Species of the subgenus Bodilus (genus Aphodius) from Russia and adjacent countries (Coleoptera: Scarabaeidae)
A.V. Frolov. Laboratory of Insect Systematics, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab., 1, 199034 St.Petersburg, Russia. E-mail: email@example.com
Species of the subgenus Bodilus Mulsant et Rey (genus Aphodius), of Russia and adjacent countries are reviewed and keyed. Two new species are described: A. zarudnyi sp. n. (Iran) and A. inylchekensis sp. n. (Kirgizia). Three new synonymies are established: Bodilus Mulsant et Rey, 1869 = Paramelinopterus Rakovič, 1984, syn. n., A. lugens Creutzer, 1799 = A. apiceopacus Petrovitz, 1959, syn. n., A. punctipennis Erichson, 1848 = A. vartiani Petrovitz, 1965, syn. n.
During examination of the material of Aphodius species from the collection of the Zoological Institute (St. Petersburg), I found two species of the subgenus Bodilus Mulsant et Rey that proved to be new. The goal of the present paper is to describe the new species and clarify the structure and diagnosis of the subgenus Bodilus. Since the Aphodius fauna of Russia and adjacent countries is still poorly studied and there is no comprehensive revision of the subgenus Bodilus, a key and notes on distribution of the Bodilus species inhabiting this area are given.
The material examined, including type specimens, is deposited at the Zoological Institute, unless indicated otherwise. Author's remarks are in square brackets.
Subgenus Bodilus Mulsant et Rey, 1869
Type species: A. lugens Creutzer, 1799 (type designation: Reitter, 1892).
Length 4.0—8.5 mm. Head and pronotum shiny, brown to dark brown on disc with fore angles or sides lighter; in some species pronotum yellowish brown with dark brown macula in center, or pronotum entirely yellowish brown. Clypeus feebly sinuate at middle, rounded at sides; head more or less densely punctate, not rugose or granulate. Frontal suture with tubercles or not. Sides and base of pronotum bordered, hind angles obtusely rounded. Scutellum triangular, shiny, punctate in basal part. Elytra yellowish brown, shiny or shagreened, more or less pubescent; striae fine, punctate (in ♂ of A. zarudnyi sp. n. striae wider and mat, impunctate); elytral intervals flat or slightly convex. Fore tibiae of ♂ with spur located opposite medial to basal outer tooth, in some species spur short and apical outer tooth of tibia with brush of dense setae on ventral side. Apical spurs of middle tibia slender and acute; lower spur longer than 2/3 of upper. Adjoining apical setae of hind tibiae of equal length, except for some outermost, or unequal. Ventral surface of body brown, legs slightly lighter. Disc of metasternum flat, more or less pubescent in both sexes. Width of eye in ventral view approximately 2 times wider than minimal interval between eye and gula. Aedeagus with long basal sclerite, therefore it is comparatively larger than in most other Aphodius subgenera (length of aedeagus up to 35 per cent length of body in A. gregarius Harold). Apices of parameres more or less rounded in lateral view, rarely acute.
Sexual dimorphism, except for shape of fore tibia and its spur, as follows: male with wider pronotum, sparser punctures on pronotum and head, denser pubescence of elytra and disc of metasternum, more pronounced tubercles on frontal suture. In ♂ of A. zarudnyi sp. n. surface of genae and pronotum, except for disc, pubescent with distinct setae.
