Akhmetova, L.A., Frolov, A.V. 2014. A review of the scarab beetle tribe Aphodiini (Coleoptera, Scarabaeidae) of the fauna of Russia. Entomological Review, 94, 846-879. 

Original Russian text: Ахметова, Л. А. & Фролов, А. В. (2014) Обзор пластинчатоусых жуков трибы Aphodiini (Coleoptera, Scarabaeidae) фауны России. Энтомологическое обозрение, 93, 403-497.

DOI: 10.1134/S0013873814060074

A review of the scarab beetle tribe Aphodiini (Coleoptera, Scarabaeidae) of the fauna of Russia

L. A. Akhmetova and А. V. Frolov

Zoological Institute, Russian Academy of Sciences, St. Petersburg, 199034 Russia

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 faunas 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.

Introduction

Scarab beetles of the tribe Aphodiini is a large, world-wide distributed group of insects. The world fauna comprises about 30 genera and over 2000 species, including about 13 genera and 700 species in the Palearctic Region (Dellacasa, 1988; Dellacasa, Dellacasa, 2006). Most aphodiines are coprophages feeding on herbivore dung. In temperate regions including Russia, aphodiines dominate dung-beetle communities both in terms of species richness and abundance. The type genus of the aphodiines, Aphodius Hellwig (sensu lato), is the largest scarab beetle genus in Russia and one of the largest genera of insects.

The earliest records of the aphodiines of the Russian Empire are available in works of Faldermann (1835a, 1835b, 1838), Gebler (1848), Ménétriés (1832, 1849), and Motschulsky (1860). In addition to the lists of known species collected in the Caucasus, Transcaucasus, and West Siberia, they include descriptions of new species. Reitter (1893) summarized the data about the Palearctic scarab beetles. His work contains identification keys and numerous descriptions of new species largely from the Caucasus and Middle Asia. Aphodiines of Southern Russia and Middle Asia have been actively studied by Russian coleopterologists in the late XIX and the early XX century (D. Koshantschikow, 1891, 1894a, 1894b, 1894c, 1894d; Semenov, 1898a, 1899, 1904, 1905; Semenov and Medvedev, 1928; W. Koshantschikov, 1911а, 1911b, 1912, 1913, 1916). Olsoufieff (1918) summarized the available data on the Caucasian coprophagous species.

The regional faunistic lists, new distribution records of the aphodiines in Russia and adjacent territories, new species descriptions, and diagnostic keys to some groups and for some regions are available in the works of Medvedev (1965, 1976, 1979), Medvedev and Ermolenko (1969), Nikritin (1969а, 1969б), Medvedev and Nikritin (1974), Berlov (1979, 1985, 1989), Nikolajev (1987, 1998), Isajev (1995), Arzanov et al. (1996), Kabakov and Frolov (1996), Nikitsky et al. (1996), Novikov (1996, 1998), Kabakov (1998), Zinchenko (1999, 2003а, 2003б, 2011), Zinchenko et al. (2002), Shokhin (2002, 2005, 2007), Gusakov (2004, 2006), Bezborodov and Berlov (2005), Makarov et al. (2009), Bezborodov (2009), Tarasov (2008), Shabalin and Berlov (2009), Zinchenko and Kyzyl-ool (2010), Shabalin and Bezborodov (2012), Zinchenko and Bezborodov (2013), as well as the authors of the present work (Frolov, 1995, 1996, 1997, 1999, 2001a, 2001b, 2001c, 2002, 2009; Akhmetova and Frolov, 2008a, b; 2009; Akhmetova, 2006, 2009; Frolov and Akhmetova, 2006, 2013).

Despite the rather large number of publications about the aphodiines of Russia, there has been no work providing a comprehensive account of the taxonomic composition and geographical distribution of the species of our fauna. Available monographs (Schmidt, 1922; Balthasar, 1964) are obsolete in terms of classification, nomenclature, and distribution data. Furthermore, they are largely compilations of older literature. Extensive materials from the collection of the Zoological Institute of Russian Academy of Sciences (ZIN), accumulated during many decades, remained mostly unstudied.

Material and methods

This work is based on the rich collection of ZIN, including vast material of more than 60 species collected by the authors in Lower Volga Region and the southern Far East. In addition, the materials from the following institutions and private collections have been studied: Moscow Pedagogical State University, Moscow, Zoological museum, Lomonosov Moscow State University, Moscow, A.V. Ivanov’s collection, Ekaterinburg, Soil biology institute FEB RAS, Vladivostok. The numbers of specimens of many studied species are approximate since not all the materials available in the Zoological Institute, specifically those stored on cotton layers and in vials with alcohol, have been processed and integrated into the main collection.

The map of distribution of relative species richness in the territory of Russia (Fig. 1) was generated with ArcGIS software. For special zoogeographic regionization, the scheme of the general zoogeographic subdivisions of the Palearctic Region (Emeljanov, 1974) was used. As the operational zoogeographic units, provinces were used. In the cases when a province completely fits the studied area, its complete fauna was included in the analysis. In the cases when only a part of a province was situated in the territory of Russia, only whose the species occurring within this part were utilized. Provincial aphodiine faunas were compared by constructing similarity dendrograms based on pairwise calculations of Jaccard and correlation coefficients. Average linkage was used as a clustering method (Fig. 7.)

Classification and nomenclature of the Aphodiini follow Medvedev and Nikritin (1970), Nikolaev (1979), and Kabakov and Frolov (1996): Aphodius is treated sensu lato, including Heptaulacus Mulsant, Sugrames Reitterand Mothon Semenov et Medvedev as subgenera.

Genus Aphodius Hellwig, 1798

— Aphodius Illiger, 1798.

Subgenus Acanthobodilus Dellacasa, 1983

1. Aphodius (Acanthobodilus) immundus Creutzer, 1799.

The species occurs throughout Europe, in Morocco, Egypt, the Caucasus, the Transcaucasus, Asia Minor, Syria, Kazakhstan, Middle and Central Asia. In Russia, it occurs from western borders up to southern Yakutia. In the ZIN collection, there are 500 spm. from the majority of the regions of the country.

The species is common in the major part of its range and prefers open biotopes. The beetles feed on horse, cow and donkey dung; they are attracted to light and occur from April to September.

2. Aphodius (Acanthobodilus) languidulus A. Schmidt, 1922.

The distribution range of this species includes North-Eastern China, Korean Peninsula and Japan. In Russia, the species occurs in southern Khabarovsk Terr. and Primorye. It was also reported from Amur Region (Berlov, 1989). In ZIN collection, there are 20 spm.

The species occurs in open biotopes, pastures, in cow dung. The beetles fly in July and August.

Subgenus Acrossus Mulsant, 1842

3. Aphodius (Acrossus) bimaculatus (Laxmann, 1770).

The species occurs in Central and Eastern Europe, North and Eastern Kazakhstan, Kyrgyzstan. In Russia, it is known from a few localities in the European Part and West Siberia (up to Krasnoyarsk in the north-east) (Kabakov and Frolov, 1996). In the ZIN collection, there are 200 spm. from Leningrad, Pskov, Smolensk, Yaroslavl, Vladimir, Moscow, Saratov, Volgograd, Rostov provinces, Krasnodar Terr., Astrakhan, Samara, Orenburg, Novosibirsk, Tomsk provinces, Altai Terr., and Khakassia.

The species is included in the Red Book of the Russian Federation (Nikitsky, 2001). The beetles and larvae feed on horse dung (Frolov and Akhmetova, 2006).

4. Aphodius (Acrossus) binaevulus Heyden, 1887.

The species is known from North Korea and North-Eastern China. In Russia, it occurs in Amur Region and Primorye. In the ZIN collection, there are 60 spm. from Amur Prov., Khabarovsk and Primorsky territories.

5. Aphodius (Acrossus) depressus (Kugelann, 1792).

The species is widely distributed in Europe; it also occurs in the Caucasus, Asia Minor, Kazakhstan, Middle Asia, Mongolia, and China. In Russia, it occurs throughout the country. In the ZIN collection, there are 300 spm. from the majority of regions.

The species occurs in cow, horse and sheep dung, less frequently in the dung of wild pigs, deer, roedeer, and hares. It prefers forest biotopes.

6. Aphodius (Acrossus) luridus (Fabricius, 1775).

The species occurs in Europe, North Africa, the Transcaucasus, Asia Minor, Kazakhstan, the mountains of Middle Asia; it was imported to North America. In Russia, it is distributed throughout the European Part and occurs in south of West Siberia. In the ZIN collection, there are 100 spm. from Leningrad, Yaroslavl, Nizhny Novgorod, Moscow, Kursk, Lipetsk, Saratov, Rostov provinces, Krasnodar Terr., Adygea, Stavropol Terr., Dagestan, Kirovsk Prov., Tatarstan and Bashkiria.

A coprophagous species feeding on cow, horse, donkey and sheep dung.

7. Aphodius (Acrossus) planicollis Reitter, 1890.

This species is endemic to The Caucasus. In Russia, it is known from a few localities in the North Caucasus. In the ZIN collection, there are 10 spm. from Krasnodar and Stavropol territories and Severnaya Osetia.

Forest dweller, the beetles are active from May to July.

8. Aphodius (Acrossus) rufipes (Linnaeus, 1758).

The species occurs in most of Europe, except for extreme north, in North Africa (Tunisia), the Transcaucasus, Kazakhstan, and Middle Asia; it was imported to North America. In Russia, it is widespread. In the ZIN collection, there are 400 spm. from majority of regions of Russia.

The species occurs in cow and horse dung and prefers forest biotopes. It is common in the major part of its range, and is often attracted to light.

9. Aphodius (Acrossus) superatratus Nomura et Nakane, 1951.

A. arsenjevi Berlov, 1989.

The species occurs in North Korea and Japan. In Russia, it is known from a few localities in Amur Region and Primorye. In the ZIN collection, there are 8 spm. from Primorye.

Subgenus Aganocrossus Reitter, 1895

10. Aphodius (Aganocrossus) urostigma Harold, 1862.

The species is widely distributed outside Russia in South and South-Eastern Asia. In Russia, it is known from a few findings from the southernmost Primorsky Terr. (Shabalin and Berlov, 2008). The beetles were collected in cow dung in September.

Subgenus Agoliinus A. Schmidt, 1913

11. Aphodius (Agoliinus) amurensis Iablokov, 1972.

The species is known from two localities in Southern Sikhote Alin: Shkotovo Distr., Pejshula Village, and Lazo Nature Reserve, the pass to Uglovaja Bay (Yablokov-Khnzorian, 1972; Akhmetova, Frolov, 2009). In the ZIN collection, there are 10 spm. from the Lazo Nature Reserve.

The species occurs in Japanese deer dung in mixed forests.

12. Aphodius (Agoliinus) guttatus Eschscholtz, 1823.

The species is known from Canada (Alberta, British Columbia, Newfoundland, Ontario, Quebec) and USA (Alaska, Massachusetts) (Gordon, Skelley, 2007). It was recently found in Kamchatka (Akhmetova, Frolov, 2009): Karaginsky Isl. (3 spm. in ZIN collection).

13. Aphodius (Agoliinus) piceus Gyllenhal, 1808.

The species range is not yet well known. In the literature, there are records of it from Central and South Europe (Dellacasa, Dellacasa, 2006), Ulyanovsk Prov. (Isajev, 1995), Tuva, Irkutsk Prov., Buryatia and Zabaykalsky Terr. (Berlov 1989). In the ZIN collection, there are 500 spm. from Murmansk, Arkhangelsk, and Tyumen provinces.

This coprophagous species inhabits forests and occurs in the dung of horses and wild ungulates from March to July.

14. Aphodius (Agoliinus) satunini Olsoufiev, 1918.

The species is known only from the original description from the Sochi environs (Olsoufieff, 1918). Its type depository is unknown. Shokhin (2007) suggested that this name might be a synonym of A. (Agrilinus) fasciatus (Olivier, 1789).

Subgenus Agolius Mulsant et Rey, 1870

15. Aphodius (Agolius) falcispinis W. Koshantschikov, 1912.

The species is known from East Kazakhstan, North-Western China, South Siberia, and Mongolia. In the ZIN collection, there are 10 spm. from Altai and Zabaykalsky Terr.

16. Aphodius (Agolius) montanus Erichson, 1848.

The species is distributed in the mountains of Central and Eastern Europe, and it was recently found in the South Urals (Zinchenko, 2011). In the ZIN collection, there are no specimens of this species from Russia.

