Abstract
The taxonomy and systematics of the genus Digitonthophagus Balthasar (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini) is revised. A detailed study of the male genitalia combined with external morphology suggests that the variability, previously recognized, for D. gazella is hiding a species complex within the Afrotropical region and the Arabian Peninsula. The current study recognizes 16 species; 13 from the Afrotropical region and Arabian Peninsula and three from the eastern portion of the Saharo-Arabian region and the continental Indomalayan region. Species are organized into six species groups based on the results of the morphology-based phylogenetic analysis. The following 12 species are described as new: D. aksumensis Génier new species; D. biflagellatus Génier new species; D. dilatatus Génier new species; D. eucatta Génier new species; D. falciger Génier new species; D. fimator Génier new species; D. namaquensis Génier new species; D. petilus Génier new species; D. sahelicus Moretto new species; D. uks Génier new species; D. ulcerosus Génier new species; and D. viridicollis Génier new species. In order to stabilize nomenclature, lectotypes are designated for Scarabaeus bonasus Fabricius, 1775; Scarabaeus catta Fabricius, 1787, and Onthophagus gazella lusinganus d’Orbigny. A neotype is designated for Scarabaeus dorcas Olivier, 1789 whose status and synonymy need to be altered in order to clarify the status of Scarabaeus gazella auctorum, the widely introduced species with economic importance. A naming scheme is presented for the sclerites of the internal sac. External and male genitalia are illustrated and distribution maps are provided for each species.
References
- A
lfieri, A. (1976) The Coleoptera of Egypt. Mémoires de la Société Entomologique d’Égypte, 5. Atlas Press, Cairo, Egypt, xvi + 362 pp.
Arrow, G.J. (1931) Coleoptera Lamellicornia Part III. (Coprinae). The Fauna of British India, including Ceylon and Burma. Taylor and Francis, London, United Kingdom, 428 pp.
Balthasar, V. (1941) Gli Scarabeidi coprofagi e Trogini dell'Africa Orient. Ital. del museo di Milano. Atti della Società Italiana di Scienze Naturali e del Museo Civico di Storia Naturale di Milano, 80, 103–154.
Balthasar, V. (1956) Scarabaeidae (Coleoptera) der Afghanistan-Expedition (1952 u. 1953) J. Klapperichs. Sborník entomologického Oddelení Národního Musea v Praze, 30, 409–439.
Balthasar, V. (1959) Beitrag zur kenntnis der Gattung Onthophagus Latr. Sborník entomologického Oddelení Národního Musea v Praze, 33, 461–471.
Balthasar, V. (1963) Monographie der Scarabaeidae und Aphodiidae der palaearktischen und orientalischen Region. Coleoptera: Lamellicornia. Band, 2. Coprinae (Onitini, Oniticellini, Onthophagini). Monographie der Scarabaeidae und Aphodiidae der palaearktischen und orientalischen Region, 2. Verlag der Tschechoslowakischen Akademie der Wissenschaften, Prague, Czech Republic, 627 pp., 16 plates.
Balthasar, V. (1969) Contribution à la connaissance de la faune de la Côte-d’Ivoire (J. Decelle, 1961–1964), XXXII. - Coleoptera Scarabaeidae et Aphodiidae. Annales du Musée Royal du Congo Belge, Nouvelle Série in-8°, Sciences Zoologiques, 175, 59–73.
Balthasar, V. (1972) Coleoptera aus Nordostafrika. Scarabaeoidea: Scarabaeidae und Aphodiidae. Notulae Entomologicae, 52, 18–32.
Barbero, E. & Palestrini, C. (2003) Coleoptera Trogidae, Geotrupidae, Hybosoridae, Scarabaeidae (Scarabaeinae and Dynastinae) collected in Australia. Result of the Zoological Missions to Australia of the Regional Museum of Natural Sciences of Turin, Italy. I. Monografia, Museo Regionale di Scienze Naturali, Torino, 35, 303–328.
