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

Full text

ISSN 0013-8738, Entomological Review, 2014, Vol. 94, No. 6, pp. 846–879. © Pleiades Publishing, Inc., 2014.
Original Russian Text © L.A. Akhmetova, А.V. Frolov, 2014, published in Entomologicheskoe Obozrenie, 2014, Vol. 93, No. 2, pp. 403–447.
846
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
e-mail: codocera@rambler.ru
Received January 10, 2014
Abstract—An annotated list of 183 species of the dung-beetle tribe Aphodiini of the Russian fauna is given. The
highest species diversity is characteristic of the southern mountainous regions of the country, the richest local fau-
nas being known from the Lower Volga area. The greatest numbers of species are known from the steppe and
broadleaved forest zones and from the nemoral types of altitudinal zonation of vegetation. The bulk of the fauna is
formed by the species with wide ranges; 111 species occur in two or more landscape zones or altitudinal zonation
types. Six groups of faunistically similar zoogeographic provinces are distinguished within the territory of Russia.
DOI: 10.1134/S0013873814060074
Scarab beetles of the tribe Aphodiini is a large,
world-wide distributed group of insects. The world
fauna comprises about 30 genera and over 2000 spe-
cies, including about 13 genera and 700 species in the
Palaearctic Region (Dellacasa, 1988; Dellacasa, Della-
casa, 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 Rus-
sian 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,
Transcucasus, and West Siberia, they include descrip-
tions of new species. Reitter (1893) summarized the
data about the Palaearctic 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 coleo-
pterologists 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) summa-
rized the available data on the Caucasian copro-
phagous 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 Er-
molenko (1969), Nikritin (1969а, 1969б), Medvedev
and Nikritin (1974), Berlov (1979, 1985, 1989), Niko-
lajev (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 au-
thors 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 taxo-
nomic 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

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847
Sciences (ZIN), accumulated during many decades,
remained mostly unstudied.
MATERIALS AND METHODS
This work is based on the rich collection of ZIN, in-
cluding vast material of more than 60 species collected
by the authors in Lower Volga Region and the south-
ern Far East. In addition, the materials from the fol-
lowing institutions and private collections have been
studied: Moscow Pedagogical State University, Mos-
cow, Zoological museum, Lomonosov Moscow State
University, Moscow, A.V. Ivanov’s collection, Ekate-
rinburg, 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 regioni-
zation, the scheme of the general zoogeographic sub-
divisions of the Palaearctic 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 correla-
tion coefficients. Average linkage was used as a clus-
tering 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 Reitter and Mothon Semenov et Medvedev
as subgenera.
Genus Aphodius Hellwig, 1798
Aphodius Illiger, 1798.
Subgenus Acanthobodilus Dellacasa, 1983
1. Aphodius (Acanthobodilus) immundus Creu-
tzer, 1799.
The species occurs throughout Europe, in Morocco,
Egypt, the Caucasus, the Transcaucasus, Asia Minor,
Syria, Kazakhstan, Middle and Central Asia. In Rus-
sia, it occurs from western borders up to southern
Yakutia. In the ZIN collection, there are 500 spms.
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
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.

AKHMETOVA, FROLOV
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Russia, the species occurs in southern Khabarovsk
Terr. and Primorskii Territory. It was also reported
from Amur Region (Berlov, 1989). In ZIN collection,
there are 20 spms.
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 spms. from Leningrad, Pskov, Smolensk,
Yaroslavl, Vladimir, Moscow, Saratov, Volgograd,
Rostov provinces, Krasnodar Terr., Astrakhan, Sa-
mara, Orenburg, Novosibirsk, Tomsk provinces, Altai
Terr., and Khakassia.
The species is included in the Red Book of the Rus-
sian Federation (Nikitsky, 2001). The beetles and lar-
vae 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 Primorskii Territory. In the ZIN collection, there
are 60 spms. from Amur Prov., Khabarovsk and Pri-
morsky 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 oc-
curs throughout the country. In the ZIN collection,
there are 300 spms. 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 moun-
tains of Middle Asia; it was imported to North Amer-
ica. In Russia, it is distributed throughout the Euro-
pean Part and occurs in south of West Siberia. In the
ZIN collection, there are 100 spms. from Leningrad,
Yaroslavl, Nizhny Novgorod, Moscow, Kursk, Li-
petsk, Saratov, Rostov provinces, Krasnodar Terr.,
Adygea, Stavropol Terr., Daghestan, 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 Cauca-
sus. In the ZIN collection, there are 10 spms. 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 Tran-
scaucasus, Kazakhstan, and Middle Asia; it was im-
ported to North America. In Russia, it is widespread.
In the ZIN collection, there are 400 spms. from major-
ity of regions of Russia.
The species occurs in cow and horse dung and pre-
fers 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 Re-
gion and Primorskii Territory. In the ZIN collection,
there are 8 spms. from Primorskii Territory.
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.

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Subgenus Agoliinus A. Schmidt, 1913
11. Aphodius (Agoliinus) amurensis Iablokov,
1972.
The species is known from two localities in South-
ern 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 spms. 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 spms. in ZIN collection).
13. Aphodius (Agoliinus) piceus Gyllenhal, 1808.
The species range is not yet well known. In the lit-
erature, there are records of it from Central and South
Europe (Dellacasa, Dellacasa, 2006), Ulyanovsk Prov.
(Isajev, 1995), Tuva, Irkutsk Prov., Buryatia and Za-
baykalskii Terr. (Berlov 1989). In the ZIN collection,
there are 500 spms. from Murmansk, Arkhangelsk, and
Tyumen provinces.
This coprophagous species inhabits forests and oc-
curs 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 de-
scription from the Sochi environs (Olsoufieff, 1918).
Its type depository is unknown. Shokhin (2007) sug-
gested that this name might be a synonym of A. (Agri-
linus) fasciatus (Olivier, 1789).
Subgenus Agolius Mulsant et Rey, 1870
15. Aphodius (Agolius) falcispinis W. Ko-
shantschikov, 1912.
The species is known from East Kazakhstan, North-
Western China, South Siberia, and Mongolia. In the
ZIN collection, there are 10 spms. from Altai and Za-
baykalskii Terr.
16. Aphodius (Agolius) montanus Erichson, 1848.
The species is distributed in the mountains of Cen-
tral and Eastern Europe, and it was recently found in
the South Urals (Zinchenko, 2011). In the ZIN collec-
tion, there are no specimens of this species from Rus-
sia.
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 collec-
tion, 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 spms. 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 Motschul-
sky, 1866.
The species is known from the Kuril islands (Itu-
rup), 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 Mi-
nor, and Turkmenistan. It was reported from Rostov
Prov. of Russia (Shokhin, 2007).
21. Aphodius (Agrilinus) convexus Erichson,
1848.
—A. emerichi Reitter, 1892.

