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A Review of the Scarab Beetle Tribe Aphodiini (Coleoptera, Scarabaeidae) of the Fauna of Russia L

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A Review of the Scarab Beetle Tribe Aphodiini (Coleoptera, Scarabaeidae) of the Fauna of Russia L ISSN 0013-8738, Entomological Review, 2014, Vol. 94, No. 6, pp. 846–879. © Pleiades Publishing, Inc., 2014. Original Russian Text © L.A. Akhmetova, А.V. Frolov, 2014, published in Entomologicheskoe Obozrenie, 2014, Vol. 93, No. 2, pp. 403–447. A Review of the Scarab 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: [email protected] Received January 10, 2014 Abstract—An annotated list of 183 species of the dung-beetle tribe Aphodiini of the Russian fauna is given. The highest species diversity is characteristic of the southern mountainous regions of the country, the richest local fau- nas being known from the Lower Volga area. The greatest numbers of species are known from the steppe and broadleaved forest zones and from the nemoral types of altitudinal zonation of vegetation. The bulk of the fauna is formed by the species with wide ranges; 111 species occur in two or more landscape zones or altitudinal zonation types. Six groups of faunistically similar zoogeographic provinces are distinguished within the territory of Russia. DOI: 10.1134/S0013873814060074 Scarab beetles of the tribe Aphodiini is a large, The regional faunistic lists, new distribution records world-wide distributed group of insects. The world of the aphodiines in Russia and adjacent territories, fauna comprises about 30 genera and over 2000 spe- new species descriptions, and diagnostic keys to some cies, including about 13 genera and 700 species in the groups and for some regions are available in the works Palaearctic Region (Dellacasa, 1988; Dellacasa, Della- of Medvedev (1965, 1976, 1979), Medvedev and Er- casa, 2006). Most aphodiines are coprophages feeding molenko (1969), Nikritin (1969а, 1969б), Medvedev on herbivore dung. In temperate regions including and Nikritin (1974), Berlov (1979, 1985, 1989), Niko- Russia, aphodiines dominate dung-beetle communities lajev (1987, 1998), Isajev (1995), Arzanov et al. both in terms of species richness and abundance. The (1996), Kabakov and Frolov (1996), Nikitsky et al. type genus of the aphodiines, Aphodius Hellwig (sensu (1996), Novikov (1996, 1998), Kabakov (1998), lato), is the largest scarab beetle genus in Russia and Zinchenko (1999, 2003а, 2003б, 2011), Zinchenko one of the largest genera of insects. et al. (2002), Shokhin (2002, 2005, 2007), Gusakov (2004, 2006), Bezborodov and Berlov (2005), The earliest records of the aphodiines of the Rus- Makarov et al. (2009), Bezborodov (2009), Tarasov sian Empire are available in works of Faldermann (2008), Shabalin and Berlov (2009), Zinchenko and (1835a, 1835b, 1838), Gebler (1848), Ménétriés Kyzyl-ool (2010), Shabalin and Bezborodov (2012), (1832, 1849), and Motschulsky (1860). In addition to Zinchenko and Bezborodov (2013), as well as the au- the lists of known species collected in the Caucasus, thors of the present work (Frolov, 1995, 1996, 1997, Transcucasus, and West Siberia, they include descrip- 1999, 2001a, 2001b, 2001c, 2002, 2009; Akhmetova tions of new species. Reitter (1893) summarized the and Frolov, 2008a, b; 2009; Akhmetova, 2006, 2009; data about the Palaearctic scarab beetles. His work Frolov and Akhmetova, 2006, 2013). contains identification keys and numerous descriptions of new species largely from the Caucasus and Middle Despite the rather large number of publications Asia. Aphodiines of Southern Russia and Middle about the aphodiines of Russia, there has been no Asia have been actively studied by Russian coleo- work providing a comprehensive account of the taxo- pterologists in the late XIX and the early XX century nomic composition and geographical distribution of (D. Koshantschikow, 1891, 1894a, 1894b, 1894c, the species of our fauna. Available monographs 1894d; Semenov, 1898a, 1899, 1904, 1905; Semenov (Schmidt, 1922; Balthasar, 1964) are obsolete in terms and Medvedev, 1928; W. Koshantschikov, 1911а, of classification, nomenclature, and distribution data. 1911b, 1912, 1913, 1916). Olsoufieff (1918) summa- Furthermore, they are largely compilations of older rized the available data on the Caucasian copro- literature. Extensive materials from the collection of phagous species. the Zoological Institute of Russian Academy of 846 A REVIEW OF THE SCARAB BEETLE TRIBE 847 Fig. 1. Map of distribution of relative species richness of Russian Aphodiini fauna. Mean number of species per territory unit: 1, 1–10; 2, 11–20; 3, 21–30; 4, 31–40; 5, 41–50; 6, 51–60; 7, 61–70; 8, 71–80. Sciences (ZIN), accumulated during many decades, tion coefficients. Average linkage was used as a clus- remained mostly unstudied. tering method (Fig. 7.) MATERIALS