Barbero, E., Borghesio, L., Dellacasa, G. & Dellacasa, M. (1998) Contribution to the knowledge of Scarabaeidae and Aphodiidae from southern Ethiopia (Coleoptera Scarabaeoidea). Bollettino della Società Entomologica Italiana, 130, 233–254.
Barbero, E., Palestrini, C. & Roggero, A. (2003) Revision of the genus Phalops Erichson, 1848 (Coleoptera: Scarabaeidae: Onthophagini). Monografie XXXVIII, Museo Regionale di Scienze Naturali, Region Piemonte, 38, Museo Regionale di Scienze Naturali, Torino, Italy, 378 pp.
Bertoloni, J. (1849) Illustratio rerum naturalium Mozambici. Dissertatio II. de Coleopteris. Novi Commentarii Academiae Scientiarum Instituti Bononiensis, 10. Instituti Bononiensis, Bologna, Italy, pp. 407–422.
Blume, R.R. & Aga, A. (1975) Onthophagus gazella: mass rearing and laboratory biology. Experimental Entomology, 4, 735–736.
https://doi.org/10.1093/ee/4.5.735
Blume, R.R. & Aga, A. (1978) Onthophagus gazella F.: progress of experimental release in south Texas. Folia Entomológica Mexicana, 38–39, 190–191.
Bornemissza, G.F. (1970) Insectary studies on the control of dung breeding flies by the activity of the dung beetle, Onthophagus gazella F. (Coleoptera: Scarabaeinae). Journal of the Australian Entomological Society, 9, 31–41.
https://doi.org/10.1111/j.1440-6055.1970.tb00767.x
Bornemissza, G.F. (1976) The Australian dung beetle project 1965–1975. Australian Meat Research Commettee, 30, 1–30.
Boucomont, A. (1914) Onthophagus asiatiques nouveaux ou peu connus. Annali del Museo civico di storia naturale di Genova Serie 3.a, Volune 6, 46, 210–243.
Boucomont, A. (1921) [species treatment] In: Boucomont, A. & Gillet, J.J.E. (Eds.), Fam. Scarabaeidae, Laparosticti (Coléoptères). Faune entomologique de l’Indochine française. Fascicule 4, 1–76.
Boucomont, A. (1923) Scarabéides recueillis par le Marquis S. Patrizi dans l'Afrique Orientale Tropicale. Annali del Museo civico di storia naturale Giacomo Doria, 51, 86–100.
Boucomont, A. (1934) Mission J. de Lépiney au Soudan français 1933-34 (Quatrième note). Bulletin de la Société des Sciences Naturelles du Maroc, 14, 25–27.
Boucomont, A. (1935) Coleoptera. VI. Scarabaeidae Scarabaeini. In: Jeannel, R. (Ed.), Mission Scientifique de l’Omo. Tome 2 Zoologie, Fascicule 16. Paul Chevalier & Fils, Paris, France, pp. 279–290.
Boucomont, A. (1937) Coprinae. In: Burgeon, L. (Ed.), Liste des Coléoptères récoltés au cours de la mission belge au Ruwenzori. Institut Royal Colonial Belge, Section Sciences Naturelles et Médicales. Collection in-8º, 6, 125–133.
Bousquet, Y. (2016) Litteratura Coleopterologica (1758–1900): a guide to selected books related to the taxonomy of Coleoptera with publication dates and notes. ZooKeys, 583, 1–776.
https://doi.org/10.3897/zookeys.583.7084
Brancsik, C. (1893) Beiträge zur Kenntnis Nossibés und dessen Fauna nach Sendungen und Mittheilungen des Herrn P. Frey. Jahresheft des naturwissenschaftlichen Vereines des Trencséner Comitates, 15–16, 202–258.
