Paleontological Journal, Vol. 30, No. 2, 1996, pp. 217Ð224. Translated from Paleontologicheskii Zhurnal, No. 2, 1996, pp. 91Ð99. Original Russian Text Copyright © 1996 by Nikolajev. English Translation Copyright © 1996 by åÄàä ç‡Û͇ /Interperiodica Publishing (Russia).

Lamellicorn Species (Coleoptera, Scarabaeidae) with Multistriate Elytra from the Lower Cretaceous Baisa Locality, Transbaikalia G. V. Nikolajev Kazakh State National University, Alma-Ata, Kazakhstan Received February 23, 1995

Abstract—Two new species of the Mesozoic genus Proteroscarabaeus, P. magnus and P. nikritini, and three new genera with one new species each, Cretanaides trogopterus (Hybosorinae), Cretobolbus rohdendorfi (Bol- boceratinae), and Hybosorites fissuratus (subfamily incertae sedis), are described from the Lower Cretaceous of Transbaikalia. A new tribe is created within the Geotrupinae for Cretogeotrupes. One new tribe is established within the Hybosorinae, and two more within the extant Bolboceratinae. Cenozoic Bolboceras tertiarium is transferred to Phaeochrous; doubt is expressed regarding the placement of the monobasic Cenozoic genus Coprologus in the Bolboceratinae.

Among lamellicorn specimens collected in the in that the middle coxae are rather broadly separated, and Lower Cretaceous Baisa locality, Transbaikalia, six from Proteroscarabaeus in that the elytra are broadly belong to the species possessing more than 10 elytral rounded lateroapically and the fore tibia is tridentate. striae. So far as numerous striae are characteristic of All the specimens described below are from the few lamellicorn taxa, an attempt is made to elucidate Baisa locality (Transbaikalia, Buryatia, Eravnenskii the systematic position of these species. One of them Region, left bank of the Vitim River downstream of the was described in the genus Cretogeotrupes (Nikolajev, mouth of Baisa River; Lower Cretaceous, ValanginianÐ 1992), five more are described below as new. Four Hauterivian, Zaza Formation) and deposited in the specimens are preserved as isolated elytra (Fig. 1), Paleontological Institute, Russian Academy of Sci- in the fifth one the body and legs are preserved as well. ences (PIN). In addition to Cretogeotrupes, more than 10 elytral striae are characteristic of the Mesozoic genus Prot- Family Scarabaeidae Laicharting, 1781 eroscarabaeus Grabau and several genera of the extant ?Subfamily Pleocominae Leconte, 1861 subfamilies Bolboceratinae, Geotrupinae, Hybosori- Genus Proteroscarabaeus Grabau, 1923 nae, and Ceratocanthinae. Due to the rather large size Proteroscarabaeus magnus Nikolajev, sp. nov. of the specimens in question, one could exclude the Ceratocanthinae from consideration. Two elytrae are of Pl. XI, fig. 1 a peculiar shape, somewhat narrowed towards the apex Etymology. From Latin magnus (large). (Figs. 1a, 1b), similar to those of the type species of Holotype. PIN 1989/3026, right elytron (part and Proteroscarabaeus (Fig. 2a) and in contrast to Creta- counterpart); Baisa locality, bed 15 (Martinson, 1961). geotrupes and all recent taxa, having broadly rounded Description (Fig. 1a). Large, weakly convex elytrae apically. The systematic position of Proteros- elytra distinctly narrowed apically, with 13Ð14 striae carabaeus is still uncertain; it is closest to the monoba- nearly equidistant from each other. Striae not deep, sic Pleocominae (Nikolajev, 1992) and could be weakly impressed, with few points. Intervals weakly assigned provisionally to the subfamily until a reexam- convex; epipleura rather narrow, distinctly narrower than ination of the type species. One more elytra (Fig. 1c) adjacent interval. Humeral tubercle poorly developed. bears a very complicated sculpture found only in some Measurements, mm. Elytra length 20; maxi- genera of the Hybosorinae. Another elytron (Fig. 1d), mal width 11. poorly preserved along the sutural margin and clearly Comparison. Distinct from other species in the distinct from that of Proteroscarabaeus, has insuffi- larger size. cient diagnostic characters and could belong to any of the three subfamilies (Bolboceratinae, Geotrupinae, or Material. Holotype. Hybosorinae). The last of the specimens discussed, a nearly com- Proteroscarabaeus nikritini Nikolajev, sp. nov. plete beetle, could be assigned to the recent Bolbocera- Etymology. In honor of the entomologist tinae; it differs from the Hybosorinae and Geotrupinae L.M. Nikritin.

