Abstract
The external and internal male genitalia of 327 species of 11 tribes of the subfamily Scarabaeinae, including species of Deltochilini, Scarabaeini, Gymnopleurini, Ateuchini, and Coprini, among others, were examined. Descriptions of the variations in the genital segment, the aedeagus, the internal sac, and its sclerites and raspules are presented. An exhaustive comparison of structures, names, and terminology used in literature for Scarabaeinae male genitalia are discussed. The internal sac of the aedeagus is divided in areas for an easer comparison of its internal structures; basal, submedial, medial, and apical areas are described in detail and compared. The variation of apical and medial sclerites, as well as the raspules of the submedial area, are described and compared in detail among all the taxa studied.
References
Anichtchenko, A. & Verdugo, A. (2004) Iberodorcadion (Hispanodorcadion) zenete, nueva especie Ibérica de cerambícido (Coleoptera, Cerambycidae, Lamiinae) procedente de Sierra Nevada (Andalucía, España). Boletín de la Sociedad Andaluza de Entomología, 11, 31–42.
Barbero, E., Palestrini C. & Zunino M. (1991) Filogenesi, biogeografia e sistematica generica dei Sisyphini (Coleoptera: Scarabaeidae), pp. 837–844. Atti XVI Congresso Nazionale Italiano di Entomologia Bari-Martina Franca (TA), 23/28 settembre. Verona, Italy.
Barbero, E., Palestrini, C. & Roggero, A. (1998) Phylogenetic relationships in the genus Scarabaeus Linnaeus. Atti 1. Colloquio Nazionale di Sistematica Cladistica. Memorie del Museo civico di storia naturale di Verona, 13, 87–96.
Berti, N. & Mariau, D. (1999) Coelaenomenodera lameensis n. sp., ravager du palmier à huile (Coleoptera, Chrysomelidae). Nouvelle Revue d’Entomologie, 16, 253–267.
Binaghi, G., Dellacasa, G. & Poggi, R. (1969) Nuovi Caratteri diagnostici per la determinazione degli Onthophagus del gruppo ovatus (L.) e geonemia controllata delle specie Italiane del grupo. Memorie de lla Società Entomologica Italiana. Genova, 48, 29–46.
Bouchard, Y., Davies, A.E., Alonso-Zarazaga, M.A., Lawrence, J.F., Lyal, C.H.C., Newton, A.F., Reid, C.A.M., Schmitt, M., Ślipiński, S.A. & Smith, A.B.T. (2011) Family-group names in Coleoptera (Insecta). Zookeys, 88, 1–972.
http://dx.doi.org/10.3897/zookeys.88.807Bovo, B. & Zunino, M. (1983) Nouvi generi di Geotrupini (Coleoptera, Scarabaeoidea: Geotrupidae) asiatici. Bullettino del Museo Regionale di Scienze Naturali. Torino, 1, 397–416.
Carlson, D.C. (1975) Taxonomic characters of the genus Ochodaeus Serville with descriptions of two new species in the O. pectoralis Leconte species complex (Coleoptera: Scarabaeidae). Bulletin Southern California Academy of Sciences, 74, 49–65.
Carlson, D.C. & Ritcher, P.O. (1974) A new genus of Ochodaeidae and a description of the larvae of Pseudochodaeus estriatus (Schaeffer) (Coleoptera: Scarabaeidae). The Pan-Pacific Entomologist, 50, 99–110.
Coca-Abia, M. & Martín-Piera, F. (1998) Revisión taxonomica del género Rhizotrogus Berthold, 1827 (Coleoptera, Scarabaeidae, Melolonthidae). Coleopterological Monographs. Asociación Europea de Entomología, 2, 1–140.
D’Hotman, D. & Scholtz, C.H. (1990) Comparative morphology of the male genitalia of derived groups of Scarabaeoidea (Coleoptera). Elytron, 4, 3–39.
Forgie, S.A., Grebennikov, V.V. & Scholtz, C.H. (2002) Revision of Sceliages Westwood, a millipede-eating genus of southern Africa dung beetles (Coleoptera: Scarabaeidae). Invertebrate Systematics, 16, 931–955.
http://dx.doi.org/10.1071/IT01025Frolov, A.V. & Scholtz, C.H. (2004) Revision of the southern African genus Macroderes Westwood (Coleoptera: Scarabaeidae: Scarabaeinae). Annales de la Société Entomologique de France, 40, 373–393.