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850
According to Dellacasa and Dellacasa (2006), this
species is widely distributed in the Palaearctic 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 spms. from
Karelia, Leningrad, Yaroslavl, Penza, Kirovsk, Chely-
abinsk, Tyumen, Tomsk provinces, Altai and Kras-
noyarsk Territories, Tuva, Irkutsk Prov., Buryatia,
Yakutia, Amur and Sakhalin provinces, Khabarovsk
Ter.
The species mostly inhabits forest biotopes. It oc-
curs 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 re-
ported from South Sakhalin and the Kunashir islands
(Shabalin and Bezborodov, 2012). In the ZIN collec-
tion, there are no specimens from Russia.
24. Aphodius (Agrilinus) inexpectatus Balthasar,
1935.
The species occurs in North Korea and Japan (Hon-
shu Island) (Stebnicka and Galante, 1991). In Russia,
it is known from Amur Region (Bezborodov and Ber-
lov, 2005) and Primorskii Territory. In the ZIN collec-
tion, there are 8 spms. from South Primorskii Terri-
tory.
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 (Mar-
mota 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 coun-
try 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 spms. from Kare-
lia, Murmansk, and Arkhangelsk provinces, the Re-
public 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 spms. from Lenin-
grad, Moscow, and Ulyanovsk provinces.
A forest dweller feeding on the dung of wild ungu-
lates. The adults are active from April to June.
28. Aphodius (Agrilinus) nikolajevi Berlov, 1989.
The species was described from South Primorskii
Territory and also reported from Zabaykalskii Terr.
and Amur Prov. (Berlov 1989; Bezborodov and Ber-
lov, 2005). In the ZIN collection, there are no speci-
mens of this species.
29. Aphodius (Agrilinus) pratensis Nomura et Na-
kane, 1951.
Reported from Sakhalin (Berlov, 1989) and Primor-
skii Territory (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 collec-
tion, there are 100 spms. from Russia.
The species occurs in open biotopes, pastures,
in cow dung; it is common in the major part of its
range.

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Subgenus Alocoderus A. Schmidt, 1913
32. Aphodius (Alocoderus) digitalis D. Koshan-
tschikow, 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 (Fabri-
cius, 1798).
—A. hydroсhoeris (Fabricius, 1798).
The distribution range of this species includes Cen-
tral and South Europe, North Africa, Armenia, Azer-
baijan, 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 spms. from Voronezh, Rostov,
Volgograd, Astrakhan provinces, Adygea and Da-
ghestan.
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 Transcau-
casus, and Western Kazakhstan. In Russia, it occurs
throughout the country up to Irkutsk Prov. in the east.
In the ZIN collection, there are 200 spms. from Kare-
lia, Leningrad, Novgorod, Pskov, Tver, Kirovsk, Yaro-
slavl provinces, Chuvashia, Moscow, Ryazan, Penza,
Volgograd provinces, Adygea, Karachay-Cherkessia,
Daghestan, Ulyanovsk, Samara provinces, Bashkiria,
Sverdlovsk, Chelyabinsk, Tomsk provinces, the Re-
public 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 spms. 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 (Tara-
sov, 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 spms. from Krasnodar
Terr., Adygea, Karachay-Cherkessia, Kabardino Bal-
kariya, and Daghestan.
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 Cen-
tral, Southern and Eastern Europe, the Caucasus, the
Transcaucasus, and Asia Minor. Tarasov (2008) re-
ported it from Krasnodar Terr., North Osetia and
Daghestan. In the ZIN collection, there are 6 spms.
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 Rus-
sia, it occurs in the European Part and South Siberia
(up to Transbaikal in the east). In the ZIN collection,
there are 70 spms. 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 Medve-
dev, 1968.
This species is endemic to the Caucasus. It was de-
scribed from Abkhazia. In the ZIN collection, there are
2 spms. from North Osetia (Alagir Distr., Yu.E. Ko-
marov leg.).

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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
spms. 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 Primorskii Territory,
Amur Region and North-Eastern China. In the ZIN
collection, there are 11 spms. from Primorskii Terri-
tory.
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 Primorskii Territory. In the
ZIN collection, there are 72 spms. from Zabaykalskii,
Khabarovsk and Primorsky territories.
Subgenus Aphodiellus A. Schmidt, 1913
44. Aphodius (Aphodiellus) impunctatus Water-
house, 1875.
The species occurs in Japan, the Korean Peninsula,
in North-Eastern China (from Beijing to Harbin). In
Russia, it is distributed in South Primorskii Territory.
In the ZIN collection, there are 17 spms.
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 Rus-
sia, it is known from the North Caucasus. In the ZIN
collection, there are 15 spms. from Rostov Prov.,
Krasnodar Terr. and Daghestan.
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, im-
ported 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 spms.
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 Mi-
nor, 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 spms.
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 Octo-
ber.
48. Aphodius (Aphodius) swaneticus Reitter,
1892.
The species is endemic to the Caucasus. In the ZIN
collection, there are 9 spms. 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 Mid-
dle Asia. In the ZIN collection, there are 10 spms.
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 Rus-
sia, it is mainly distributed in North the Caucasus and
the Volga Region. In the ZIN collection, there are
40 spms. from Voronezh, Saratov, Volgograd, Rostov