Breeschoten, T., Doorenweerd, C., Tarasov, S. & Vogler, A.P. (2016) Phylogenetics and biogeography of the dung beetle genus Onthophagus inferred from mitochondrial genomes. Molecular Phylogenetics and Evolution, 105, 86–95.
https://doi.org/10.1016/j.ympev.2016.08.016
Bremer, K. (1994) Branch support and tree stability. Cladistic, 10, 295–304.
https://doi.org/10.1111/j.1096-0031.1994.tb00179.x
Cabral-de-Mello, D.C., Gomes de Oliveira, S., Ramos, I.C. & Cássia de Moura, R. de (2008) Karyotype differentiation patterns in species of the subfamily Scarabaeinae (Scarabaeidae, Coleoptera). Micron, 39, 1243–1250.
https://doi.org/10.1016/j.micron.2008.04.002
Cambefort, Y. (1972) Scarabaeinae récoltés en Mauritanie par J.-L. Amiet. Nouvelle Revue d’Entomologie, 2, 243–251.
Cambefort, Y. (2006) Des coléoptères des collection & des hommes. Muséum national d’Histoire naturelle, Paris, France, 375 pp.
Cambefort, Y. & Bordat, P. (2003) Coléoptères Scarabaeidae s. str., Aphodiidae et Ceratocanthinae du mont Nimba et des régions limitrophes. In: Lamotte, M. & Roy, R. (Eds.), Le peuplement animal du mont Nimba (Guinée, Côte d’Ivoire, Liberia). Mémoires du Muséum national d’Histoire naturelle, 190, 551–580.
Davis, A.L.V. & Scholtz, C.H. (2004) Local and regional species ranges of a dung beetle assemblage from the semi-arid Karoo/Kalahari margins, South Africa. Journal of Arid Environments 57, 61–85.
https://doi.org/10.1016/S0140-1963(03)00094-6
Dechambre, R.-P. (2002) Néalloptype ou Alloréférent : de l’utilité de ce concept et son bon usage. Le Coléopteriste, 43, 163–164
Dejean, P.F.M.A. (1821) Catalogue de la collection de coléoptères de M. le Baron Dejean. Crevot, Paris, France, viii + 136 + [2] pp.
Dejean, P.F.M.A. (1836) Catalogue des coléoptères de la collection de M. le Comte Dejean. Troisième édition, revue, corrigée et augmentée. Méquignon-Marvis Père et Fils, Paris, France, xiv + 503 pp.
Deloya, C. (2000) Escarabajos exóticos (Coleoptera: Scarabaeidae) para la fauna de los estados de Morelos y Oaxaca, Mexico. Folia Entomológica Mexicana, 108, 125–126.
d’Orbigny, H. (1900a) Synopsis des Onthophagides paléarctiques. L’Abeille, 29, 117–254.
d’Orbigny, H. (1900b) Supplément au synopsis des Onthophagides paléarctiques. L’Abeille, 29, 289–300.
d’Orbigny, H. (1904) Onthophagides africains de la collection du Musée civique de Gêne. Annali del Museo civico di storia naturale di Genova Serie 3.a, Volume 1, 41, 253–331.
d’Orbigny, H. (1905a) Onthophagides provenant du voyage de M. Alluaud dans l’Afrique orientale et description d’autres espèces africaines nouvelles. Annales de la Société Entomologique de France, 74, 381–536.
d’Orbigny, H. (1905b) Onthophagides provenant du voyage de M. L. Fea dans l’Afrique occidentale. Annali del Museo civico di storia naturale di Genova Serie 3.a, Volume 1, 41, 417–448.
d’Orbigny, H. (1909) Collections recueillis dans le Sahara et les régions voisines par M. René Chudeau. Onthophagides (déterminés par M. H. d'Orbigny). Bulletin du Muséum National d’Histoire Naturelle, 15, 523–525.
d’Orbigny, H. (1911) Mission Niger-Tchad, dirigée par M. le Capitaine Tilho. Collections recueillis par M. le Dr Gaillard : Coléoptères. Onthophagides. Bulletin du Muséum National d’Histoire Naturelle, 17, 17–21.