AND METHODS Classification and nomenclature of the Aphodiini follow Medvedev and Nikritin (1970), Nikolaev This work is based on the rich collection of ZIN, in- (1979), and Kabakov and Frolov (1996): Aphodius is cluding vast material of more than 60 species collected treated sensu lato, including Heptaulacus Mulsant, by the authors in Lower Volga Region and the south- Sugrames Reitter and Mothon Semenov et Medvedev ern Far East. In addition, the materials from the fol- as subgenera. lowing institutions and private collections have been studied: Moscow Pedagogical State University, Mos- Genus Aphodius Hellwig, 1798 cow, Zoological museum, Lomonosov Moscow State University, Moscow, A.V. Ivanov’s collection, Ekate- Aphodius Illiger, 1798. rinburg, Soil biology institute FEB RAS, Vladivostok. Subgenus Acanthobodilus Dellacasa, 1983 The numbers of specimens of many studied species are approximate since not all the materials available in the 1. Aphodius (Acanthobodilus) immundus Creu- Zoological Institute, specifically those stored on cotton tzer, 1799. layers and in vials with alcohol, have been processed The species occurs throughout Europe, in Morocco, and integrated into the main collection. Egypt, the Caucasus, the Transcaucasus, Asia Minor, The map of distribution of relative species richness Syria, Kazakhstan, Middle and Central Asia. In Rus- in the territory of Russia (Fig. 1) was generated with sia, it occurs from western borders up to southern ArcGIS software. For special zoogeographic regioni- Yakutia. In the ZIN collection, there are 500 spms. zation, the scheme of the general zoogeographic sub- from the majority of the regions of the country. divisions of the Palaearctic Region (Emeljanov, 1974) was used. As the operational zoogeographic units, The species is common in the major part of its range provinces were used. In the cases when a province and prefers open biotopes. The beetles feed on horse, completely fits the studied area, its complete fauna cow and donkey dung; they are attracted to light and was included in the analysis. In the cases when only occur from April to September. a part of a province was situated in the territory of 2. Aphodius (Acanthobodilus) languidulus Russia, only whose the species occurring within this A. Schmidt, 1922. part were utilized. Provincial aphodiine faunas were compared by constructing similarity dendrograms The distribution range of this species includes based on pairwise calculations of Jaccard and correla- North-Eastern China, Korean Peninsula and Japan. In ENTOMOLOGICAL REVIEW Vol. 94 No. 6 2014 848 AKHMETOVA, FROLOV Russia, the species occurs in southern Khabarovsk tains of Middle Asia; it was imported to North Amer- Terr. and Primorskii Territory. It was also reported ica. In Russia, it is distributed throughout the Euro- from Amur Region (Berlov, 1989). In ZIN collection, pean Part and occurs in south of West Siberia. In the there are 20 spms. ZIN collection, there are 100 spms. from Leningrad, Yaroslavl, Nizhny Novgorod, Moscow, Kursk, Li- The species occurs in open biotopes, pastures, in petsk, Saratov, Rostov provinces, Krasnodar Terr., cow dung. The beetles fly in July and August. Adygea, Stavropol Terr., Daghestan, Kirovsk Prov., Subgenus Acrossus Mulsant, 1842 Tatarstan and Bashkiria. A coprophagous species feeding on cow, horse, 3. Aphodius (Acrossus) bimaculatus (Laxmann, donkey and sheep dung. 1770). The species occurs in Central and Eastern Europe, 7. Aphodius (Acrossus) planicollis Reitter, 1890. North and Eastern Kazakhstan, Kyrgyzstan. In Russia, This species is endemic to the Caucasus. In Russia, it is known from a few localities in the European Part it is known from a few localities in the North Cauca- and West Siberia (up to Krasnoyarsk in the north-east) sus. In the ZIN collection, there are 10 spms. from (Kabakov and Frolov, 1996). In the ZIN collection, Krasnodar and Stavropol territories and Severnaya there are 200 spms. from Leningrad, Pskov, Smolensk, Osetia. Yaroslavl, Vladimir, Moscow, Saratov, Volgograd, Forest dweller, the beetles are active from May to Rostov provinces, Krasnodar Terr., Astrakhan, Sa- July. mara, Orenburg, Novosibirsk, Tomsk provinces, Altai Terr., and Khakassia. 8. Aphodius (Acrossus) rufipes (Linnaeus, 1758). The species is included in the Red Book of the Rus- The species occurs in most of Europe, except for sian Federation (Nikitsky, 2001). The beetles and lar- extreme north, in North Africa (Tunisia), the Tran- vae feed on horse dung (Frolov and Akhmetova, scaucasus, Kazakhstan, and Middle Asia; it was im- 2006). ported to North America. In Russia, it is widespread. In the ZIN collection, there are 400 spms. from major- 4. Aphodius (Acrossus) binaevulus Heyden, ity of regions of Russia. 1887. The species occurs in cow and horse dung and pre- The species is known from
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