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provinces, Krasnodar and Stavropol territories, Astra-
khan and Samara provinces, and Daghestan.
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 spms. 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
spms. from Leningrad, Novgorod, Tver, Moscow,
Ryazan, Kursk, Lipetsk, Saratov, Volgograd, Rostov
provinces, Krasnodar Terr., Adygea, Astrakhan, Ki-
rovsk, Samara provinces, Chechnia and Daghestan.
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 for-
est-steppe subzone of the deciduous forest zone from
the western border to the Trans-Urals. In the ZIN col-
lection, there are 100 spms. from Tver, Kursk, Bel-
gorod, Voronezh, Penza, Saratov, Volgograd, Rostov,
Astrakhan provinces, Krasnodar and Stavropol territo-
ries, Daghestan, Samara, Orenburg and Kurgan prov-
inces.
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 spms. from Ryazan, Kursk, Belgorod,
Penza, Volgograd, Rostov, Astrakhan, Samara prov-
inces, Stavropol Terr., Daghestan 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 spms. from Chelyabinsk, Tomsk, and Kemerovo
provinces, Altai and Krasnoyarsk territories, Khakas-
sia, Tuva, Buryatia, Zabaykalskii Terr.
Subgenus Calamosternus Motschulsky, 1859
57. Aphodius (Calamosternus) granarius (Linna-
eus, 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 Re-
gion. In the ZIN collection, there are 200 spms. from
Kaliningrad, Leningrad, Tver, Nizhny Novgorod, Rya-
zan, Belgorod, Voronezh, Volgograd provinces, Kras-
nodar Terr., Kalmykia, Astrakhan Prov., Daghestan,
Kirovsk and Samara provinces, Bashkiria, Tyumen
Prov. and Zabaykalskii 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 Mo-
tschulsky, 1860.
The distribution range of this species includes
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Taiwan Island. In Russia, it is distributed in Primorskii
Territory and Amur Region. In the ZIN collection,
there are 100 spms. 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 spe-
cies.
59. Aphodius (Calamosternus) trucidatus Harold,
1863.
—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 spms. 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 Primorskii Territory. In the ZIN
collection, there are 8 spms. from the Amur Prov. and
Primorskii Territory.
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 Tran-
scaucasus, Turkey, Iran, and Middle Asia (Nikolajev,
1987). Its distribution range needs clarification be-
cause 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 spms. from Tyumen, Novosi-
birsk, Tomsk, and Kemerovo provinces, Altai and
Krasnoyarsk territories, the Republic of Altai, Khakas-
sia, Tuva, Irkutsk Prov., Buryatia, Zabaykalskii Terr.,
the Amur Prov., Yakutia, and Khabarovsk Terr.
64. Aphodius (Chilothorax) conspurcatus (Lin-
naeus, 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
spms. from Leningrad, Novgorod, and Yaroslavl prov-
inces.
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 spms. from the
majority of the regions of the country.
The species occurs in cow and horse dung, in mar-
mot holes, and also is attracted to light. In the territory
of Russia the beetles are active from March to Octo-
ber. It is a common, locally abundant species.
66. Aphodius (Chilothorax) exilimanus Kabakov,
1994.
The species is known from a type series from Uly-
anovsk Prov. (15 spms. 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 collec-

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tion, there are 100 spms. 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 spms. from Penza and Ulyanovsk prov-
inces.
The species occurs on sandy riversides.
70. Aphodius (Chilothorax) jacobsoni W. Ko-
shantschikov, 1911.
The species is distributed in Eastern Kazakhstan
and Mongolia. In Russia, it is known from South Sibe-
ria. In the ZIN collection, there are 18 spms. 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 Sibe-
ria. In the ZIN collection, there are 22 spms. from
Altai.
The species occurs in gray marmot holes (Nikola-
jev, Puntsagdulam, 1984; Zinchenko, 1999).
72. Aphodius (Chilothorax) logunovi Zinchenko,
2003.
The species is known only from the type series from
South-East Altai (3 spms., 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 spms.
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 Octo-
ber. The species is common throughout its range.
74. Aphodius (Chilothorax) mongolaltaicus Niko-
lajev, 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 spms. 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 spms. from Kaliningrad and
Leningrad provinces.
76. Aphodius (Chilothorax) planus D. Koshan-
tschikow, 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 Ciscau-
casus to the south of West Siberia. In the ZIN collec-
tion, there are 16 spms. 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. Ko-
shantschikow, 1894.
The species is distributed in the deserts of the North
Caspian Region. In the ZIN collection, there are 100
spms. from Astrakhan Prov.
A coprophagous species, feeding on horse dung.
Beetles are active in late fall.
78. Aphodius (Chilothorax) subpolitus Motschul-
sky, 1860.
The species is known only from the original de-
scription from Siberia (without a more precise local-
ity) (Motschulsky, 1860). The type depository is un-
known.

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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 Mi-
nor, and Northern Iran. In Russia, it is widely distrib-
uted in the European part up to the South Urals Moun-
tains. In the ZIN collection, there are 100 spms. 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 spms.
from Chuya steppe.
81. Aphodius (Chilothorax) variicolor D. Koshan-
tschikow, 1894.
The species is distributed in the Caspian lowland
deserts. In the ZIN collection, there are 28 spms. 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 spms. from Krasnodar Terr.,
Adygea, Karachay-Cherkessia and North Osetia.
83. Aphodius (Colobopterus) erraticus (Linnaeus,
1758).
The species occurs in Europe (except for the ex-
treme 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 spms. 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 Manner-
heim, 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 Primorskii Territory. In the ZIN
collection, there are 45 spms. from Republic of Altai,
Irkutsk Prov., Buryatia, Zabaykalskii Terr., Amur
Prov. and Primorsky Terr
85. Aphodius (Colobopterus) notabilipennis Pet-
rovitz, 1972.
The species is known from the Amur Region and
Primorskii Territory. In the ZIN collection, there are
100 spms. from Zabaykalskii Terr., the Jewish
Autonomous Province, Khabarovsk and Primorsky
territories.
86. Aphodius (Colobopterus) propraetor Baltha-
sar, 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 spms. 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 spms. 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 collec-
tion, there are 10 specimens from Krasnodar, Stavro-
pol territories and Daghestan.
The beetles mostly occur in open biotopes, in cow
dung.