Subgenus Agrilinus Mulsant et Rey, 1870

17. Aphodius (Agrilinus) aleutus Eschscholtz, 1822.

A. ursinus Motschulsky, 1845.

The species was recorded by Berlov (1989) from Yakutia and Kamchatka as A. aleutus ursinus,and by Shabalin and Bezborodov (2012) from the northern Kuril Islands. The range of the nominative subspecies includes the western part of Canada (South-West Alberta, South British Columbia) and the West USA (West Washington, Oregon, and the northern coast of California) (Gordon, Skelley, 2007). In the ZIN collection, there are no specimens from Russia.

18. Aphodius (Agrilinus) ater (De Geer, 1774).

The species is distributed in Europe, North Africa, Kazakhstan, Middle Asia (except for deserts), and Mongolia. In Russia, it occurs throughout the country. In the ZIN collection, there are 100 spm. from the majority of the regions of Russia.

The species occurs in open biotopes and in forests, in cow and sheep dung. It was also found in gopher (Nikolajev, 1987) and marmot (Zinchenko, 1999) holes.

19. Aphodius (Agrilinus) breviusculus Motschulsky, 1866.

The species is known from the Kuril islands (Iturup), as well as from Japan and the Korean Peninsula (Berlov 1989). In the ZIN collection, there are no specimens of this species from Russia.

20. Aphodius (Agrilinus) constans Duftschmid, 1805.

The species occurs in Central and South Europe, Ukraine, the Caucasus, the Transcaucasus, Asia Minor, and Turkmenistan. It was reported from Rostov Prov. of Russia (Shokhin, 2007).

21. Aphodius (Agrilinus) convexus Erichson, 1848.

A. emerichi Reitter, 1892.

According to Dellacasa and Dellacasa (2006), this species is widely distributed in the Palearctic Region; however its range requires clarification.

22. Aphodius (Agrilinus) fasciatus (Olivier, 1789).

A. putridus (Herbst, 1789);

A. uliginosus Hardy, 1847.

The species occurs in Northern, Central and Eastern Europe, Eastern Kazakhstan, Mongolia, and it was also imported to North America. In Russia, it occurs throughout the country except for steppe and desert zones. In the ZIN collection, there are 100 spm. from Karelia, Leningrad, Yaroslavl, Penza, Kirovsk, Chelyabinsk, Tyumen, Tomsk provinces, Altai and Krasnoyarsk Territories, Tuva, Irkutsk Prov., Buryatia, Yakutia, Amur and Sakhalin provinces, Khabarovsk Ter.

The species mostly inhabits forest biotopes. It occurs in wild ungulate dung, as well as in cow and horse dung.

23. Aphodius (Agrilinus) hasegawai Nomura et Nakane, 1951.

The species is distributed in Japan. It was also reported from South Sakhalin and the Kunashir islands (Shabalin and Bezborodov, 2012). In the ZIN collection, there are no specimens from Russia.

24. Aphodius (Agrilinus) inexpectatus Balthasar, 1935.

The species occurs in North Korea and Japan (Honshu Island) (Stebnicka and Galante, 1991). In Russia, it is known from Amur Region (Bezborodov and Berlov, 2005) and Primorye. In the ZIN collection, there are 8 spm. from South Primorye.

25. Aphodius (Agrilinus) isajevi Kabakov, 1994.

The species is known from a few localities in the lower Volga and Don basins. In the ZIN collection, there is a type series from Ulyanovsk Prov.

The species is a specialist nidicolous occurring in steppe biotopes in native colonies of marmots (Marmota bobac); it is not found in the reacclimatized marmot colonies (Isajev, 1995).

26. Aphodius (Agrilinus) lapponum Gyllenhal, 1806.

The species is distributed from North Europe to Kamchatka. In Russia, it occurs throughout the country in the taiga zone; in Dzungarian Ala Tau and Tien-Shan it occurs in the coniferous forest altitudinal zone. In the ZIN collection, there are 300 spm. from Karelia, Murmansk, and Arkhangelsk provinces, the Republic of Komi, Tyumen Prov., the Republic of Altai, Krasnoyarsk Terr., Tuva, Buryatia, Yakutia, Amur, Magadan and Kamchatka provinces and Khabarovsk Terr.

A coprophagous species occurring in ungulate dung.

27. Aphodius (Agrilinus) nemoralis Erichson, 1848.

The species occurs in mixed and deciduous forests in Northern, Central and Eastern Europe. In Russia, it is known from the western border up to Volga Region. In the ZIN collection, there are 30 spm. from Leningrad, Moscow, and Ulyanovsk provinces.

A forest dweller feeding on the dung of wild ungulates. The adults are active from April to June.

28. Aphodius (Agrilinus) nikolajevi Berlov, 1989.

The species was described from South Primorye and also reported from Zabaykalsky Terr. and Amur Prov. (Berlov 1989; Bezborodov and Berlov, 2005). In the ZIN collection, there are no specimens of this species.

29. Aphodius (Agrilinus) pratensis Nomura et Nakane, 1951.

Reported from Sakhalin (Berlov, 1989) and Primorye (Gusakov, 2009). In the ZIN collection, there are no specimens of this species from Russia.

30. Aphodius (Agrilinus) tenax Balthasar, 1932.

The species was described from Sino-Tibetan Mountains. It was reported from Vladivostok environs (Balthasar, 1964) and the lower Amur River (Berlov, 1979). In the ZIN collection, there are no specimens of this species from Russia.

31. Aphodius (Agrilinus) uniformis Waterhouse, 1875.

The species occurs in south Primorsky Terr., as well as in Sakhalin and Kunashir islands. In the ZIN collection, there are 100 spm. from Russia.

The species occurs in open biotopes, pastures, in cow dung; it is common in the major part of its range.

Subgenus Alocoderus A. Schmidt, 1913

32. Aphodius (Alocoderus) digitalis D. Koshantschikow, 1894.

In Russia, it is known from only 2 localities in the vicinity of Dosang Station in Astrakhan Prov. (Shokhin, 2007; Akhmetova and Frolov, 2008).

33. Aphodius (Alocoderus) hydroсhaeris (Fabricius, 1798).

A. hydroсhoeris (Fabricius, 1798).

The distribution range of this species includes Central and South Europe, North Africa, Armenia, Azerbaijan, Middle East, and Kazakhstan. In Russia, it is mostly distributed in the steppe and desert zones of the North Caucasus and the Volga Region. In the ZIN collection, there are 50 spm. from Voronezh, Rostov, Volgograd, Astrakhan provinces, Adygea and Dagestan.

This coprophagous species mostly occurs in horse dung.

34. Aphodius (Alocoderus) rufus (Moll, 1782).

A. scybalarius (Fabricius, 1781).

The species is distributed in Europe, the Transcaucasus, and Western Kazakhstan. In Russia, it occurs throughout the country up to Irkutsk Prov. in the east. In the ZIN collection, there are 200 spm. from Karelia, Leningrad, Novgorod, Pskov, Tver, Kirovsk, Yaroslavl provinces, Chuvashia, Moscow, Ryazan, Penza, Volgograd provinces, Adygea, Karachay-Cherkessia, Dagestan, Ulyanovsk, Samara provinces, Bashkiria, Sverdlovsk, Chelyabinsk, Tomsk provinces, the Republic of Altai, and Irkutsk Prov.

The species occurs from May to September in open biotopes, pastures, in cow and horse dung.

35. Aphodius (Alocoderus) sordidus (Fabricius, 1775).

The species is known from North and Central Europe, North Kazakhstan, Mongolia, China, Korean Peninsula, and Japan. In Russia, it occurs throughout the country from Leningrad Prov. to Kunashir Island. In the ZIN collection, there are 300 spm. from the majority of the regions of the country.

The species occurs in cow dung in open biotopes.

Subgenus Amidorus Mulsant et Rey, 1870

36. Aphodius (Amidorus) alagoezi Olsoufiev, 1918.

The species was described from Armenia (Aragats Mountain). In Russia, it is known from Elbrus (Tarasov, 2008).

37. Aphodius (Amidorus) obscurus (Fabricius, 1792).

The species occurs in Central and South Europe, the Carpathian Mountains, the Caucasus, and Asia Minor. In Russia, it is known from the North Caucasus. In the ZIN collection, there are 20 spm. from Krasnodar Terr., Adygea, Karachay-Cherkessia, Kabardino Balkariya, and Dagestan.

The species occurs in cow dung in the mountains, up to 2000-3000 m a.s.l. in the Caucasus.

38. Aphodius (Amidorus) thermicola Erichson, 1848.

The distribution range of this species includes Central, Southern and Eastern Europe, the Caucasus, the Transcaucasus, and Asia Minor. Tarasov (2008) reported it from Krasnodar Terr., North Osetia and Dagestan. In the ZIN collection, there are 6 spm. from Krasnodar Terr.

Subgenus Ammoecius Mulsant, 1842

39. Aphodius (Ammoecius) brevis Erichson, 1848.

The species is distributed in Central and Eastern Europe, the Caucasus and the Transcaucasus. In Russia, it occurs in the European Part and South Siberia (up to Transbaikal in the east). In the ZIN collection, there are 70 spm. from Leningrad, Saratov provinces, Krasnodar Terr., Samara, Kurgan, Tyumen, Tomsk, Irkutsk provinces, the Republic of Altai.

The species occurs in dry cow, horse and, rarely, elk dung.

Subgenus Aparammoecius Petrovitz, 1958

40. Aphodius (Aparammoecius) leisteri Medvedev, 1968.

This species is endemic to the Caucasus. It was described from Abkhazia. In the ZIN collection, there are 2 spm. from North Osetia (Alagir Distr., Yu.E. Komarov leg.).

Subgenus Aphodaulacus W. Koshantschikov, 1911

41. Aphodius (Aphodaulacus) kizeritskyi Frolov, 2002.

The species was described from Turkmenistan. It is known from a few localities in Karakum and Caspian Lowland deserts. In the ZIN collection, there are 150 spm. from Astrakhan Prov. (Dosang environs).

The species occurs in horse dung.

42. Aphodius (Aphodaulacus) koltzei Reitter, 1892.

A. medvedevi Nikritin, 1969;

A. kurenzovi Nikritin, 1969.

The species is known from Primorye, Amur Region and North-Eastern China. In the ZIN collection, there are 11 spm. from Primorye.

43. Aphodius (Aphodaulacus) nigrotessellatus Motschulsky, 1866.

The distribution range of this species includes North-Eastern China and Japan. In Russia, it is known from Amur Region and Primorye. In the ZIN collection, there are 72 spm. from Zabaykalsky, Khabarovsk and Primorsky territories.

Subgenus Aphodiellus A. Schmidt, 1913

44. Aphodius (Aphodiellus) impunctatus Waterhouse, 1875.

The species occurs in Japan, the Korean Peninsula, in North-Eastern China (from Beijing to Harbin). In Russia, it is distributed in South Primorye. In the ZIN collection, there are 17 spm.

The species occurs in pastures, in cow dung. A few specimens were collected in a forest under rotten mushrooms.

Subgenus Aphodius Hellwig, 1798

45. Aphodius (Aphodius) conjugatus (Panzer, 1795).

The species occurs in Central and South Europe, the Caucasus, the Transcaucasus, and Asia Minor. In Russia, it is known from the North Caucasus. In the ZIN collection, there are 15 spm. from Rostov Prov., Krasnodar Terr. and Dagestan.

A coprophagous species occurring in cow dung.

46. Aphodius (Aphodius) fimetarius (Linnaeus, 1758).

The species is widely distributed in Europe, North Africa, Kazakhstan, Middle and Central Asia, imported to North America and Australia. In Russia, it occurs throughout the country up to Eastern Siberia in the east; in the ZIN collection, there are 300 spm. from the majority of the regions of the country.

In Russia, imagoes are active from April to October occurring in open biotopes, pastures, in cow, sheep and horse dung.

47. Aphodius (Aphodius) foetens (Fabricius, 1787).

A. aestivalis Stephens, 1839.

The species is widely distributed in Europe, also occurs in the Caucasus, the Transcaucasus, Asia Minor, and North Kazakhstan. In Russia, it is known from the European part and Siberia (up to Yakutia in the east). In the ZIN collection, there are 100 spm. from the majority of the regions of the country.

In Russia, the beetles occur in open biotopes, pastures, in cow and horse dung, from May to October.

48. Aphodius (Aphodius) swaneticus Reitter, 1892.

The species is endemic to the Caucasus. In the ZIN collection, there are 9 spm. from Krasnodar Terr. and Adygea.

