krasnodar Terr. and Adygea. there are 100 spms. from Moscow, Ryazan, Kursk, Belgorod, Lipetsk, Saratov, Volgograd, Astrakhan The species occurs in forests, in deer dung. provinces, Krasnodar Terr., Tatarstan, Ulyanovsk and Samara provinces. Subgenus Motschulsky, 1859 In the Volga Region, the beetles are often found in 105. Aphodius (Liothorax) kraatzi Harold, 1868 cow and horse dung, on sandy soils, in open biotopes. The distribution range of this species includes The larvae were found in soil on the left riverside of Southern Europe, the Caucasus, the Transcaucasus, the Akhtuba River (Astrakhan Prov.). Asia Minor, West Kazakhstan, Middle Asia, and Af- ghanistan. In Russia, the species occurs mostly in 101. Aphodius (Heptaulacus) testudinarius (Fab- steppe and desert zones, from the Azov Sea to the ricius, 1775). Lower Volga region. In the ZIN collection, there are The species occurs almost throughout Europe (ex- 100 spms. from Nizhny Novgorod, Volgograd, Rostov, cept for the extreme north), and in West Kazakhstan. and Astrakhan provinces, and Kalmykia.

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This species does not occur in dung. We collected Syria, Iran (Dellacasa, 1983). Indicated for Rostov specimens from sand and in light traps. Province (Arzanov et al., 1996), and Chechnia (Shokhin, 2007). 106. Aphodius (Liothorax) linearis Reiche et Saul- cy, 1856. Subgenus Loraspis Mulsant et Rey, 1870 The distribution range of this species includes 112. Aphodius (Loraspis) frater Mulsant et Rey, Southern Europe (Portugal, Spain, Greece), France, 1870. Syria, Palestine. In Russia, the species is known from the Volga Region. In the ZIN collection, there are The distribution range of this species includes Cen- 7 spms. from Volgograd, Kurgan provinces, and Tatar- tral and South Europe, the Caucasus, the Transcauca- stan. sus, Asia Minor, Kazakhstan, and Middle Asia. In Russia, the species is mostly distributed in the Vorga 107. Aphodius (Liothorax) niger (Panzer, 1797). Region and the Ciscaucasus. In the ZIN collection, The species range includes almost the whole of there are 40 spms. from Saratov, Volgograd, Samara, Europe, except for the extreme north, the Caucasus, Kurgan, Astrakhan provinces, Kalmykia, and Dagh- Asia Minor, Syria, and Middle Asia. In the ZIN collec- estan. tion, there are 10 spms. from Leningrad, Nizhny Nov- The species occurs in diverse biotopes, mostly in gorod, Moscow, Volgograd provinces and Krasnodar dry cow and horse dung. It is rather rare throughout its Terr. range. 108. Aphodius (Liothorax) plagiatus (Linnaeus, 1767). Subgenus Mecynodes (Mulsant et Rey, 1870) The distribution range of this species includes al- 113. Aphodius (Mecynodes ) kisilkumi Solsky, most the whole of Europe, except for the extreme 1876. north, North Africa (Tunisia), the Transcaucasus, Asia Minor, Syria, Kazakhstan, Middle Asia, Mongolia, The distribution range of this species includes the and North China. In the ZIN collection, there are 100 Transcaucasus, South-West Kazakhstan, and Middle spms. from Leningrad, Tver, Nizhny Novgorod, Sara- Asia. In Russia, the species is known from the Lower tov, Volgograd, Astrakhan provinces, the Republic of Volga region. In the ZIN collection, there are 20 spms. Altai, Irkutsk Prov., Buryatia, Yakutia, and the Amur from Astrakhan Prov. and Kalmykia. Prov. Subgenus Melaphodius Reitter, 1892 The species occurs in riverside debris. 114. Aphodius (Melaphodius) caspius Ménétriés, 109. Aphodius (Liothorax) rusakovi Gusakov, 1832. 2004. The species is distributed in the steppe zone from The species is known from the original description the Caucasus to West Siberia, in Kazakhstan and Mid- from Rostov and Orenburg provinces of Russia, Kharkov Prov. of Ukraine and Ural Prov. of Kazakh- dle Asia. In the ZIN collection, there are 100 spms. stan. from Penza, Saratov, Volgograd provinces, Stavropol Terr., Astrakhan, Ulyanovsk, Samara provinces, Bash- Subgenus Loraphodius Reitter, 1892 kiria, Chelyabinsk, Tomsk, Kemerovo provinces, Khakassia, Krasnoyarsk Terr., and Irkutsk Prov. 110. Aphodius (Loraphodius) latisulcus Reitter, 1892. The species occurs in cow dung in open biotopes The species is known from the Caucasus and from with meadow and steppe vegetation. the Crimea. It was reported from Krasnodar Terr. 115. Aphodius (Melaphodius) circumcinctus (Shokhin, 2007). W. Schmidt, 1840. 111. Aphodius (Loraphodius) suarius Faldermann, The species is distributed in South-Eastern Europe, 1836. Kazakhstan, Turkmenistan, and Kyrgyzstan. In Russia, The species occurs in South-Eastern Europe, North it mostly occurs in the steppe zone from the Ciscauca- Africa, the Caucasus, the Transcaucasus, Asia Minor, sus to West Siberia. In the ZIN collection, there are

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22 spms. from Ulyanovsk, Volgograd, Rostov prov- gorod, Arkhangelsk, Yaroslavl, Nizhny Novgorod inces, Stavropol Terr., Kalmykia, and Altai Terr. provinces, Chuvashia, Moscow, Kaluga, Ryazan, Orel, Penza, Saratov, Astrakhan provinces, Daghestan, Ki- Subgenus Melinopterus Mulsant, 1842 rovsk Prov., Tatarstan, Orenburg, Tyumen, Tomsk provinces, Altai Terr., the Republic of Altai, Kras- 116. Aphodius (Melinopterus) consputus (Creu- noyarsk Terr., Khakassia, Irkutsk Prov., Buryatia, Za- tzer, 1799). baykalskii Terr., and Yakutia. The species occurs in Central and South Europe, North Africa (Morocco, Algeria, Tunisia), Ukraine, 120. Aphodius (Melinopterus) scuticollis Se- the Caucasus, in West Asia. It was reported from menov, 1898. Krasnodar Terr. (Arzanov et al., 1996). The distribution range of this species includes South-Eastern Kazakhstan, Middle Asia and North- 117. Aphodius (Melinopterus) meuseli Reitter, West China. In Russia, it is known from a few speci- 1906. mens from Altai Terr. and Khakassia. The species is known only from the type series from Tomsk environs. In the ZIN collection, there are no 121. Aphodius (Melinopterus) sphacelatus (Pan- specimens of this species. zer, 1798). The species occurs in the greatest part of Europe, in 118. Aphodius (Melinopterus) prodromus (Brahm, North Africa, the Caucasus, Asia Minor, and Middle 1790). Asia. In Russia, it is known mostly from the European The species occurs throughout Europe, in North Af- part. In the ZIN collection, there are 80 spms. from rica (Morocco, Algeria), the Caucasus, the Transcau- Leningrad, Novgorod, Pskov, Vologda, Yaroslavl, casus, Asia Minor, Syria, Lebanonе, Israel, Kazakh- Vladimir, Nizhny Novgorod, Moscow, Kaluga, Li- stan, Middle Asia, and Mongolia; it was imported to petsk provinces, and Krasnodar Terr. USA and Canada. In the ZIN collection, there are 300 spms. from Karelia, Kaliningrad, Leningrad, Nov- Subgenus Mendidaphodius Reitter, 1901 gorod, Bryansk, Arkhangelsk, Vologda, Yaroslavl, 122. Aphodius (Mendidaphodius) brancsiki Reit- Nizhny Novgorod provinces, Chuvashia, Moscow, ter, 1899. Kaluga, Ryazan, Belgorod, Lipetsk, Saratov, Volgo- grad provinces, Krasnodar and Stavropol territories, The species is widely distributed in deserts of Mid- Adygea, Karachay-Cherkessia, North Osetia, Dagh- dle Asia and West Kazakhstan (Nikolajev, 1987). It estan, Kirovsk Prov., Udmurtia, Tatarstan, Samara was reported from Astrakhan Prov. (Shokhin, 2007). Prov., Bashkiria, Chelyabinsk, Tyumen, Novosibirsk provinces, Altai Terr., the Republic of Altai, Kras- Subgenus Mendidius Harold, 1868 noyarsk Terr., Khakassia, Irkutsk Prov., Zabaykalskii 123. Aphodius (Mendidius) baigakumi (W. Ko- Terr., and Yakutia. shantschikov, 1911). A coprophagous species feeding on dung of cows, The species was described from Turkmenistan horses and wild ungulates. In Russia, this species is (Baigakum Desert). In the ZIN collection, there are no common; in southern regions, it occurs from March to specimens from Russia. Shokhin (2007) reported it December. from Astrakhan Prov.

