Akhmetova, L.A., Frolov, A.V. 2008. A Review of Scarabs of the Subgenus Nobius Mulsant et Rey, Genus Aphodius Illiger (Coleoptera, Scarabaeidae), of the Fauna of Russia and Neighboring Countries. Entomological Review, 88(4): 421-434.

Original Russian text: Ахметова, Л. А., Фролов, А. В. 2008. Обзор пластинчатоусых жуков подрода Nobius Mulsant et Rey рода Aphodius Illiger (Coleoptera, Scarabaeidae) фауны России и сопредельных стран. Энтомологическое обозрение, 87, 397-410.

A review of scarabs of the subgenus Nobius Mulsant et Rey, genus Aphodius Illiger (Coleoptera, Scarabaeidae), of the fauna of Russia and neighboring countries

L.A. Akhmetova & A.V. Frolov

Laboratory of Insect Systematics, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, St.-Petersburg, 199034 Russia. Email: codocera@rambler.ru

Abstract

Scarab beetles of the subgenus Nobius Mulsant et Rey, genus Aphodius Illiger, distributed in Russia and neighboring countries are reviewed. A key to the species and notes on their biology are given. The tree new synonymies are established: A. serotinus (Panz.),1799 = A. x-signum Reitter, 1892, syn. n.; A. gresseri Sem. 1898 = A. korgaldzhensis Nikolajev, 1987, syn. n.; A. chaldaeus Petr., 1971 = A. moreleti Baraud, 1980, syn. n. Lectotypes of A. inclusus Reitt., 1892 and A. gresseri are designated.

Subgenus Nobius Mulsant et Rey, 1869, of genus Aphodius Illiger, 1798, was described on the basis of the presence of a single character, the dark brown elytra with small pale macula near the apex of each elytron. However, the coloration of elytra in Aphodius species is subject to much intraspecific and interspecific variability. A number of species unrelated to A. serotinus Panzer, 1799, (the type species of the subgenus Nobius)were previously placed in this subgenus because of using the elytral coloration without consideration of other characters. Among the species occurring in Russia and neighbouring countries, it concerns A. (Chilothorax) flammulatus Harold, 1876, and A. (Carinaulus) costatellus A. Schmidt, 1916 (authors data). At the same time some species with similar shape of the aedeagus and other structures were placed in the different subgenera: A. pustulifer Reitter, 1892, and A. chaldaeus Petrovitz, 1971, in Volinus Mulsant, 1870; A. moreleti Baraud, 1980, in Esymus Mulsant et Rey, 1869.

Incomplete descriptions of most species, absence of illustrations of characters, and rarity of the majority of these species in collections did not allow to identify them reliably. The goal of the present study is to clarify the diagnosis of the subgenus Nobius, diagnostic characters of the species included in this subgenus, nomenclature of the names, and distribution of the species in Russia and neighbouring countries. During examination of the collection of Zoological Institute RAS (ZIN) a new species was found. Its description is given below.

 This study is based on the large material deposited in ZIN including the specimens collected by the authors over the past years. Also, the material from the Hungarian Museum of Natural History in Budapest (HNHM) and Department of Zoology of Moscow Pedagogical State University (MGPU) was examined. Coordinates of the collecting localities for the distribution map were taken from the specimen labels, available atlases, and the GNS database (http://earth-info.nga.mil/gns/html/index.html). Scheme of measurements of the aedeagus is shown on the fig. 25. Examined material is deposited in ZIN, if not indicated otherwise.

 The authors would like to thank K.V. Makarov and A.V. Matalin (MGPU) for the loan of material and Otto Merkl (HNHM) for the opportunity to study the type specimens of E.Reitter's species. We are also indebted to V.P. Bulychev (Dosang Antiplague Station) for logistics assistance. This research was supported by the RFBR grant 07-04-00482-a.

Genus Aphodius Illiger

Subgenus Nobius Mulsant and Rey, 1869

Type species A. serotinus Panzer, 1799.

Body length 3.0-5.5 mm. Elytra shiny, glabrous, colored from yellowish-straw to dark brown.  Most species have an obscure macula occupying the most part of each elytron, or only a few small maculae and dark sutural intervals. The shape of the maculae and intensity of the elytral coloration is subject to some interspecific variability. Scutellum small, triangular. Lateral sides and base of pronotum with fine border. Head slightly convex or flat. Dorsal surface of head without setae. Frontoclypeal suture indistinct. Clypeus sinuate at middle, rounded at sides. Genae slightly protruding past eyes. Adjacent apical setae of middle and hind tibiae of different length. Parameres without feebly sclerotized processes, more or less acute apically.

The most reliable diagnostic characters for the members of the subgenus are the shape and relative size of the parameres and the coloration of elytra. Reliable identification of females, especially of single specimens, can be difficult in same cases. We have studied spermathecae of about 30 specimens of all Nobius species included in this review. Most species have curved and evenly thickened spermatheca with the largest part of its surface folded (fig. 33). Spermatheca of  A. serotinus differs in being pear-shaped basally, less curved, and having less folded surface (fig. 34). Spermatheca of  A. chaldaeus Petrovitz, 1971, is similar to that of  A. serotinus, but more curved (fig. 35). However, diagnostic reliability of the shape of spermateca cannot be appreciated without morphometric analysis which is beyond the scope of this work.

 

Key to the Nobius species occurring in Russia and adjacent countries

1. Elytra yellowish-straw, with brown sutural intervals and 1 - 2 small brown maculae on each elytron (fig. 1 - 4, see inset). Metasternum of male pubescent ... 2

- Elytra entirely brown (fig. 9 - 12) or pale with large dark obscure macula occupying most part of each elytron (fig. 5 - 8, 15, 16, see inset). Metasternum of male pubescent or not ... 3

 2. Clypeus granulate. Each elytron with 2 brown maculae (one longitudinal macula located approximately in the middle of elytron closer to the lateral margin, and another near the apex) (fig. 1, 2). Diameter of eye in ventral view approximately equal to the minimum interval between eye and gula. Middle and hind femora in male with relatively sparse, thin setae ventrally (fig. 17). Basal sclerite of aedeagus approximately 1.8 times the length of parameres (fig. 25) ... A. pustulifer Reitt.