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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 Primorskii Territory; In
the ZIN collection, there are 200 spms.
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 col-
lection, 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 bee-
tles 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 Penin-
sula and Himalaya). It was reported from Rostov, Vol-
gograd, 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 (Ar-
zanov 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 spms. from Lenin-
grad, Novgorod, Tver, Vologda, Yaroslavl, Nizhny
Novgorod, Moscow, Ryazan, Lipetsk, Volgograd,
Samara provinces, Krasnodar Terr., Daghestan 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 al-
most 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 spms. 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 Ciscauca-
sus. In the ZIN collection, there are 100 spms. 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 al-
most the whole of Europe (except for the extreme
north), the Transcaucasus, Asia Minor, West Kazakh-
stan, and Tajikistan. In the ZIN collection, there are
40 spms. from Leningrad, Nizhny Novgorod, Vo-
ronezh, 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 Mon-
golia. In the ZIN collection, there are 50 spms. from

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Altai and Krasnoyarsk territories, the Republic of Al-
tai, Khakassia, Buryatia, Irkutsk Prov., Zabaykalskii
Terr., Yakutia, Amur Prov., and the Jewish Autono-
mous Province.
98. Aphodius (Eupleurus) subterraneus (Lin-
naeus, 1758).
The distribution range of this species includes al-
most the whole of Europe (except for the extreme
north), North Africa, the Caucasus, the Transcaucasus,
Kazakhstan, Asia Minor and Middle Asia, Afghani-
stan, Mongolia, China, North the Korean Peninsula; it
was imported in North America. In Russia, it is dis-
tributed throughout the country. In the ZIN collection,
there are 300 spms. 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, Kazakh-
stan, Mongolia, and North China. In Russia, it is dis-
tributed in the steppe zone and the forest-steppe sub-
zone of the deciduous forest zone from the Ciscauca-
sus to the Amur Region. In the ZIN collection, there
are 23 spms. from North Osetia, Daghestan, Kurgan
Prov., Altai and Krasnoyarsk territories, Irkutsk and
the Amur provinces.
100. Aphodius (Heptaulacus) sus (Herbst, 1783).
The distribution range of this species includes al-
most the whole of Europe, North Africa, the Caucasus,
the Transcaucasus, Asia Minor, North Iran, West Ka-
zakhstan, and Turkmenistan. In the ZIN collection,
there are 100 spms. 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 (Fab-
ricius, 1775).
The species occurs almost throughout Europe (ex-
cept 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 collec-
tion, there are 30 spms. 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 spms. from Leningrad and Ulyanovsk prov-
inces, 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, ex-
cept for the extreme north, in North Africa, the Cauca-
sus, Asia Minor, Kazakhstan, and Middle Asia. In the
ZIN collection, there are 100 spms. from Belgorod,
Volgograd, Rostov, Astrakhan provinces, Krasnodar
Terr., Kalmykia, and Daghestan.
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 Transcauca-
sus, 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 Af-
ghanistan. 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 spms. from Nizhny Novgorod, Volgograd, Rostov,
and Astrakhan provinces, and Kalmykia.

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This species does not occur in dung. We collected
specimens from sand and in light traps.
106. Aphodius (Liothorax) linearis Reiche et Saul-
cy, 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 spms. from Volgograd, Kurgan provinces, and Tatar-
stan.
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 collec-
tion, there are 10 spms. from Leningrad, Nizhny Nov-
gorod, Moscow, Volgograd provinces and Krasnodar
Terr.
108. Aphodius (Liothorax) plagiatus (Linnaeus,
1767).
The distribution range of this species includes al-
most 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
spms. from Leningrad, Tver, Nizhny Novgorod, Sara-
tov, 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 Kazakh-
stan.
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 Cen-
tral and South Europe, the Caucasus, the Transcauca-
sus, 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 spms. from Saratov, Volgograd, Samara,
Kurgan, Astrakhan provinces, Kalmykia, and Dagh-
estan.
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 spms.
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 Mid-
dle Asia. In the ZIN collection, there are 100 spms.
from Penza, Saratov, Volgograd provinces, Stavropol
Terr., Astrakhan, Ulyanovsk, Samara provinces, Bash-
kiria, 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 Ciscauca-
sus to West Siberia. In the ZIN collection, there are