119. Aphodius (Melinopterus) punctatosulcatus 124. Aphodius (Mendidius) bidens (Solsky, 1876). Sturm, 1805. The species is known from eastern West Asia, from —A. sabulicola Thomson, 1868. Middle and Central Asia. It was reported from Bury- The species is distributed in Europe, except for the atia and Zabaykalskii Terr. (Berlov 1989). extreme north, North Africa (Morocco, Tunisia), the Trancaucasus, West Asia, Kazakhstan, and Middle 125. Aphodius (Mendidius) bispinifrons Reitter, Asia. In Russia, it is common in the European part, 1889. reaching the Transbaikal Region to the east. In the ZIN The species is known from Armenia, the North collection, there are 500 spms. from Leningrad, Nov- Caspian Region, Middle Asia, and North China (Niko-

ENTOMOLOGICAL REVIEW Vol. 94 No. 6 2014 A REVIEW OF THE SCARAB BEETLE TRIBE 861 lajev, 1987). It was reported from Volgograd Prov. The species occurs in riverine sands, among grass (Shokhin, 2007). roots (Medvedev, 1965).

126. Aphodius (Mendidius) curtulus (Harold, Subgenus Neagolius W. Koshantschikov, 1912 1866). 132. Aphodius (Neagolius) abchasicus Reitter, Medvedev and Nikritin (1974) reported it from 1892. South and South-Eastern Ukraine. In the ZIN collec- tion, there are 14 spms. from Astrakhan Prov. of Rus- The species occurs in the alpine and subalpine alti- sia (Dosang environs). tudinal zones of the Greater and Lesser Caucasus. It is known from Krasnodar Terr., Karachay-Cherkessia, We collected the beetles in October and April North Osetia, and Daghestan (Shokhin, 2007). among cereal grass roots, in fixed sands and small patches of barchan sands. One specimen was also col- Subgenus Nialus Mulsant et Rey, 1870 lected in horse dung, and a few specimens were col- lected in a light trap. 133. Aphodius (Nialus) varians Duftschmid, 1805.

127. Aphodius (Mendidius) endroedii Balthasar, The distribution range of this species includes Cen- 1967. tral and Southern Europe, North Africa, the Transcau- casus, Asia Minor, Kazakhstan, and Middle Asia. In The species is known only from the original de- Russia, it is mostly distributed in the steppe zone and scription from the Transbaikal Region. It is possible forest-steppe subzone of the zone of deciduous forest that this name is a junior synonym of A. fimbriolatus. from the western border to South Siberia. In the ZIN 128. Aphodius (Mendidius) fimbriolatus Manner- collection, there are 50 spms. from Nizhny Novgorod, heim, 1849. Ryazan, Belgorod, Voronezh, Saratov, Volgograd, Rostov provinces, Krasnodar Terr., Astrakhan Prov., The species is known from a few localities in Cen- Daghestan, Samara Prov., Bashkiria, Orenburg Prov., tral Asia. In the ZIN collection, there are 3 spms. from and Khakassia. Irkutsk Prov. and Buryatia. The species occurs in diverse biotopes, in cow and 129. Aphodius (Mendidius) multiplex Reitter, horse dung, and in riverside debris. 1897. This species is widely distributed in the deserts and Subgenus Nimbus Mulsant et Rey, 1870 semideserts of Middle Asia, in the Lower Volga Re- 134. Aphodius (Nimbus) circassicus Reitter, 1892. gion, the Ciscaucasus. In the ZIN collection, there are 50 spms. from Volgograd and Astrakhan provinces. The species is endemic to the Caucasus. It is known from a few localities in the Greater Caucasus Range. It The species occurs in horse and cow dung in arid biotopes. was reported from Krasnodar Terr. (Kabakov and Frolov, 1996; Shokhin, 2007). 130. Aphodius (Mendidius) nelsinae (Medvedev, 1968). 135. Aphodius (Nimbus) lederi Harold, 1876. The species is known from Middle Asia. Shokhin The species is endemic to the Caucasus. In the ZIN (2007) reported it from Astrakhan Prov. of Russia collection, there are 4 spms. from Krasnodar Terr. (Dosang environs). In the ZIN collection, there are no The biology of this species is poorly known. Proba- specimens from Russia. bly it occurs in foothill and mid-altitude forests (S. I. Tarasov, pers. comm.). Subgenus Mothon Semenov et Medvedev, 1927 131. Aphodius (Mothon) sarmaticus Semenov et 136. Aphodius (Nimbus) obliteratus Panzer, 1823. Medvedev, 1927. The species is widely distributed in Central and The species was described from South Ukraine (riv- South Europe, occurring also in Ukraine (Transcarpa- erside of the Lower Dnepr). In the ZIN collection, thian Region), the Transcaucasus, and Asia Minor. In there are 2 spms. from Krasnodar Terr. the ZIN collection, there are 6 spms. from Daghestan.