d’Orbigny, H. (1913a) Synopsis des Onthophagides d’Afrique. Annales de la Société Entomologique de France, 82, 1–608.
d’Orbigny, H. (1913b) Insectes Coléoptères II. Scarabaeidae Onthophagini. In: Voyage de Ch. Alluaud et R. Jeannel en Afrique Orientale (1911–1912). Résultats scientifiques. Librairie Albert Schulz, Paris, pp. 65–128.
d’Orbigny, H. (1914) Synopsis des Onthophagides d’Afrique. Catalogue bibliographique et synonymique des Onthophagides d’Afrique. Annales de la Société Entomologique de France, 82, 609–742.
d’Orbigny, H. (1916) Insectes coléoptères Scarabaeidae Onthophagini et Oniticellini. In: Voyage de M. Guy Babault dans l’Afrique orientale anglaise. Résultats scientifiques. Paris, France, pp. 1–33, 3 plates.
Edmonds, W.D. (1972) Comparative skeletal morphology, systematics and evolution of the phanaeine dung beetles (Coleoptera: Scarabaeidae). University of Kansas Science Bulletin, 49 (11), 731–874.
Endrödi, S. (1971) The scientific results of the Hungarian zoological expeditions to Tanganyika. 16. Coleoptera Lamellicornia (partim). Folia Entomologica Hungarira, Series Nova, 24 (26), 289–311.
Endrödi, S. (1973a) Zoological collecting trips of the Hungarian Natural History Museum in North Vietnam (D. R. V.). No. 4. Neue Arten aus der Gattung Onthophagus Latr. Acta Musei Silesiae, Series A, 22, 119–125.
Endrödi, S. (1973b) Entomological explorations in Ghana by Dr. S. Endrödy-Younga. 14. Lamellicornia (Coleoptera), I. Annales Historico-Naturales Musei Nationalis Hungarici, 65, 195–229.
Endrödi, S. (1974) The scientific results of Dr. Gy. Topál's collecting in India. Nr. 5. Sammelergebnisse aus der Superfamilie Lamellicornia (Coleoptera). Acta Musei Silesiae, Series A, 23, 1–5.
Endrödi, S. (1976) Lamellicornia (Coleoptera) aus Ghana, II. Annales Historico-Naturales Musei Nationalis Hungarici, 68, 155–164.
Erichson, W.F. (1843) Beitrag zur Insecten-Fauna von Angola, in besonder Beziehung zur geographischen Verbreitung der Insecten Afrika. Archiv für Naturgeschichte, 9, 199–267.
Evenhuis, N.L. (1997) Litteratura taxonomica dipterorum (1758–1930); being a selected list of the books and prints of Diptera taxonomy from the beginning of Linnean zoological nomenclature to the end of the year 1930; containing information on the biographies, bibliographies, types, collections, and patronymic genera of the authors listed in this work; including detailed information on publication dates, original and subsequent editions, and other ancillary data concerning the publications listed herein. Volume I: A–K. Backhuys Publishers, Leiden, Netherlands, vii + 426 pp.
Evenhuis, N. (2007) The insects and spider collections of the world website. Available from: hbs.bishopmuseum.org/codens/codens-r-us.html (accessed 15 July 2016)
Fabricius, J.C. (1775) Systema Entomologiae sistens Insectorum Classes, Ordines, Genera, Species adiectis Synonymis, Locis Descriptionibus, Observationibus. Officina Libraria Kortii, Flensburg & Leipzig, Germany, xxxii + 832 pp.
Fabricius, J.C. (1781) Species insectorum exhibens eorum differentias specificas, synonyma auctorum, loca natalia, metamorphosis, adiecitis observationibus, descriptionibus. Tomus I. Carol Ernest Bohnii, Hamburg & Kiel, Germany, viii + 552 pp.