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(b)

(a) (c)

(d)

Fig. 1. Scarabaeid elytrae from Baisa locality: (a) Proteroscarabaeus magnus sp. nov., holotype PIN 1989/3026; (b) P. nikritini sp. nov., holotype PIN 3064/7154; (c) Cretanaides trogopterus sp. nov., holotype PIN 4210/706; (d) Hybosorites fissuratus sp. nov., holotype PIN 4210/709. Scale unit in all figures 1 mm.

Holotype. PIN 3064/7154, right elytra (part and Comparison. Distinct from the type species, counterpart); Baisa locality, bed 31 (Martinson, 1961). P. yeni Grabau in the relatively shorter elytrae (length/width ratio of P. yeni elytron is 2.3 : 1) (Fig. 2a). Description (Fig. 1b). Large, weakly convex Material. Holotype. elytra distinctly narrowed apically; its length/width ratio 1.7 : 1. 11 striae between suture and humeral tubercle. Striae deep, with numerous points; first three Subfamily Hybosorinae Lacordaire, 1856 striae possibly terminate against scutellum. Intervals ?Tribe Anaidini Nikolajev, trib. nov. convex; sutural interval somewhat broader than the Genus Cretanaides Nikolajev, gen. nov. next one, remaining of nearly equal width; epipleura somewhat narrower than adjacent interval. Humeral Etymology. From Latin cretaceus (Cretaceous) tubercle well developed. Measurements, mm. Elytra and genus Anaides. length 17.0; maximal width 9.9. Type species. C. trogopterus sp. nov.

PLATE XI Fig. 1. Proteroscarabaeus magnus sp. nov.; holotype PIN 1989/3026 (×4.4): (a) cast; (b) counterpart. Fig. 2. Cretanaides trogopterus sp. nov.; holotype PIN 4210/706 (×10). Fig. 3. Hybosorites fissuratus sp. nov., holotype PIN 4210/709 (×7.1).

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(a) (b)

Fig. 2. Reconstructions of Mesozoic Scarabaeidae: (a) Proteroscarabaeus yeni Grabau; (b) Cretogeotrupes convexus Nikolajev.

Diagnosis. Middle-sized convex beetle. Elytra Nel, 1987) demonstrates a similar sculpture of the ely- with numerous (more than 15) punctate striae and tral intervals. In these genera the borders of the elytral weakly convex intervals (some of them bearing elon- striae are vertical (the same is true of at least the first gate tubercles). Fine sutural stria not developed. stria in Cretanaides), middle and hind tibiae lack trans- Species composition. Monobasic. verse carinae, fore tibiae with three large teeth at outer margin, and the wings (studied in Anaides laticollis Comparison. Distinct from all Mesozoic scara- Harold) with a single free vein between the cubital and baeid genera with multistriate elytra, in the complex first anal vein reaching the wing base. The above char- sculpture of intervals between striae. acters are sufficient for separation of the genera listed Systematic position. Strongly convex, rela- into the new tribe Anaidini Nikolajev, trib. nov. The tively large elytron with complicated sculpture character states could be considered as derived in the excludes the genus from the Ceratocanthinae. Deeply Anaidini relative to the Hybosorini s. str. rounded outer apical margin of elytron prevents associ- ation of Cretanaides with the Mesozoic Proteroscara- baeus, provisionally included in the Pleocominae. The Cretanaides trogopterus Nikolajev, sp. nov. very complex elytral sculpture is unknown in the Geotrupinae. No single character known contradicts Pl. XI, fig. 2 the placement of the genus into the Hybosorinae. Etymology. From Greek pteron (wing) and Within the latter subfamily, the group of American gen- genus Trox. era (Anaides Westwood, Cryptogenius Westwood, Cre- mastochiloides Krikken, Callosides Howden) Holotype. PIN 4210/706, left elytra (part and (Howden, 1971; Krikken, 1975; Scholtz, D’Hotman, counterpart); Baisa locality, bed 4 (Martinson, 1961).