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137. Aphodius (Nimbus) affinis Panzer, 1823. of July to October; larvae were collected in Astrakhan Prov. in April and May. —A. hoberlandti Tesar, 1945. The species is known from a few localities in East Subgenus Orodaliscus Reitter, 1900 Europe (Juřena et al., 2008; Král, 2008) and the South 142. Aphodius (Orodaliscus) rotundangulus Reit- Urals (A. hoberlandti). In the ZIN collection, there are ter, 1900. no specimens from Russia. The distribution range of this species includes South Subgenus Nobius Mulsant et Rey, 1870 Ukraine and Kazakhstan (except for its southern part). In Russia, the species is mostly distributed in the 138. Aphodius (Nobius) dosangi Akhmetova steppe zone of the Ciscaucasus and the Volga Region. et Frolov, 2008. In the ZIN collection, there are 42 spms. from Vo- The species is distributed in the Caspian Depres- ronezh, Saratov, Volgograd, Rostov, Kurgan prov- sion. In the ZIN collection, there are 200 spms. from inces, and Stavropol Terr. Astrakhan Prov. A specialized nidicolous species occuring in gopher The beetles feed on cow and horse dung, the larvae and marmot holes. were found in horse dung in Dosang environs. 143. Aphodius (Orodaliscus) spalacophilus No- 139. Aphodius (Nobius) gresseri Semenov, 1898. vikov, 1996. —A. korgaldzhensis Nikolajev, 1987. The species was described from East Ukraine. In the ZIN collection, there is 1 spm. from Kursk Prov. The species is known from West and Central Ka- zakhstan. In the ZIN collection, there are 43 spms. The species occurs in mole rat holes (Novikov, from Vladimir, Voronezh, Volgograd and Astrakhan 1996). provinces of Russia. 144. Aphodius (Orodaliscus) zangi A. Schmidt, A coprophagous species. All the beetles and larvae 1906. we collected were found in horse dung on sandy soils. The distribution range of this species includes North Kazakhstan and North Kirgizstan. In Russia, the spe- 140. Aphodius (Nobius) inclusus Reitter, 1892. cies is known from a few localities in the steppe zone from the Volga Region to the Ural Mountains. In the The species is distributed in the Caucasus, the Tran- ZIN collection, there are 80 spms. from Astrakhan, scaucasus, and Middle Asian mountains. In the ZIN Samara, Orenburg, and Kurgan provinces. collection, there are 5 spms. from Daghestan. A specialized nidicolous species occuring in mar- 141. Aphodius (Nobius) serotinus (Panzer, 1799). mot holes. —A. x-signum Reitter, 1892. Subgenus Otophorus Mulsant, 1842 The distribution range of this species includes Cen- 145. Aphodius (Otophorus) haemorrhoidalis (Lin- tral, South and Eastern East Europe, the Transcauca- naeus, 1758). sus, Asia Minor, Kazakhstan, and Middle Asia. In the ZIN collection, there are 200 spms. from Leningrad, The distribution range of this species includes the Novgorod, Yaroslavl, Nizhny Novgorod, Moscow, whole of Europe, North Africa, the Caucasus, the Ryazan, Voronezh, Penza, Volgograd, Rostov prov- Transcaucasus, Kazakhstan, Middle Asia, Afghani- inces, Karachay-Cherkessia, Astrakhan Prov., Tatar- stan, Mongolia, North and South-Western China, the stan, Ulyanovsk, Samara, Sverdlovsk, Tyumen, Ke- Korean Peninsula, and Japan; it was imported to North merovo provinces, Republic of Altai, Krasnoyarsk America. In the ZIN collection, there are 500 spms. Terr., Khakassia, Irkutsk Prov., and Buryatia. from the majority of the regions of Russia. The species occurs in cow and horse dung, mostly The species mostly occurs in cow and horse dung, on open biotopes. The adults are active from the end in open biotopes.

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Subgenus Parammoecius Seidlitz, 1891 from Rostov and Volgograd provinces (Shokhin, 2007). 146. Aphodius (Parammoecius) asphaltinus Kole- nati, 1846. 152. Aphodius (Phaeaphodius) rectus Motschul- The species is endemic to the Caucasus. In the ZIN sky, 1866. collection, there are 25 spms. from Krasnodar Terr., The species is widely distributed in the Central and Adygea, Karachay-Cherkessia, Kabardino-Balkaria, Eastern Palaearctic Region from the Volga Region to North Osetia, and Daghestan. the Japan islands. The western border of its range is A coprophagous species occuring in cow dung at unclear. In 2008 г., N.V. Prasolov collected a series of 1800–3000 m a.s.l. 14 spms. in Leningrad Prov. In the ZIN collection, there are 1000 spms. from Leningrad, Astrakhan, Uly- 147. Aphodius (Parammoecius) brevithorax Su- anovsk, Samara, Chelyabinsk, Novosibirsk provinces, makov, 1903. Altai Terr., the Republic of Altai, Irkutsk Prov., Bury- atia, Zabaykalskii Terr., the Amur Prov., the Jewish The species is known from Abkhasia, Georgia and Autonomous Province, Khabarovsk and Primorsky North-Eastern Turkey (Ziani, 1999). It was reported territories, and Sakhalin Prov. from Krasnodar Terr. (Shokhin, 2007). A common species, abundant locally in the Far A specialized nidicolous species occuring in the East. It occurs from March to September in open bio- holes of long-clawed mole-vole (Prometheomys topes, pastures, in cow and horse dung. schaposchnikowi). 153. Aphodius (Phaeaphodius) roddi W. Ko- Subgenus Phaeaphodius Reitter, 1892 shantschikov, 1911. 148. Aphodius (Phaeaphodius) costalis Gebler, The species is known from a type series from Altai 1848. Mountains. It was reported from North-Eastern Ka- zakhstan (Nikolajev, 1998). The species is known from the steppe zone from the Volga Region to the Altai Mountains. In the ZIN col- 154. Aphodius (Phaeaphodius) scoparius Harold, lection, there are 7 spms. from Astrakhan Prov. and 1877. the Republic of Altai (Chuya Steppe). The species is known from South Siberia, Mongolia 149. Aphodius (Phaeaphodius) dauricus Harold, and North-Eastern China. In the ZIN collection, there 1863. are 7 spms. from Buryatia, Zabaykalskii Terr. and the Amur Prov. The species is known from Asia Minor, Iran, Ka- zakhstan, Middle Asia, Afghanistan, Mongolia, West Subgenus Phalacronothus Motschulsky, 1859 China (Kabakov, 1998). It was reported from Irkutsk 155. Aphodius (Phalacronothus) biguttatus Ger- Prov. and Khabarovsk Terr. (Berlov 1989). mar, 1824. 150. Aphodius (Phaeaphodius) jouravliowi Reit- The species is distributed in Central, South and ter, 1907. Eastern Europe, the Transcaucasus, Asia Minor, Ka- zakhstan, and Turkmenistan. In Russia, it is known The species is known from the type specimens from from Ulyanovsk (Isajev, 1995) and Novosibirsk Uralsk (North-Western Kazakhstan) and from the Al- (Zinchenko, 1999) provinces. In the ZIN collection, tai Mountains (without more precise indication of there are 11 spms. from Volgograd, Rostov, Orenburg locality). The taxonomic status of this nominal species provinces and Krasnoyarsk Terr. requires clarification. In the ZIN collection, there are 156. Aphodius (Phalacronothus) citellorum Se- no specimens of this species. menov et Medvedev, 1928. 151. Aphodius (Phaeaphodius) novikovi Kabakov, The species is distributed in the steppe zone of Cen- 1998. tral and Eastern Europe and Kazakhstan. The species is known from a type series from East A nidicolous steppe species occuring in gopher and Ukraine and the Lower Volga region. It was reported marmot holes (Zinchenko et al., 2002; Isajev, 1995).