- Clypeus punctate. Elytra with small brown macula located on intervals 2 - 4 near apex of each elytron (fig. 3, 4). Diameter of eye in ventral view approximately 2 times larger than minimum interval between eye and gula. Middle and hind femora in male with only single thin setae ventrally (fig. 18, see inset). Basal sclerite of aedeagus approximately 2.8 times longer than parameres (fig. 26). Spermatheca smooth on most part of surface, folded  apically, and slightly widened basally (fig. 35) ... A. chaldaeus Petr.

3. Metasternum of male glabrous. Middle and hind femora with single thin setae ventrally (fig. 19, see inset). Spermatheca smooth on convex side, folded on concave side and apically, and pear-shaped basally (fig. 34). Parameres approximately as long as basal sclerite of aedeagus (fig. 27). Elytra reddish-orange, with darken apices and large dark obscure macula occupying almost entire surface of each elytron (fig. 5, 6) ... A. serotinus (Panz.)

- Metasternum of male pubescent. Middle and hind femora of male with dense setae ventrally (fig. 20, 21) or with only single setae (fig. 22 – 24, see inset). Spermatheca folded, elongate, not pear-shaped basally (fig. 33). Parameres shorter than basal sclerite ... 4

4.  Middle and hind femora of male with dense setae ... 5

-  Middle and hind femora of male with only single thin setae ... 6

5. Elytra orange-yellow, with brown or dark brown sutural intervals, apices, and large longitudinal macula on each elytron. Sometimes apices of elytra pale with dark macula or elytra entirely dark brown with paler humeral umbones and part of interval 2 near the base (fig. 7, 8). Parameres more curved, acute apically. Basal sclerite of aedeagus approximately 1.5 times longer than parameres (fig. 28) ... A. inclusus Reitt.

- Elytra yellowish-brown, with dark brown suture and brown sutural intervals (fig. 9, 10). Parameres less curved, smoothly tapering and acute apically. Basal sclerite of aedeagus approximately 2 times longer than parameres (fig. 29) ... A. bonnairei Reitt.

6. Elytra brown or dark brown, almost uniformly colored (fig. 11, 12). Basal sclerite of aedeagus approximately 1.8 times longer than parameres (fig. 30) ... A. gresseri Sem.

- Elytra with more or less contrast pattern (fig. 13 – 16). Basal sclerite of aedeagus more than 3 times longer than parameres (fig. 31, 32) ... 7

7. Elytra with very contrast pattern, yellowish-straw with dark brown sutural intervals, apices, and large longitudinal macula not occupying interval 2 and sometimes interval 3 (fig. 13, 14). Apices of parameres less curved, straight angled to basal sclerite (fig. 31) ... A. circumductus Solsky.

- Elytra with less contrast pattern, with dark brown suture, brown apices and large brown obscure macula occupying almost entire surface of elytron except for small part of  intervals 2 and 3 near the base, humeral umbones, and area near apices of each elytron (fig. 15, 16). Apices of parameres more curved, not straight angled to basal sclerite (fig. 32) ... А. dosangi sp. n.

Figs. 1–8. Aphodius Ill., general view: (1, 2) A. pustulifer Reitt., (3, 4) A. chaldaeus Petr., (5, 6) A. serotinus (Panz.), (7, 8) A. inclusus Reitt. 
Figs. 1–8. Aphodius Ill., general view: (1, 2) A. pustulifer Reitt., (3, 4) A. chaldaeus Petr., (5, 6) A. serotinus (Panz.), (7, 8) A. inclusus Reitt. [(1, 3, 5, 7) male, (2, 4, 6, 8) female].

 

Aphodius (Nobius) inclusus Reitter, 1892 (fig. 7, 8, 20, 28, 36).

- inclusum Reitter, 1892 : 97, 1906 : 441. - inclusus Reitter (incorrect subsequent spelling): Schmidt, 1910 : 33; Yablokov-Khnzoryan, 1967 : 110; Nikolajev, 1987 : 118. - glasunowi  D. Koshantschikow, 1894 : 108-109 (synonymy by: Reitter, 1906);  Nikolajev, 1987 : 117.

Differential diagnosis. The species is most similar to A. bonnairei Reitter, 1892, but can be separated from it by the elytral pattern and the shape and size of parameres. Males of this species differ from A. serotinus Panzer, 1799 in having setae on the perimeter of metasternal disc and on the ventral surface of middle and hind femora, and also by the shape and length of the parameres.

Male (fig. 7). Head with black disc and lighter anterior margin. Frontoclypeal suture indistinct. Frons not tuberculate. Clypeus sinuate at middle, rounded at sides. Genae rounded, almost not separated from lateral margins of clypeus, slightly protruding past eyes. Diameter of eye in ventral view approximately as wide as minimum interval between eye and gula.

Pronotum shiny, dark brown becoming nearly black on disc, with paler lateral margins. Its surface with irregular and sparse punctation (punctures separated by 1 - 3 puncture diameters on disc). Lateral sides and base of pronotum with fine border; hind angles rounded. Scutellum triangular, dark brown, becoming nearly black laterally.

Elytra without humeral teeth, glabrous, shiny on disc and shagreened apically, colored orange-yellow with brown or dark brown sutural intervals, apices, and large longitudinal macula on each elytron. Sometimes apices of elytra pale with dark macula or elytra entirely dark brown with paler humeral umbones and part of interval 2 near the base. Striae relatively deep, punctures of striae wider in diameter than width of stria. Elytral intervals flat, very finely and sparsely punctate.

Venter of body dark brown. Perimeter of metasternal disc with dense, long setae. Legs brown. Hind femora with relatively dense setae ventrally. Middle femora with sparser setae. Outer teeth of fore tibia relatively long. Spur of fore tibia slightly curved downward and acute apically. Lower spur of middle tibia shorter than the first segment of tarsi and as long as 1/2 length of upper spur. First segment of hind tarsi slightly longer than upper spur and as long as 3 following segments together. Adjacent apical setae of middle and hind tibiae of different length.

Parameres relatively strongly curved, acute apically, 1.5 times shorter than length of basal sclerite.

Female (fig. 8) differs from male in glabrous disc of metasternum and in having only a few setae on middle and hind femora. Spermatheca folded, elongate, not pear-shaped basally.

Body length 3.7 - 4 mm.