Fabricius, J.C. (1787a) Mantissa insectorum sistens eorum species nuper detectas adiectis characteribus generici, differentiis specificis, emendationibus, observationibus. Tom. I. Christ. Gottl. Proft., Copenhagen, Denmark, xx + 348 pp.
https://doi.org/10.5962/bhl.title.36471
Fabricius, J.C. (1787b) Mantissa insectorum sistens species nuper detectas adiectis synonymis, observationibus, descriptionibus, emendationibus. Tom. II. Christ. Gottl. Proft., Copenhagen, Denmark, [2] + 382 pp.
Fabricius, J.C. (1792) Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species adjictis synonimis, locis, observationibus, descriptionibus. Tom. I. Christ. Gottl. Proft., Copenhagen, Denmark, xx + 330 pp.
Fabricius, J.C. (1798) Supplementum entomologicae systematicae. Proft et Storch, Copenhagen, Denmark, [4] + 572 pp.
Fabricius, J.C. (1801) Systema Eleutheratorum secundum ordines, genera, species: adiectis synonymis, locis, observationibus, descriptionibus. Tomus I. Bibliopoli academici novi, Kiel, Germany, xxiv + 506 pp.
Fåhraeus, O.I. (1857) [new taxa]. In: Boheman, C.H. (Ed.), Insecta Caffrariae annis 1838–1845 a J. A. Wahlberg collecta amici auxilio suffultus descripsit. Pars II. Coleoptera (Scarabaeidae). Officina Norstedtiana, Stockholm, Sweden, 395 pp.
Fairmaire, L. (1893) Quelques coléoptères de la République Argentine. Annales de la Société Entomologique de Belgique, 37, 9–50.
Ferreira, M.C. (1958) Coleoptera (Scarabaeidae): Coprinae. In: South African Animal Life. Results of the Lund University Expedition in 1950–1951, 5. Almqvist & Wiksells Boktryckeri AB, Uppsala, Sweden, pp. 479–516.
Ferreira, M.C. (1959) A contribution to the study of the Coleoptera from the Kalahari Gemsbok National Park (Scarabaeidae, Buprestidae, Cerambycidae). Koedoe, Journal for Scientific Research in the National Parks of South Africa, 2, 77–86.
https://doi.org/10.4102/koedoe.v2i1.856
Ferreira, M.C. (1962) Contribuição para o conhecimento dos Escarabídeos de Angola. Publicações Culturais da Companhia de Diamentes de Angola-Diamag, 54, 121–190.
Ferreira, M.C. (1967) Catálogo dos Coleópteros de Angola. Revista de Entomologia de Moçambique, 8, 417–1317.
Ferreira, M.C. (1968) Os escarabídeos de Moçambique. I. Revista de Entomologia de Moçambique, 10, 5–778.
Ferreira, M.C. (1972) Os escarabídeos de África (sul do Sáara). I. Revista de Entomologia de Moçambique, 11, 5–1088.
Fincher, G.T., Blume, R.R., Hunter III, J.S. & Beerwinkle, K.R. (1986) Seasonal distribution and diel flight activity of dung-feeding scarabs in open and wooded pasture in east-central Texas. The Southwestern Entomologist, supplement, 10, 1–35.
Frey, G. (1956) Liste der von S. Nathan in der Wüste Thar im Jahre 1955 gesammelten Lamellicorniern mit der Beschreibung einer neuen Catharsius-Art (Col.). Entomologische Arbeiten aus dem Museum G. Frey, Tutzing Bei München, 7, 654–656.
Frey, G. (1958) Onthophagini (Coleoptera Polyphaga) (Fam. Scarabaeidae). Exploration du Parc National de l’Upemba, I. Mission G.F. de Witte en collaboration avec W. Adam, A. Janssens, L. van Meel et R. Verheyen (1946–1949), 54, 69–90.
Frey, G. (1961) Onthophagini (Coleoptera Lamellicornia). Exploration du Parc National de la Garamba, Mission H. de Saeger en collaboration avec P. Baert, G. Demoulin, I. Denisoff, J. Martin, M. Micha, A. Noirfalise, P. Schoemaker, G. Troupin et J. Verschuren (1949–1952), 21, 69–98.