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157. Aphodius (Phalacronothus) quadrimacula- zakhstan, and Mongolia. In Russia, it is distributed in tus (Linnaeus, 1761). the forest zone, from the western border to Kam- chatka. In the ZIN collection, there are 30 spms. from The species is widely distributed in Central and Karelia, Murmansk and Leningrad provinces, Adygea, South Europe; it is also known from North Africa, the Buryatia, Irkutsk Prov., Zabaykalskii Terr., Amur and Transcaucasus, Asia Minor, Syria and Turkmenistan. Magadan provinces, Khabarovsk and Primorsky terri- In the ZIN collection, there are 11 spms. from Astra- tories. khan Prov. A coprophagous species occuring in forest biotopes. The species occurs in cow dung. 162. Aphodius (Planolinus) vittatus Say, 1825. Subgenus Pharaphodius Reitter, 1892 The species occurs in South and Eastern Europe, the 158. Aphodius (Pharaphodius) rugosostriatus Transcaucasus, Turkey, Syria, Kazakhstan, Middle Waterhouse, 1875. Asia, Mongolia, and China (Dellacasa, Dellacasa, —A. raddei Berlov, 1989 2006). In Russia, it is known from the Volga Region, the Altai Mountains, and the Transbaikal Region. In The distribution range of this species includes North the ZIN collection, there are 100 spms. from Astra- and Central China, Southern Primorskii Territory, the khan and Saratov provinces, the Republic of Altai, Korean Peninsula, and Japan. In the ZIN collection, Tuva, Buryatia, and Zabaykalskii Terr. there are 50 spms. from Primorskii Territory. The species occurs in cattle dung. Distribution records of this species from the Palae- arctic Region refer to A. vittatus mundus Reitter, 1892, Subgenus Plagiogonus Mulsant, 1842 A . vittatus sellatus Mannerheim, 1852, and A. vittatus tjanshanicus Balthasar, 1956. The nominative subspe- 159. Aphodius (Plagiogonus) arenarius Olivier, cies is distributed in South Canada, most of the USA 1789. and in North Mexico (Gordon and Skelley, 2007). —A. putridus (Fourcroy, 1785). Maté (2003: cited from Wilson and Angus, 2006) sug- gested that the American subspecies differs from the This species is widely distributed in Europe, occurs Eurasian ones in the molecular characters; however also in the Caucasus, the Transcaucasus, Turkey, Ka- the stutus of these nominal taxa needs clarification. zakhstan, and Turkmenistan. In Russia, it is known from the steppe zone from Kursk Prov. to West Sibe- Subgenus Platyderides A. Schmidt, 1916 ria. In the ZIN collection, there are 100 spms. from Chuvashia, Nizhny Novgorod, Kursk, Saratov, Volgo- 163. Aphodius (Platyderides) suvorovi Kabakov, grad, Rostov provinces, Daghestan, Samara Prov., the 1996 Republic of Altai, and Krasnoyarsk Terr. The species is known from a single specimen from The species occurs in rodent holes (mostly of mar- the Amur Prov., housed in the ZIN collection. mots and gophers). Subgenus Pseudacrossus Reitter, 1892 160. Aphodius (Plagiogonus) culminarius Reitter, 1900. 164. Aphodius (Pseudacrossus) grebenschikovi Balthasar, 1961. The distribution range of this species includes North-Eastern China and Primorskii Territory. In the The species was described from Mongolia and also ZIN collection, there are 7 spms. from Primorskii Ter- recorded from Zabaykalskii Terr. (Berlov 1989). In the ritory. ZIN collection, there are no specimens of this species.

Subgenus Planolinus Mulsant et Rey, 1870 165. Aphodius (Pseudacrossus) nasutus Reitter, 1887. 161. Aphodius (Planolinus) borealis Gyllenhal, The species is known from Mongolia and North 1827. China (up to East Tibet in the south) (Král, 1997). In The species is widely distributed in Europe, it also Russia, it is distributed in South Siberia and Amur occurs in the Transcaucasus, Asia Minor, Eastern Ka- Region. In the ZIN collection, there are 50 spms. from

ENTOMOLOGICAL REVIEW Vol. 94 No. 6 2014 A REVIEW OF THE SCARAB BEETLE TRIBE 865 the Altai Mountains, Krasnoyarsk Terr., Irkutsk Prov., 171. Aphodius (Sinodiapterna) troitzkyi Jacobson, Zabaykalskii and Khabarovsk territories. 1897. 166. Aphodius (Pseudacrossus) tenebricosus The species was recorded from China, North Korea, A. Schmidt, 1916. Taiwan, and Japan (Dellacasa, 1986). In Russia, it is known from several findings in southern Far East. In The species is known from East Kazakhstan and the ZIN collection, there are 10 spms. from Kha- Mongolia (Zinchenko et al., 2002). In Russia, it was barovsk and Primorsky territories. found in Novosibirsk Prov. (Zinchenko, 1999) and Tyva (Zinchenko et al., 2002). A coprophagous species occurring in cow dung. The species occurs in gray marmot holes (Zin- Subgenus Stenotothorax A. Schmidt, 1913 chenko et al., 2002). 172. Aphodius (Stenotothorax) hibernalis Nakane Subgenus Pubinus Mulsant et Rey, 1870 et Tsukamoto, 1956. 167. Aphodius (Pubinus) tomentosus (Müller, The species was described from Japan. In Russia, it 1776). is known from Amur Prov. (Kabakov and Frolov, The species is known from Northern, Central and 1996), Khabarovsk Terr. and Sakhalin Island (Berlov Eastern Europe, Kazakhstan (Nikolajev, 1998). In 1989). In the ZIN collection, there is 1 spm. from Pri- Russia, it is distributed from western border to Trans- morskii Territory, collected by O.N.Kabakov from soil baikal Region. In the ZIN collection, there are 100 under rotten mushrooms. spms. from Leningrad, Novgorod, Pskov, Tver, Yaro- slavl, Vladimir, Nizhny Novgorod, Moscow, Rostov Subgenus Subrinus Mulsant et Rey, 1870 provinces, Tatarstan, Samara, Orenburg, Tyumen, 173. Aphodius (Subrinus) subtilis D. Koshantschi- Omsk, Novosibirsk, Irkutsk provinces, Altai and Kras- kow, 1894. noyarsk territories. It was reported from Stavropol Terr. (Arzanov et al., 1996) and Karachay-Cherkessia The species is known only from the original de- (Shokhin, 2007). scription from Astrakhan (Koshantschikow, 1894c). The type depository is unknown. The species feeds on cow and horse dung. 174. Aphodius (Subrinus) sturmi Harold, 1870. Subgenus Sigorus Mulsant et Rey, 1870 The species occurs in Central and South Europe, 168. Aphodius (Sigorus) porcus (Fabricius, 1792) North Africa, the Caucasus, the Transcaucasus, Asia The species is widely distributed in Central and Minor, Kazakhstan, Middle Asia, Iran, Mongolia, South Europe, known from Ukraine (Carpathian North-Eastern China, the Korean Peninsula, and Ja- Mountains, Kherson Prov.), the Caucasus, Asia Minor pan. In Russia, it mostly occurs in the steppe zone and and Turkmenistan (Kopet Dagh). In Russia, it is the forest-steppe subzone of the deciduous forest zone known from Rostov Prov. (Shokhin, 2007). from the Azov Sea to Sakhalin Island. In the ZIN col- lection, there are 40 spms. from Volgograd and Rostov Subgenus Sinodiapterna Dellacasa, 1986 provinces, Stavropol Terr., Astrakhan and Samara 169. Aphodius (Sinodiapterna) gorodinskiyi Gusa- provinces, Daghestan, Primorsky Terr., and Sakhalin kov, 2006. Prov. The species is known from single specimen from A coprophagous species mostly occurring in cow foothills of Pogranichny Range in Promorsky Terr. In dung in open biotopes. the ZIN collection, there are no specimens of this spe- cies. Subgenus Sugrames Reitter, 1894 170. Aphodius (Sinodiapterna) hammondi Della- 175. Aphodius (Sugrames) hauseri (Reitter, casa, 1986. 1894). The species was described from North China. In the The species is widely distributed in sand deserts of ZIN collection, there are 2 spms. from Primorskii Ter- Kazakhstan and Middle Asia; it was also found in ritory. Afghanistan. In Russia, it is known from the Lower