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Description (Fig. 1c). Medium-sized, oblong, antennal club, and middle and hind tibiae with spurs convex elytra, slightly narrowed apically. Striae rather displaced to the same side from tarsal base. The only deeply punctate; the points separated by a little more derived character shared by the Bolboceratinae with than their diameter. First stria (from suture) not deep, Geotrupinae–Scarabaeinae “clade”, 3-segmented anten- with points broader than stria, and borders sharply nal club, could be regarded as a synapomorphy uniting impressed (vertical); it is uncertain, whether or not these taxa in one group (Nikolajev, 1995). remaining striae are of the same structure. No less than The Bolboceratinae is now separated into the five 15 striae, but exact number is unknown, since outer tribes: monobasic Bolboceratini Mulsant, Athyreini striae are faint. Intervals of different width; some of Howden et Martinez (4 genera; Howden, Martinez, them with tubercles, large and prominent in the 3rd, 1963), Bolbochromini Nikolajev (3 genera; Nikolajev, 7th, 11th and 15th intervals, and superficial but longer 1970), monobasic Gilletinini Nikolajev (Nikolajev, in the 5th, 9th and 13th intervals; tubercles well devel- 1990), and Eubolbitini Nikolajev with remaining gen- oped on disc but decreasing both basally and apically. era (Nikolajev, 1970). So far as the trend of the outer Epipleura rather broad (preserved only in basal third of teeth number to decrease on fore tibia occurs in several elytron). tribes of the subfamily, the tridentate condition is not Measurements, mm. Length 9.5; width 4.4 sufficient to separate the genus into one more tribe. The (perhaps a little larger, since outer margin and apex are broad scutellum and rather narrowly separated middle poorly preserved). coxae preclude the assignment of Cretobolbus to the Material. Holotype. Athyreini or Gilletinini. Contiguous middle coxae dis- tinguish the Bolboceratini from Cretobolbus. Almost contiguous middle coxae and also (in most genera) sev- Subfamily Bolboceratinae Mulsant, 1842 eral transverse carinae on middle and hind tibiae in the Genus Cretobolbus Nikolajev, gen. nov. Bolbochromini separate the tribe from Cretobolbus as Etymology. From Latin cretaceus (Cretaceous) well. It is only within the Eubolbitini that all the char- and Latin bolbus (bulb), a root of many generic names acters of Cretobolbus (except for tridentate fore tibia) in the subfamily. could be met. However, in my opinion, the tribe Eubol- Type species. C. rohdendorfi sp. nov. bitini is a conglomeration and deserves separation into Diagnosis. Middle-sized convex beetle. Elytra at least three tribes. with numerous (no less than 14) punctate striae and The genera Bolbelasmus Boucomont, Bolbotrypes weakly convex intervals; fine sutural stria developed. Olsoufieff and Bolbogonium Boucomont share the mul- Fore tibiae with three teeth at outer margin. Middle tidentate fore tibiae, numerous transverse carinae on coxae not broadly separated, forming a right angle to middle and hind tibiae (their number tending to each other. Middle tibiae transversely unicarinate. decrease), unmodified clypeus (discal tubercle not Species composition. Monobasic. tending to fuse with raised anterior border), and basally emarginate wing with radial vein lacking apical extra Comparison. Distinct from other genera of the branches. The tribe Bolbelasmini Nikolajev, trib. nov. is subfamily in the tridentate fore tibiae, and from all created for these three genera. Cretobolbus is distinct Mesozoic scarabaeids with more than 10 elytral striae, from the tribe in the unicarinate middle tibiae. in the fine sutural stria present. Systematic position. A combination of fine The Australian genera Australobolbus Howden et sutural stria and separated middle coxae indicates that Cooper, Bolbobaineus Howden et Cooper, Bolborach- the genus could belong only to the Bolboceratinae. ium Boucomont, Blackburnium Boucomont, Blackbol- bus Howden et Cooper, Bolboleaus Howden et Cooper Most authors treat the latter taxon as a tribe within (Howden and Cooper, 1977), North American Bolboce- the Geotrupinae (Howden, 1982; Zunino, 1984a, b) rastes Cartwright and Bradycinetulus Cockerell (Cart- rather than a subfamily. However, the character analy- wright, 1953), African Prototrupes Krikken (Krikken, sis shows the both are sister groups of equal rank. Char- 1977), and Asiatic Indobolbus Nikolajev (Nikolajev, acters shared by the Bolboceratinae and Geotrupinae 1979) are characterized by the base of elytra emargin- (11-segmented antennae, fore tibiae with many teeth ated, and discal tubercle of clypeus fused with its ante- along outer margin, wings with two free veins between rior border. These genera are separated into the Austral- cubital and first anal vein reaching wing base) are prob- obolbini Nikolajev, trib. nov. (a paper with more ably symplesiomorphic for the whole superfamily. detailed discussion of taxonomic structure of the Bolbo- A combination of the middle coxae separated (plesio- ceratinae is in preparation). morphy) and the absence of a distinct silky hair spot at the fore femora (apomorphy) in the Bolboceratinae The only specimen of Cretobolbus is too insuffi- forces us to consider it as a sister group to the Geotrupi- ciently preserved to include the genus into either nae, having the middle coxae contiguous (apomorphy) Eubolbitini or Australobolbini. and fore femora with distinct silky hair spot (plesio- R e m a r k s. The descriptions and illustrations of morphy). Apomorphic characters shared by the Bolbo- the Cenozoic species from Europe assigned to the Bol- ceratinae and the Hybosorinae are the 3-segmented boceratinae (Heer, 1848; Deichmüller, 1881) show that