Distribution range: Russia (Dagestan), Georgia, Armenia, Tadzhikistan, Azerbaijan, Afghanistan.

Type material. A. inclusus: lectotype (designated here): female with labels "Caucasus. Arexesthal. Leder. Reitter.", "inclusum m.", "coll. Reitter", "Paratypus, 1891, Aphodius (Melinopterus) inclusus Reitter" (HNHM); paralectotype female with labels "Caucasus. Arexesthal. Leder. Reitter.", "coll. Reitter" and "Holotypus, 1891, Aphodius (Melinopterus) inclusus Reitter" (HNHM). A. glasunowi, syntypes: female with labels "Turkestan Lac. Kulikulan Glasunov 1892." and "Glasunowi Kosh. type!"; 4 males and 4 females with labels "Turkestan, Lac. Kulikulan, Glasunov, 1892." and "Aphodius glasunowi"; 3 males and 4 females with labels "Seravschan Kshtut. Artutsh, Glasunov, 1892" and "Aphodius glasunowi".

Additional material. Russia. Dagestan, Derbent, 5 spm. Georgia. Tbilisi, 29.XII.1927, 2 spm. (Ya. Kirshenblatt). Armenia. Erevan, 10-25.XI.12, 3 spm. Iran. Lorestan, 1 spm.; Teheran, 8.V.1915, 1 spm.; Teheran, 4.VI.1915, 1 spm.; Khurasan, 5-9.XI.1900, 1 spm. (N.А. Zarudny). Afghanistan. Herat, NE of Adraskan , 1300 m, 20.XI.1971, 1 spm. (О.N. Kabakov); Gnor Tulak, 2400 m, 4.X.1969, 1 spm. (О.N. Kabakov).

Remark. It can be inferred from the original description of A. inclusus that the author examined several specimens of this species, but their exact number is not mentioned. The label "Holotypus [...]" was added to one of the specimens by the HNHM staff later and does not indicate the holotype. To insure stability of the nomenclature we designate the specimen with the Reitter's handwritten label the lectotype.

Schmidt (1910) was probably the first author who used incorrect subsequent spelling "inclusus" instead of original "inclusum". This spelling has been commonly used since then and attributed to Reitter (1892).

 

Aphodius (Nobius) serotinus Panzer, 1799 (fig. 5, 6, 19, 27, 34, 36).

Panzer, 1799; Nikolajev, 1987 : 117; Dzhambazishvili, 1979 : 97 – 98; Medvedev, 1965 : 177. -  x-signum Reitter, 1892 : 97, syn. n.

Differential diagnosis. The species is most similar to A. inclusus, but can be separated from it by glabrous metasternal disc in both sexes, presence of only a few setae on the ventral surface of middle and hind femora, and by the shape and length of parameres.

Male (fig. 5). Head black on disc, sometimes with paler margins. Frontoclypeal suture indistinct. Frons not tuberculate. Clypeus slightly sinuate at middle, rounded at sides. The shape and size of genae vary from rounded, almost not separated from lateral margins of clypeus and slightly protruding past eyes to obtuse-angled, distinctly protruding past eyes. Diameter of eye in ventral view approximately as wide as minimum interval between eye and gula.

Pronotum shiny, dark on disc, sometimes with reddish-brown lateral margins or fore angles. Its surface with irregular and dense punctation. Sides and base of pronotum with fine border; hind angles rounded. Scutellum triangular, dark brown, sometimes nearly black.

Elytra without humeral teeth, glabrous, shiny. Elytra colored orange-red, with darken apices and large dark obscure macula occupying almost entire surface of each elytron. Size and shape of the macula vary, but normally it doesn't reach humeral umbones and the base of elytron except for interval 5. Striae relatively deep, wide, punctures of striae wider in diameter than width of striae. Elytral intervals flat, very finely and sparsely punctate.

 Venter of body dark brown. Metasternal disc paler, reddish-brown or orange-red, shiny, glabrous or, rarely, with a few thin setae.

Legs reddish-brown to orange-red, orange in paler specimens. Middle and hind femora glabrous or with a few thin setae ventrally. Outer teeth of fore tibia relatively long and thin. Spur of fore tibia slightly curved downward and acute apically. Lower spur of middle tibia as long as or slightly shorter then first segment of tarsi, acute apically, and as long as 1/2 length of upper spur. First segment of hind tarsi longer than upper spur and as long as or a bit shorter than 3 following segments together. Adjacent apical setae of middle and hind tibiae of different length.

Parameres feebly curved, taperring and slightly acute apically, approximately as long as basal sclerite of the aedeagus.

Female (fig. 6) differs from male in slightly narrower pronotum with denser punctation. Spermatheca smooth on the convex side, folded on the concave side and apically, pear-shaped basally.

Body length 3.5 — 5.5 mm.

Distribution range: the species is widespread in Europa and Asia to Siberia. Records from North-Eastern China need for confirmation. 

Type material. A. x-signum: holotype (?), male with labels "Quellgebiet des Irkut Leder.", "coll. Reitter", "x-signum" (HNHM).