Gahan, C.J. & Arrow, G.J. (1900) On a collection of insects and arachnids made in 1895 and 1897 by Mr. C. V. A. Peel in Somaliland, with descriptions of new species. Proceedings of the Zoological Society of London, 1900, 21–25.
Gaikwad, A.R. & Bhawane, G.P. (2015) Study of nidification behavior in three dung beetle species (Scarabaeidae: Scarabaeinae) from south-western Maharashtra, India. International Journal of Science and Research, 4, 1538–1542.
Génier, F. (2013) Transfert d’Onthophagus bubalus Harold, 1867, dans le genre Phalops Erichson, 1847, et notes sur sa position phylogénétique (Coleoptera, Scarabaeidae, Scarabaeinae). Catharsius, La Revue, 7, 1–4.
Génier, F. & Davis, A.L.V. (2017) Digitonthophagus gazella auctorum: an unfortunate case of mistaken identity for a widely introduced species (Coleoptera: Scarabaeidae: Scarabaeinae: Onthophagini). Zootaxa, 4221 (4), 497–500.
https://doi.org/10.11646/zootaxa.4221.4.8
Génier, F. & Krell, F.T. (2017) Case 3722. Scarabaeus gazella Fabricius (currently Digitonthophagus gazella or Onthophagus gazella; Insecta, Coleoptera, SCARABAEIDAE): proposed conservation of usage of the specific name by designation of a neotype. Bulletin of Zoological Nomenclature, in press.
Gerstaecker, A. (1871) Beitrag zur Insektenfauna von Zanzibar III. Coleoptera. Archiv für Naturgeschichte, 37, 42–86.
Gerstaecker, C.E.A. (1873) Gliederthier-Fauna des Sansibar-Gebietes (Insekten, Arachniden, Myriopoden und Isopoden). Baron Carl Claus von der Decken’s Reisen in Ost-Afrika in den Jahren 1859–1865. Bd 3 Abt. 2. Wissenschaftliche-Ergebnisse, Adt. 2, C.F. Winter, Liepzig & Heidelberg, Germany, xii + 542 + [2] pp.
Gillet J.J.E. & Boucomont A. (1927) Scarabaeidae: Coprinae II, Termitotroginae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterorum Catalogus. Partes 38 et 90, 38, 90. W. Junk, Berlin, Germany, pp. 101–264.
Gmelin, J.F. (1790) Caroli a Linné, systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, loci. Editio decima tertia, aucta, reformata. Tom. I, Pars IV. Classis V. Insecta. G.E. Beer, Leipzig, Germany, pp. 1517–2224.
Goeze, J.A.E. (1777) Entomologische Beyträge zu des Ritter Linné zwölften Ausgabe des Natursystems. Erster Teil. Weidmanns Erben und Reich, Leipzig, Germany, xvi + 736 pp.
https://doi.org/10.5962/bhl.title.45974
Goloboff, P.A. Farris, J.S. & Nixon, K. (2003) TNT: tree analysis using new technology. Version 1.0, Beta test v. 0.2. Program and documentation. Available from: www.lillo.org.ar/phylogeny/tnt (accessed 1 February 2017)
Gomes Alves, M.L. (1956) Entomofauna da Guiné portuguesa e S. Tomé e Príncipe (Contribuição para o seu conhecimento), Coleópteros (Lucanidae, Scarabaeidae, Cicindelidae, Buprestidae). Boletim Cultural da Guiné Portuguesa, 44, 87–105.
Gomes Alves, M.L.M. (1944) Contribuições para o estudo dos Onthophagus de Angola e Moçambique. Memórias e Estudos do Museu Zoológico da Universidade de Coimbra, 157, 1–20.