Frolov, A.V., Akhmetova, L.A. & Scholtz, C.H. (2008) Revision of the obligate mushroom-feeding African ‘‘dung beetle’’ genus Coptorhina Hope (Coleoptera: Scarabaeidae: Scarabaeinae). Journal of Natural History, 42, 1477–1508.
http://dx.doi.org/10.1080/00222930802002535Génier, F. (1996) A revision of the Neotropical genus Ontherus. Memoirs of the Entomological Society of Canada, 170, 1–169.
http://dx.doi.org/10.4039/entm128170fvGénier, F. (2000) New North American Ateuchus Weber (Coleoptera: Scarabaeidae, Scarabaeinae). The Coleopterists Bulletin, 54, 341–346.
http://dx.doi.org/10.1649/0010-065X(2000)054[0341:ANNAAW]2.0.CO;2Génier F. (2012) A new species and notes on the subgenus Deltochilum (Deltochilum) Eschscholtz, 1822 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini). Zootaxa, 3357, 25–36.
González, A., Molano, F., & Medina, C.A. (2009) Los subgéneros Calhyboma Kolbe 1893, Hybomidium Shipp 1897 y Telhyboma kolbe 1893 de Deltochilum (Coleoptera: Scarabaeidae: Scarabaeinae) en Colombia. Revista Colombiana de Entomología, 35, 253–274.
House, C.M. & Simmons, L.W. (2003) Genital morphology and fertilization success in the dung beetle Onthophagus taurus: an example of sexual selected male genitalia. Proceedings the Royal Society London, 270, 447–455.
http://dx.doi.org/10.1098/rspb.2002.2266House, C.M. & Simmons, L.W. (2005) The evolution of male genitalia: patterns of genetic variation and covariation in the genital sclerites of the dung beetle Onthophagus taurus. Journal of Evolutionary Biology, 18, 1281–1292.
http://dx.doi.org/10.1111/j.1420-9101.2005.00926.xKohlmann, B. (2000) New species and distribution records of Mesoamerican Ateuchus (Coleoptera: Scarabaeidae). Revista de Biología Tropical, 48, 235–246.
Kral, D. (2000) A review of Chinese Aphodius species (Coleoptera: Scarabaeidae). Part 2: revision of the subgenus Brachiapodius. Acta Societatis Zoologicae Bohemicae, 64, 65–92.
Krikken, J. & Huijbregts, J. (2007) Taxonomic diversity of the genus Ochicanthon in Sundaland (Coleoptera: Scarabaeidae: Scarabaeinae). Tijdschrift voor Entomologie, 150, 421–479.
http://dx.doi.org/10.1163/22119434-900000241López-Guerrero, I. (1999) Morphological comparison of the male genitalia of Eurysternus Dalman and Sisyphus Latreille (Coleoptera, Scarabaeidae, Scarabaeinae). The Coleopterists Bulletin, 53, 201–207.
López-Guerrero, I. (2005) Los Dichotomius (Coleoptera: Scarabaeidae: Dichotomiinae) de la Fauna de México. Boletín de la Sociedad Entomológica Aragonesa, 36, 195–209.
López-Guerrero, I., Zunino, M., & Halffter, G. (2009) Taxonomic use of the characters in Mexican Copris (Coleoptera: Scarabaeidae, Scarabaeinae): the case of Copris klugi sierrensis Matthews and the C. armatus species complex. The Coleopterists Bulletin, 63, 203–212. http://dx.doi.org/10.1649/1145.1
Márquez, J. (2001) Systematic review of Heterolinus Sharp (Coleoptera: Staphylinidae: Xantholinini). The Coleopterists Bulletin, 55, 317–329.
http://dx.doi.org/10.1649/0010-065X(2001)055[0317:SROHSC]2.0.CO;2Martín-Piera, F. (1987) Revision of the genus Chironitis Lansberge 1875. Entomologische Arbeiten aus dem Museum Georg Frey, 35/36, 203–245.
Martínez, A. & Pereira, F.S. (1956) Dois generos novos de Deltochilini Americanos. Papéis Avulsos do departamento de Zoologia, 12, 363–388.