Additional material. Hungary. 1 spm. without  exact data. Moldova. Orhei distr., Furceni, 29.IV.1978, 1 spm. (Kabakov). Ukraine. Zakarpatskaya Prov.,  15.X.1971, 1 spm. (Stepanov); Krivoj Rog, 15.IX.1987, 1 spm. (Kiselev); Crimea, Ishun', 9.X.1990, 1 spm. (Gusarov); Mount pass Baidarskie vorota 15.IX.1903, 16 spm.; Lastochkino, 10.IV.1907, 1 spm.; Kerch, 5.VI.1902, 4 spm., X.1906, 1 spm.; Sevastopol, 1.X.1911, 1 spm. (V. Pliginskiy); Simferopol Distr., Balka Dzhalman, 19-29.IX.1906, 4 spm., 21.IX.1910, 2 spm. (Kakhovskiy); Poltava Prov., 1908, 1 spm.; Vilkovo,  Odessa Prov., 2.X.1990, 1 spm. (V. Gusarov); Kharkov Prov., Zmiev, 11.IX.1926, 1 spm. (Arnoldi); Merefa, 11.IX.1956, 1 spm. (Stepanov). Russia. Leningrad Prov.: Sankt-Petersburg, 1 spm. (E. Koenig); Luga, 1 spm. (Yakobson). Novgorod Prov.: Valday, 5 spm.  Nizhni Novgorod Prov.: Sarov, 15.IX.1872, 1 spm. Voronezh Prov.: Bobrov, 21.VIII.1912, 1 spm. (B. Karavaev). Ryazan Prov.: Dubrovka, 14 spm., 7.VIII.1903, 1 spm. (Olsufiev); Gremyachka, 14.VIII.1903, 2 spm., 20.VIII.1899, 1 spm. (A. Semenov); Saraevskiy Distr., Kozlovka, 20.VIII.1897, 2 spm. (Olsufiev); Grotovskiy, 12.VIII.1898, 1 spm. (A. Semenov); Kemerovo Prov.: 27.VII.1911, 1 spm. (Gortchakovskiy). Yaroslavl Prov.: 10.VIII.1895, 4 spm., 31.VIII.1899, 5 spm., 5-11.IX.1899, 7 spm., 29.VIII.1896, 2 spm.; Yaroslavskiy Distr., Berditsyno village, 17.VIII.1895, 2 spm., 8.IX.1898, 5 spm., 1 spm. (A.I. Yakovlev); Danilovkiy Distr., 5 spm. (Shestakov). Penza Prov.: environs of Penza, 25.VIII.1918, 1 spm. (V. Strogashevitch). Rostov Prov.: Nokolaj -Polje, 26.XI.1906, 3 spm. (Fisher). Samara Prov.: Samara, 1 spm.; Volzhskiy Distr., village Kurumotch, 12.IX.2004, 6 spm. (L. Akhmetova). Ulyanovsk Prov.: Maynskiy Distr., 2.IX.1990, 2 spm. (Isajev). Volgograd Prov.: Sarepta (Volgograd), 1 spm.; village Kotluban, 13.IX.1991, 2 spm. (K. Grebennikov). Astrakhan Prov.: village Dosang, flood-plain of Akhtuba River, 6.X.2006, 3 spm., 7.X.2006, 19 spm., 17.IV.2007, 8 spm., 17.V.2007, 9 spm., 18.V.2007, 8 spm. (A. Frolov, L. Akhmetova). Republic of Tatarstan: Kazan, 12.IV.1915, 2 spm., 22.VIII.1917, 2 spm. (I. Ivanov). Karachaevo-Cherkessia: plato Bechasyn, 14.VIII.1911, 1 spm.; Teberda, 19.VIII.1897, 1 spm. (N. Schukin). Sverdlovskaya Province: environs of Ekaterinburg, 15.VI.1893, 1 spm. Republic of Altai: Katanda, 26.VIII.1902, 1 spm.; Kyrsay, 11.IX.1901, 2 spm. Irkutsk Prov.: Irkutsk, 9 spm. (V.E. Yakovlev), 21.VIII.1976, 5 spm. (Berlov). Tyumen Prov.: Tobolsk, 30.IX.1926, 1 spm. (Samko). Khakassia: environs of  Minusinsk, Beya, VIII.1916, 19 spm. (Kozhantchikov). Krasnoyarsk Terr.: Krasnoyarsk, 1 spm. (V.E. Yakovlev); Achinskiy Distr., Novaya Elovka, 1912, 5 spm. (Uspenskaya). Georgia. Tbilisi, 3-5 X 1910, 2 spm. (K. Satunin), 30 IX 1916, 1 spm. (Olsufiev), 2 spm. (E. Kenig); Mtskheta, 24.X.1965, 2 spm. (Kryzhanovskiy). Abkhazia, Chkhalty river, 25.VIII.1897, 1 spm. (N. Schukin). Armenia. Erevan, 10-25.XI.1912, 3 spm. Azerbaidzhan. Baku, 1.IV.1930, 1 spm. Kazakhstan. environs of Uralsk, 22.VIII.1906, 17 spm. Turkmenia. Repetek, 1 spm. (A. Krikheldorff). Tadzhikistan. Petr Velikij Range, 16.VI.1969, 1 spm. (G. S. Medvedev).

Remark. A. x-signum was described from environs of Irkutsk. According to the original description, specimens of this species differ from A. serotinus in longer spurs of  hind tibiae, denser punctation of elytra and in the body slightly widened posteriorly. Examination of the type specimen showed that the first two characters are within the limits of variability of A. serotinus. Body relatively widened posteriorly is typical for females of most Aphodius species. We suppose that the diagnoses of A. x-signum and A. serotinus (Reitter, 1892) were based on  females and males, respectively, of same species treated here as A. serotinus.

Figs. 9–16. Aphodius Ill.: (9, 10) A. bonnairei Reitt., (11, 12) A. gresseri Sem., (13, 14) A. circumductus Solsky, (15, 16) A. dosangi sp. n.
Figs. 9–16. Aphodius Ill.: (9, 10) A. bonnairei Reitt., (11, 12) A. gresseri Sem., (13, 14) A. circumductus Solsky, (15, 16) A. dosangisp. n. [(15) holotype]. (9, 11, 13, 15) male, (10, 12, 14, 16) female.

 

Aphodius (Nobius) circumductus Solsky, 1876 (fig. 13, 14, 23, 31, 36).

Solsky, 1876 : 338; Nikolajev, 1987 : 117.

Differential diagnosis. The species is most similar to A. dosangi sp. n., but differs from it in contrast elytral pattern, less curved apices of parameres, and in angle between parameres and basal sclerite.

Male (fig. 13). Head with dark brown disc and paler margins. Frontoclypeal suture indistinct. Frons not tuberculate. Clypeus slightly sinuate at middle, with rounded fore angles. Genae rounded, almost not separated from lateral margins of clypeus, slightly protruding past eyes. Diameter of eye in ventral view approximately 1.5 times as wide as minimum interval between eye and gula.

Pronotum shiny, dark brown to almost black on disc, with paler, brown lateral margins. Its surface with relatively dense punctation (punctures approximately equal in diameter, separated by 1-2 puncture diameters). Lateral sides and base of pronotum with fine border; hind angles rounded. Scutellum wide, triangular, dark brown, sometimes nearly black.

 Elytra glabrous, shiny on disc and laterally, shagreened apically. Elytra colored yellowish-straw, with dark brown sutural intervals, apices, and large dark brown longitudinal macula, which does not occupy interval 2 and at least the part of interval 3 of each elytron. Striae shallow, punctures of striae larger than width of striae. Elytral interstices flat, very finely and sparsely punctate on  disc of elytron, coarser and denser punctate apically.