Subgenus Biralus Mulsant et Rey, 1870

49. Aphodius (Biralus) menetriesi Ménétriés, 1849.

The species is distributed in South-Eastern Europe, the Transcaucasus, South Kazakhstan, West and Middle Asia. In the ZIN collection, there are 10 spm. from Volgograd and Astrakhan provinces.

The species occurs in open biotopes, in cow dung; it reaches middle altitudes in the mountain regions.

50. Aphodius (Biralus) satellitus (Herbst, 1789).

The species occurs in Central and South Europe, North Africa, the Transcaucasus, Middle Asia. In Russia, it is mainly distributed in North the Caucasus and the Volga Region. In the ZIN collection, there are 40 spm. from Voronezh, Saratov, Volgograd, Rostov provinces, Krasnodar and Stavropol territories, Astrakhan and Samara provinces, and Dagestan.

The species occurs in cow and horse dung, mostly in open biotopes.

Subgenus Bodilus Mulsant et Rey, 1870

51. Aphodius (Bodilus) gregarius Harold, 1871.

The species is distributed in South-Eastern Europe, Kazakhstan, Middle Asia, Mongolia and North-West China. In Russia, it occurs in the steppe zone and the forest-steppe subzone of the broadleaf forest zone from Rostov Prov. up to southern West Siberia. In the ZIN collection, there are 60 spm. from Volgograd, Astrakhan, Orenburg provinces and Khakassia.

In the territory of Russia, the beetles occur in cow and horse dung from May to August.

52. Aphodius (Bodilus) ictericus (Laicharting, 1781).

A. nitidulus (Fabricius, 1792).

The distribution range of this species includes the whole of Europe (except for the extreme north), North Africa, the Transcaucasus, Asia Minor, Iran and North Kazakhstan. In Russia, it is widely distributed in the European Part. In the ZIN collection, there are 200 spm. from Leningrad, Novgorod, Tver, Moscow, Ryazan, Kursk, Lipetsk, Saratov, Volgograd, Rostov provinces, Krasnodar Terr., Adygea, Astrakhan, Kirovsk, Samara provinces, Chechnia and Dagestan.

The species occurs in open biotopes, pastures in cow and horse dung from spring to late autumn.

53. Aphodius (Bodilus) longeciliatus Reitter, 1887.

The species occurs throughout Mongolia and in China (Qinghai, Inner Mongolia provinces). It was reported from Buryatia (Berlov, 1989).

54. Aphodius (Bodilus) lugens Creutzer, 1799.

The species occurs in Central, South and Eastern Europe, North Africa, the Transcaucasus, Asia Minor, Iran, Afghanistan, Kazakhstan and Middle Asia. In Russia, it is distributed in the steppe zone and the forest-steppe subzone of the deciduous forest zone from the western border to the Trans-Urals. In the ZIN collection, there are 100 spm. from Tver, Kursk, Belgorod, Voronezh, Penza, Saratov, Volgograd, Rostov, Astrakhan provinces, Krasnodar and Stavropol territories, Dagestan, Samara, Orenburg and Kurgan provinces.

The species occurs in open biotopes, in cow dung. In Russia, the beetles are active from April to October.

55. Aphodius (Bodilus) punctipennis Erichson, 1848.

The species occurs in Central, South and Eastern Europe, Egypt, the Transcaucasus, Iran, Afghanistan, South Kazakhstan and Middle Asia. In Russia, it is distributed in the steppe zone and the forest-steppe subzone of the broadleaf forest zone from the western border to the Volga Region. In the ZIN collection, there are 90 spm. from Ryazan, Kursk, Belgorod, Penza, Volgograd, Rostov, Astrakhan, Samara provinces, Stavropol Terr., Dagestan and Bashkiria.

56. Aphodius (Bodilus) sordescens Harold, 1869.

The species occurs in North-Eastern Kazakhstan, Mongolia, and North China. In Russia, it is known from the steppe zone of South Siberia, from the Trans-Urals to Transbaikal. In the ZIN collection, there are 100 spm. from Chelyabinsk, Tomsk, and Kemerovo provinces, Altai and Krasnoyarsk territories, Khakassia, Tuva, Buryatia, Zabaykalsky Terr.

Subgenus Calamosternus Motschulsky, 1859

57. Aphodius (Calamosternus) granarius (Linnaeus, 1767).

The species is distributed throughout Europe (except for the extreme north), West Asia, Kazakhstan, and Middle Asia; imported to North America. In Russia, it occurs from the western border up to Transbaikal Region. In the ZIN collection, there are 200 spm. from Kaliningrad, Leningrad, Tver, Nizhny Novgorod, Ryazan, Belgorod, Voronezh, Volgograd provinces, Krasnodar Terr., Kalmykia, Astrakhan Prov., Dagestan, Kirovsk and Samara provinces, Bashkiria, Tyumen Prov. and Zabaykalsky Terr.

The species is common in all the parts of its range. It occurs in open biotopes in the dung of different animals, in carrion, and in marmot holes.

58. Aphodius (Calamosternus) sublimbatus Motschulsky, 1860.

The distribution range of this species includes North-Eastern China, the Korean Peninsula, Japan and Taiwan Island. In Russia, it is distributed in Primorye and Amur Region. In the ZIN collection, there are 100 spm. from the Jewish Autonomous Province, Khabarovsk, and Primorsky territories.

The species occurs in open biotopes, in cow and horse dung. This is a common, locally abundant species.

59. Aphodius (Calamosternus) trucidatus Harold, 1863.jh

A. suturifer Reitter, 1892.

The species occurs in South Europe, North Africa, Transcaucasus, Middle East (Syria, Lebanon, Israel, Iran, Iraq), Kazakhstan, Middle Asia, and Mongolia. In Russia, the species is known from the Ciscaucasus and Lower Volga Region (Arzanov et al., 1996; Shokhin, 2007). In the ZIN collection, there are 2 spm. from Astrakhan Prov. (Lake Baskunchak and Dosang).

Subgenus Carinaulus Tesar, 1945

60. Aphodius (Carinaulus) costatellus A. Schmidt, 1916.

The species is known from several findings in the Amur Region and Primorye. In the ZIN collection, there are 8 spm. from the Amur Prov. and Primorye.

Subgenus Chilothorax Motschulsky, 1859

 – Volinus Mulsant, 1870.

61. Aphodius (Chilothorax) badenkoi Nikolajev, 1987.

The species is distributed in deserts of Middle Asia, and the Northern Caspian Region. In Russia, it is known from single specimen from Astrakhan Prov. (Dosang environs).

62. Aphodius (Chilothorax) clathratus Reitter, 1892.

The species is known from the Caucasus, the Transcaucasus, Turkey, Iran, and Middle Asia (Nikolajev, 1987). Its distribution range needs clarification because some records of A. clathratus may refer to the very similar A. melanostictus. In Russia, it was found in Dosang environs (Astrakhan Prov.), in horse dung, in fixed sands.

63. Aphodius (Chilothorax) comma Reitter, 1892.

The species occurs in North Kazakhstan, eastern Middle Asia, and Mongolia. In Russia, it is known from South Siberia and the Amur Region. In the ZIN collection, there are 300 spm. from Tyumen, Novosibirsk, Tomsk, and Kemerovo provinces, Altai and Krasnoyarsk territories, the Republic of Altai, Khakassia, Tuva, Irkutsk Prov., Buryatia, Zabaykalsky Terr., the Amur Prov., Yakutia, and Khabarovsk Terr.

64. Aphodius (Chilothorax) conspurcatus (Linnaeus, 1758).

The species is distributed in North and Central Europe; it locally occurs in South Europe (Dellacasa, Dellacasa, 2006). In the ZIN collection, there are 30 spm. from Leningrad, Novgorod, and Yaroslavl provinces.

65. Aphodius (Chilothorax) distinctus (Müller, 1776).

A. inquinatus Herbst, 1783.

The distribution range of this species includes the whole of Europe (except for the extreme north), North Africa, the Transcaucasus, Asia Minor, Kazakhstan, Middle Asia, Mongolia. It was imported to North America. In Russia, it occurs throughout the country in the European part, reaching Baikal region in the east. In the ZIN collection, there are 300 spm. from the majority of the regions of the country.

The species occurs in cow and horse dung, in marmot holes, and also is attracted to light. In the territory of Russia the beetles are active from March to October. It is a common, locally abundant species.

66. Aphodius (Chilothorax) exilimanus Kabakov, 1994.

The species is known from a type series from Ulyanovsk Prov. (15 spm. in the ZIN collection). It is a specialist nidicolous species. All the specimens were collected in the holes of Marmota bobac.

67. Aphodius (Chilothorax) grafi Reitter, 1901.

A. kryzhanovskii Nikritin, 1969;

А. schutovae Nikritin, 1969.

This species is widely distributed in Mongolia. In Russia, it occurs in South Siberia from the Altai Mountains to Transbaikal Region. In the ZIN collection, there are 100 spm. from the Republic of Altai, Khakassia, and Buryatia.

68. Aphodius (Chilothorax) hahni Reitter, 1907.

The species is known from a few localities in North-West Kazakhstan. In the ZIN collection, there is 1 spm. from Volgograd Prov.

69. Aphodius (Chilothorax) ivanovi Lebedev, 1912.

The species is distributed in the Middle Volga and Don basins (Medvedev, 1965). In the ZIN collection, there are 27 spm. from Penza and Ulyanovsk provinces.

The species occurs on sandy riversides.

70. Aphodius (Chilothorax) jacobsoni W. Koshantschikov, 1911.

The species is distributed in Eastern Kazakhstan and Mongolia. In Russia, it is known from South Siberia. In the ZIN collection, there are 18 spm. from Altai and Khakassia.

71. Aphodius (Chilothorax) kerzhneri Nikolajev, 1984.

The species is distributed in Eastern Kazakhstan and Mongolia. In Russia, it is known from South Siberia. In the ZIN collection, there are 22 spm. from Altai.

The species occurs in gray marmot holes (Nikolajev, Puntsagdulam, 1984; Zinchenko, 1999).

72. Aphodius (Chilothorax) logunovi Zinchenko, 2003.

The species is known only from the type series from South-East Altai (3 spm., one of which is housed in the ZIN collection) (Zinchenko, 2003a).

73. Aphodius (Chilothorax) melanostictus W. Schmidt, 1840.

The species occurs in Central, South and Eastern Europe, North Africa, West Asia, Kazakhstan, and Middle Asia. In Russia, it is distributed from western border to Transbaikal Region, mostly in the steppe zone and the forest-steppe subzone of the deciduous forest zone. In the ZIN collection, there are 300 spm.

The species occurs in open biotopes, in the dung of domestic animal and in marmot holes. On the territory of Russia, the beetles are active from March to October. The species is common throughout its range.

74. Aphodius (Chilothorax) mongolaltaicus Nikolajev, 1984.

The species is known from East Kazakhstan and Mongolia. In Russia, it was found in South Altai. In the ZIN collection, there are 3 spm. from Chuya Steppe.

75. Aphodius (Chilothorax) paykulli (Bedel, 1798).

A. tessulatus Duftschmid, 1805.

The species occurs in the greatest part of Europe, the Caucasus, the Transcaucasus and Asia Minor. In Russia, it is known from a few findings in North-Western Region and the North Caucasus. In the ZIN collection, there are 3 spm. from Kaliningrad and Leningrad provinces.

76. Aphodius (Chilothorax) planus D. Koshantschikow, 1894.

A. transvolgensis Semenov, 1898.

The species is known from South Ukraine, North and Central Kazakhstan and Turkmenistan. In Russia, the species occurs in the steppe zone from the Ciscaucasus to the south of West Siberia. In the ZIN collection, there are 16 spm. from Saratov and Volgograd provinces, Krasnodar Terr., Orenburg and Novosibirsk provinces.

The species occurs in gopher and marmot holes (Isajev, 1995; Zinchenko, 1999).

77. Aphodius (Chilothorax) plustschewskii D. Koshantschikow, 1894.

The species is distributed in the deserts of the North Caspian Region. In the ZIN collection, there are 100 spm. from Astrakhan Prov.

A coprophagous species, feeding on horse dung. Beetles are active in late fall.

78. Aphodius (Chilothorax) subpolitus Motschulsky, 1860.

The species is known only from the original description from Siberia (without a more precise locality) (Motschulsky, 1860). The type depository is unknown.

79. Aphodius (Chilothorax) sticticus (Panzer, 1798).

A. equestris (Panzer, 1798).

The species occurs throughout Europe (except for the extreme north), in the Trans caucasus, Asia Minor, and Northern Iran. In Russia, it is widely distributed in the European part up to the South Urals Mountains. In the ZIN collection, there are 100 spm. from Yaroslavl Prov., Chuvashia, Moscow, Ryazan, Kursk, Belgorod, Penza, Samara provinces, Krasnodar Terr., and Bashkiria.