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22 spms. from Ulyanovsk, Volgograd, Rostov prov-
inces, Stavropol Terr., Kalmykia, and Altai Terr.
Subgenus Melinopterus Mulsant, 1842
116. Aphodius (Melinopterus) consputus (Creu-
tzer, 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 Af-
rica (Morocco, Algeria), the Caucasus, the Transcau-
casus, Asia Minor, Syria, Lebanonе, Israel, Kazakh-
stan, Middle Asia, and Mongolia; it was imported to
USA and Canada. In the ZIN collection, there are 300
spms. from Karelia, Kaliningrad, Leningrad, Nov-
gorod, Bryansk, Arkhangelsk, Vologda, Yaroslavl,
Nizhny Novgorod provinces, Chuvashia, Moscow,
Kaluga, Ryazan, Belgorod, Lipetsk, Saratov, Volgo-
grad provinces, Krasnodar and Stavropol territories,
Adygea, Karachay-Cherkessia, North Osetia, Dagh-
estan, Kirovsk Prov., Udmurtia, Tatarstan, Samara
Prov., Bashkiria, Chelyabinsk, Tyumen, Novosibirsk
provinces, Altai Terr., the Republic of Altai, Kras-
noyarsk Terr., Khakassia, Irkutsk Prov., Zabaykalskii
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 spms. from Leningrad, Nov-
gorod, Arkhangelsk, Yaroslavl, Nizhny Novgorod
provinces, Chuvashia, Moscow, Kaluga, Ryazan, Orel,
Penza, Saratov, Astrakhan provinces, Daghestan, Ki-
rovsk Prov., Tatarstan, Orenburg, Tyumen, Tomsk
provinces, Altai Terr., the Republic of Altai, Kras-
noyarsk Terr., Khakassia, Irkutsk Prov., Buryatia, Za-
baykalskii Terr., and Yakutia.
120. Aphodius (Melinopterus) scuticollis Se-
menov, 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 speci-
mens from Altai Terr. and Khakassia.
121. Aphodius (Melinopterus) sphacelatus (Pan-
zer, 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 spms. from
Leningrad, Novgorod, Pskov, Vologda, Yaroslavl,
Vladimir, Nizhny Novgorod, Moscow, Kaluga, Li-
petsk provinces, and Krasnodar Terr.
Subgenus Mendidaphodius Reitter, 1901
122. Aphodius (Mendidaphodius) brancsiki Reit-
ter, 1899.
The species is widely distributed in deserts of Mid-
dle Asia and West Kazakhstan (Nikolajev, 1987). It
was reported from Astrakhan Prov. (Shokhin, 2007).
Subgenus Mendidius Harold, 1868
123. Aphodius (Mendidius) baigakumi (W. Ko-
shantschikov, 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 Bury-
atia and Zabaykalskii Terr. (Berlov 1989).
125. Aphodius (Mendidius) bispinifrons Reitter,
1889.
The species is known from Armenia, the North
Caspian Region, Middle Asia, and North China (Niko-

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lajev, 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 collec-
tion, there are 14 spms. from Astrakhan Prov. of Rus-
sia (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 col-
lected in horse dung, and a few specimens were col-
lected in a light trap.
127. Aphodius (Mendidius) endroedii Balthasar,
1967.
The species is known only from the original de-
scription from the Transbaikal Region. It is possible
that this name is a junior synonym of A. fimbriolatus.
128. Aphodius (Mendidius) fimbriolatus Manner-
heim, 1849.
The species is known from a few localities in Cen-
tral Asia. In the ZIN collection, there are 3 spms. 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 Re-
gion, the Ciscaucasus. In the ZIN collection, there are
50 spms. 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 (riv-
erside of the Lower Dnepr). In the ZIN collection,
there are 2 spms. 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 alti-
tudinal zones of the Greater and Lesser Caucasus. It is
known from Krasnodar Terr., Karachay-Cherkessia,
North Osetia, and Daghestan (Shokhin, 2007).
Subgenus Nialus Mulsant et Rey, 1870
133. Aphodius (Nialus) varians Duftschmid, 1805.
The distribution range of this species includes Cen-
tral and Southern Europe, North Africa, the Transcau-
casus, 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 spms. from Nizhny Novgorod,
Ryazan, Belgorod, Voronezh, Saratov, Volgograd,
Rostov provinces, Krasnodar Terr., Astrakhan Prov.,
Daghestan, 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 spms. from Krasnodar Terr.
The biology of this species is poorly known. Proba-
bly 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 (Transcarpa-
thian Region), the Transcaucasus, and Asia Minor. In
the ZIN collection, there are 6 spms. from Daghestan.

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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 Depres-
sion. In the ZIN collection, there are 200 spms. 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 Ka-
zakhstan. In the ZIN collection, there are 43 spms.
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 Tran-
scaucasus, and Middle Asian mountains. In the ZIN
collection, there are 5 spms. from Daghestan.
141. Aphodius (Nobius) serotinus (Panzer, 1799).
—A. x-signum Reitter, 1892.
The distribution range of this species includes Cen-
tral, South and Eastern East Europe, the Transcauca-
sus, Asia Minor, Kazakhstan, and Middle Asia. In the
ZIN collection, there are 200 spms. from Leningrad,
Novgorod, Yaroslavl, Nizhny Novgorod, Moscow,
Ryazan, Voronezh, Penza, Volgograd, Rostov prov-
inces, Karachay-Cherkessia, Astrakhan Prov., Tatar-
stan, Ulyanovsk, Samara, Sverdlovsk, Tyumen, Ke-
merovo 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 Reit-
ter, 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 spms. from Vo-
ronezh, Saratov, Volgograd, Rostov, Kurgan prov-
inces, and Stavropol Terr.
A specialized nidicolous species occuring in gopher
and marmot holes.
143. Aphodius (Orodaliscus) spalacophilus No-
vikov, 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 spe-
cies 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 spms. from Astrakhan,
Samara, Orenburg, and Kurgan provinces.
A specialized nidicolous species occuring in mar-
mot holes.
Subgenus Otophorus Mulsant, 1842
145. Aphodius (Otophorus) haemorrhoidalis (Lin-
naeus, 1758).
The distribution range of this species includes the
whole of Europe, North Africa, the Caucasus, the
Transcaucasus, Kazakhstan, Middle Asia, Afghani-
stan, Mongolia, North and South-Western China, the
Korean Peninsula, and Japan; it was imported to North
America. In the ZIN collection, there are 500 spms.
from the majority of the regions of Russia.
The species mostly occurs in cow and horse dung,
in open biotopes.