Legs and venter of body brown. Perimeter of metasternal disc with dense, long setae.

Hind and middle femora with single thin setae ventrally. Outer teeth of fore tibia relatively long. Spur of fore tibiae slightly curved downward and acute apically. Lower spur of middle tibia as long as or slightly shorter than first segment of tarsi and as long as 1/2 length of upper spur. First segment of hind tarsi longer than upper spur and slightly shorter than 3 following segments together. Adjacent apical setae of middle and hind tibiae of different length.

Parameres very short, tapering and acute apically. In lateral view, parameres 3.3 times shorter than length of basal sclerite.

Female (fig. 14) differs from male in denser punctation of head and pronotum, narrower pronotum and glabrous disc of metasternum. Spermatheca folded, elongate, not pear-shaped basally.

Body length 3.1-4.5 mm.

Distribution range: Kazakhstan, Uzbekistan, Turkmenia.

Examined material. Kazakhstan. Moinkum Desert, III.1907, 2 spm. (E. Fischer); 15.V.1910, 2 spm. (Golbek); Taraz, 1 spm.; Aidar-Kuduk, 12.V.1912, 10 spm. (Zarudny); Baygakum, III.1912, 29 spm., 6.IV.1913, 1 spm. (V. Kozhantchikov), 15-18.V.1908, 24 spm. (Glazunov). Turkmenia. Krasnovodsk, 30.IV-12.V.1899, 2 spm. (K.O. Anger); Kara-Bogaz, 17.V.1953, 1 spm. (Odintsova), 30.IV.1953, 1 spm. (Shteinberg); Pereval Station, 26.IV.1889, 7 spm. (A. Semenov). Uzbekistan. Samarkand, 1 spm.; Ul'yanovo, IV.1908, 18 spm. (Golbek); Kyzylkum desert, Baimakhan, 16.V.1995, 4 spm. (Ovchinnikov); Gulistan station, 15-20.V.1903, 2 spm. (G. Yakobson); Kyzylkum desert, 1 spm. (F. Moravits); border between Kyzylkum Desert and Golodnaya Step near Syr-Darya, 10.V.1903, 1 spm. (G. Yakobson).

Figs. 17–24. Aphodius Ill., underside of male body: (17) A. pustulifer Reitt., (18) A. chaldaeus Petr., (19) A. serotinus (Panz.), (20) A. inclusus Reitt., (21) A. bonnairei Reitt., (22) A. gresseri Sem., (23) A. circumductus Solsky, (24) A. dosangi sp. n.
Figs. 17–24. Aphodius Ill., underside of male body: (17) A. pustulifer Reitt., (18) A. chaldaeus Petr., (19) A. serotinus (Panz.), (20) A. inclusus Reitt., (21) A. bonnairei Reitt., (22) A. gresseri Sem., (23) A. circumductus Solsky, (24) A. dosangi sp. n.

 

Aphodius (Nobius) gresseri Semenow, 1898 (fig. 11, 12, 22, 30, 36).

Semenow, 1898 : 611 - 613; Yablokov-Khnzoryan, 1967 : 110; Medvedev, 1965 : 177. - korgaldzhensis Nikolajev, 1987 : 118, syn. n.

Differential diagnosis. The species is most similar to A. circumductus and A. serotinus but differs from them in almost uniformly brown or dark brown coloration of elytra. Males can be easily separated by the shape and length of parameres; from A. serotinus  they also differ in pubescent perimeter of metasternal disc.

Male (fig. 11). Head dark brown. Frontoclypeal suture indistinct. Clypeus slightly sinuate at middle, with rounded fore angles, finely punctate (punctures separated by approximately 2 puncture diameters, punctation becoming denser laterally). The shape and size of genae vary from rounded, almost not separated from lateral margins of clypeus and slightly protruding past eyes to obtusely angled, separated from lateral margins of clypeus and distinctly protruding past eyes. Diameter of eye in ventral view approximately as wide as or slightly larger than minimum interval between eye and gula.

Pronotum shiny, dark brown to nearly black on disc, with paler sides or fore angles. Its surface with irregular, relatively dense punctation (adjascent punctures clearly deffer in size). Lateral sides and base of pronotum with fine border; hind angles rounded. Scutellum wide, triangular, brown.

Elytra without humeral teeth, glabrous, shiny, brown or dark brown, almost uniformly colored (some specimens have slightly paler intervals 1 and 2 near the base and humeral umbones, or the the entire sutural interval slightly paler). Striae relatively deep, punctures of striae wider in diameter than width of striae. Elytral intervals flat, very finely and sparsely punctate.

Legs and venter of body brown or dark brown. Perimeter of metasternal disc with dense, long setae. Hind and middle femora with a few thin setae ventrally. Outer teeth of fore tibia relatively long. Spur of fore tibia slightly curved downward and acute apically. Lower spur of middle tibia as long as the first segment of tarsi and longer than 1/2 upper spur. First segment of hind tarsi slightly longer than upper spur and slightly shorter than 3 following segments together. Adjacent apical setae of middle and hind tibiae of different length.

Parameres relatively curved, acute apically, 1.8 times shorter than basal sclerite.

Female (fig. 12) differs from male in more convex, rugose clypeus, narrower pronotum with denser punctation, and glabrous metasternal disc. Spermatheca folded, elongate, not pear-shaped basally.  

Body length 4 - 5 mm.

Distribution range: Russia (from the Center of European Region to the Lower Volga territory), Western Kazakhstan.

Type material. A. gresseri: lectotype (designated here), male with labels "Olgino, Vladim. gub. 1.X.1898, T.S. Chicherin" and "Aphod. Gresseri male. Typ. m. XII.98. A. Semenow det.". A. korgaldzhensis: holotype, male with labels "Kurgaldzhino, Sultan-Keldy 18.04.1982 Tarantul" and "Holotypus Aphodius korgaldzhensis Nikolajev 1985"; paratypes: 2 males and 3 females with the same data, that holotype; male and 5 females with the same data but the date (15.IV.1982).