A forest dweller, feeding mostly on cow and horse dung.

80. Aphodius (Chilothorax) tanhensis Frolov, 2001.

The species is known from Mongolia and the Altai Mountains. In the ZIN collection, there are 5 spm. from Chuya steppe.

81. Aphodius (Chilothorax) variicolor D. Koshantschikow, 1894.

The species is distributed in the Caspian lowland deserts. In the ZIN collection, there are 28 spm. from Astrakhan Prov.

The beetles feed on horse dung; the larvae occur in barchan sands and apparently feed on the roots of cheat grass.

Subgenus Colobopterus Mulsant, 1842

82. Aphodius (Colobopterus) brignolii Carpaneto, 1973.

The species is known from the North Caucasus, the Transcaucasus and North-Eastern Turkey. In the ZIN collection, there are 11 spm. from Krasnodar Terr., Adygea, Karachay-Cherkessia and North Osetia.

83. Aphodius (Colobopterus) erraticus (Linnaeus, 1758).

The species occurs in Europe (except for the extreme north), West, Middle and Central Asia; it was imported to North America. In Russia, it is distributed throughout the country. In the ZIN collection, there are 1000 spm. from the majority of the regions of Russia.

This eurybiont species is abundant throughout its range occurring mostly in open biotopes, in the dung of domestic and wild animals.

84. Aphodius (Colobopterus) indagator Mannerheim, 1849.

The distribution range of this species includes Mongolia and North-Eastern China. In Russia, it is known from Altai Mountains, south of East Siberia, Amur Region and Primorye. In the ZIN collection, there are 45 spm. from Republic of Altai, Irkutsk Prov., Buryatia, Zabaykalsky Terr., Amur Prov. and Primorsky Terr

85. Aphodius (Colobopterus) notabilipennis Petrovitz, 1972.

The species is known from the Amur Region and Primorye. In the ZIN collection, there are 100 spm. from Zabaykalsky Terr., the Jewish Autonomous Province, Khabarovsk and Primorsky territories.

86. Aphodius (Colobopterus) propraetor Balthasar, 1932.

The distribution range of this species includes North-Eastern China and the Korean Peninsula. In Russia, it is widely distributed in southern Far East. In the ZIN collection, there are 200 spm. from the Amur Prov., the Jewish Autonomous Province, Khabarovsk, and Primorsky territories, Sakhalin Prov. (Sakhalin and Kunashir islands).

A common, locally abundant coprophagous species occurring from April to August in cow and horse dung.

87. Aphodius (Colobopterus) quadratus Reiche, 1847.

A. haroldianus Balthasar, 1932.

The species was described from Japan. In the ZIN collection, there are 6 spm. from Kunashir Island.

Subgenus Coprimorphus Mulsant, 1842

88. Aphodius (Coprimorphus) scrutator (Herbst, 1789).

The species is distributed in Central and South Europe, Azores, the Caucasus, the Transcaucasus, South-West Asia. In Russia, it is known from the northern foothills of the Caucasus. In the ZIN collection, there are 10 specimens from Krasnodar, Stavropol territories and Dagestan.

The beetles mostly occur in open biotopes, in cow dung.

Subgenus Coptochiroides Balthasar, 1938

89. Aphodius (Coptochiroides) subcostatus Kolbe, 1886

The distribution range of this species includes North-Eastern China, Korean Peninsula and Japan. In Russia, it occurs in southern Primorye; In the ZIN collection, there are 200 spm.

Subgenus Erytus Mulsant et Rey, 1870

90. Aphodius (Erytus) aequalis A. Schmidt, 1907.

The species occurs in South Europe, North Africa, The Caucasus, Transcaucasus, Turkey, Iran, Syria, Kazakhstan, Middle and Central Asia. In the ZIN collection, there are 30 specimens from Volgograd, Astrakhan provinces and Stavropol Terr. Russia.

The beetles of this species mostly occur in cow dung in late spring. In the Lower Volga Terr. The beetles are active at nights.

91. Aphodius (Erytus) klugi A. Schmidt, 1910.

The distribution range of this species includes Southern Europe, North Africa, South-Western Asia, Kazakhstan, Middle Southern Asia (Hindustan Peninsula and Himalaya). It was reported from Rostov, Volgograd, Astrakhan provinces (Arzanov et al., 1996) and Kalmykia (Shokhin, 2007).

92. Aphodius (Erytus) pruinosus Reitter, 1892.

The distribution range of this species includes North Africa, South-Western Asia, Southern Kazakhstan, and Middle Asia. It was reported from Kalmykia (Arzanov et al., 1996) and Astrakhan Prov. (Shokhin, 2007).

Subgenus Esymus Mulsant et Rey, 1870

93. Aphodius (Esymus) merdarius (Fabricius, 1775).

The species is distributed in Europe (Dellacasa, Dellacasa, 2006), North Africa, the Caucasus, the Transcaucasus, Asia Minor, Iran, Kazakhstan, and Middle Asia. In Russia, the species occurs from the European part to the South Urals Mountains. In the ZIN collection, there are 100 spm. from Leningrad, Novgorod, Tver, Vologda, Yaroslavl, Nizhny Novgorod, Moscow, Ryazan, Lipetsk, Volgograd, Samara provinces, Krasnodar Terr., Dagestan and Bashkiria.

A coprophagous species occurring mostly in cow dung, in open biotopes, from April to September.

94. Aphodius (Esymus) pusillus (Herbst, 1789).

The distribution range of this species includes almost whole Europe (except for extreme north), the Caucasus, the Transcaucasus, Kazakhstan, Middle Asia, Asia Minor, Mongolia, North China, Korean Peninsula, and Japan. In Russia, it is distributed throughout the country. In the ZIN collection, there are 300 spm. from the majority of the regions.

It is a common, locally abundant species feeding on cow, horse, and sheep dung.

Subgenus Eudolus Mulsant et Rey, 1870

95. Aphodius (Eudolus) quadriguttatus (Herbst, 1783).

The species is distributed in Europe, North Africa, West Asia, Kazakhstan, and Middle Asia. In Russia, it mostly occurs in the Vorga Region and the Ciscaucasus. In the ZIN collection, there are 100 spm. from Saratov, Volgograd, Astrakhan, Samara provinces and Krasnodar Terr.

A coprophagous species occurring in the dung of different animals. It prefers arid biotopes with sandy and sandy clay soils.

Subgenus Euorodalus Dellacasa, 1983

96. Aphodius (Euorodalus) coenosus (Panzer, 1798).

The distribution range of this species includes almost the whole of Europe (except for the extreme north), the Transcaucasus, Asia Minor, West Kazakhstan, and Tajikistan. In the ZIN collection, there are 40 spm. from Leningrad, Nizhny Novgorod, Voronezh, Saratov, Volgograd and Astrakhan provinces.

A coprophagous species occurring in the dung of cows, horses, and wild ungulates. The species occurs in different biotopes: in Eastern Europe, in forests; Isajev (1995) recorded it from steppes; we collected it on sandy soils on the Akhtuba riverside in Dosang environs (Astrakhan Prov.).

Subgenus Eupleurus Mulsant, 1842

97. Aphodius (Eupleurus) antiquus Faldermann, 1835.

The species is known from South Siberia and Mongolia. In the ZIN collection, there are 50 spm. from Altai and Krasnoyarsk territories, the Republic of Altai, Khakassia, Buryatia, Irkutsk Prov., Zabaykalsky Terr., Yakutia, Amur Prov., and the Jewish Autonomous Province.

98. Aphodius (Eupleurus) subterraneus (Linnaeus, 1758).

The distribution range of this species includes almost the whole of Europe (except for the extreme north), North Africa, the Caucasus, the Transcaucasus, Kazakhstan, Asia Minor and Middle Asia, Afghanistan, Mongolia, China, North the Korean Peninsula; it was imported in North America. In Russia, it is distributed throughout the country. In the ZIN collection, there are 300 spm. from the majority of the regions.

A common, locally abundant species occurring in open biotopes, pastures, in cow and horse dung.

Subgenus Heptaulacus Mulsant, 1842

99. Aphodius (Heptaulacus) carinatus (Germar, 1824).

The species occurs in Central and South-Eastern Europe, the Caucasus, Turkey, North Iran, Kazakhstan, Mongolia, and North China. In Russia, it is distributed in the steppe zone and the forest-steppe subzone of the deciduous forest zone from the Ciscaucasus to the Amur Region. In the ZIN collection, there are 23 spm. from North Osetia, Dagestan, Kurgan Prov., Altai and Krasnoyarsk territories, Irkutsk and the Amur provinces.

100. Aphodius (Heptaulacus) sus (Herbst, 1783).

The distribution range of this species includes almost the whole of Europe, North Africa, the Caucasus, the Transcaucasus, Asia Minor, North Iran, West Kazakhstan, and Turkmenistan. In the ZIN collection, there are 100 spm. from Moscow, Ryazan, Kursk, Belgorod, Lipetsk, Saratov, Volgograd, Astrakhan provinces, Krasnodar Terr., Tatarstan, Ulyanovsk and Samara provinces.

In the Volga Region, the beetles are often found in cow and horse dung, on sandy soils, in open biotopes. The larvae were found in soil on the left riverside of the Akhtuba River (Astrakhan Prov.).

101. Aphodius (Heptaulacus) testudinarius (Fabricius, 1775).

The species occurs almost throughout Europe (except for the extreme north), and in West Kazakhstan. In Russia, the species is mostly distributed in the steppe zone to the Cis-Urals Region. In the ZIN collection, there are 30 spm. from Samara and Astrakhan provinces.

In the Volga Region, the beetles occur in dry horse dung.

102. Aphodius (Heptaulacus) villosus Gyllenhal, 1806.

The species range includes Ukraine (the Carpathian Mountains), North and Central Europe. The species is known from Nizhny Novgorod, Volgograd, Rostov provinces (Kabakov and Frolov, 1996) and Karachay-Cherkessia (Shokhin, 2007). In the ZIN collection, there are 9 spm. from Leningrad and Ulyanovsk provinces, North Osetia, Tomsk Prov. And the Republic of Altai.

Subgenus Labarrus Mulsant et Rey, 1870

103. Aphodius (Labarrus) lividus (Olivier, 1789).

The species occurs almost throughout Europe, except for the extreme north, in North Africa, the Caucasus, Asia Minor, Kazakhstan, and Middle Asia. In the ZIN collection, there are 100 spm. from Belgorod, Volgograd, Rostov, Astrakhan provinces, Krasnodar Terr., Kalmykia, and Dagestan.

A coprophagous species occurring in horse and cow dung.

Subgenus Limarus Mulsant et Rey, 1870

104. Aphodius (Limarus) maculatus Sturm, 1800.

The species occurs in Central and South Europe, Ukraine (the Carpathian Mountains), the Transcaucasus, and Asia Minor. In the ZIN collection, there are specimens from Krasnodar Terr. and Adygea.

The species occurs in forests, in deer dung.

Subgenus Liothorax Motschulsky, 1859

105. Aphodius (Liothorax) kraatzi Harold, 1868

The distribution range of this species includes Southern Europe, the Caucasus, the Transcaucasus, Asia Minor, West Kazakhstan, Middle Asia, and Afghanistan. In Russia, the species occurs mostly in steppe and desert zones, from the Azov Sea to the Lower Volga region. In the ZIN collection, there are 100 spm. from Nizhny Novgorod, Volgograd, Rostov, Astrakhan provinces, and Kalmykia.

This species does not occur in dung. We collected specimens from sand and in light traps.

106. Aphodius (Liothorax) linearis Reiche et Saulcy, 1856.

The distribution range of this species includes Southern Europe (Portugal, Spain, Greece), France, Syria, Palestine. In Russia, the species is known from the Volga Region. In the ZIN collection, there are 7 spm. from Volgograd, Kurgan provinces, and Tatarstan.

107. Aphodius (Liothorax) niger (Panzer, 1797).

The species range includes almost the whole of Europe, except for the extreme north, the Caucasus, Asia Minor, Syria, and Middle Asia. In the ZIN collection, there are 10 spm. from Leningrad, Nizhny Novgorod, Moscow, Volgograd provinces and Krasnodar Terr.