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(a) (b)

Fig. 3. Cretobolbus rohdendorfi sp. nov.: (a) holotype PIN 3064/7137; (b) reconstruction. they are incorrectly assigned. Bolboceras tertiarium impressed only laterally) and a deep sutural stria; lat- Deichmüller should be assigned to the Hybosorinae on eral elytral margin weakly evenly rounded. Fore femur account of the middle tibiae lacking transverse carina; (only right leg preserved) with sparse points at anterior the species most probably could be included with margin (silky hair spot probably “blurred”). Fore tibia Phaeochrous Castelnau. Coprologus gracilis Heer pre- with a long longitudinal carina and three large, equidis- sumably belongs to the Hybosorinae as well (judging tant teeth on outer margin. Middle coxae clearly sepa- from the description of head shape and labrum), but the rated, forming almost right angle to each other. Middle holotype should be reexamined to confirm this. tibia (left one preserved) with single transverse carina near midlength. Radial vein (traceable in apical part of right wing) possibly branched. Cretobolbus rohdendorfi Nikolajev, sp. nov. Measurements, mm. Length from anterior Etymology. In memory of paleoentomologist pronotal margin to wing apex 22; median length of B.B. Rohdendorf. pronotum 4.5, width 8.1; elytron length 12.5, width 6.6; Holotype. PIN 3064/7137, beetle lacking head fore tibia length 3.5; middle tibia length 3.6; space and most of legs, and the wings projecting from beyond between middle coxae 1.0. elytra (cast and counterpart); Baisa locality, bed 15 Material. Holotype. (Martinson, 1961).

Description (Fig. 3). Rather large oval beetle. Subfamily Geotrupinae Latreille, 1804 Pronotum with leathery margin anteriorly, emarginated also laterally and posteriorly. Anterior angles of prono- Tribe Cretogeotrupini Nikolajev, trib. nov. tum obtuse; pronotum slightly emarginate laterally Diagnosis. Relatively small . Elytrae near acute posterior angles. Scutellum large, triangular. concealing abdomen entirely, each with numerous Elytra with numerous punctate striae (deeply (more than 11) striae; wings well developed. Fore tibiae