Additional material. Russia. Voronezh Prov.: Korotoyak, 23.IX.1911, 5 spm. (B. Karavaev). Volgograd Prov.: environs of Elton village, Sorochya Balka, 8.IV.2007, 1 spm. (K. Makarov, A. Matalin) (MGPU); Northern environs of Elton village, 10.IV.2007, 2 spm. (K. Makarov, A. Matalin) (MGPU); environs of Elton Lake, right bank of Khara River, 2.IV.2007, 12 spm. (K. Makarov, A. Matalin) (MGPU); environs of Elton village, Utinaya Balka, 9.IV.2007, 3 spm. (K. Makarov, A. Matalin) (MGPU). Astrakhan Prov.: environs of Dosang village, 10.IV.2007, 2 spm., 17.IV.2007, 12 spm., 16.IV.2007, 2 spm., 21.IV.2007, 2 spm. (A. Frolov, L. Akhmetova); 1.5-5 km NE  Dosang village, 14.IV.2007, 1 spm. (K. Makarov, A. Matalin) (MGPU). Kazakhstan: environs of Uralsk, 4.IV.1906, 1 spm. (B. Uvarov).

Remark. A. korgaldzhensis was described from Kurgaldzhino Nature Reserve (Central Kazakhstan). Comparison of the type specimens of this species with the types of A. gresseri has shown that they are similar in all characters except for darker, almost homogeneously colored elytra in A. korgaldzhensis. We found, however, specimens with somewhat intermediate coloration (paler elytra with obscure dark macula) in the series from environs of Elton Lake and Dosang. Therefore we think that all these color aberrations are within the range of variability of one species.

 Figs. 25–35. Aphodius Ill.: (25–32) aedeagus, dorsal and lateral view; (33–35) spermatheca [(25) A. pustulifer Reitt., (26, 35) A. chaldaeus Petr., (27, 34) A. serotinus (Panz.), (28) A. inclusus Reitt., (29) A. bonnairei Reitt., (30) A. gresseri Sem., (31) A. circumductus Solsky, (32, 33) A. dosangi sp. n
Figs. 25–35. Aphodius Ill.: (25–32) aedeagus, dorsal and lateral view; (33–35) spermatheca [(25) A. pustulifer Reitt., (26, 35) A. chaldaeus Petr., (27, 34) A. serotinus (Panz.), (28) A. inclusus Reitt., (29) A. bonnairei Reitt., (30) A. gresseri Sem., (31) A. circumductusSolsky, (32, 33) A. dosangi sp. n.]. A, scale to Figs. 25–32; B, to Figs. 33–35.

 

Aphodius (Nobius) pustulifer Reitter, 1892 (fig. 1, 2, 17, 25, 36).

Reitter, 1892 : 90; Nikolajev, 1987 : 114.

Differential diagnosis. The species is most similar to A. chaldaeus, but differ from it in the granulated clypeus, number of elytral maculae and smaller diameter of eyes.

Male (fig. 1). Head dark brown, with paler margins, granulated on the clypeus and coarsely punctate on the disc. Frons not tuberculate. Frontoclypeal suture indistinct. Clypeus sinuate at middle, rounded at sides. Genae rounded, almost not separated from lateral margins of clypeus, slightly protruding past eyes. Diameter of eye in ventral view approximately as wide as minimum interval between eye and gula.

 Pronotum shiny, dark brown on  disc, with paler, brown lateral margins. Its surface with relatively dense punctation (approximately equal punctures separated by 1-2 puncture diameters). Sides and base of pronotum with fine border; hind angles rounded. Scutellum brown, wide, triangular, punctate.

Elytra without humeral teeth, glabrous, shiny on disc and laterally, slightly shagreened apically, colored yellowish-straw with brown sutural intervals and 2 brown maculae on each elytron (one longitudinal macula located approximately in the middle of elytron closer to the lateral margin, and another – near the apex). Size of maculae vary, one of them may be absent. Striae shallow, punctures of striae larger than width of striae. Elytral intervals flat, finely and sparsely punctate  except for apices where punctation is coarser and denser.

Venter of body brown. Perimeter of metasternal disc with dense, long setae.

Legs yellowish-brown. Hind and middle femora with relatively sparse, thin setae ventrally. Outer teeth of fore tibia relatively long. Spur of fore tibia slightly curved downward and acute apically. Lower spur of middle tibia as long as the first segment of tarsi and longer than 1/2 length of upper spur. First segment of hind tarsi longer than upper spur and slightly shorter than 3 following segments together. Adjacent apical setae of middle and hind tibiae of different length.

Parameres curved, taperring and acute apically, 1.8 times shorter than length of basal sclerite.

Female (fig. 2) differs from male in narrower pronotum, glabrous disc of metasternum and having only a few setae on the middle and hind femora. Spermatheca of specimens studied by the authors folded, elongate, not pear-shaped basally.

Body length 3 - 4.5 mm.

Distribution range: The species is distributed in the arid regions of Middle Asia. Record from south-eastern part of European Russia (Sarepta Station, Rеitter, 1992 : 90), is probably based on  mislabeled material and was not confirmed by subsequent findings.

Examined material. Kazakhstan. Balamurun, 6.IV.1913, 2 spm. (V. Kozhantchikov). Turkmenia. Krasnovodsk, 1899, 1 spm. (K.O. Anger); Krasnovodsk, 30.IV-12.V.1899, 4 spm. (K.O. Anger); Annau, 26.IV.1902, 5 spm. (K.O. Anger); 20 km south than Kerki village, 23.IV.1984, 6 spm. (T.N. Vereschagina); Ashkhabad, 3.V.1889, 15 spm. (A. Semenow); Repetek, 18.XI.1938, 17 spm. (Vinogradov); Yolatan, 5-14.V.1926, 3 spm. (V. Kizeritskiy); Imambaba, 6-14.IV.1912, 1 spm. (Kozhantchikov); Kopet-Dag, 1892, 1 spm. Uzbekistan. Samarkand, 1892, 8 spm. (Glazunov), 9-11.V.1904, 7 spm. (G. Suvorov), 1 spm.; Nurata, 1892, 1 spm. (Glazunov); Ul'yanovo, IV.1908, 86 spm. (Golbek); Saray, 1892, 5 spm. (Glazunov); Baba-Tau mauntains [Babatag], 18.V.1897, 1 spm. (Kaznakov); Kyzylkum desert, Baimakhan, 16.V.1995, 1 spm. (Ovchinnikov); "Kisil-Kum mer. Jgam-Berdy", 1892, 13 spm. (Glazunov);  Tadzhikistan. Shartuzskiy district, Chilichor-Chamma, desert, 21.IV.1959, 3 spm. (I. Lopatin); Artuch, 1892, 1 spm. (Glazunov); Pendzhikent, 1892, 3 spm. (Glazunov). 