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Subgenus Parammoecius Seidlitz, 1891
146. Aphodius (Parammoecius) asphaltinus Kole-
nati, 1846.
The species is endemic to the Caucasus. In the ZIN
collection, there are 25 spms. from Krasnodar Terr.,
Adygea, Karachay-Cherkessia, Kabardino-Balkaria,
North Osetia, and Daghestan.
A coprophagous species occuring in cow dung at
1800–3000 m a.s.l.
147. Aphodius (Parammoecius) brevithorax Su-
makov, 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 col-
lection, there are 7 spms. from Astrakhan Prov. and
the Republic of Altai (Chuya Steppe).
149. Aphodius (Phaeaphodius) dauricus Harold,
1863.
The species is known from Asia Minor, Iran, Ka-
zakhstan, Middle Asia, Afghanistan, Mongolia, West
China (Kabakov, 1998). It was reported from Irkutsk
Prov. and Khabarovsk Terr. (Berlov 1989).
150. Aphodius (Phaeaphodius) jouravliowi Reit-
ter, 1907.
The species is known from the type specimens from
Uralsk (North-Western Kazakhstan) and from the Al-
tai 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 Motschul-
sky, 1866.
The species is widely distributed in the Central and
Eastern Palaearctic 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 spms. in Leningrad Prov. In the ZIN collection,
there are 1000 spms. from Leningrad, Astrakhan, Uly-
anovsk, Samara, Chelyabinsk, Novosibirsk provinces,
Altai Terr., the Republic of Altai, Irkutsk Prov., Bury-
atia, Zabaykalskii 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 bio-
topes, pastures, in cow and horse dung.
153. Aphodius (Phaeaphodius) roddi W. Ko-
shantschikov, 1911.
The species is known from a type series from Altai
Mountains. It was reported from North-Eastern Ka-
zakhstan (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 spms. from Buryatia, Zabaykalskii Terr. and the
Amur Prov.
Subgenus Phalacronothus Motschulsky, 1859
155. Aphodius (Phalacronothus) biguttatus Ger-
mar, 1824.
The species is distributed in Central, South and
Eastern Europe, the Transcaucasus, Asia Minor, Ka-
zakhstan, and Turkmenistan. In Russia, it is known
from Ulyanovsk (Isajev, 1995) and Novosibirsk
(Zinchenko, 1999) provinces. In the ZIN collection,
there are 11 spms. from Volgograd, Rostov, Orenburg
provinces and Krasnoyarsk Terr.
156. Aphodius (Phalacronothus) citellorum Se-
menov et Medvedev, 1928.
The species is distributed in the steppe zone of Cen-
tral and Eastern Europe and Kazakhstan.
A nidicolous steppe species occuring in gopher and
marmot holes (Zinchenko et al., 2002; Isajev, 1995).

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157. Aphodius (Phalacronothus) quadrimacula-
tus (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 spms. from Astra-
khan 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 Primorskii Territory, the
Korean Peninsula, and Japan. In the ZIN collection,
there are 50 spms. from Primorskii Territory.
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, Ka-
zakhstan, and Turkmenistan. In Russia, it is known
from the steppe zone from Kursk Prov. to West Sibe-
ria. In the ZIN collection, there are 100 spms. from
Chuvashia, Nizhny Novgorod, Kursk, Saratov, Volgo-
grad, Rostov provinces, Daghestan, Samara Prov., the
Republic of Altai, and Krasnoyarsk Terr.
The species occurs in rodent holes (mostly of mar-
mots and gophers).
160. Aphodius (Plagiogonus) culminarius Reitter,
1900.
The distribution range of this species includes
North-Eastern China and Primorskii Territory. In the
ZIN collection, there are 7 spms. from Primorskii Ter-
ritory.
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 Ka-
zakhstan, and Mongolia. In Russia, it is distributed in
the forest zone, from the western border to Kam-
chatka. In the ZIN collection, there are 30 spms. from
Karelia, Murmansk and Leningrad provinces, Adygea,
Buryatia, Irkutsk Prov., Zabaykalskii Terr., Amur and
Magadan provinces, Khabarovsk and Primorsky terri-
tories.
A coprophagous species occuring in forest biotopes.
162. Aphodius (Planolinus) vittatus Say, 1825.
The species occurs in South and Eastern Europe, the
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 spms. from Astra-
khan and Saratov provinces, the Republic of Altai,
Tuva, Buryatia, and Zabaykalskii Terr.
Distribution records of this species from the Palae-
arctic Region refer to A. vittatus mundus Reitter, 1892,
A. vittatus sellatus Mannerheim, 1852, and A. vittatus
tjanshanicus Balthasar, 1956. The nominative subspe-
cies 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) sug-
gested 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 Zabaykalskii 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 spms. from

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the Altai Mountains, Krasnoyarsk Terr., Irkutsk Prov.,
Zabaykalskii 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 (Zin-
chenko 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 Trans-
baikal Region. In the ZIN collection, there are 100
spms. from Leningrad, Novgorod, Pskov, Tver, Yaro-
slavl, Vladimir, Nizhny Novgorod, Moscow, Rostov
provinces, Tatarstan, Samara, Orenburg, Tyumen,
Omsk, Novosibirsk, Irkutsk provinces, Altai and Kras-
noyarsk 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 Gusa-
kov, 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 spe-
cies.
170. Aphodius (Sinodiapterna) hammondi Della-
casa, 1986.
The species was described from North China. In the
ZIN collection, there are 2 spms. from Primorskii Ter-
ritory.
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 spms. from Kha-
barovsk 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 Pri-
morskii Territory, collected by O.N.Kabakov from soil
under rotten mushrooms.
Subgenus Subrinus Mulsant et Rey, 1870
173. Aphodius (Subrinus) subtilis D. Koshantschi-
kow, 1894.
The species is known only from the original de-
scription 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 Ja-
pan. 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 col-
lection, there are 40 spms. from Volgograd and Rostov
provinces, Stavropol Terr., Astrakhan and Samara
provinces, Daghestan, 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

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Volga region and Daghestan. In the ZIN collection,
there are 10 spms. from Astrakhan Prov.
A psammophillous species. We collected a few lar-
vae 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 Cen-
tral and Eastern China, Korean Peninsula and Japan.
In Russia, the species is known from Amur Region,
Primorskii Territory and Sakhalin Isl. In the ZIN col-
lection, there are 100 spms. 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 al-
most the whole of Europe (except for the extreme
north), North Africa (Algeria), the Caucasus, Tran-
scaucasus, West Asia, Kazakhstan, Middle Asia and
Northern Mongolia. It was imported to North America.
In Russia, it is distributed throughout the country up to
Zabaykalskii Terr. (Nerchinsk) in the east. In the ZIN
collection, there are 300 spms. from Karelia, Lenin-
grad, 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, Daghestan,
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 Zabaykalskii Terr.
The species occurs in open biotopes, pastures, in
cow and horse dung. In Russia, it is a common spe-
cies; 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 spms.
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 spe-
cies 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
spms. 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 Sibe-
ria 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 Puntsag-
dulam, 1984). Berlov and Anishchenko (1998) re-
ported 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 Depres-
sion. In the ZIN collection, there are 80 spms. from
Astrakhan Prov.