PALEONTOLOGICAL JOURNAL Vol. 30 No. 2 1996 LAMELLICORN BEETLE SPECIES (COLEOPTERA, SCARABAEIDAE) 223 tridentate on outer margin; middle coxae contiguous; Subfamily incertae sedis middle and hind tibiae transversely bicarinate (see Fig. 2b). Genus Hybosorites Nikolajev, gen. nov. Generic composition. Cretogeotrupes Etymology. From recent genus . Nikolajev, 1992, Lower Cretaceous of Asia. Type species. H. fissuratus sp. nov. Systematic position. Up to now only three Diagnosis. Medium-sized beetle. Elytra with tribes were established within the subfamily: Geotru- numerous striae (more than 11), outer apical margin pini, Lethrini Mulsant et Rey, 1871 and Taurocerastini broadly rounded. Germain, 1897. The nominotypical one includes both winged and apterous species, having multistriate elytra Species composition. Monobasic. and transversely tricarinate (sometimes bi- or even uni- Comparison. Distinct from Mesozoic genera: carinate) middle and hind tibiae. The monobasic from Proteroscarabaeus in the elytra broadly rounded Lethrini is composed only of apterous species (apomor- apically; from Cretanaides and Cretogeotrupes in the phy) with elytra bearing 10 striae (plesiomorphy) and less convex elytra; from Cretobolbus in the somewhat middle and hind tibiae transversely unicarinate (apo- smaller size. morphy). Therefore, these two tribes could be only in Systematic position. The fossil is too large sister group relationship. The Taurocerastini, with only to belong in the Ceratocanthinae, and insufficiently pre- two genera included, demonstrates along with apomor- served to show diagnostic characters of the Hybosori- phic characters (10-segmented antennae, middle and nae, Bolboceratinae, or Geotrupinae. hind tibiae transversely unicarinate), one plesiomor- phy, the eyes only partly subdivided by the genal pro- jection (completely divided eyes in Geotrupini and Hybosorites fissuratus Nikolajev, sp. nov. Lethrini is an apomorphy). Thus, the Taurocerastini Pl. XI, fig. 3 could be only a sister taxon to one of two remaining Etymology. From Latin fissuratus (cracked). recent tribes. Holotype. PIN 4210/709, right elytron; Baisa The new tribe is certainly a sister group to Lethrini. locality, bed 22 (Martinson, 1961). Lack of data on the head structure prevent the analysis of relationships of two other tribes to the Cretogeotru- Description (Fig. 1d). Epipleura broad, ending pini. If Cretogeotrupes had the structure of eyes, anten- at about 3/4 of elytron length. Striae deeply impressed, nae and mouthparts comparable to those of the Tauro- with large points. Humeral tubercle well developed. cerastini, it could be ancestral to this tribe; if as in the Measurements, mm. Length (as preserved) Geotrupini, it presumably would be derivative of the 11.5, width 6.5 (estimated length of complete elytron latter tribe. below 13). R e m a r k s. The Geotrupinae constitutes one phyl- Material. Holotype. etic lineage with the Troginae, Aphodiinae, and Scara- baeinae. That of all these taxa originate from a common ancestor is beyond doubt. The Aphodiinae (Nikolajev, ACKNOWLEDGMENTS 1993) and Scarabaeinae are more closely related to I am grateful to A.G. Ponomarenko and V.V. Zherikhin each other than to remaining subfamilies of the (Moscow) for loan their of the fossil specimens and to “branch”. It is still possible that contrary to the earlier H.F. Howden (Canada) for the loan of the recent suggestion (Nikolajev, 1993) the Scarabaeinae is rather Anaides species, and to F.-T. Krell (Germany) who a derivative of one of the Aphodiinae subgroups than its helped with the literature on fossil species. This study sister group. Such a possibility was considered by some was supported by the International Science Foundation authors (e.g., Cambefort, 1987) (for family group taxa, and Russian Academy of Natural Sciences. the name Scarabaeinae has priority). Avoiding a discus- sion on the rank of the three taxa, one could try to elu- cidate their relationships. REFERENCES The Geotrupinae and Troginae share the middle Cambefort, Y., Insectes Coleopteres Aulonocnemidae, coxae contiguous and middle and hind tibial spurs Faune de Madagascar, 1987, vol. 69, pp. 1Ð87. always close-set. These apomorphies often appear in Cartwright, O.L., Scarabaeid Beetles of the Genus several lamellicorn groups (Nikolajev, 1993) and can Bradycinetulus and Closely Related Genera in the not be considered sufficient for uniting the two subfam- United States, Proc. U. S. Nat. Mus., 1953, vol. 103, ilies. The Troginae and Scarabaeinae share the other no. 3318, pp. 95Ð120. apomorphies, the reduced number of both antennal seg- Deichmüller, J.V., Fossile Insekten aus dem Diatomen- ments and abdominal sterna. However, these characters schiefer von Kutschlin bei Bilin, Böhmen, Nova Acta appear in many lamellicorn groups as well. Therefore, Acad. Leopold., 1881, vol. 42, no. 6, pp. 1Ð39. it is reasonable to treat these three groups as taxa of Heer, O., Die Insektenfauna der Tertiärgebilde von equal rank, until the relative “weight” of the above Oeningen und von Radoboj in Croatien, Neue Denkschr. characters is determined. Allg. Schweiz. Ges. Naturwiss., 1848, vol. 8, pp. 1Ð230.