Remark. A. pustulifer was described in the subgenus Volinus Mulsant, but the shape of the aedeagus and other characters suggest close relationship with other species treated here.

Fig. 36. Aphodius Ill., distribution of species of the subgenus Nobius in the territory of Russia and adjacent countries.
Fig. 36. Aphodius Ill., distribution of species of the subgenus Nobius in the territory of Russia and adjacent countries.

 

Aphodius (Nobius) chaldaeus Petrovitz, 1971 (fig. 3, 4, 18, 26, 35, 36).

Petrovitz, 1971 : 217. -  moreleti Baraud, 1980 : 140, syn. n.

Differential diagnosis. The species is most similar to A. pustulifer, but differs from it in the not granulated clypeus, presence of only one macula on each elytron, and larger diameter of eyes.

Male (fig. 3). Head brown, lateral sides paler than disc. Frons not tuberculate. Frontoclypeal suture indistinct. Clypeus slightly sinuate at middle, with rounded fore angles, finely punctate (punctures on disc separated approximately by 2 - 4 puncture diameters, on sides punctation denser). Genae rounded, almost not separated from lateral margins of clypeus, slightly protruding past eyes. Diameter of eye in ventral view approximately 2 times larger than minimum interval between eye and gula.

Pronotum shiny, brown. Its surface with irregular punctation. Lateral sides and base of pronotum with fine border; hind angles rounded. Scutellum brown, triangular.

Elytra without humeral teeth, glabrous, shiny on disc and shagreened apically, colored yellowish-straw with brown sutural intervals and small brown macula located on 2 - 4 intervals near the apex of each elytron. Striae shallow, thin, punctures of striae larger than width of striae. Elytral interstices flat, punctate.

Legs and venter of body brown. Perimeter of metasternal disc with dense setae.

Hind and middle femora with single, thin setae ventrally. Outer teeth of fore tibia relatively long. Spur of fore tibia slightly curved downward and acute apically. Lower spur of middle tibia acute, as long as the first segment of tarsi and longer than 1/2 length of upper spur. First segment of hind tarsi longer than upper spur and as long as 3 following segments together. Adjacent apical setae of middle and hind tibiae of different length.

Parameres short, evenly taperring and acute apically. Basal sclerite 2.8 times longer than length of parameres.

Female (fig. 4) differs from male in denser punctation of head and pronotum, narrower pronotum and glabrous metasternal disc. Spermatheca smooth on most part of surface and folded on the apex, slightly widened basally (fig. 35).

Body length 4 - 5 mm.

Distribution range: Turkmenia, Iran, Iraq.

Additional material. Turkmenia. Kara-Bogaz, XII 1894, 3 spm. (Maksimovich); Kizyl-Arvat, 30.IV.1896, 1 spm. (Anger); Kopetdag, 10-11.X.1902, 1 spm. (K.O. Anger); environs of Uyar well (30 km W of  Yaskhan), 7.V.1952, 1 spm. (Kryzhanovskiy); Serakhs, 14.V.1903, 8 spm. (K.O. Anger); Pereval station, 17.IV.1903, 1 spm. (K.O. Anger), 26.IV.1889, 8 spm. (A. Semenow); Tersakan, 21.X.1902, 2 spm. (K.O. Anger); Chat, 1-3.XI.1903, 1 spm. (K.O. Anger). Iran. Kerman, 23-25.II.1928, 1 spm. (V. Kuznetsov); Atrek, 13.IV.1916, 1 spm. (Il'in); "Persia", 2 spm.

Remark. Comparison of the type specimens of A. moreleti and A. chaldaeus has shown that they belong to the same species.

 

Aphodius (Nobius) bonnairei Reitter, 1892 (fig. 9, 10, 21, 29, 36).

Reitter, 1892 : 96.

Differential diagnosis. The species is most similar to A. inclusus, but differs from it in the elytral pattern and the shape and size of parameres.

Male (fig. 9). Head dark brown, sometimes almost black. Frons not tuberculate. Frontoclypeal suture indistinct. Clypeus sinuate at middle, with rounded fore angles, irregularly punctate. Genae rounded, almost not separated from lateral margins of clypeus, slightly protruding past eyes. Diameter of eye in ventral view approximately as wide as minimum interval between eye and gula.

 Pronotum shiny, dark brown becoming nearly black on the disc, with paler lateral margins. Lateral sides and base of pronotum with fine border; hind angles rounded. Scutellum triangular, dark brown.

 Elytra without humeral teeth, glabrous, shiny, colored yellowish-brown, with dark brown suture and brown sutural intervals. Striae relatively deep, punctures of striae wider in diameter than width of striae. Elytral intervals flat, finely and sparsely punctate.

Legs and venter of body brown. Perimeter of metasternal disc and middle and hind femora with dense, long setae ventrally.

 Outer teeth of fore tibia relatively long. Spur of fore tibia slightly curved downward and acute apically. Lower spur of middle tibia as long as the first segment of tarsi and as long as or slightly longer than 1/2 length of upper spur. First segment of hind tarsi longer than upper spur and as long as 3 following segments together. Adjacent apical setae of middle and hind tibiae of different length.

Parameres less curved, evenly tapering and acute apically. Basal sclerite of aedeagus approximately 2 times longer than the length of parameres.

Female (fig. 10) differs from male in coarser punctation of head, more convex, rugose clypeus, narrower pronotum with denser punctation, glabrous metasternal disc, and in having only sparse setae on the middle and hind femora. Spermatheca folded, elongate, not pear-shaped basally.  

Body length 3.1 - 4 mm.

Distribution range. Range of this species is poorly known. It was found in Southern Europe (Italy, Spain) and Northern Africa (Algeria, Morocco).

Additional material. Afghanistan: Gereshk, 6.XII.1969, 1 spm. (O.N. Kabakov). "Gallia", 2 spm.