Gutierrez, J., Macqueen, A. & Brun, L.O. (1988) Essais d'introduction de quatre espèces de bousiers Scarabaeinae en Nouvelle Calédonie et au Vanuatu. Acta Œcologica, 9, 39–53.
Harold, E. von (1867) Beiträge zur Kenntniss der Gattung Onthophagus. Coleopterologische Hefte, 2, 23–59.
Harold, E. von (1869) Tom. IV. Scarabaeidae. In: Gemminger. M. & Harold, E. v. (Eds.), Catalogus Coleopterorum hucusque descriptorum synonymicus et systematicus autoribus Dr. Gemminger et B. de Harold. E.H. Gummi, Munich, Germany, pp. 979–1346, [1347–1353].
Harold, E. von (1871) Berichtigungen und Zusätze zum Catalogus Coleopterorum synonymicus et systematicus. Coleopterologische Hefte, 8, 117–123.
Herbst, J.F.W. (1786) Erste Mantisse zum Verzeichniss der ersten Klasse meiner Insektensammlung. Archiv der Insectengeschichte, 7–8, 153–182, plates 43–48.
Herbst, J.F.W. (1789) Natursystem aller bekannten in- und ausländischen Insekten, als eine Fortsetzung der von Büffonschen Naturgeschichte. Nach dem System des Ritters von Linné und Fabricius zu bearbeiten angefangen von Carl Gustav Jablonsky von Johan Friedrich Wilhem Herbst. Der Käfer zweyter Theil. Joachim Pauli, Berlin, Germany, lxiv + 330 + [6] pp., plates 7–20.
Hope, F.W. (1837) The Coleopterist’s Manual, containing the lamellicorn insect of Linneus and Fabricius. Henry G. Bohn, London, United Kingdom, xiii + [2] + 15–121 + [4] pp., 3 plates.
International Commission on Zoological Nomenclature (1999) International Code of Zoological Nomenclature. Fourth Edition. The International Trust for Zoological Nomenclature, London, United Kingdom. 306 pp.
Illiger, J.C.W. (1800) Olivier’s Entomologie oder Naturgeschichte der Insekten mit ihren Gattungs- und ArtMerkmalen, ihrer Beschreibung und Synonymie. Käfer. Uebersetzt und mit Zusätzen und Anmerkungen durchgängig begleitet. Erster Theil. Karl Reichard, Braunschweig, Germany, xvi + 309 pp., 2 plates.
Illiger, K. (1804) Zusätze, Berichtigungen und Bemerkungen zu Fabricii. Systema Eleutheratorum. Tomus II. Magazin für Insektenkunde, 3, 146–180.
Ivie, M.A. & Philips, T.K. (2008) Three new species of Canthonella Chapin from Hispaniola, with new records and nomenclatural changes for West Indian dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). Zootaxa, 1701, 1–14.
Janssens, A. (1953) Contribution à l’étude de la zone d’inondation du Niger (mission G. Remaudière, 1950). V - Coléoptères Scarabaeinae. Bulletin de l'Institut Français de l’Afrique Noire, 15, 1548–1551.
Kabakov, O.N. (1979) Obzor plastichnatoystykh zhukov podsemeystva Coprinae (Scarabaeidae, Coleoptera) Dalnego Vostoka i Sopredelnykh territor. In: Krivolutskaya, G.O. (Ed.), Zhuki Dal’nego Vostoka i vostochnoi Sibirii. Akademiya Nauk SSSR, Vladivostok, Russia, pp. 58–98.
Kim, J.I. & Lumaret, J.-P. (1989) Les larves du genre Onthophagus Latreille, 1802. 5 – Les espèces des sous-genres Digitonthophagus, Trichonthophagus et « Incetae Sedis » (Coleoptera Scarabaeidae). Bulletin Mensuel de la Société Linnéenne de Lyon, 58, 335-344.
Klug, J.C.F. (1833) Bericht über eine auf Madagascar veranstaltete Sammlung von Insecten aus der Ordnung Coleoptera. Eine in der Königl. Akademie der Wissenschaften am 29. März 1832 gelesene Abhandlung. Mit fünf illuminirten Tafeln. Königlichen Akademie der Wissenschaften, Berlin, Germany. 135 pp, 5 plates.