Key to the Bodilus species from Russia and adjacent countries
1. Hind tarsi longer than hind tibiae. Head not tuberculate or with traces of tubercles. Pronotum light brown with dark macula in middle; macula occupies less than half the area of pronotum, its margin more or less distinct (Fig. 3) (except for A. inylchekensis sp. n., where some specimens have pronotum dark brown with yellowish brown sides (Fig. 1)) ..... 2
― Hind tarsi shorter or equal in length to hind tibiae. Head tuberculate in ♂, with distinct traces of tubercles in ♀. Pronotum brown to dark brown with lighter fore angles or sides; if pronotum with dark macula in middle, then macula occupies more than half the area of pronotum and its margins indistinct (Fig. 4) ..... 7
2. Apical tooth of fore tibia of ♂ with brush of dense setae on ventral side (Fig. 5, 6). Elytra entirely pubescent with dense setae in ♂ and sparser in ♀ (except for A. insperatus, which has elytral disc pubescent with sparse setae in ♂ and smooth in ♀) ..... 3
― Apical tooth of fore tibia of ♂ without brush of dense setae on ventral side. Elytral disc smooth ..... 5
3. Elytra pubescent with dense setae in ♂ and with sparser but distinct setae in ♀ ..... 4
― Elytral disc pubescent with sparse setae in ♂ and smooth in ♀. Aedeagus ― Fig. 10 ..... A. insperatus Petrovitz
4. Apices of parameres curved upwards (Fig. 11) ..... A. rudii Endroedi
― Apices of parameres not curved upwards (Fig. 12) ..... A. gregarius Harold
5. Fore tibial spur of ♂ attached opposite basal tooth of tibia, apical tooth of fore tibia acute (Fig. 8, 9) ..... 6
― Fore tibial spur of ♂ attached opposite middle tooth of tibia, apical tooth of fore tibia widened (Fig. 7). Aedeagus ― Fig. 13 ..... A. hastatus Reitter
6. Inner margin of fore tibia of ♂ more or less convex (Fig. 8). Head and disc of pronotum dark brown. Aedeagus ― Fig. 14 ..... A. inylchekensis Frolov, sp. n.
—Inner margin of fore tibia of ♂ straight or slightly concave (Fig. 9). Head and disc of pronotum light brown. Aedeagus ― Fig. 15 ..... A. sordescens Harold
7. Elytral intervals densely punctate and entirely pubescent with dense, long setae in ♂ and sparser, shorter setae in ♀ ..... 8
― Elytral intervals sparsely punctate; elytral disc smooth ..... 9
8. Elytral striae fine and shiny, distinctly punctate (diameter of punctures bigger than width of striae). Surface of pronotum smooth or with minute setae on sides. Aedeagus ― Fig. 16 ..... A. punctipennis Erichson
― Elytral striae of ♂ wider and mat, almost impunctate. Sides and base of pronotum of ♂ pubescent with distinct setae (setae may be abraded in some specimens). Aedeagus ― Fig. 17 ..... A. zarudnyi Frolov, sp. n.
9. Smaller (4.0―5.5 mm). Apices of parameres broadly rounded in lateral view (Fig. 18) ..... A. ictericus (Laicharting)
― Longer (6.0―8.0 mm). Parameres of different shape ..... 10
10. Frontal suture with 3 tubercles in ♂. Apices of parameres narrow in lateral view (Fig. 19). Central Asia (up to Altai mts. on west) ..... A. longeciliatus Reitter
― Frontal suture with central tubercle in ♂. Apices of parameres wide in lateral view (Fig. 20). Europe, Fore and Middle Asia (up to south-western Kazakhstan on east) ..... A. lugens Creutzer
Aphodius (Bodilus) longeciliatus Reitter, 1887
Central Asian species. In the collection of the Zoological Institute there are specimens from almost all of the territory of Mongolia and from China (Burhan Budai Shan Mts. in Qinghai, Alashan Desert in Nei Mongol autonomous region). This species is most similar to A. lugens and can be separated from it by the paramere shape, stronger frontal tubercles, and by the smaller average size, more convex body, and reddish color.
Unfortunately, I could not study the type of A. crassus A. Schmidt, described from northern Mongolia, but all the characters mentioned in the original description (Schmidt, 1916) completely fit the limits of variability of A. longeciliatus.
Aphodius (Bodilus) lugens Creutzer, 1799
(Figs 4, 20)
= A. apiceopacus Petrovitz, 1959, syn. n.
This species is widely distributed in the western Palearctic region, extending to eastern Kazakhstan in the northeast and the Tien Shan mts. in the southeast. It can be easily separated from other species of the subgenus Bodilus by the shape of the parameres and relatively large size.
Aphodius (Bodilus) ictericus (Laicharting, 1781)
This species is widely distributed in the western Palearctic region, and its natural habitat includes all of Europe, Northern Africa, Asia Minor, Iran, Transcaucasia, northern Kazakhstan, and South Siberia. It can be separated from other Bodilus species by its smaller size and by the shape of the parameres.
Aphodius (Bodilus) punctipennis Erichson, 1848
= A. vartiani Petrovitz, 1965, syn. n.