 

Aphodius (Nobius) dosangi Akhmetova et Frolov sp. n. (fig. 15, 16, 24, 32, 33, 36).

Male, holotype (fig. 15). Body length 3.9 mm. Head shiny, dark brown, almost black on disc and paler on sides. Its surface densely punctate (punctures separated approximately by a puncture diameter). Clypeus slightly sinuate at middle, with rounded fore angles. Genae rounded, almost not separated from lateral margins of clypeus, slightly protruding past eyes. Frontoclypeal suture indistinct. Frons not tuberculate. Diameter of eye in ventral view slightly larger than minimum interval between eye and gula.

Pronotum shiny, dark brown, almost black on disc, with paler, brown lateral margins. Fore margin not bordered, lateral margins and base of pronotum with fine border. Disc of pronotum relatively densely, irregularly punctate (punctures separated by 1-3 times a puncture diameter); hind angles rounded. Scutellum wide, triangular, shiny, dark brown, sparsely punctate.

Elytra shiny, yellowish-straw, with dark brown suture, brown apices and large brown obscure macula, which occupy almost entire surface of elytron, except for small part of the intervals 2 and 3 near the base, humeral umbones and area near the apices of each elytron. Humeral teeth absent. Striae relatively deep, punctures of striae larger than width of striae. Elytral intervals flat, very finely and sparsely punctate.

Venter of body dark brown. Perimeter of metasternal disc with dense setae.

Legs brown. Hind and middle femora with a few thin setae ventrally.

Outer teeth of fore tibia relatively long. Spur of fore tibia slightly curved downward and acute apically. Lower spur of middle tibia as long as the first segment of tarsi and slightly longer than 1/2 length of upper spur. First segment of hind tarsi longer than upper spur and slightly shorter than 3 following segments together. Adjacent apical setae of middle and hind tibiae of different length.

Parameres very short, tapering and acute apically. In lateral view, basal sclerite approximately 3 times longer than length of parameres.  

Female (fig. 16) differs from male in coarser punctation of head, rugose clypeus, narrower pronotum and glabrous metasternal disc. Spermatheca folded, elongate, not pear-shaped basally.  

Variability. Body coloration,  punctation of pronotum and head, size of genae slightly vary in the examined specimens. Body length of paratypes 3.1 - 4.2 mm. 

Differential diagnosis. The new species is most similar to A. circumductus, but differ from it in the less contrast elytral pattern, more curved apices of parameres and the angle between the parameres and basal sclerite.

Etymology. The species name is derived from Dosang Village (Russia, Astrakhan Province), where holotype and larger part of the paratypes were collected.

Type material. Holotype, male: Russia, Astrakhan Prov.: Dosang village, at night near railway  station, attracted to light, 24.V.2007 (A. Frolov, L. Akhmetova). Paratypes (27 males and 32 females): the same data as the holotype, 5 females, flood-plain of Akhtuba River, cattle dung, 6.X.2006, 2 females, sands, horse dung, 11.X.2006, male and female (A. Frolov, L. Akhmetova); environs of Dosang, fixed sands, horse dung, 17.IV.2007, 12 males and 6 females reared from larvae, old horse dung, 13-15.IV.2007, female, horse dung, 17.IV.2007, 3 females (A. Frolov, L. Akhmetova). Kazakhstan. Chelkar, 28.IV.1984, male and female (G. Nikolajev); Malye Barsuki, 2.VII.1907, male (Andreev); Uralsk Prov., Temirskiy Distr., 1.VI.1908, male and 2 females (Uvarov), IV-V.1908 2 males,  Astau-Saldy River , 5.VI.08, 4 males and 7 females, Daumchar cemetery, 1.IV.1908, 2 males and female, Kuzha-Tugai, IV.V.1908, 3 males and 3 females (Borodin and Uvarov).  

Specimen from Georgia (Marneuli environs, 15.V.1960, Kabakov) probably belong to A. dosangi sp. n., but it is damaged and therefore not included in the type series.                   

 

References

Dzhambazishvili, Ya.S., Scarab Beetles of Georgia, Tbilisi: Metsniereba, 1979 [in Russian].

Medvedev, S.I., "Scarabaeidae" in Keys to Insects of European Part of the USSR (Nauka, Moscow, 1965), Vol. 2, pp. 166–208 [in Russian].

Nikolajev, G.V., Scarab Beetles (Coleoptera, Scarabaeoidea) of Kazakhstan and Middle Asia, Alma-Ata: Nauka, 1987 [in Russian].

Yablokov-Khnzoryan, S.M., Coleoptera. Scarab Beetles. (Izd-vo AN ArmSSR, Erevan, 1967). Vol. 6, 225 pp. (Fauna Armyanskoy SSR) [in Russian].

Baraud, J., "Deux nouveaux Coleopteres Scarabaeoidea d'Irak", Bull. Mens. Soc. Linn., 49 : 139 — 142 (1980).

Koshantschikow, D., "Verzeichniss der von Herrn D. Glasunow in Turkestan gesammelten coprophagen Lamellicornen", Horae Soc. Ent. Ross., 28 : 103 — 119 (1894).

Petrovitz, R., "Scarabaeidae from the Near East", Israel J. Ent., 6 : 215-238 (1971).

Reitter, E., "Bestimmungs-Tabelle der Lucaniden un coprophanen Lamellicornen des palaearctischen Faungebietes", Verh. Naturf. Ver., 30 : 140 — 262 (1892).

Reitter, E., "Uebersicht der Coleopteren-Arten der Gattung Aphodius Illig. aus dem nachsten Verwandtschaftskreise des A. prodromus Brahm, des Sbg. Melinopterus s. str.", Deut. Ent. Zeitschr., 435-442 (1906).

Schmidt, A., Aphodiinae in Coleopterum Catalogus auspiciis et auxilio W. Junk (editus a S. Schenkling), Berlin, 1910, vol. 20, pp. 1-111.

Semenow, A., "Coleoptera nova Rossiae Europaeae Caucasique", Horae Soc. Ent. Ross., 32, 604 — 615 (1898).

Solsky, M., Kauffman herausgegeben von der Kaiserlichen Gesandtschaft. Die Reise in Turkestan auf Veranlassung des Genneral-Gouverneurs K.P., St. Petersburg, 2: 290 – 398 (1876).