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The species occurs in barchan sands, under salt-
worts 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 Amer-
ica. In Russia, it occurs from North-Western border to
Kirovsk and Samara provinces. In ZIN collection,
there are 24 spms. from Leningrad Prov., Chuvashia,
Kaluga, Ryazan provinces, Krasnodar Terr., North
Osetia, Daghestan, 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
Palaearctic ranges, Caucasian endemics and suben-
demics, 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).
With the advance to the north, the aphodiine diver-
sity gradually declines. At the latitude of 65°N, less
than 10 species occur. The north border of the distri-
bution 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 chaf-
ers 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 ex-
plained 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 fac-
tors. 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 vegeta-
tion altitudinal zonality types (National Atlas of Rus-
sia, 2007) was used. According to the map, the terri-
tory of Russia can be devided into 5 lanscape zones
and 6 main altitudinal zonation types. Aphodiines
occur in the taiga, broadleaved forest, steppe and de-
sert zones, and in the boreal (taiga), nemoral (decidu-
ous) and subarid altitudinal zonation types. Aphodi-
ines are apparently absent from tundra zone, and arc-
tic, 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 zona-
tion 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 land-
scapes. Almost the same number of species occurs in
the nemoral altitudinal zonation types. However, as
opposed to the continuous steppe zone, these altitud-
inal 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.

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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 land-
scape zones or altitudinal zonation types.
Typology of the Ranges of the Aphodiines
of the Russian Fauna
Cluster analysis (Jaccard index as similarity coeffi-
cient, 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 Palaearctic Region (Emeljanov,
1974).
The distribution ranges of the aphodiines occurring
in Russia can be grouped into the following types:
Wide Palaearctic (Fig. 2а). This type comprises the
widest ranges. The following species have the ranges
that can be classed as Wide Palaearctic: Aphodius
depressus, A. sus, A. plagiatus, A. erraticus, A. fascia-
tus, A. ater, A. rufipes, A. haemorrhoidalis, A. subter-
raneus, A. pusillus, A. scrofa, A. luridus, A. piceus,
A. borealis, A. villosus, A. rufus, A. brevis, A. tomento-
sus, A. foetens, A. fossor, A. carinatus, A. distinctus,
A. fimetarius, A. granarius, A. immundus, A. melanos-
tictus, 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 Palaearctic Region. Most of them
occur at least in 4 landscape and altitudinal zones.
Wide West-Palaearctic (Fig. 2b). The ranges of the
following species can be classed as Wide West-
Palaearctic: A. arenarius, A. biguttatus, A. coenosus,
A. frater, A. ictericus, A. kraatzi, A. linearis,
A. lividus, A. lugens, A. merdarius, A. niger, A. pay-
kulli, A. punctipennis, A. quadriguttatus, A. satellitus,
A. serotinus, A. sphacelatus, A. sticticus, A. testudi-
narius, A. varians, Oxyomus silvestris. These species
are also eurybiont and rather widely distributed in the
Palaearctic 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. 2c). The ranges of
the two European species, A. conspurcatus and A. ne-
moralis, can be placed in this group. Their ranges sub-
stantially 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. 3a)
comprises the ranges of A. conjugatus, A. constans,
A. maculatus, A. obliteratus, A. obscurus, A. porcus,
A. quadrimaculatus, A. scrutator, A. suarius, A. ther-
micola. In the territory of Russia, most of these species
occur in the steppe zone and the nemoral altitudinal
zones.
The Ancient Mediterranean type (Fig. 3b) com-
prises 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 Rus-
sia, most of these species occur in the steppe and de-
sert zones.
The Westscythian-Northturanian type (Fig. 3c)
comprises the ranges of A. bimaculatus, A. caspius,
A. circumcinctus, A. citellorum, A. costalis, A. gre-
garius, A. gresseri, A. hahni, A. planus, A. rotundan-
gulus, and A. zangi. All these species inhabit steppe
biotopes, and a part of them are specialized nidicols.
The Conblacksean Plane type (Fig. 4a) comprises
the ranges of A. curtulus, A. exilimanus, A. isajevi,
A. ivanovi, A. novikovi, A. sarmaticus, A. spalacophi-
lus. In the territory of Russia, most of these species
occur in the steppe zone and forest-steppe subzone of
the deciduous forest zone.
The Caucasian type (Fig. 4b) comprises the ranges
of A. swaneticus, A. planicollis, A. lederi, A. circas-
sicus, A. latisulcus, A. brignolii, A. leisteri, A. bre-
vithorax, A. asphaltinus, A. alagoezi and A. abcha-
sicus. All these species are endemic to the Caucasus
and are distributed within Euxine Mountain Province
of the European Nemoral Region.
The Sethian type (Fig. 5a) 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 situ-

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ated largely within the Irano-Turanian and Centralasi-
atic subregions of the Sethian Desert Region.
The Caspian type (Fig. 4b) comprises the ranges of
A. dosangi, A. plustschewskii, A. variicolor, and Cne-
misus rufescens. In the territory of Russia, these spe-
cies inhabit the desert zone. Their ranges almost com-
pletely 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. 5b) comprises the
ranges of A. antiquus, A. bidens, A. comma, A. cras-
sus, A. falcispinis, A. fimbriolatus, A. grafi, A. greben-
schikovi, 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 kaz-
nakovi. 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 Palaearctic 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. 5c) comprises
the ranges of A. amurensis, A. binaevulus, A. brachy-
somus, A. breviusculus, A. costatellus, A. comatus,
A. culminarius, A. hammondi, A. koltzei, A. hibernalis,
A. impunctatus, A. inexpectatus, A. languidulus, A. ni-
grotesselatus, A. notabilipennis, A. pratensis, A. pro-
prietor, A. quadratus, A. rugosostriatus, A. sub-
costatus, 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 Primorskii Territory, North-
Fig. 2. Types of the ranges of the Aphodiini: (a) Wide Palaearctic, (b) Wide West-Palaearctic, (c) Wide European Nemoral.