This species is distributed in southeastern Europe, Middle Asia, Iran, Afghanistan, southern Kazakhstan up to the Dzhungar Mts. in the northeast. In the collection of the Zoological Institute there are specimens from Russia (Stavropol Terr., Rostov, Volgograd, Astrakhan' provinces, Dagestan), Ukraine (Crimea, Poltava Prov.), Azerbaijan (Zuvant), Turkmenistan (Kara-Kala, Karlyuk, Ashkhabad), Uzbekistan (Kashkadar'ya and Surkhandar'ya provinces), Kazakhstan (Karaganda, Ural'sk, Aktyubinsk, Dzhambul, Akmola, Kyzyl-Orda, and Alma-Ata provinces). The species is most similar to A. zarudnyi Frolov, sp. n., and can be separated from it by the shape of aedeagus, smooth surface of pronotum (only minute setae can be found on fore angles) and shiny elytral striae in males, and by the smaller average size and darker color.
Aphodius (Bodilus) zarudnyi Frolov,sp. n.
(Figs 2, 17)
Type material. Holotype ♂ labeled "Kerman: str. Sargad, 1.V.1901. N. Zarudnyi [in Cyrillic letters]". Paratypes: 6 ♂♂ and 11 ♀♀ with same data; ♂ with same data, but collected 5―6.V.1901; ♂ labeled "Persia, Vost. Khorasan, s. Charakhs, 25.IV.1898, N.A. Zarudnyi [in Cyrillic letters]"; 3 ♂♂ and 2 ♀♀ labeled " Persia, Vost. Khorasan, s. Avaz, 28.IV.1898, N.A. Zarudnyi [in Cyrillic letters]".
Holotype. Male. Head light brown, shiny, frons slightly darker than clypeus. Clypeus sparsely punctate (punctures separated by 3―4 times a puncture diameter, clypeus feebly sinuate at middle, rounded at sides. Frontal suture with central tubercle. Genae rectangular, protruding past eyes. Surface of head pubescent with minute, indistinct setae, except for genae, which are pubescent with distinct, relatively long setae. Pronotum brown on disc with sides and base lighter, shiny, densely, regularly punctate (punctures separated by 1―2 times a puncture diameter on disc, becoming denser on sides). Surface of pronotum, except for disc, pubescent with distinct setae. Anterior margin of pronotum not bordered, sides and base finely bordered; hind angles obtuse. Scutellum triangular, light brown, finely punctate, pubescent in basal part. Elytra yellowish-brown, without humeral teeth. Intervals flat, moderately shiny, densely punctate, uniformly pubescent with dense, long setae. Striae relatively wide, more mat than intervals, almost impunctate. Apical spur of fore tibia relatively short and wide, curved downward, attached opposite middle tibial tooth. Apical spurs of middle tibia slender and acute; lower spur longer than 2/3 of upper. First segment of hind tarsi shorter than upper spur of tibia and nearly as long as 3 following segments together. Adjoining apical setae of hind tibiae of equal length, except for some outermost. Venter of body yellowish-brown. Disc of metasternum flat, almost entirely pubescent with dense long setae. Apices of parameres dilate and flat, slightly curved downward (Fig. 14). Body length 8.5 mm.
Female can be separated from male by the narrower pronotum, denser punctures on pronotum and head, smooth surface of pronotum and head, shorter and sparser setae on elytra, thinner shiny striae with distinct punctures, and smaller frontal tubercles.
Paratypes. Body length 7.0 to 8.0 mm. Pubescence of surface of pronotum and genae sparser in some specimens: only sides of pronotum possess distinct setae.
Differential diagnosis. The species is most similar to the widely distributed A. punctipennis and can be reliably separated from it only by the characters, pronounced in males: more pubescent body (well visible, relatively long setae on the perimeter of pronotum and on the genae; longer setae on elytra), wider elytral striae (they are mat, indistinctly punctate). The new species differs from A. punctipennis by lager average size and lighter color.
Etymology. The species is named in honor of N.A. Zarudny, who led (on behalf of Russian Geographical Society) a few expeditions to Iran at the end of the nineteenth and beginning of the twentieth centuries, and who gathered vast entomological collections, including the type series of the new species.
Distribution. Known from south-eastern (Serkhed upland region) and north-eastern (Zirkukh Mts.) Iran. This species is probably distributed throughout eastern Iran and replaces A. punctipennis there.
Aphodius (Bodilus) sordescens Harold, 1869
(Figs 9, 15)
The species is widely distributed in Central Asia, extending to north-eastern Kazakhstan in the northwest and the Kunlun mountains in the southwest. It is known from all the territory of Mongolia and from adjacent regions of South Siberia. In the collection of Zoological Institute there are also specimens from China (Alashan desert, Burhan Budai Shan Mts., Karasai, Sharagoldzhin river valley in Nan-Shan, Landzou). This species is most similar to A. inylchekensis Frolov, sp. n. and can be separated from it by the color of head and pronotum, sparser and shorter setae on elytra, and paramere shape.