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Volga region and Daghestan. In the ZIN collection, known only from southern Primorsky Terr.; in the ZIN there are 10 spms. from Astrakhan Prov. collection there are 8 spms. A psammophillous species. We collected a few lar- A coprophagous species occurring in cow and horse vae among grass roots on a small patch of barchans dung. sands in Astrakhan Prov. Subgenus Trichonotulus Bedel, 1911 Subgenus Teuchestes Mulsant, 1842 179. Aphodius (Trichonotulus) scrofa (Fabricius, 176. Aphodius (Teuchestes) brachysomus Solsky, 1787). 1874. The species occurs in Europe (up to Finland in the The distribution range of this species includes Cen- north), North Africa (Morocco), the Caucasus, the tral and Eastern China, Korean Peninsula and Japan. Transcaucasus, Asia Minor, Kazakhstan, Middle Asia, In Russia, the species is known from Amur Region, Afghanistan, Mongolia, China, North Korea. It was Primorskii Territory and Sakhalin Isl. In the ZIN col- imported to Canada and the USA. In Russia, the spe- lection, there are 100 spms. from Khabarovsk and cies is mostly distributed in the forest-steppe subzone Primorsky territories. of the deciduous forest zone from the western border A coprophagous species mostly occuring in cow to the Amur Region. In the ZIN collection, there are 21 dung from March to July. spms. from Leningrad, Nizhny Novgorod, Volgograd, Rostov provinces, Karachay-Cherkessia, Astrakhan 177. Aphodius (Teuchestes) fossor (Linnaeus, and Irkutsk provinces. 1758). A coprophagous species occuring in ungulate dung. The distribution range of this species includes al- most the whole of Europe (except for the extreme Subgenus Vladimirellus Dellacasa, Dellacasa north), North Africa (Algeria), the Caucasus, Tran- et Bordat, 2002 scaucasus, West Asia, Kazakhstan, Middle Asia and Northern Mongolia. It was imported to North America. 180. Aphodius (Vladimirellus) socors Balthasar, In Russia, it is distributed throughout the country up to 1967. Zabaykalskii Terr. (Nerchinsk) in the east. In the ZIN The species is known from 3 findings in South Sibe- collection, there are 300 spms. from Karelia, Lenin- ria and the Amur Region (Balthasar, 1967; Zinchenko grad, Pskov, Tver, Smolensk, Vologda, Yaroslavl, and Bezborodov, 2013). In the ZIN collection, there Kostroma, Nizhny Novgorod provinces, Chuvashia, are no speimens of this species. Moscow, Ryazan, Belgorod, Lipetsk, Voronezh, Penza, Saratov provinces, Krasnodar and Stavropol Genus CNEMISUS Motschulsky, 1868 territories, Adygea, Kabardino Balkariya, Daghestan, Kirovsk, Samara provinces, Bashkiria, Sverdlovsk, The genus includes 4 species distributed in the sand Chelyabinsk, Tyumen, Omsk, Novosibirsk, Tomsk, deserts and semideserts of Eurasia. In the territory of Kemerovo provinces, Altai Terr., Republic of Altai, Russia, 2 species occur. Krasnoyarsk Terr., Khakassia, Tuva, Irkutsk Prov., 181. Cnemisus kaznakovi (Semenov, 1903). Buryatia, and Zabaykalskii Terr. The species was described from North China and The species occurs in open biotopes, pastures, in also reported from Mongolia (Nikolajev and Puntsag- cow and horse dung. In Russia, it is a common spe- dulam, 1984). Berlov and Anishchenko (1998) re- cies; the beetles are active from April to September. ported it from Buryatia. In the ZIN collection, there are no specimens of this species from Russia. Subgenus Trichaphodius A. Schmidt, 1913 178. Aphodius (Trichaphodius) comatus 182. Cnemisus rufescens (Motschulsky, 1845). A. Schmidt, 1920. The species is distributed in the Caspian Depres- The distribution range of this species includes sion. In the ZIN collection, there are 80 spms. from China, the Korean Peninsula and Japan. In Russia, it is Astrakhan Prov.