108. Aphodius (Liothorax) plagiatus (Linnaeus, 1767).

The distribution range of this species includes almost the whole of Europe, except for the extreme north, North Africa (Tunisia), the Transcaucasus, Asia Minor, Syria, Kazakhstan, Middle Asia, Mongolia, and North China. In the ZIN collection, there are 100 spm. from Leningrad, Tver, Nizhny Novgorod, Saratov, Volgograd, Astrakhan provinces, the Republic of Altai, Irkutsk Prov., Buryatia, Yakutia, and the Amur Prov.

The species occurs in riverside debris.

109. Aphodius (Liothorax) rusakovi Gusakov, 2004.

The species is known from the original description from Rostov and Orenburg provinces of Russia, Kharkov Prov. of Ukraine and Ural Prov. of Kazakhstan.

Subgenus Loraphodius Reitter, 1892

110. Aphodius (Loraphodius) latisulcus Reitter, 1892.

The species is known from the Caucasus and from the Crimea. It was reported from Krasnodar Terr. (Shokhin, 2007).

111. Aphodius (Loraphodius) suarius Faldermann, 1836.

The species occurs in South-Eastern Europe, North Africa, the Caucasus, the Transcaucasus, Asia Minor, Syria, Iran (Dellacasa, 1983). Indicated for Rostov Province (Arzanov et al., 1996), and Chechnia (Shokhin, 2007).

Subgenus Loraspis Mulsant et Rey, 1870

112. Aphodius (Loraspis) frater Mulsant et Rey, 1870.

The distribution range of this species includes Central and South Europe, the Caucasus, the Transcaucasus, Asia Minor, Kazakhstan, and Middle Asia. In Russia, the species is mostly distributed in the Vorga Region and the Ciscaucasus. In the ZIN collection, there are 40 spm. from Saratov, Volgograd, Samara, Kurgan, Astrakhan provinces, Kalmykia, and Dagestan.

The species occurs in diverse biotopes, mostly in dry cow and horse dung. It is rather rare throughout its range.

Subgenus Mecynodes (Mulsant et Rey, 1870)

113. Aphodius (Mecynodes) kisilkumi Solsky, 1876.

The distribution range of this species includes the Transcaucasus, South-West Kazakhstan, and Middle Asia. In Russia, the species is known from the Lower Volga region. In the ZIN collection, there are 20 spm. from Astrakhan Prov. and Kalmykia.

Subgenus Melaphodius Reitter, 1892

114. Aphodius (Melaphodius) caspius Ménétriés, 1832.

The species is distributed in the steppe zone from the Caucasus to West Siberia, in Kazakhstan and Middle Asia. In the ZIN collection, there are 100 spm. from Penza, Saratov, Volgograd provinces, Stavropol Terr., Astrakhan, Ulyanovsk, Samara provinces, Bashkiria, Chelyabinsk, Tomsk, Kemerovo provinces, Khakassia, Krasnoyarsk Terr., and Irkutsk Prov.

The species occurs in cow dung in open biotopes with meadow and steppe vegetation.

115. Aphodius (Melaphodius) circumcinctus W. Schmidt, 1840.

The species is distributed in South-Eastern Europe, Kazakhstan, Turkmenistan, and Kyrgyzstan. In Russia, it mostly occurs in the steppe zone from the Ciscaucasus to West Siberia. In the ZIN collection, there are 22 spm. from Ulyanovsk, Volgograd, Rostov provinces, Stavropol Terr., Kalmykia, and Altai Terr.

Subgenus Melinopterus Mulsant, 1842

116. Aphodius (Melinopterus) consputus (Creutzer, 1799).

The species occurs in Central and South Europe, North Africa (Morocco, Algeria, Tunisia), Ukraine, the Caucasus, in West Asia. It was reported from Krasnodar Terr. (Arzanov et al., 1996).

117. Aphodius (Melinopterus) meuseli Reitter, 1906.

The species is known only from the type series from Tomsk environs. In the ZIN collection, there are no specimens of this species.

118. Aphodius (Melinopterus) prodromus (Brahm, 1790).

The species occurs throughout Europe, in North Africa (Morocco, Algeria), the Caucasus, the Transcaucasus, Asia Minor, Syria, Lebanonе, Israel, Kazakhstan, Middle Asia, and Mongolia; it was imported to USA and Canada. In the ZIN collection, there are 300 spm. from Karelia, Kaliningrad, Leningrad, Novgorod, Bryansk, Arkhangelsk, Vologda, Yaroslavl, Nizhny Novgorod provinces, Chuvashia, Moscow, Kaluga, Ryazan, Belgorod, Lipetsk, Saratov, Volgograd provinces, Krasnodar and Stavropol territories, Adygea, Karachay-Cherkessia, North Osetia, Dagestan, Kirovsk Prov., Udmurtia, Tatarstan, Samara Prov., Bashkiria, Chelyabinsk, Tyumen, Novosibirsk provinces, Altai Terr., the Republic of Altai, Krasnoyarsk Terr., Khakassia, Irkutsk Prov., Zabaykalsky Terr., and Yakutia.

A coprophagous species feeding on dung of cows, horses and wild ungulates. In Russia, this species is common; in southern regions, it occurs from March to December.

119. Aphodius (Melinopterus) punctatosulcatus Sturm, 1805.

A. sabulicola Thomson, 1868.

The species is distributed in Europe, except for the extreme north, North Africa (Morocco, Tunisia), the Trancaucasus, West Asia, Kazakhstan, and Middle Asia. In Russia, it is common in the European part, reaching the Transbaikal Region to the east. In the ZIN collection, there are 500 spm. from Leningrad, Novgorod, Arkhangelsk, Yaroslavl, Nizhny Novgorod provinces, Chuvashia, Moscow, Kaluga, Ryazan, Orel, Penza, Saratov, Astrakhan provinces, Dagestan, Kirovsk Prov., Tatarstan, Orenburg, Tyumen, Tomsk provinces, Altai Terr., the Republic of Altai, Krasnoyarsk Terr., Khakassia, Irkutsk Prov., Buryatia, Zabaykalsky Terr., and Yakutia.

120. Aphodius (Melinopterus) scuticollis Semenov, 1898.

The distribution range of this species includes South-Eastern Kazakhstan, Middle Asia and North-West China. In Russia, it is known from a few specimens from Altai Terr. and Khakassia.

121. Aphodius (Melinopterus) sphacelatus (Panzer, 1798).

The species occurs in the greatest part of Europe, in North Africa, The Caucasus, Asia Minor, and Middle Asia. In Russia, it is known mostly from the European part. In the ZIN collection, there are 80 spm. from Leningrad, Novgorod, Pskov, Vologda, Yaroslavl, Vladimir, Nizhny Novgorod, Moscow, Kaluga, Lipetsk provinces, and Krasnodar Terr.

Subgenus Mendidaphodius Reitter, 1901

122. Aphodius (Mendidaphodius) brancsiki Reitter, 1899.

The species is widely distributed in deserts of Middle Asia and West Kazakhstan (Nikolajev, 1987). It was reported from Astrakhan Prov. (Shokhin, 2007).

Subgenus Mendidius Harold, 1868

123. Aphodius (Mendidius) baigakumi (W. Koshantschikov, 1911).

The species was described from Turkmenistan (Baigakum Desert). In the ZIN collection, there are no specimens from Russia. Shokhin (2007) reported it from Astrakhan Prov.

124. Aphodius (Mendidius) bidens (Solsky, 1876).

The species is known from eastern West Asia, from Middle and Central Asia. It was reported from Buryatia and Zabaykalsky Terr. (Berlov 1989).

125. Aphodius (Mendidius) bispinifrons Reitter, 1889.

The species is known from Armenia, the North Caspian Region, Middle Asia, and North China (Nikolajev, 1987). It was reported from Volgograd Prov. (Shokhin, 2007).

126. Aphodius (Mendidius) curtulus (Harold, 1866).

Medvedev and Nikritin (1974) reported it from South and South-Eastern Ukraine. In the ZIN collection, there are 14 spm. from Astrakhan Prov. of Russia (Dosang environs).

We collected the beetles in October and April among cereal grass roots, in fixed sands and small patches of barchan sands. One specimen was also collected in horse dung, and a few specimens were collected in a light trap.

127. Aphodius (Mendidius) endroedii Balthasar, 1967.

The species is known only from the original description from the Transbaikal Region. It is possible that this name is a junior synonym of A. fimbriolatus.

128. Aphodius (Mendidius) fimbriolatus Mannerheim, 1849.

The species is known from a few localities in Central Asia. In the ZIN collection, there are 3 spm. from Irkutsk Prov. and Buryatia.

129. Aphodius (Mendidius) multiplex Reitter, 1897.

This species is widely distributed in the deserts and semideserts of Middle Asia, in the Lower Volga Region, the Ciscaucasus. In the ZIN collection, there are 50 spm. from Volgograd and Astrakhan provinces.

The species occurs in horse and cow dung in arid biotopes.

130. Aphodius (Mendidius) nelsinae (Medvedev, 1968).

The species is known from Middle Asia. Shokhin (2007) reported it from Astrakhan Prov. of Russia (Dosang environs). In the ZIN collection, there are no specimens from Russia.

Subgenus Mothon Semenov et Medvedev, 1927

131. Aphodius (Mothon) sarmaticus Semenov et Medvedev, 1927.

The species was described from South Ukraine (riverside of the Lower Dnepr). In the ZIN collection, there are 2 spm. from Krasnodar Terr.

The species occurs in riverine sands, among grass roots (Medvedev, 1965).

Subgenus Neagolius W. Koshantschikov, 1912

132. Aphodius (Neagolius) abchasicus Reitter, 1892.

The species occurs in the alpine and subalpine altitudinal zones of the Greater and Lesser Caucasus. It is known from Krasnodar Terr., Karachay-Cherkessia, North Osetia, and Dagestan (Shokhin, 2007).

Subgenus Nialus Mulsant et Rey, 1870

133. Aphodius (Nialus) varians Duftschmid, 1805.

The distribution range of this species includes Central and Southern Europe, North Africa, the Transcaucasus, Asia Minor, Kazakhstan, and Middle Asia. In Russia, it is mostly distributed in the steppe zone and forest-steppe subzone of the zone of deciduous forest from the western border to South Siberia. In the ZIN collection, there are 50 spm. from Nizhny Novgorod, Ryazan, Belgorod, Voronezh, Saratov, Volgograd, Rostov provinces, Krasnodar Terr., Astrakhan Prov., Dagestan, Samara Prov., Bashkiria, Orenburg Prov., and Khakassia.

The species occurs in diverse biotopes, in cow and horse dung, and in riverside debris.

Subgenus Nimbus Mulsant et Rey, 1870

134. Aphodius (Nimbus) circassicus Reitter, 1892.

The species is endemic to the Caucasus. It is known from a few localities in the Greater Caucasus Range. It was reported from Krasnodar Terr. (Kabakov and Frolov, 1996; Shokhin, 2007).

135. Aphodius (Nimbus) lederi Harold, 1876.

The species is endemic to the Caucasus. In the ZIN collection, there are 4 spm. from Krasnodar Terr.

The biology of this species is poorly known. Probably it occurs in foothill and mid-altitude forests (S. I. Tarasov, pers. comm.).

136. Aphodius (Nimbus) obliteratus Panzer, 1823.

The species is widely distributed in Central and South Europe, occurring also in Ukraine (Transcarpathian Region), the Transcaucasus, and Asia Minor. In the ZIN collection, there are 6 spm. from Dagestan.

137. Aphodius (Nimbus) affinis Panzer, 1823.

A. hoberlandti Tesar, 1945.

The species is known from a few localities in East Europe (Juřena et al., 2008; Král, 2008) and the South Urals (A. hoberlandti). In the ZIN collection, there are no specimens from Russia.

Subgenus Nobius Mulsant et Rey, 1870

138. Aphodius (Nobius) dosangi Akhmetova et Frolov, 2008.

The species is distributed in the Caspian Depression. In the ZIN collection, there are 200 spm. from Astrakhan Prov.

The beetles feed on cow and horse dung, the larvae were found in horse dung in Dosang environs.

139. Aphodius (Nobius) gresseri Semenov, 1898.

A. korgaldzhensis Nikolajev, 1987.

The species is known from West and Central Kazakhstan. In the ZIN collection, there are 43 spm. from Vladimir, Voronezh, Volgograd and Astrakhan provinces of Russia.

A coprophagous species. All the beetles and larvae we collected were found in horse dung on sandy soils.

140. Aphodius (Nobius) inclusus Reitter, 1892.

The species is distributed in the Caucasus, the Transcaucasus, and Middle Asian mountains. In the ZIN collection, there are 5 spm. from Dagestan.