Aphodius (Bodilus) inylchekensis Frolov, sp. n.
(Figs 1, 8, 14)
Type material. Holotype ♂ labeled "Kirgiztan, 5 km S Inylchek, 2100 m., 5./10.8.1994 leg. Saldaitis". Paratypes: 2 ♂♂ and ♀ with the same data; ♂ and ♀ labeled "SU Kirgizia, E lower Inylchek vall., 2600―2700 m., 22―25.7.1989 Businsky".
Holotype. Male. Head dark brown, shiny, not tuberculate; frontal suture indistinct. Clypeus feebly sinuate at middle, rounded at sides, sparsely punctate (punctures separated by 2―3 times a puncture diameter). Genae rectangular, protruding past eyes. Pronotum brown on disc with sides and part of base lighter, shiny, densely, irregularly punctate. Anterior margin of pronotum not bordered, sides and base finely bordered; hind angles obtuse. Scutellum triangular, shiny, light brown, finely punctate. Elytra yellowish-brown, shiny, without humeral teeth. Striae fine, distinctly punctate. Intervals flat, sparsely punctate. Elytra pubescent with long setae on apices and sides and with short, sparse setae on disc. Apical spur of fore tibia relatively slender and acute, attached opposite basal tibial tooth, reaching middle of second tarsal segment (Fig. 8). Apical spurs of middle tibia slender and acute; lower spur longer than 2/3 of upper. First segment of hind tarsi as long as upper spur of tibia and slightly shorter than 3 following segments together. Adjoining apical setae of hind tibiae relatively long and unequal. Venter of body brown. Disc of metasternum with sparse setae on the perimeter. Aedeagus ― Fig. 14. Body length 6.0 mm.
Female can be separated from male by the narrower pronotum, denser punctures on the pronotum and head, and by the shorter apical spur of the fore tibia, attached opposite the middle tibial tooth.
Paratypes. Body length 7.0 to 8.0 mm. Color of pronotum varies slightly; one specimen with base of pronotum yellowish-brown.
Differential diagnosis. The species is most similar to A. sordescens, which is widely distributed in Central Asia, and it can be separated from it by the darker color of the head and pronotum, longer and denser setae on the elytra, and by the shape of the parameres.
Etymology. The name is derived from Inylchek Ridge in eastern Kirgizstan where the type specimens were collected.
Aphodius (Bodilus) hastatus Reitter, 1892
(Figs 7, 13)
Material examined: holotype ♂ labeled "Buchara", "A. hastatus m. 1893"; ♂ labeled "Buchara", "Type", "A. hastatus Rtt." [not the name-bearing type]; Turkmenistan: Ashkhabad, 3 spm., Imam-Baba, 26.IV.1912, 13 spm. (W. Koshantschikov); Uzbekistan: 70 km S Tamdybulak, 14.V.1965, 1 spm. (G. Medvedev); Kazakhstan: Dzhambul. This rare species can easily be separated from other Bodilus species by the widened apical tooth of the fore tibia in males.
Aphodius (Bodilus) gregarius Harold, 1871
(Figs 3, 5, 6, 12)
This species was described from Sarepta (outskirts of Volgograd) and is widely distributed in Kazakhstan and Middle Asia. In the collection of Zoological Institute there are specimens from Russia (Volga region), Kazakhstan (Gur'ev, Ural'sk, Aktyubinsk, Kyzyl-Orda, Karaganda, Semipalatinsk, Alma-Ata and Taldy-Kurgan provinces), Mongolia (Gov'-Altaj, Bajanhongor, Hovd aimaks) and Xinjiang Uygur autonomous region of China (1 spm. from outskirts of Hami). The species was recorded from Pakistan (Swat, Rawalpindi) by Stebnicka (1989). However, judging from the figures of the aedeagus in her article, this reference belongs rather to A. rudii or to undescribed species.
Aphodius (Bodilus) rudii Endroedi, 1968
This species was described from Mongolia (Dornod aimak). It is distributed throughout Central Asia and to northeastern Tibet. I have examined numerous specimens from Mongolia (Dornod, Uverhangaj, Umnegov, Bajanhongor aimaks) and northern China (Xinjiang Uygur and Nei Mongol autonomous regions).