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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. uro-
stigma are essentially different from those of other
species and cannot be classed into any of the above-
mentioned types. Special distribution types can be
proposed for these species:
The Canada-Kamchatian type (Fig. 6a) is character-
istic of A. guttatus. This species is widely distributed
in the Nearctic Region and was recently found in
Kamchatka.
The East-Chinese-Indomalayan type (Fig. 6b) is
characteristic of A. urostigma which is the only repre-
sentative of tropical subgenus Aganocrossus in Russia.
The range of this species mostly lies outside the Palae-
arctic Region. Only the north-easternmost part of its
range includes Southern Primorskii Territory 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).
The European group of provinces comprises Mid-
dleuropean 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
Fig. 3. Types of the ranges of the Aphodiini: (a) Wide Mediterranean Euxinian, (b) Ancient Mediterranean, (c) Westscythian-
N
orthturanian.

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group of provinces. 65 species occur in the Middleu-
ropean mixed province, 57, in the Easteuropean Plain,
80, in the Euxinian mountain, 73 species, in the Con-
blacksean Plain provinces. 29 species (A. paykulli,
A. conspurcatus, A. conjugatus, A. consputus, A. con-
stans, A. maculatus, A. obliteratus, A. porcus, A. scru-
tator, A. suarius, A. thermicola, A. abchasicus, A. ala-
goezi, 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
Fig. 4. Types of the ranges of the Aphodiini: (a) Conblacksean Plane , (b) Caucasian, (c) Caspian.

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Steppe and Sethian Desert regions. 88 aphodiine spe-
cies are known (86 Aphodius, 1 Cnemisus and 1 Oxyo-
mus) from this territory. In the studied part of North-
turanian 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. ki-
zeritskyi, A. menetriesi, A. plustschewskii, A. varii-
color, 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).
Fig. 5. Types of the ranges of the Aphodiini: (a) Sethian , (b) Centralasiatic, (c) Stenopean Nemoral.

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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 dis-
tribution 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 East-
mongolian 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 calcu-
lated 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. 7а). In the simi-
larity dendrogram calculated with the correlation coef-
ficient, they belong to a cluster with the Eastsiberian-
Altaian-Eastmongolian group (Fig. 7b); 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. nasu-
tus, 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 Prov-
ince, 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 Estmongo-
lian 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. jacob-
soni, A. kerzhneri, A. logunovi, A. mongolaltaicus, and
Cnemisus kaznakovi) are not known from other prov-
inces 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 Eurosibe-
rian Boreal Region. This territory is inhabited by
47 aphodiine species which all belong to Aphodius.
Fig. 6. Types of the ranges of the Aphodiini: (a) Canada-Kamchatian, (b) East-Chinese-Indomalayan.

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ENTOMOLOGICAL REVIEW Vol. 94 No. 6 2014
874
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. uro-
stigma, A. breviusculus, A. inexpectatus, A. proprae-
tor, A. binaevulus, A. hammondi, A. tenax, A. hiber-
nalis, A. pratensis, A. quadratus, A. amurensis,
A. costatellus, A. culminarius, A. gorodinskiyi, A. kol-
tzei, and A. suvorovi) are unknown from other
provinces of the studied area.
The Northpacific Mixed Province of the Circumpo-
lar Tundra Region differs most from the other prov-
inces 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 Apho-
dius: A. lapponum, A. borealis, A. aleutus, and A. gut-
tatus. The two former species have wide Palaearctic
ranges. The uniqueness of this province is due to
Fig. 7. Similarity dendrograms of provincial aphodiine faunas: (a) similarity coefficient—Jaccard index, clustering method—average
values; (b) similarity coefficient—correlation, clustering method—average values). Biogeographic provinces: I-4, Northpacific mixed;
II-1, Bothnian mixed; II-2, Zyrianian plane; II-3, Uralian mountain; II-4, Obean plane; II-6, Angaran mixed; II-7, Subokhotian moun-
tain; II-8, Okhotian mountain; II-9, Westmongolian complex; II-10, Eastmongolian complex province of Eurosiberian taiga Region;
III-2, Middleuropean mixed; III-3, Easteuropean plane; III-4, Euxine mountain; IV-1, Eaststenopean mixed; IV-2, Korean mountain;
IV-3, North Japanese mountain; VII-2, Conblacksean plane; VII-3, Kazakhstanian plane; VII-4, Altaian complex; VII-6, Eastmongolian
complex province of Scythian steppe Region; VIII-15, Northturanian.

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875
A. guttatus which is widely distributed in the Nearctic
Region and was recently found in Kamchatka.
ACKNOWLEDGMENTS
We are gfrateful to V.K. Zinchenko (Siberian Zoo-
logical Museum, Institute of Animal Systematics and
Ecology, Siberian Branch of the Russian Academy of
Sciences, 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. Sha-
balin (Soil Biology Institute FEB RAS, Vladivostok),
K.V. Makarov and A.V. Matalin (Moscow Pedagogi-
cal State University, Moscow), A.A. Gusakov (Zoo-
logical Museum, Lomonosov Moscow State Univer-
sity, Moscow), G.V. Nikolaev (Al-Farabi Kazakh Na-
tional University, Almaty), S.I.Tarasov (Natural His-
tory Museum, Oslo University, Oslo), and O.A. Khru-
leva (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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