141. Aphodius (Nobius) serotinus (Panzer, 1799).

A. x-signum Reitter, 1892.

The distribution range of this species includes Central, South and Eastern East Europe, the Transcaucasus, Asia Minor, Kazakhstan, and Middle Asia. In the ZIN collection, there are 200 spm. from Leningrad, Novgorod, Yaroslavl, Nizhny Novgorod, Moscow, Ryazan, Voronezh, Penza, Volgograd, Rostov provinces, Karachay-Cherkessia, Astrakhan Prov., Tatarstan, Ulyanovsk, Samara, Sverdlovsk, Tyumen, Kemerovo provinces, Republic of Altai, Krasnoyarsk Terr., Khakassia, Irkutsk Prov., and Buryatia.

The species occurs in cow and horse dung, mostly on open biotopes. The adults are active from the end of July to October; larvae were collected in Astrakhan Prov., in April and May.

Subgenus Orodaliscus Reitter, 1900

142. Aphodius (Orodaliscus) rotundangulus Reitter, 1900.

The distribution range of this species includes South Ukraine and Kazakhstan (except for its southern part). In Russia, the species is mostly distributed in the steppe zone of the Ciscaucasus and the Volga Region. In the ZIN collection, there are 42 spm. from Voronezh, Saratov, Volgograd, Rostov, Kurgan provinces, and Stavropol Terr.

A specialist nidicolous species occuring in gopher and marmot holes.

143. Aphodius (Orodaliscus) spalacophilus Novikov, 1996.

The species was described from East Ukraine. In the ZIN collection, there is 1 spm. from Kursk Prov.

The species occurs in mole rat holes (Novikov, 1996).

144. Aphodius (Orodaliscus) zangi A. Schmidt, 1906.

The distribution range of this species includes North Kazakhstan and North Kirgizstan. In Russia, the species is known from a few localities in the steppe zone from the Volga Region to the Ural Mountains. In the ZIN collection, there are 80 spm. from Astrakhan, Samara, Orenburg, and Kurgan provinces.

A specialized nidicolous species occuring in marmot holes.

Subgenus Otophorus Mulsant, 1842

145. Aphodius (Otophorus) haemorrhoidalis (Linnaeus, 1758).

The distribution range of this species includes the whole of Europe, North Africa, the Caucasus, the Transcaucasus, Kazakhstan, Middle Asia, Afghanistan, Mongolia, North and South-Western China, the Korean Peninsula, and Japan; it was imported to North America. In the ZIN collection, there are 500 spm. from the majority of the regions of Russia.

The species mostly occurs in cow and horse dung, in open biotopes.

Subgenus Parammoecius Seidlitz, 1891

146. Aphodius (Parammoecius) asphaltinus Kolenati, 1846.

The species is endemic to the Caucasus. In the ZIN collection, there are 25 spm. from Krasnodar Terr., Adygea, Karachay-Cherkessia, Kabardino-Balkaria, North Osetia, and Dagestan.

A coprophagous species occuring in cow dung at 1800–3000 m a.s.l.

147. Aphodius (Parammoecius) brevithorax Sumakov, 1903.

The species is known from Abkhasia, Georgia and North-Eastern Turkey (Ziani, 1999). It was reported from Krasnodar Terr. (Shokhin, 2007).

A specialized nidicolous species occuring in the holes of long-clawed mole-vole (Prometheomys schaposchnikowi).

Subgenus Phaeaphodius Reitter, 1892

148. Aphodius (Phaeaphodius) costalis Gebler, 1848.

The species is known from the steppe zone from the Volga Region to the Altai Mountains. In the ZIN collection, there are 7 spm. from Astrakhan Prov. and the Republic of Altai (Chuya Steppe).

149. Aphodius (Phaeaphodius) dauricus Harold, 1863.

The species is known from Asia Minor, Iran, Kazakhstan, Middle Asia, Afghanistan, Mongolia, West China (Kabakov, 1998). It was reported from Irkutsk Prov. and Khabarovsk Terr. (Berlov 1989).

150. Aphodius (Phaeaphodius) jouravliowi Reitter, 1907.

The species is known from the type specimens from Uralsk (North-Western Kazakhstan) and from the Altai Mountains (without more precise indication of locality). The taxonomic status of this nominal species requires clarification. In the ZIN collection, there are no specimens of this species.

151. Aphodius (Phaeaphodius) novikovi Kabakov, 1998.

The species is known from a type series from East Ukraine and the Lower Volga region. It was reported from Rostov and Volgograd provinces (Shokhin, 2007).

152. Aphodius (Phaeaphodius) rectus Motschulsky, 1866.

The species is widely distributed in the Central and Eastern Palearctic Region from the Volga Region to the Japan islands. The western border of its range is unclear. In 2008 г., N.V. Prasolov collected a series of 14 spm. in Leningrad Prov. In the ZIN collection, there are 1000 spm. from Leningrad, Astrakhan, Ulyanovsk, Samara, Chelyabinsk, Novosibirsk provinces, Altai Terr., the Republic of Altai, Irkutsk Prov., Buryatia, Zabaykalsky Terr., the Amur Prov., the Jewish Autonomous Province, Khabarovsk and Primorsky territories, and Sakhalin Prov.

A common species, abundant locally in the Far East. It occurs from March to September in open biotopes, pastures, in cow and horse dung.

153. Aphodius (Phaeaphodius) roddi W. Koshantschikov, 1911.

The species is known from a type series from Altai Mountains. It was reported from North-Eastern Kazakhstan (Nikolajev, 1998).

154. Aphodius (Phaeaphodius) scoparius Harold, 1877.

The species is known from South Siberia, Mongolia and North-Eastern China. In the ZIN collection, there are 7 spm. from Buryatia, Zabaykalsky Terr. and the Amur Prov.

Subgenus Phalacronothus Motschulsky, 1859

155. Aphodius (Phalacronothus) biguttatus Germar, 1824.

The species is distributed in Central, South and Eastern Europe, the Transcaucasus, Asia Minor, Kazakhstan, and Turkmenistan. In Russia, it is known from Ulyanovsk (Isajev, 1995) and Novosibirsk (Zinchenko, 1999) provinces. In the ZIN collection, there are 11 spm. from Volgograd, Rostov, Orenburg provinces and Krasnoyarsk Terr.

156. Aphodius (Phalacronothus) citellorum Semenov et Medvedev, 1928.

The species is distributed in the steppe zone of Central and Eastern Europe and Kazakhstan.

A nidicolous steppe species occuring in gopher and marmot holes (Zinchenko et al., 2002; Isajev, 1995).

157. Aphodius (Phalacronothus) quadrimaculatus (Linnaeus, 1761).

The species is widely distributed in Central and South Europe; it is also known from North Africa, the Transcaucasus, Asia Minor, Syria and Turkmenistan. In the ZIN collection, there are 11 spm. from Astrakhan Prov.

The species occurs in cow dung.

Subgenus Pharaphodius Reitter, 1892

158. Aphodius (Pharaphodius) rugosostriatus Waterhouse, 1875.

A. raddei Berlov, 1989

The distribution range of this species includes North and Central China, Southern Primorye, the Korean Peninsula, and Japan. In the ZIN collection, there are 50 spm. from Primorye.

The species occurs in cattle dung.

Subgenus Plagiogonus Mulsant, 1842

159. Aphodius (Plagiogonus) arenarius Olivier, 1789.

A. putridus (Fourcroy, 1785).

This species is widely distributed in Europe, occurs also in the Caucasus, the Transcaucasus, Turkey, Kazakhstan, and Turkmenistan. In Russia, it is known from the steppe zone from Kursk Prov. to West Siberia. In the ZIN collection, there are 100 spm. from Chuvashia, Nizhny Novgorod, Kursk, Saratov, Volgograd, Rostov provinces, Dagestan, Samara Prov., the Republic of Altai, and Krasnoyarsk Terr.

The species occurs in rodent holes (mostly of marmots and gophers).

160. Aphodius (Plagiogonus) culminarius Reitter, 1900.

The distribution range of this species includes North-Eastern China and Primorye. In the ZIN collection, there are 7 spm. from Primorye.

Subgenus Planolinus Mulsant et Rey, 1870

161. Aphodius (Planolinus) borealis Gyllenhal, 1827.

The species is widely distributed in Europe, it also occurs in the Transcaucasus, Asia Minor, Eastern Kazakhstan, and Mongolia. In Russia, it is distributed in the forest zone, from the western border to Kamchatka. In the ZIN collection, there are 30 spm. from Karelia, Murmansk and Leningrad provinces, Adygea, Buryatia, Irkutsk Prov., Zabaykalsky Terr., Amur and Magadan provinces, Khabarovsk and Primorsky territories.

A coprophagous species occuring in forest biotopes.

162. Aphodius (Planolinus) vittatus Say, 1825.

The species occurs in South and Eastern Europe, Transcaucasus, Turkey, Syria, Kazakhstan, Middle Asia, Mongolia, and China (Dellacasa, Dellacasa, 2006). In Russia, it is known from the Volga Region, the Altai Mountains, and the Transbaikal Region. In the ZIN collection, there are 100 spm. from Astrakhan and Saratov provinces, the Republic of Altai, Tuva, Buryatia, and Zabaykalsky Terr.

Distribution records of this species from the Palearctic Region refer to A. vittatus mundus Reitter, 1892, A. vittatus sellatus Mannerheim, 1852, and A. vittatus tjanshanicus Balthasar, 1956. The nominative subspecies is distributed in South Canada, most of the USA and in North Mexico (Gordon and Skelley, 2007). Maté (2003: cited from Wilson and Angus, 2006) suggested that the American subspecies differs from the Eurasian ones in the molecular characters; however the stutus of these nominal taxa needs clarification.

Subgenus Platyderides A. Schmidt, 1916

163. Aphodius (Platyderides) suvorovi Kabakov, 1996

The species is known from a single specimen from the Amur Prov., housed in the ZIN collection.

Subgenus Pseudacrossus Reitter, 1892

164. Aphodius (Pseudacrossus) grebenschikovi Balthasar, 1961.

The species was described from Mongolia and also recorded from Zabaykalsky Terr. (Berlov 1989). In the ZIN collection, there are no specimens of this species.

165. Aphodius (Pseudacrossus) nasutus Reitter, 1887.

The species is known from Mongolia and North China (up to East Tibet in the south) (Král, 1997). In Russia, it is distributed in South Siberia and Amur Region. In the ZIN collection, there are 50 spm. from the Altai Mountains, Krasnoyarsk Terr., Irkutsk Prov., Zabaykalsky and Khabarovsk territories.

166. Aphodius (Pseudacrossus) tenebricosus A. Schmidt, 1916.

The species is known from East Kazakhstan and Mongolia (Zinchenko et al., 2002). In Russia, it was found in Novosibirsk Prov. (Zinchenko, 1999) and Tyva (Zinchenko et al., 2002).

The species occurs in gray marmot holes (Zinchenko et al., 2002).

Subgenus Pubinus Mulsant et Rey, 1870

167. Aphodius (Pubinus) tomentosus (Müller, 1776).

The species is known from Northern, Central and Eastern Europe, Kazakhstan (Nikolajev, 1998). In Russia, it is distributed from western border to Transbaikal Region. In the ZIN collection, there are 100 spm. from Leningrad, Novgorod, Pskov, Tver, Yaroslavl, Vladimir, Nizhny Novgorod, Moscow, Rostov provinces, Tatarstan, Samara, Orenburg, Tyumen, Omsk, Novosibirsk, Irkutsk provinces, Altai and Krasnoyarsk territories. It was reported from Stavropol Terr. (Arzanov et al., 1996) and Karachay-Cherkessia (Shokhin, 2007).

The species feeds on cow and horse dung.

Subgenus Sigorus Mulsant et Rey, 1870

168. Aphodius (Sigorus) porcus (Fabricius, 1792)

The species is widely distributed in Central and South Europe, known from Ukraine (Carpathian Mountains, Kherson Prov.), the Caucasus, Asia Minor and Turkmenistan (Kopet Dagh). In Russia, it is known from Rostov Prov. (Shokhin, 2007).

Subgenus Sinodiapterna Dellacasa, 1986

169. Aphodius (Sinodiapterna) gorodinskiyi Gusakov, 2006.

The species is known from single specimen from foothills of Pogranichny Range in Promorsky Terr. In the ZIN collection, there are no specimens of this species.

170. Aphodius (Sinodiapterna) hammondi Dellacasa, 1986.

The species was described from North China. In the ZIN collection, there are 2 spm. from Primorye.