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The species occurs in barchan sands, under salt- than 10 species occur. The north border of the distri- worts of the genus Salsola. The beetles are active in bution of the members of the tribe is not clear. late fall. However, in the areas with the permafrost lying near the surface horizon, aphodiines probably do not occur Genus Stephens, 1839 or are only represented by a few boreal species. The genus comprises up to 27 species distributed in Medvedev (1951) suggested that the absence of chaf- all the zoogeographical regions except for Australian. ers in most parts of the West Siberian taiga is due to In Russia, only the type species of the genus occurs. its heavy bogginess. Chafer larvae live in soil and cannot tolerate very high moisture. The absence of 183. Oxyomus silvestris (Scopoli, 1763). chafers from North-Eastern Siberia Medvedev ex- The species is distributed in Europe, North Africa plained by the severity of climate and by permafrost. (Tunisia), the Caucasus, the Transcaucasus, Middle Probably, the very low diversity of the Aphodiini in Asia, and Asia Minor; it was imported to North Amer- the northern taiga may be explained by the same fac- ica. In Russia, it occurs from North-Western border to tors. Aphodiine larvae develop in dung or soil and Kirovsk and Samara provinces. In ZIN collection, high moisture of the substrate, combined with low there are 24 spms. from Leningrad Prov., Chuvashia, temperatures, is unfavorable to the development of all Kaluga, Ryazan provinces, Krasnodar Terr., North the scarabeids. Osetia, Daghestan, Kirovsk and, Samara provinces. Distribution of the Aphodiini of the Fauna of Russia The species occurs in rotten plant residues and in According to Landscape zones and Altitudinal dung. Zonation Types Pattern of the Aphodiini Species Diversity For the analysis of the zonal distribution of the in the Territory of Russia Aphodiini, the map of the landscape zones and vegeta- In Russia, aphodiines are the most diverse in the tion altitudinal zonality types (National Atlas of Rus- southern European Part (Fig. 1). This can be explained sia, 2007) was used. According to the map, the terri- by a variety of natural conditions (e.g., altitudinal tory of Russia can be devided into 5 lanscape zones zonation of the Caucasus) and a long vegetative season and 6 main altitudinal zonation types. Aphodiines with a high sum of effective temperatures. The total occur in the taiga, broadleaved forest, steppe and de- number of species is high due to the species with Wide sert zones, and in the boreal (taiga), nemoral (decidu- Palaearctic ranges, Caucasian endemics and suben- ous) and subarid altitudinal zonation types. Aphodi- demics, and the species having most of their ranges in ines are apparently absent from tundra zone, and arc- South Europe. tic, hypoarctic tundra and hypoarctic taiga altitudinal The accuracy of our map (Fig. 1) is probably higher zonation types. for plain regions than for the mountainous ones. The Aphodiines are the most diverse in the steppe and large average number of species per area unit in the broadleaved forest zones as well as the nemoral zona- mountainous regions (the Caucasus, South Siberia and, tion types. In the steppe zone, 54% of the species of to a lesser degree, the southern Far East) can be the the Russian fauna occur. Probably, the high diversity result of a loose extrapolation of the ranges of separate of the aphodiines in the steppe zone is due to their species and rather reflects the total number of species association with animals largely inhabiting open land- of the regional faunas. The richest local faunas are scapes. Almost the same number of species occurs in found in the Lower Volga Region: 47 species are the nemoral altitudinal zonation types. However, as known from the vicinity of Elton Lake (Makarov et al., opposed to the continuous steppe zone, these altitud- 2009), 45, from Dosang environs (Shokhin, 2007; inal zonation types are situated in distant regions and Frolov and Akhmetova, 2013). In the latter case, at have a drastically different fauna of the aphodiines. least 43 species have stable populations in the territory 50% of species are known from the broadleaved forest of about 3 square kilometers (Frolov, Akhmetova, zone. A reasonable part of these species inhabit open 2013). landscapes of the forest-steppe subzone. In the taiga With the advance to the north, the aphodiine diver- and desert zones, as well as in the boreal and subarid sity gradually declines. At the latitude of 65°N, less types of altitudinal zonation, fewer species occur.

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32% of the species of Russian fauna are known from Wide European Nemoral (Fig. 2c). The ranges of the taiga zone, and only 4% from the subarid zone. the two European species, A. conspurcatus and A. ne- The relatively low species richness in these zones and moralis, can be placed in this group. Their ranges sub- altitudinal zonation types is due to the climate and stantially differ from those of other species and are environmental conditions being less suitable for the situated within the European Nemoral Region (except group. for Euxine Mountain Province) and transitional zones between the Eurosiberian Boreal and European The bulk of the fauna is formed by the species with Nemoral regions and between European Nemoral and wide ranges; 111 species occur in two or more land- the Scythian Steppe regions. In Russia, both species scape zones or altitudinal zonation types. inhabit the boreal (taiga) and broadleaf forest zones. Typology of the Ranges of the Aphodiines The Wide Mediterranean Euxinian type (Fig. 3a) of the Russian Fauna comprises the ranges of A. conjugatus, A. constans, Cluster analysis (Jaccard index as similarity coeffi- A. maculatus, A. obliteratus, A. obscurus, A. porcus, cient, average clustering) yielded a range similarity A. quadrimaculatus, A. scrutator, A. suarius, A. ther- dendrogram which allowed us to group the distribution micola. In the territory of Russia, most of these species ranges of the Aphodiini of Russia into 14 main types occur in the steppe zone and the nemoral altitudinal (Fig. 2–5). For the nomenclature of the types of the zones. ranges we used the schema of general biogeographic The Ancient Mediterranean type (Fig. 3b) com- subdivisions of the Palaearctic Region (Emeljanov, prises the ranges of A. trucidatus, A. pruinosus, A. 1974). hydrochaeris, A. brancsiki, A. consputus, A. aequalis, The distribution ranges of the aphodiines occurring A. inclusus, and A. klugi, which are situated within the in Russia can be grouped into the following types: Ancient Mediterranean Region. In the territory of Rus- sia, most of these species occur in the steppe and de- Wide Palaearctic (Fig. 2а). This type comprises the sert zones. widest ranges. The following species have the ranges that can be classed as Wide Palaearctic: Aphodius The Westscythian-Northturanian type (Fig. 3c) depressus, A. sus, A. plagiatus, A. erraticus, A. fascia- comprises the ranges of A. bimaculatus, A. caspius, tus, A. ater, A. rufipes, A. haemorrhoidalis, A. subter- A. circumcinctus, A. citellorum, A. costalis, A. gre- raneus, A. pusillus, A. scrofa, A. luridus, A. piceus, garius, A. gresseri, A. hahni, A. planus, A. rotundan- A. borealis, A. villosus, A. rufus, A. brevis, A. tomento- gulus, and A. zangi. All these species inhabit steppe sus, A. foetens, A. fossor, A. carinatus, A . distinctus, biotopes, and a part of them are specialized nidicols. A. fimetarius, A. granarius, A. immundus, A. melanos- The Conblacksean Plane type (Fig. 4a) comprises tictus, A. prodromus, A. punctatosulcatus, A. vittatus, the ranges of A. curtulus, A. exilimanus, A. isajevi, A. sordidus, A. rectus, A. sturmi, A. lapponum, A. ivanovi, A. novikovi, A. sarmaticus, A. spalacophi- A. dauricus. All these eurybiont species are widely lus. In the territory of Russia, most of these species distributed in the Palaearctic Region. Most of them occur in the steppe zone and forest-steppe subzone of occur at least in 4 landscape and altitudinal zones. the deciduous forest zone. Wide West-Palaearctic (Fig. 2b). The ranges of the The Caucasian type (Fig. 4b) comprises the ranges following species can be classed as Wide West- of A. swaneticus, A. planicollis, A. lederi, A. circas- Palaearctic: A. arenarius, A. biguttatus, A. coenosus, sicus, A. latisulcus, A. brignolii, A. leisteri, A. bre- A. frater, A. ictericus, A. kraatzi, A. linearis, vithorax, A. asphaltinus, A. alagoezi and A. abcha- A. lividus, A. lugens, A. merdarius, A. niger, A. pay- sicus. All these species are endemic to the Caucasus kulli, A. punctipennis, A. quadriguttatus, A. satellitus, and are distributed within Euxine Mountain Province A. serotinus, A. sphacelatus, A. sticticus, A. testudi- of the European Nemoral Region. narius, A. varians, Oxyomus silvestris. These species are also eurybiont and rather widely distributed in the The Sethian type (Fig. 5a) comprises the ranges of Palaearctic Region. More than a half of them occur at A. badenkoi, A. bispinifrons, A. clathratus, A. digitalis, least in 4 landscape and altitudinal zones. The ranges A. hauseri, A. kisilkumi, A. kizeritskyi , A. menetriesi, of this type are situated within different provinces of a A. multiplex. These species mostly occur in the desert few zoogeographical regions. zone and inhabit arid biotopes. Their ranges are situ-