Matalin, A.V. (1998) The tiger-beetles of “hybrida”-species group (Coleoptera, Carabidae, Cicindelinae). III. A taxonomic review of the Iberian Cicindela lagunensis Gautier, 1872 complex. Graellsia, 54, 75–96.
http://dx.doi.org/10.3989/graellsia.1998.v54.i0.346Matalin, A.V. (1999) The tiger-beetles of the hybrida species-group. II. A taxonomic review of subspecies of Cicindela sahlbergii Fischer von Waldheim, 1824 (Coleoptera Carabidae Cicindelini). In: A.Zamotajilov & R. Sciaky (eds.), Advances in Carabidology. Muiso publishers, Krasnodar, Russia, pp. 1–473.
Matthews, E.G. (1974) A revision of the Scarabaeine dung beetles of Australia II. Tribe Scarabaeini. Australian Journal of Zoology, Supplementary Series, 24, 1–211.
http://dx.doi.org/10.1071/AJZS024Medina, C. A., Scholtz, C. & Gill, B. D. (2003) Morphological variation and systematics of Canthon Hoffmannsegg 1817, and related genera of new world Deltochilini dung beetles (Coleoptera, Scarabaeidae). Deutsche Entomologische Zeitschrift, 50, 23–68.
http://dx.doi.org/10.1002/mmnd.20030500105Medina, C.A. & Scholtz, C. (2005) Systematics of the southern African genus Epirinus Reiche (Coleoptera: Scarabaeinae: Deltochilini): descriptions of new species and phylogeny. Insects Systematic and Evolution, 36, 145–160.
http://dx.doi.org/10.1163/187631205788838500Molano, F. & Medina, C.A. (2010) Especie nueva de Scybalocanthon (Coleoptera: Scarabaeinae: Deltochilini) y descripción de la variación del órgano genital masculino. Revista Mexicana de Biodiversidad, 81, 689–699.
Monaghan, M.T., Inward, D. J.G., Hunt, T. & Vogler, A.P. (2007) A molecular phylogenetic analysis of the Scarabaeinae (dung beetles). Molecular Phylogenetics and Evolution, 45, 674–692.
http://dx.doi.org/10.1016/j.ympev.2007.06.009Montreuil, O. (1998) Analyze phylogénétique et paraphylie des Coprini et Dichotomiini (Coleoptera: Scarabaeidae). Scénario biogéographique. Annales de la Société Entomologique de France (N.S.), 34, 135–148.
Ocampo, F.C. & Molano, F. (2011) Revision and Biogeography of the Neotropical dung beetle genus Scybalophagus (Coleoptera: Scarabaeidae). Revista de la Sociedad Entomológica Argentina, 70 (3-4), 231–253.
Ochi, T. & Masahiro, K. (1996) Studies on the coprophagous scarab beetles from East Asia: IV. (Coleoptera, Scarabaeidae). Giornale Italiano di Entomologia, 8, 17–28.
Ochi, T., Masahiro, K. & Kikuta, T. (1997) Studies of the family Scarabaeidae (Coleoptera) from Borneo. The genera Paragymnopleurus and Sisyphus. Giornale Italiano di Entomologia, 8, 239–250.
Pereira, F. (1941) Pinotus de la república de Argentina (Col. Lam. Copridae). Anales de la Sociedad Científica Argentina, 131, 262–267.
Pereira, F.S. & D´Andretta, M.A.V. (1955) The species of Deltochilum of the subgenus Calhyboma Kolbe. Revista Brasileira de Entomologia, 4, 7–50.
Pereira, F.S. & Martínez, A. (1956) Os generos de Canthonini Americanos (Col. Scarabaeidae). Revista Brasiliera Entomologica, 6, 91–192.
Pereira, F. S. & Martínez, A. (1960) Notas escarabeidológicas – II. Revista Brasileira de Entomologia, 9, 37–55.
Philips, T.K., Scholtz, C.H. & Ocampo, F.C. (2002) A phylogenetic analysis of the Eucraniini (Coleoptera: Scarabaeidae: Scarabaeinae). Insects Systematic and Evolution, 33, 241–252.
http://dx.doi.org/10.1163/187631202X00145Philips, K., Pretorius, E. & Scholtz, C.H. (2004) A phylogenetic analysis of dung beetles (Scarabaeinae: Scarbaeidae) unrolling an evolutionary history. Invertebrate Systematics, 18, 53–88.
http://dx.doi.org/10.1071/IS03030Price, D.L. (2005) Descriptions of the male and female genitalia of Phanaeus (MacLeay) (Scarabaeidae: Scarabaeinae): The vindex species group. The Coleopterists Bulletins, 59, 197–203. http://dx.doi.org/10.1649/743
Reid, C.A.M. (2000) A complex of cryptic species in the genus Coptodactyla Burmeister (Coleoptera: Scarabaeidae: Coprini). Memoirs of the Queensland Museum, 46, 231–251.