171. Aphodius (Sinodiapterna) troitzkyi Jacobson, 1897.

The species was recorded from China, North Korea, Taiwan, and Japan (Dellacasa, 1986). In Russia, it is known from several findings in southern Far East. In the ZIN collection, there are 10 spm. from Khabarovsk and Primorsky territories.

A coprophagous species occurring in cow dung.

Subgenus Stenotothorax A. Schmidt, 1913

172. Aphodius (Stenotothorax) hibernalis Nakane et Tsukamoto, 1956.

The species was described from Japan. In Russia, it is known from Amur Prov. (Kabakov and Frolov, 1996), Khabarovsk Terr. and Sakhalin Island (Berlov 1989). In the ZIN collection, there is 1 spm. from Primorye, collected by O.N.Kabakov from soil under rotten mushrooms.

Subgenus Subrinus Mulsant et Rey, 1870

173. Aphodius (Subrinus) subtilis D. Koshantschikow, 1894.

The species is known only from the original description from Astrakhan (Koshantschikow, 1894c). The type depository is unknown.

174. Aphodius (Subrinus) sturmi Harold, 1870.

The species occurs in Central and South Europe, North Africa, the Caucasus, the Transcaucasus, Asia Minor, Kazakhstan, Middle Asia, Iran, Mongolia, North-Eastern China, the Korean Peninsula, and Japan. In Russia, it mostly occurs in the steppe zone and the forest-steppe subzone of the deciduous forest zone from the Azov Sea to Sakhalin Island. In the ZIN collection, there are 40 spm. from Volgograd and Rostov provinces, Stavropol Terr., Astrakhan and Samara provinces, Dagestan, Primorsky Terr., and Sakhalin Prov.

A coprophagous species mostly occurring in cow dung in open biotopes.

Subgenus Sugrames Reitter, 1894

175. Aphodius (Sugrames) hauseri (Reitter, 1894).

The species is widely distributed in sand deserts of Kazakhstan and Middle Asia; it was also found in Afghanistan. In Russia, it is known from the Lower Volga region and Dagestan. In the ZIN collection, there are 10 spm. from Astrakhan Prov.

A psammophillous species. We collected a few larvae among grass roots on a small patch of barchans sands in Astrakhan Prov.

Subgenus Teuchestes Mulsant, 1842

176. Aphodius (Teuchestes) brachysomus Solsky, 1874.

The distribution range of this species includes Central and Eastern China, Korean Peninsula and Japan. In Russia, the species is known from Amur Region, Primorye and Sakhalin Isl. In the ZIN collection, there are 100 spm. from Khabarovsk and Primorsky territories.

A coprophagous species mostly occuring in cow dung from March to July.

177. Aphodius (Teuchestes) fossor (Linnaeus, 1758).

The distribution range of this species includes almost the whole of Europe (except for the extreme north), North Africa (Algeria), The Caucasus, Transcaucasus, West Asia, Kazakhstan, Middle Asia and Northern Mongolia. It was imported to North America. In Russia, it is distributed throughout the country up to Zabaykalsky Terr. (Nerchinsk) in the east. In the ZIN collection, there are 300 spm. from Karelia, Leningrad, Pskov, Tver, Smolensk, Vologda, Yaroslavl, Kostroma, Nizhny Novgorod provinces, Chuvashia, Moscow, Ryazan, Belgorod, Lipetsk, Voronezh, Penza, Saratov provinces, Krasnodar and Stavropol territories, Adygea, Kabardino Balkariya, Dagestan, Kirovsk, Samara provinces, Bashkiria, Sverdlovsk, Chelyabinsk, Tyumen, Omsk, Novosibirsk, Tomsk, Kemerovo provinces, Altai Terr., Republic of Altai, Krasnoyarsk Terr., Khakassia, Tuva, Irkutsk Prov., Buryatia, and Zabaykalsky Terr.

The species occurs in open biotopes, pastures, in cow and horse dung. In Russia, it is a common species; the beetles are active from April to September.

Subgenus Trichaphodius A. Schmidt, 1913

178. Aphodius (Trichaphodius) comatus A. Schmidt, 1920.

The distribution range of this species includes China, the Korean Peninsula and Japan. In Russia, it is known only from southern Primorsky Terr.; in the ZIN collection there are 8 spm.

A coprophagous species occurring in cow and horse dung.

Subgenus Trichonotulus Bedel, 1911

179. Aphodius (Trichonotulus) scrofa (Fabricius, 1787).

The species occurs in Europe (up to Finland in the north), North Africa (Morocco), the Caucasus, the Transcaucasus, Asia Minor, Kazakhstan, Middle Asia, Afghanistan, Mongolia, China, North Korea. It was imported to Canada and the USA. In Russia, the species is mostly distributed in the forest-steppe subzone of the deciduous forest zone from the western border to the Amur Region. In the ZIN collection, there are 21 spm. from Leningrad, Nizhny Novgorod, Volgograd, Rostov provinces, Karachay-Cherkessia, Astrakhan and Irkutsk provinces.

A coprophagous species occuring in ungulate dung.

Subgenus Vladimirellus Dellacasa, Dellacasa et Bordat, 2002

180. Aphodius (Vladimirellus) socors Balthasar, 1967.

The species is known from 3 findings in South Siberia and the Amur Region (Balthasar, 1967; Zinchenko and Bezborodov, 2013). In the ZIN collection, there are no speimens of this species.

Genus Cnemisus Motschulsky, 1868

The genus includes 4 species distributed in the sand deserts and semideserts of Eurasia. In the territory of Russia, 2 species occur.

181. Cnemisus kaznakovi (Semenov, 1903).

The species was described from North China and also reported from Mongolia (Nikolajev and Puntsagdulam, 1984). Berlov and Anishchenko (1998) reported it from Buryatia. In the ZIN collection, there are no specimens of this species from Russia.

182. Cnemisus rufescens (Motschulsky, 1845).

The species is distributed in the Caspian Depression. In the ZIN collection, there are 80 spm. from Astrakhan Prov.

The species occurs in barchan sands, under saltworts of the genus Salsola. The beetles are active in late fall.

Genus Oxyomus Stephens, 1839

The genus comprises up to 27 species distributed in all the zoogeographical regions except for Australian. In Russia, only the type species of the genus occurs.

183. Oxyomus silvestris (Scopoli, 1763).

The species is distributed in Europe, North Africa (Tunisia), the Caucasus, the Transcaucasus, Middle Asia, and Asia Minor; it was imported to North America. In Russia, it occurs from North-Western border to Kirovsk and Samara provinces. In ZIN collection, there are 24 spm. from Leningrad Prov., Chuvashia, Kaluga, Ryazan provinces, Krasnodar Terr., North Osetia, Dagestan, Kirovsk and, Samara provinces.

The species occurs in rotten plant residues and in dung.

Pattern of the Aphodiini species diversity in the territory of Russia

In Russia, aphodiines are the most diverse in the southern European Part (Fig. 1). This can be explained by a variety of natural conditions (e.g. altitudinal zonation of the Caucasus) and a long vegetative season with a high sum of effective temperatures. The total number of species is high due to the species with Wide Palearctic ranges, Caucasian endemics and subendemics, and the species having most of their ranges in South Europe.

The accuracy of our map (Fig. 1) is probably higher for plain regions than for the mountainous ones. The large average number of species per area unit in the mountainous regions (the Caucasus, South Siberia and, to a lesser degree, the southern Far East) can be the result of a loose extrapolation of the ranges of separate species and rather reflects the total number of species of the regional faunas. The richest local faunas are found in the Lower Volga Region: 47 species are known from the vicinity of Elton Lake (Makarov et al., 2009), 45, from Dosang environs (Shokhin, 2007; Frolov and Akhmetova, 2013). In the latter case, at least 43 species have stable populations in the territory of about 3 square kilometers (Frolov, Akhmetova, 2013).

Рис. 1. Карта распределения относительного видового разнообразия Aphodiini фауны России.

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.

With the advance to the north, the aphodiine diversity gradually declines. At the latitude of 65°N, less than 10 species occur. The north border of the distribution of the members of the tribe is not clear. However, in the areas with the permafrost lying near the surface horizon, aphodiines probably do not occur or are only represented by a few boreal species. Medvedev (1951) suggested that the absence of chafers in most parts of the West Siberian taiga is due to its heavy bogginess. Chafer larvae live in soil and cannot tolerate very high moisture. The absence of chafers from North-Eastern Siberia Medvedev explained by the severity of climate and by permafrost. Probably, the very low diversity of the Aphodiini in the northern taiga may be explained by the same factors. Aphodiine larvae develop in dung or soil and high moisture of the substrate, combined with low temperatures, is unfavorable to the development of all the scarabeids.

Distribution of the Aphodiini of the fauna of Russia according to landscape zones and altitudinal zonation types

For the analysis of the zonal distribution of the Aphodiini, the map of the landscape zones and vegetation altitudinal zonality types (National Atlas of Russia, 2007) was used. According to the map, the territory of Russia can be devided into 5 lanscape zones and 6 main altitudinal zonation types. Aphodiines occur in the taiga, broadleaved forest, steppe and desert zones, and in the boreal (taiga), nemoral (deciduous) and subarid altitudinal zonation types. Aphodiines are apparently absent from tundra zone, and arctic, hypoarctic tundra and hypoarctic taiga altitudinal zonation types.

Aphodiines are the most diverse in the steppe and broadleaved forest zones as well as the nemoral zonation types. In the steppe zone, 54% of the species of the Russian fauna occur. Probably, the high diversity of the aphodiines in the steppe zone is due to their association with animals largely inhabiting open landscapes. Almost the same number of species occurs in the nemoral altitudinal zonation types. However, as opposed to the continuous steppe zone, these altitudinal zonation types are situated in distant regions and have a drastically different fauna of the aphodiines. 50% of species are known from the broadleaved forest zone. A reasonable part of these species inhabit open landscapes of the forest-steppe subzone. In the taiga and desert zones, as well as in the boreal and subarid types of altitudinal zonation, fewer species occur. 32 % of the species of Russian fauna are known from the taiga zone, and only 4% from the subarid zone. The relatively low species richness in these zones and altitudinal zonation types is due to the climate and environmental conditions being less suitable for the group.

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.

Typology of the ranges of the aphodiines of the Russian fauna

Cluster analysis (Jaccard index as similarity coefficient, average clustering) yielded a range similarity dendrogram which allowed us to group the distribution ranges of the Aphodiini of Russia into 14 main types (Fig. 2–5). For the nomenclature of the types of the ranges we used the schema of general biogeographic subdivisions of the Palearctic Region (Emeljanov, 1974).

The distribution ranges of the aphodiines occurring in Russia can be grouped into the following types:

Wide Palearctic (Fig. 2, а). This type comprises the widest ranges. The following species have the ranges that can be classed as Wide Palearctic: Aphodius depressus, A. sus, A. plagiatus, A. erraticus, A. fasciatus, A. ater, A. rufipes, A. haemorrhoidalis, A. subterraneus, A. pusillus, A. scrofa, A. luridus, A. piceus, A. borealis, A. villosus, A. rufus, A. brevis, A. tomentosus, A. foetens, A. fossor, A. carinatus, A. distinctus, A. fimetarius, A. granarius, A. immundus, A. melanostictus, A. prodromus, A. punctatosulcatus, A. vittatus, A. sordidus, A. rectus, A. sturmi, A. lapponum, A. dauricus. All these eurybiont species are widely distributed in the Palearctic Region. Most of them occur at least in 4 landscape and altitudinal zones.

Рис. 2. Типы ареалов Aphodiini.

Fig. 2. Types of the ranges of the Aphodiini.

a — Wide Palearctic, b — Wide West-Palearctic, c — Wide European Nemoral.

Wide West-Palearctic (Fig. 2, b). The ranges of the following species can be classed as Wide West-Palearctic: A. arenarius, A. biguttatus, A. coenosus, A. frater, A. ictericus, A. kraatzi, A. linearis, A. lividus, A. lugens, A. merdarius, A. niger, A. paykulli, A. punctipennis, A. quadriguttatus, A. satellitus, A. serotinus, A. sphacelatus, A. sticticus, A. testudinarius, A. varians, Oxyomus silvestris. These species are also eurybiont and rather widely distributed in the Palearctic Region. More than a half of them occur at least in 4 landscape and altitudinal zones. The ranges of this type are situated within different provinces of a few zoogeographical regions.

Wide European Nemoral (Fig. 2, c). The ranges of the two European species, A. conspurcatus and A. nemoralis, can be placed in this group. Their ranges substantially differ from those of other species and are situated within the European Nemoral Region (except for Euxine Mountain Province) and transitional zones between the Eurosiberian Boreal and European Nemoral regions and between European Nemoral and the Scythian Steppe regions. In Russia, both species inhabit the boreal (taiga) and broadleaf forest zones.