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Fig. 2. Types of the ranges of the Aphodiini: (a) Wide Palaearctic, (b) Wide West-Palaearctic, (c) Wide European Nemoral. ated largely within the Irano-Turanian and Centralasi- but we think that they should be placed in the group of atic subregions of the Sethian Desert Region. species with Wide Palaearctic ranges. In Russia, the majority of the species with this type of ranges occur The Caspian type (Fig. 4b) comprises the ranges of in the boreal altitudinal zones and the steppe zone. A. dosangi, A. plustschewskii, A. variicolor, and Cne- misus rufescens. In the territory of Russia, these spe- The Stenopean Nemoral type (Fig. 5c) comprises cies inhabit the desert zone. Their ranges almost com- the ranges of A. amurensis, A. binaevulus, A. brachy- pletely lie in north-west part of the North Turanian somus, A. breviusculus, A. costatellus, A. comatus, Lowland Province of the Iranо-Turanian Subprovince A. culminarius, A. hammondi, A. koltzei, A. hibernalis, of the Sethian Desert Region. A. impunctatus, A. inexpectatus, A. languidulus, A. ni- The Centralasiatic type (Fig. 5b) comprises the grotesselatus, A. notabilipennis, A. pratensis, A. pro- ranges of A. antiquus, A. bidens, A. comma, A. cras- prietor, A. quadratus, A. rugosostriatus, A. sub- sus, A. falcispinis, A. fimbriolatus, A. grafi, A. greben- costatus, A. sublimbatus, A. superatratus, A. tenax, schikovi, A. indagator, A. jacobsoni, A. kerzhneri, A. troitzkyi, and A. uniformis. All these species inhabit A. longeciliatus, A. mongolaltaicus, A. nasutus, nemoral altitudinal zones and the majority of them A. roddi, A. scoparius, A. scuticollis, A. sordescens, inhabit also the broadleaf forest zone. The range of A. tanhensis, A. tenebricosus, and Cnemisus kaz- A. amurensis is the least similar to the ranges of other nakovi. The ranges of A. lapponum and A. dauricus species of this type. It is possible, that this species has belong to the Centralasiatic cluster of the dendrogram, a wider distribution in Primorskii Territory, North-

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Fig. 3. Types of the ranges of the Aphodiini: (a) Wide Mediterranean Euxinian, (b) Ancient Mediterranean, (c) Westscythian- Northturanian.

Eastern China and North Korea, but is limited to the arctic Region. Only the north-easternmost part of its areas with stable populations of Japanese deer, as both range includes Southern Primorskii Territory while beetles and larvae feed on their dung. most part is situated in South and South-Eastern Asia.

The distribution ranges of A. guttatus and A. uro- Special Zoogeographic Regionization of the Territory stigma are essentially different from those of other of Russia, Based on the Distribution of the Aphodiini species and cannot be classed into any of the above- and General Zoogeographic Subdivisions mentioned types. Special distribution types can be proposed for these species: Based on a cluster analysis, we distinguished 6 groups of zoogeographical provinces (Fig. 8) which The Canada-Kamchatian type (Fig. 6a) is character- correspond to separate clusters on the dendrogram istic of A. guttatus. This species is widely distributed (Fig. 7). in the Nearctic Region and was recently found in The European group of provinces comprises Mid- Kamchatka. dleuropean mixed, Easteuropean plane and Euxine The East-Chinese-Indomalayan type (Fig. 6b) is mountain provinces of European Nemoral Region, as characteristic of A. urostigma which is the only repre- well as Conblacksean plain province of the Scythian sentative of tropical subgenus Aganocrossus in Russia. Steppe Region. 99 aphodiine species, 98 of which The range of this species mostly lies outside the Palae- belong to Aphodius and 1 to Oxyomus, inhabit this

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Fig. 4. Types of the ranges of the Aphodiini: (a) Conblacksean Plane , (b) Caucasian, (c) Caspian. group of provinces. 65 species occur in the Middleu- A. circassicus, A. inclusus, A. latisulcus, A. lederi, ropean mixed province, 57, in the Easteuropean Plain, A. obscurus, A. planicollis, A. swaneticus, A. curtulus, 80, in the Euxinian mountain, 73 species, in the Con- A. isajevi, A. ivanovi, A. novikovi, A. sarmaticus, and blacksean Plain provinces. 29 species (A. paykulli, A. spalacophilus) are not known from other provinces A. conspurcatus, A. conjugatus, A. consputus, A. con- of the studied area. stans, A. maculatus, A. obliteratus, A. porcus, A. scru- tator, A. suarius, A. thermicola, A. abchasicus, A. ala- The Kazakhstan-Northturanian group of provinces goezi, A. asphaltinus, A. brevithorax, A. brignolii, comprises the homonymic provinces of the Scythian

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Fig. 5. Types of the ranges of the Aphodiini: (a) Sethian , (b) Centralasiatic, (c) Stenopean Nemoral.

Steppe and Sethian Desert regions. 88 aphodiine spe- color, A. exilimanus, A. hahni, A. roddi, and Cnemisus cies are known (86 Aphodius, 1 Cnemisus and 1 Oxyo- rufescens) are unknown from other provinces of the mus) from this territory. In the studied part of North- studied area. turanian Plane province, 80 aphodiine species occur; The European-Westsiberian group of provinces in Kazakhstanian Plane province 65 species occur. comprises all the provinces of the Westeurosiberian 15 species (A. zangi, A. badenkoi, A. brancsiki, Subregion of the Eurosiberian Boreal Region (except A. digitalis, A. dosangi, A. hauseri, A. kisilkumi, A. ki- for the Altaian complex province). Here, 47 aphodiine zeritskyi, A. menetriesi, A. plustschewskii, A. varii- species occur (46 Aphodius and 1 Oxyomus species).

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Fig. 6. Types of the ranges of the Aphodiini: (a) Canada-Kamchatian, (b) East-Chinese-Indomalayan.

Twenty two species occur in the Bothnian Mixed with the other provinces of the group is mainly due to province, 35, in the Zyrianian Plane province, 38, in the Mongolian-Chinese species (A. antiquus, A. nasu- the Uralian mountain province, 32 in the Obean plane tus, A. scoparius) occurring only in the southern parts province. All the species except for A. meuseli also of the provinces. The territory occupied by this group occur in other provinces of the studied area. The dis- of provinces is inhabited by 68 aphodiine species, of tribution range of A. meuseli, so far known only from which 67 belong to Aphodius and one to Cnemisus. the type series, requires clarification. It is possible that 20 species are known from the Angaran Mixed Prov- this species has a wider range. ince, 23, from the Subokhotian mountains, 48, from The Eastsiberian-Altaian-Eastmongolian group of the Westmongolian Complex, 44, from Estmongolian provinces comprises the Westmongolian and East- Complex Province of the Eurosiberian Boreal Region, mongolian complex provinces of the Eastsiberian 57, from the Altaian Complex, 38, from the Estmongo- Subregion of the Eurosiberian Boreal Region, Altaian lian Complex Province of the Scythian Steppe Region. and Eastmongolian complex provinces of the Scythian 15 species (A. antiquus, A. scoparius, A. falcispinis, Steppe Province. In the similarity dendrogram calcu- A. grafi, A. bidens, A. fimbriolatus, A. longeciliatus, lated with the Jaccard index, the Angaran mixed and A. tanhensis, A. crassus, A. grebenschikovi, A. jacob- Subokhotian mountain provinces of the Eurosiberian soni, A. kerzhneri, A. logunovi, A. mongolaltaicus, and Boreal Region form a separate cluster most similar to Cnemisus kaznakovi) are not known from other prov- the Eastsiberian-Altaian-Eastmongolian and European- inces of the studied area. Westsiberian group of provinces (Fig. 7а). In the simi- The Stenopean group of provinces comprises the larity dendrogram calculated with the correlation coef- Weststenopean Mixed, Korean and the North Japanese ficient, they belong to a cluster with the Eastsiberian- Mountain Province of the Stenopean Nemoral Region, Altaian-Eastmongolian group (Fig. 7b); therefore we and the Okhotian Mountain Province of the Eurosibe- include them into this group of provinces. However, rian Boreal Region. This territory is inhabited by the greater degree of similarity of the two provinces 47 aphodiine species which all belong to Aphodius.