Reid, C.A.M. & Storey, R.I. (2000) Revision of the dung beetle genus Temnoplectron. Memoirs of the Queensland Museum, 46, 253–297.
Roig-Juñent, S. (2000) The subtribes and genera of the tribe Broscini (Coleoptera, Carabidae): cladistic analysis, taxonomic treatment, and biogeographical considerations. Bulletin of the American Museum of Natural History, 255, 1–90.
http://dx.doi.org/10.1206/0003-0090(2000)255%3C0001:TSAGOT%3E2.0.CO;2Rubenyan, A.A. (2002) A method for making preparations of the endophallus sack in beetles of the family Cerambycidae. Entomological Review, 82, 376–377.
Snodgrass, R.E. (1935) Principles of Insect Morphology. McGraw-Hill, New York, 667 pp.
Stefano, Z. & Ivo, G. (2001) A survey of the Onthophagus (s.l.) species occurring in Syria (Coleoptera Scarabaeidae Scarabaeinae). Memories de lla Societa Entomologica Italiana, 80, 87–105.
Tarasov, S.I. & Kabakov, O.N. (2010) Two new species of Onthophagus (Coleoptera: Scarabaeidae) from Indochina, with a discussion of some problems with the classification of Serrophorus and similar subgenera. Zootaxa, 2344, 17–28.
Tarasov, S.I. & Solodovnikov, A.Y. (2011) Phylogenetic analyses reveal reliable morphological markers to classify mega-diversity in Onthophagini dung beetle (Coleoptera: Scarabaeidae: Scarabaeinae). Cladistics, 27, 1–39.
http://dx.doi.org/10.1111/j.1096-0031.2011.00351.xThompson, R.T. (1998) Revision of the weevil genus Leptostethus Waterhouse, 1853 (Coleoptera, Curculionidae, Entiminae). Cimbebasia Memoir, 7, 1–80.
Vaz-de-Mello, F. (2008) Synopsis of the new subtribe Scatimina (Coleoptera: Scarabaeidae: Scarabaeinae: Ateuchini), with descriptions of twelve new genera and review of Genieridium, new genus. Zootaxa, 1955, 1–75.
Werner, M & Simmons, L.W. (2008) The evolution of male genitalia: functional integration of genital sclerites in the dung beetle Onthophagus taurus. Biological Journal of the Linnean Society, 93, 257–266.
http://dx.doi.org/10.1111/j.1095-8312.2007.00924.xZunino, M. (1972) Revisione delle specie paleartiche del genere Onthophagus. I.Il sottogenere Euonthophagus. Bollettino del Museo di Zoologia dell’Università di Torino, 1, 1–28.
Zunino, M. (1978) La posizione sistematica del “Caccobius (Caccophilus) anomalus” (Coleoptera, Scarabaeoidea). Bollettino del Museo di Zoologia dell”Università di Torino, 3, 9–14.
Zunino, M. (1979) Gruppi artificiali e gruppi naturali negli Onthophagus (Coleoptera Scarabaeoidea. Bollettino del Museo di Zoologia dell’Università di Torino, 1, 1–18.
Zunino, M. (1981) Note su alcuni Onthophagus americani e descrizione di nuove specie (Coleoptera. Scarabaeidae). Bollettino del Museo di Zoologia dell’Università di Torino, 6, 75–86.
Zunino, M. (1983) Essai préliminaire sur 1´evolution des armures genitales des Scarabaeinae, par rapport à la taxonomie du groupe et à 1´évolution du comportement di nidification. Bulletin de la Societé Entomologique de France, 88, 531–542.
Zunino, M. (1985) Las relaciones taxonómicas de los Phanaeina (Coleoptera, Scarabaeinae) y sus implicaciones biogeográficas. Folia Entomologica Mexicana, 64, 101–115.
Zunino, M. & Halffter, G. (1987) Sobre Onthophagus Latreille, 1802 Americanos (Coleoptera: Scarabaeidae: Scarabaeinae). Elytron, 11, 157–178.
Zunino, M. & Halffter, G. (1988) Análisis taxonómico, ecológico y biogreográfico de un grupo americano de Onthophagus. Museo Regionale di Scienze Naturale Torino Monografia, 9, 1–211.