The Wide Mediterranean Euxinian type (Fig. 3, a) comprises the ranges of A. conjugatus, A. constans, A. maculatus, A. obliteratus, A. obscurus, A. porcus, A. quadrimaculatus, A. scrutator, A. suarius, A. thermicola. In the territory of Russia, most of these species occur in the steppe zone and the nemoral altitudinal zones.

The Ancient Mediterranean type (Fig. 3, b) comprises the ranges of A. trucidatus, A. pruinosus, A. hydrochaeris, A. brancsiki, A. consputus, A. aequalis, A. inclusus, and A. klugi, which are situated within the Ancient Mediterranean Region. In the territory of Russia, most of these species occur in the steppe and desert zones.

Рис. 3. Типы ареалов Aphodiini.

Fig. 3. Types of the ranges of the Aphodiini.

a — Wide Mediterranean Euxinian, b — Ancient Mediterranean, c — Westscythian-Northturanian.

The Westscythian-Northturanian type (Fig. 3, c) comprises the ranges of A. bimaculatus, A. caspius, A. circumcinctus, A. citellorum, A. costalis, A. gregarius, A. gresseri, A. hahni, A. planus, A. rotundangulus, and A. zangi. All these species inhabit steppe biotopes, and a part of them are specialized nidicols.

The Conblacksean Plane type (Fig. 4, a) comprises the ranges of A. curtulus, A. exilimanus, A. isajevi, A. ivanovi, A. novikovi, A. sarmaticus, A. spalacophilus. In the territory of Russia, most of these species occur in the steppe zone and forest-steppe subzone of the deciduous forest zone.

Рис. 4. Типы ареалов Aphodiini

Fig. 4. Types of the ranges of the Aphodiini.

a — Conblacksean Plane , b — Caucasian, c — Caspian.

The Caucasian type (Fig. 4, b) comprises the ranges of A. swaneticus, A. planicollis, A. lederi, A. circassicus, A. latisulcus, A. brignolii, A. leisteri, A. brevithorax, A. asphaltinus, A. alagoezi and A. abchasicus. All these species are endemic to the Caucasus and are distributed within Euxine Mountain Province of the European Nemoral Region.

The Sethian type (Fig. 5, a) comprises the ranges of A. badenkoi, A. bispinifrons, A. clathratus, A. digitalis, A. hauseri, A. kisilkumi, A. kizeritskyi, A. menetriesi, A. multiplex. These species mostly occur in the desert zone and inhabit arid biotopes. Their ranges are situated largely within the Irano-Turanian and Centralasiatic subregions of the Sethian Desert Region.

Рис. 5. Типы ареалов Aphodiini

Fig. 5. Types of the ranges of the Aphodiini.

a — Sethian , b — Centralasiatic , c — Stenopean Nemoral.

The Caspian type (Fig. 4, b) comprises the ranges of A. dosangi, A. plustschewskii, A. variicolor, and Cnemisus rufescens. In the territory of Russia, these species inhabit the desert zone. Their ranges almost completely lie in north-west part of the North Turanian Lowland Province of the Iranо-Turanian Subprovince of the Sethian Desert Region.

The Centralasiatic type (Fig. 5, b) comprises the ranges of A. antiquus, A. bidens, A. comma, A. crassus, A. falcispinis, A. fimbriolatus, A. grafi, A. grebenschikovi, A. indagator, A. jacobsoni, A. kerzhneri, A. longeciliatus, A. mongolaltaicus, A. nasutus, A. roddi, A. scoparius, A. scuticollis, A. sordescens, A. tanhensis, A. tenebricosus, and Cnemisus kaznakovi. The ranges of A. lapponum and A. dauricus belong to the Centralasiatic cluster of the dendrogram, but we think that they should be placed in the group of species with Wide Palearctic ranges. In Russia, the majority of the species with this type of ranges occur in the boreal altitudinal zones and the steppe zone.

The Stenopean Nemoral type (Fig. 5, c) comprises the ranges of A. amurensis, A. binaevulus, A. brachysomus, A. breviusculus, A. costatellus, A. comatus, A. culminarius, A. hammondi, A. koltzei, A. hibernalis, A. impunctatus, A. inexpectatus, A. languidulus, A. nigrotesselatus, A. notabilipennis, A. pratensis, A. proprietor, A. quadratus, A. rugosostriatus, A. subcostatus, A. sublimbatus, A. superatratus, A. tenax, A. troitzkyi, and A. uniformis. All these species inhabit nemoral altitudinal zones and the majority of them inhabit also the broadleaf forest zone. The range of A. amurensis is the least similar to the ranges of other species of this type. It is possible, that this species has a wider distribution in Primorye, North-Eastern China and North Korea, but is limited to the areas with stable populations of Japanese deer, as both beetles and larvae feed on their dung.

The distribution ranges of A. guttatus and A. urostigma are essentially different from those of other species and cannot be classed into any of the abovementioned types. Special distribution types can be proposed for these species:

The Canada-Kamchatian type (Fig. 6, a) is characteristic of A. guttatus. This species is widely distributed in the Nearctic Region and was recently found in Kamchatka.

Рис. 6. Типы ареалов Aphodiini.

Fig. 6. Types of the ranges of the Aphodiini.

a — Canada-Kamchatian , b — East-Chinese-Indomalayan.

The East-Chinese-Indomalayan type (Fig. 6, b) is characteristic of A. urostigma which is the only representative of tropical subgenus Aganocrossus in Russia. The range of this species mostly lies outside the Palearctic Region. Only the north-easternmost part of its range includes Southern Primorye while most part is situated in South and South-Eastern Asia.

Special zoogeographic regionization of the territory of Russia, based on the distribution of the Aphodiini and general zoogeographic subdivisions

Based on a cluster analysis, we distinguished 6 groups of zoogeographical provinces (Fig. 8) which correspond to separate clusters on the dendrogram (Fig. 7).

Рис. 7. Дендрограммы сходства фаун Aphodiini отдельных провинций.

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 mountain, 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.

Рис. 8. Группы провинций частного зоогеографического районирования территории России на основе распространения Aphodiini

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.

The European group of provinces comprises Middleuropean mixed, Easteuropean plane and Euxine mountain provinces of European Nemoral Region, as well as Conblacksean plain province of the Scythian Steppe Region. 99 aphodiine species, 98 of which belong to Aphodius and 1 to Oxyomus,inhabit this group of provinces. 65 species occur in the Middleuropean mixed province, 57, in the Easteuropean Plain, 80, in the Euxinian mountain, 73 species, in the Conblacksean Plain provinces. 29 species (A. paykulli, A. conspurcatus, A. conjugatus, A. consputus, A. constans, A. maculatus, A. obliteratus, A. porcus, A. scrutator, A. suarius, A. thermicola, A. abchasicus, A. alagoezi, A. asphaltinus, A. brevithorax, A. brignolii, A. circassicus, A. inclusus, A. latisulcus, A. lederi, A. obscurus, A. planicollis, A. swaneticus, A. curtulus, A. isajevi, A. ivanovi, A. novikovi, A. sarmaticus, and A. spalacophilus) are not known from other provinces of the studied area.

The Kazakhstan-Northturanian group of provinces comprises the homonymic provinces of the Scythian Steppe and Sethian Desert regions. 88 aphodiine species are known (86 Aphodius, 1 Cnemisus and 1 Oxyomus) from this territory. In the studied part of Northturanian Plane province, 80 aphodiine species occur; in Kazakhstanian Plane province 65 species occur. 15 species (A. zangi, A. badenkoi, A. brancsiki, A. digitalis, A. dosangi, A. hauseri, A. kisilkumi, A. kizeritskyi, A. menetriesi, A. plustschewskii, A. variicolor, A. exilimanus, A. hahni, A. roddi, and Cnemisus rufescens) are unknown from other provinces of the studied area.

the European-Westsiberian group of provinces comprises all the provinces of the Westeurosiberian Subregion of the Eurosiberian Boreal Region (except for the Altaian complex province). Here, 47 aphodiine species occur (46 Aphodius and 1 Oxyomus species). Twenty two species occur in the Bothnian Mixed province, 35, in the Zyrianian Plane province, 38, in the Uralian mountain province, 32 in the Obean plane province. All the species except for A. meuseli also occur in other provinces of the studied area. The distribution range of A. meuseli, so far known only from the type series, requires clarification. It is possible that this species has a wider range.

The Eastsiberian-Altaian-Eastmongolian group of provinces comprises the Westmongolian and Eastmongolian complex provinces of the Eastsiberian Subregion of the Eurosiberian Boreal Region, Altaian and Eastmongolian complex provinces of the Scythian Steppe Province. In the similarity dendrogram calculated with the Jaccard index, the Angaran mixed and Subokhotian mountain provinces of the Eurosiberian Boreal Region form a separate cluster most similar to the Eastsiberian-Altaian-Eastmongolian and European-Westsiberian group of provinces (Fig. 7, а). In the similarity dendrogram calculated with the correlation coefficient, they belong to a cluster with the Eastsiberian-Altaian-Eastmongolian group (Fig. 7, b); therefore we include them into this group of provinces. However, the greater degree of similarity of the two provinces with the other provinces of the group is mainly due to the Mongolian-Chinese species (A. antiquus, A. nasutus, A. scoparius) occurring only in the southern parts of the provinces. The territory occupied by this group of provinces is inhabited by 68 aphodiine species, of which 67 belong to Aphodius and one to Cnemisus. 20 species are known from the Angaran Mixed Province, 23, from the Subokhotian mountains, 48, from the Westmongolian Complex, 44, from Estmongolian Complex Province of the Eurosiberian Boreal Region, 57, from the Altaian Complex, 38, from the Estmongolian Complex Province of the Scythian Steppe Region. 15 species (A. antiquus, A. scoparius, A. falcispinis, A. grafi, A. bidens, A. fimbriolatus, A. longeciliatus, A. tanhensis, A. crassus, A. grebenschikovi, A. jacobsoni, A. kerzhneri, A. logunovi, A. mongolaltaicus, and Cnemisus kaznakovi) are not known from other provinces of the studied area.

The Stenopean group of provinces comprises the Weststenopean Mixed, Korean and the North Japanese Mountain Province of the Stenopean Nemoral Region, and the Okhotian Mountain Province of the Eurosiberian Boreal Region. This territory is inhabited by 47 aphodiine species which all belong to Aphodius. 40 species are known from the Weststenopean Mixed Province, 25, from the Korean Mountain Province, 23, from the North Japanese Mountain Province, 18, from the Okhotian Mountain Province. 25 Primoryan species (A. sublimbatus, A. uniformis, A. comatus, A. impunctatus, A. languidulus, A. rugosostriatus, A. subcostatus, A. superatratus, A. troitzkyi, A. urostigma, A. breviusculus, A. inexpectatus, A. propraetor, A. binaevulus, A. hammondi, A. tenax, A. hibernalis, A. pratensis, A. quadratus, A. amurensis, A. costatellus, A. culminarius, A. gorodinskiyi, A. koltzei, and A. suvorovi) are unknown from other provinces of the studied area.

The Northpacific Mixed Province of the Circumpolar Tundra Region differs most from the other provinces in its fauna. It cannot be placed in any other group. This can be explained by the extremely poor aphodiine fauna comprised of only 4 species of Aphodius: A. lapponum, A. borealis, A. aleutus, and A. guttatus. The two former species have wide Palearctic ranges. The uniqueness of this province is due to A. guttatus which is widely distributed in the Nearctic Region and was recently found in Kamchatka.

Acknowledgements

We are gfrateful to V.K. Zinchenko (Siberian zoological museum, Institute of animal systematics and ecology SB RAS, Novosibirsk), I.V. Shokhin (Azov Branch of Murmansk marine biological institute of KSC RAS, Rostov-on-Don), A.V. Ivanov (Institute of plant and animal ecology UB RAS, Ekaterinburg), S.A. Shabalin (Soil biology institute FEB RAS, Vladivostok), K.V. Makarov and A.V. Matalin (Moscow Pedagogical State University, Moscow), A.A. Gusakov (Zoological museum, Lomonosov Moscow State University, Moscow), G.V. Nikolaev (Al-Farabi Kazakh National University, Almaty), S.I.Tarasov (Natural History Museum, Oslo University, Oslo), and O.A. Khruleva (Severtsov Institute of Ecology and Evolution, Moscow) for providing material for this study and for assistance. The work was supported by the Russian Foundation for Basic Research (grant no 13-04-01002-a).

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