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Fig. 7. Similarity dendrograms of provincial aphodiine faunas: (a) similarity coefficient—Jaccard index, clustering method—average values; (b) similarity coefficient—correlation, clustering method—average values). Biogeographic provinces: I-4, Northpacific mixed; II-1, Bothnian mixed; II-2, Zyrianian plane; II-3, Uralian mountain; II-4, Obean plane; II-6, Angaran mixed; II-7, Subokhotian moun- tain; II-8, Okhotian mountain; II-9, Westmongolian complex; II-10, Eastmongolian complex province of Eurosiberian taiga Region; III-2, Middleuropean mixed; III-3, Easteuropean plane; III-4, Euxine mountain; IV-1, Eaststenopean mixed; IV-2, Korean mountain; IV-3, North Japanese mountain; VII-2, Conblacksean plane; VII-3, Kazakhstanian plane; VII-4, Altaian complex; VII-6, Eastmongolian complex province of Scythian steppe Region; VIII-15, Northturanian.

40 species are known from the Weststenopean Mixed tzei, and A. suvorovi) are unknown from other Province, 25, from the Korean Mountain Province, provinces of the studied area. 23, from the North Japanese Mountain Province, 18, from the Okhotian Mountain Province. 25 Primoryan The Northpacific Mixed Province of the Circumpo- species (A. sublimbatus, A. uniformis, A. comatus, lar Tundra Region differs most from the other prov- A. impunctatus, A. languidulus, A. rugosostriatus, inces in its fauna. It cannot be placed in any other A. subcostatus, A. superatratus, A. troitzkyi, A. uro- group. This can be explained by the extremely poor stigma, A. breviusculus, A. inexpectatus, A. proprae- aphodiine fauna comprised of only 4 species of Apho- tor, A. binaevulus, A. hammondi, A. tenax, A. hiber- dius: A. lapponum, A. borealis, A. aleutus, and A. gut- nalis, A. pratensis, A. quadratus, A. amurensis, tatus. The two former species have wide Palaearctic A. costatellus, A. culminarius, A. gorodinskiyi, A. kol- ranges. The uniqueness of this province is due to

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Fig. 8. Province groups of special zoogeographic regionization of the territory of Russia, based on the distribution of the Aphodiini: I, European; II, Kazakhstan-Northturanian; III, European-Westsiberian; IV, Eastsiberian-Altaian-Eastmongolian; V, Stenopean; VI, Northpacific.

A. guttatus which is widely distributed in the Nearctic Countries,” Entomol. Obozr. 88 (2), 391–399 (2009) Region and was recently found in Kamchatka. [Entomol. Rev. 89 (4), 428–436 (2009)]. 3. Akhmetova, L.A. and Frolov, A.V., “New Records of ACKNOWLEDGMENTS Aphodius Species from Russia (Coleoptera: Scarabaei- dae),” Zoosyst. Rossica 17 (1), 72 (2008a). We are gfrateful to V.K. Zinchenko (Siberian Zoo- 4. Akhmetova, L.A. and Frolov, A.V., “A Review of the logical Museum, Institute of Animal Systematics and Scarab Beetle Genus Aphodius Illiger Subgenus Nobius Ecology, Siberian Branch of the Russian Academy of Mulsant et Rey (Coleoptera, Scarabaeidae) of Russia Sciences, Novosibirsk), I.V. Shokhin (Azov Branch of and Adjacent Countries,” Entomol. Obozr. 87 (2), Murmansk Marine Biological Institute of KSC RAS, 397–410 (2008b) [Entomol. Rev. 88 (4), 421–434 Rostov-on-Don), A.V. Ivanov (Institute of Plant and (2008)]. Animal Ecology UB RAS, Ekaterinburg), S.A. Sha- 5. Akhmetova, L.A. and Frolov, A.V., “New to Russia and balin (Soil Biology Institute FEB RAS, Vladivostok), Little Known Species of the Genus Aphodius Illiger K.V. Makarov and A.V. Matalin (Moscow Pedagogi- (Coleoptera, Scarabaeidae),” Zoosyst. Rossica 18 (2), cal State University, Moscow), A.A. Gusakov (Zoo- 278–284 (2009). 6. Arzanov, Yu.G., Shokhin, I.V., Komarov, E.V., and logical Museum, Lomonosov Moscow State Univer- Grebennikov, S.A., “Scarab Beetles of the Subfamily sity, Moscow), G.V. Nikolaev (Al-Farabi Kazakh Na- ,” Data on the Beetle (Coleoptera) Fauna of tional University, Almaty), S.I.Tarasov (Natural His- North the Caucasus and Lower Don (Rostov-on-Don, tory Museum, Oslo University, Oslo), and O.A. Khru- 1996), vol. 3, issue 2, pp. 1–28 [in Russian] [Deposited leva (Severtsov Institute of Ecology and Evolution, in VINITI, no. 1130-В96]. Moscow) for providing material for this study and for 7. Balthasar, V., “Monographie der Scarabaeidae und assistance. Aphodiidae der Palaearktischen und Orientalischen Re- The work was supported by the Russian Foundation gion. Coleoptera Lamellicornia. Aphodiidae,” Verl. Tsch. Akad. Wiss. 3, 1–652 (1964). for Basic Research (grant no 13-04-01002-a). 8. Balthasar, V., “Neue Arten der Familie Aphodiidae (Coleoptera),” Acta Entomol. Bohem. 64, 122–139 REFERENCES (1967). 1. Akhmetova, L.A., “Contribution to the Knowledge of 9. Berlov, E.Ya., “Data on the Coprophagous Beetles of Scarab Beetles of the Genus Aphodius Illiger, 1798 Eastern Siberia and Far East,” in Beetles of the Far East (Coleoptera, Scarabaeidae) of Samara Province,” and Eastern Siberia (New Data on Fauna and Systemat- Evraziat. Entomol. Zh. 5 (3), 247–248 (2006). ics) (AS USSR, Vladivostok, 1979), pp. 102–110 [in 2. Akhmetova, L.A., “A Review of the Scarab Beetle Ge- Russian]. nus Aphodius Illiger Subgenus Plagiogonus Mulsant 10. Berlov, E.Ya., “Diagnostic Key to the Coprophagous (Coleoptera, Scarabaeidae) of Russia and Adjacent Beetles of the Genus Aphodius Illig. of Baikal Region,”

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