Nikolajev and Puntsagdulam (1984) considered A. rudii as a junior synonym of A. gregarius. However, there is a clear difference in the paramere shape between the two species and there were no individuals with intermediate form of aedeagus found. This difference cannot be accounted for deformation of parameres during mounting as the aedeagi were examined in glycerine. A. rudii can also be separated from A. gregarius by its smaller average size. Females are difficult to separate.
Aphodius (Bodilus) insperatus Petrovitz, 1967
The species was described from Pakistan (Kwetta). I have examined the holotype (♂) and the paratype (♀) deposited in Naturhistorisches Museum Wien and the following material from Iran (collected by N.A. Zarudny): East Horasan, Avaz, 26―29.IV.1898, ♂; Mazenderan, Bender-Gez, 14.IX.1903, ♂; Sistan, Serkhed upland region, Noukabad, 1.IV.1901, ♂ and ♀. The species is similar to A. gregarius and A. rudii and can be separated from them by sparser pubescence of the elytra and the shape of the parameres.
Rakovič (1984) described A. longipennis from a male specimen from Afghanistan (Kalat-Mukur) and established the monotypic subgenus Paramelinopterus for it. It can be inferred from the original description of the species that it differs from A. insperatus only by the darker color of head and pronotum, which may be within the limits of variation of one species. Unfortunately, I could not examine the type of the species, but it undoubtedly shares all the characters of subgenus Bodilus (group gregarius), and there seems to be no reasons to establish a separate subgenus for it. Therefore, the new synonymy (Bodilus Mulsant et Rey, 1869 = Paramelinopterus Rakovič, 1984, syn. n.) is here established.
Although the species involved must constitute a monophyletic group, the subgenus is not homogeneous. According to morphological characters of the species involved, they can be grouped into a few groups of species: 1) group lugens: A. lugens, A. longeciliatus, A. ictericus; 2) group punctipennis: A. punctipennis, A. zarudnyi sp. n.; 3) group sordescens: A. sordescens, A. inylchekensis sp. n., A. hastatus; and 4) group gregarius: A. gregarius, A. insperatus, A. rudii.
From examination of these species, some preliminary conclusions about the phylogeny of the subgenus can be made. If one consider the modification of fore tibia and its spur in the male (characters of sexual dimorphism, which are found in different variations in many Aphodius species) as an apomorphic state, and “normal” or “typical” state as plesiomorphic, then the ancestral form of the group might be close to the recent species A. longeciliatus. During speciation, the ancestral form diverged in a few daughter lineages, corresponding here to groups of species. In A. lugens and A. ictericus, weakly deviated form the ancestral form, the apices of the parameres are widened in lateral aspect, therefore they lost their primordial dorso-ventral flatness. In the group punctipennis, which preserves the main characters of the ancestral form, including the shape of aedeagus, a comparatively denser pubescence of the body developed (especially on the elytra, disc of metasternum, and sides of pronotum). The third lineage includes the group sordescens, which is characterized by lighter color, distinct macula on the disc of pronotum, fore tibial spur attached opposite basal tooth of tibia in male, and by the head not tuberculate. The most deviated lineage includes the group gregarius, which is characterized by the features of the previous group, and additionally by some more specific characters expressed in males: fore tibiae with heavily shortened spur and brush of dense setae on ventral surface of apical tooth.
I would like to thank Dr. Manfred Jaeh (Wien) for the opportunity to examine R. Petrovitz’s types, Dr. Brett Ratcliffe (University of Nebraska State Museum, Lincoln) for valuable suggestions and linguistic review of the manuscript, and Mr. Kirill Dowgailo (Minsk) for helping make photos of the new species.
Nikolajev G.V., Puntsagdulam Z. 1984. Plastinchatousye Mongolskoj Narodnoij Respubliki // Nasekomye Mongolii. Vol. 9. P. 90―294 [in Russian].
Rakovič M. 1984. Two interesting species of the genus Aphodius Illiger from Afghanistan // Sbor. faun. Praci ent. Mus. Vol. 17. P. 213―218.
Schmidt A. 1916. Namenanderungen und Beschreibung neuer Aphodiinen // Arch. Naturg., Abt. A. Bd. 82. S. 95―116.
Stebnicka Z. 1989. Revision of the Aphodiinae of the Western Himalayas // Stutt. Beitr. Naturk. Bd. 441. S. 1―29.