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Howden, H.F., Five Unusual Genera of New World Scar- Nikolajev, G.V., Neue Gattungen und Untergattungen abaeidae (Coleoptera), Canad. Entomol., 1971, vol. 103, der Blatthornkäfer (Coleoptera, Scarabaeidae), Reichen- pp. 1434Ð1471. bachia, 1979, vol. 17, no. 23, pp. 189Ð191. Howden, H.F., Larval and Adult Characters of Frickius Nikolajev, G.V., Use of Wing Venation Features for Germain, Its Relationships to the Geotrupini, and a Phy- Improvement of the Systematics of Lamellicorn Beetles logeny of Some Major Taxa in the (Coleoptera, Scarabaeidae), Uspekhi entomologii v (Insecta: Coleoptera), Canad. J. Zool., 1982, vol. 60, SSSR: Zhestkokrylye nasekomye. Materialy X s’ezda no. 11, pp. 2713Ð2724. Vsesoyuznogo entomologicheskogo obshchestva 11Ð15 Howden, H.F. and Cooper, J.B., The Generic Classifica- sentyabrya 1989 g., Leningrad, 1990, pp. 98Ð99. tion of the Bolboceratini of the Australian Region, with Nikolajev, G.V., Taxonomic Characters and Composi- Descriptions of Four New Genera (Scarabaeidae: tion of the Genera of Mesozoic Lamellicorn Beetles Geotrupinae), Austral. J. Zool., 1977, Suppl. Ser., no. 50, (Coleoptera, Scarabaeidae), Paleontol. Zh., 1992, no. 1, pp. 1Ð50. pp. 76Ð88. Howden, H.F. and Martinez, A., The New Tribe Nikolajev, G.V., The Taxonomic Placement in the Sub- Athyreini and Its Included Genera (Coleoptera: Scara- family Aphodiinae (Coleoptera, Scarabaeidae) of the baeidae, Geotrupinae), Canad. Entomol., 1963, vol. 95, New Genus of Lower Cretaceous Scarab Beetles from no. 4, pp. 345Ð352. Transbaykalia, Paleontol. J., 1993, vol. 27, suppl. 1A, Krikken, J., Cremastochilodius tristis, a New Scarabae- pp. 1Ð8. oid Genus and Species from Brazil, Entomol. Berichten, 1975, vol. 35, pp. 190Ð192. Nikolajev, G.V., A New Subfamily of Lamellicorn Bee- tles (Coleoptera, Scarabaeidae) from the Lower Creta- Krikken, J., New Bolboceratinae Genera from South- ceous of Transbaikalia and Its Phylogenetic Relation- western Africa (Coleoptera: Geotrupidae), Zool. Meded., ships, Paleontol. Zh., 1995, no. 2, pp. 147Ð151. 1977, vol. 52, no. 12, pp. 161Ð168. Scholtz, C.H., D’Hotman, D., and Nel, A., Systematic Martinson, G.G., Mezozoiskie i kainozoiskie mollyuski Position of the South American Genus Cryptogenius kontinental’nykh otlozhenii Sibirskoi platformy, Westwood (Coleoptera: Scarabaeoidea: Trogidae and Zabaikal’ya i Mongolii (Mesozoic and Cenozoic Mol- ), Coleopt. Bull., 1987, vol. 41, no. 2, lusca from Continental Deposits of Siberian Platform, pp. 193Ð199. Transbaikalia and Mongolia), MoscowÐLeningrad: Akad. Nauk SSSR, 1961, (Trudy Baikal. limnol. stantsii, Zunino, M., Sistematica generica dei Geotrupinae vol. 19). (Coleoptera, Scarabaeoidea: Geotrupidae), filogenesi della sottofamiglia e considerationi biogeografiche, Boll. Nikolajev, G.V., Taxonomic Rank of the Groups Mus. Reg. Sci. Nat. Torino, 1984a, vol. 2, no. 1, pp. 9Ð162. Included in the Subfamily Geotrupinae (Coleoptera, Scarabaeidae), Materialy II nauchnoi konferentsii molo- Zunino, M., Analisi sistematica e zoogeografica della dykh spetsialistov i aspirantov, posvyashchennoi sottofamiglia Taurocerastinae Germain (Coleoptera, 100-letiyu so dnya rozhdeniya V.I. Lenina i 50- letiyu Scarabaeoidea: Geotrupidae), Boll. Mus. Reg. Sci. Nat. Kazakhstana, Alma-Ata, 1970, p. 31Ð34. Torino, 1984b, vol. 2, no. 2, pp. 445Ð464.

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