Klug, J.C.F. (1835) Insekten. In: Erman, A. (Ed.), Reise um die Erde durch Nord-Asien und die beiden Oceane, in den Jahren 1828, 1829 und 1830 ausgeführt von Adolph Erman. Naturhistorischer Atlas. Reimer,G., Berlin, Germany, pp. 27–52, plates xv–xvi.
Klug, J.C.F. (1862) Insecten und Myriapoden. Bearbeitet in verbindung mit Klug, Loew, Schaum, hagen, Gerstaecker, Hopffer mit fünf und dreissig kupfertalfeln. In: Peters, W.C.H., (Ed.), Naturwissenschaftliche Reise nach Mossambique auf Befehl Seiner Majestät des Königs Friedrich Wilhelm IV in den Jahren 1842 bis 1848 ausgeführt. Zoologie V. Georg Reimer, Berlin, Germany, pp. 143–267.
Kohlmann, B. (1994) A preliminary study of the invasion and dispersal of Digitonthophagus gazella (Fabricius, 1787) in Mexico (Coleoptera: Scarabaeidae: Scarabaeinae). Acta Zoologica Mexicana (n.s.), 61, 35–42.
Kolbe, H.J. (1887) Beiträge zur Zoogeographie Westafrikas nebst einem Bericht über die während der Loango-Expedition von Herrn Dr. Falkenstein bei Chinchoxo gesammelten Coleoptera. Nova Acta Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae Curiosorum, 50, 153–364.
Kolbe, H.J. (1897a) Käfer und Netzflügler Ost-Afrikas. In: K. Möbius (Ed.), Deutsch-Ost-Afrikas, 4 (4), Reimer, D., Berlin, Germany, 367 + 42 pp., 5 plates.
Kolbe, H.J. (1897b) Über die von Hern Dr. F. Stuhlmann in Deutsch-Ostafrika und Mosambik während der Jahre 1888 bis 1890 gesammelten Coleopteren. Mitteilungen aus dem Naturhistorischen Museum in Hamburg, 14, 73–101.
Kolbe, H.J. (1908) Dynastidae, Cetoniidae und Scarabaeidae. In: Schultze L. (Ed.), Zoologische und Arthropologische ergebnisse einer forschungsreise im westlichen und zentralen südafrika, 1. Gustav Fischer, Jena, Germany, pp. 121–132.
Kolbe, H.J. (1914) Lieferung 3: Scarabaeiden. In: Shubotz, H. (Ed.), Wissenschaftliches Ergebnisse der Deutschen Zentral-Afrika-Expedition 1907 unter Führung Adolf Friedrichs, Herzog zu Mecklenburg. Zoologie III, 5. Klinkhardt & Biermann, Leipzig, Germany, pp. 261–416.
Lansberge, J.W. van (1882) Énumération des scarabaeides rapportés du pays des Somalis (Afrique Équatoriale) par M. Révoil. Avec diagnoses des espèces nouvelles. Comptes-Rendus des Séances de la Société Entomologique de Belgique, 26, xxi–xxxi.
Laporte, F. (Comte de Castelnau) (1840) Histoire Naturelle des Insectes Coléoptères. Tome deuxième. Histoire Naturelle des animaux articulées, crustacés, arachnides, myriapode et insectes. Tome troisième, 2, P. Duménil, Paris, France, 564 pp.
Latreille, P.A. (1827) Description des Insectes Figurés sur la pl. LVIII du tom. II, recueillis par M. Cailliaud, et décrits par M. Latreille, membre de l’Académie royale des sciences, Etc. In: Voyage à Méroé, au Fleuve Blanc, au-delà de Fazoql dans le midi du royaume de Sennâr, à Syouah et dans cinq autres oasis; fait dans les années 1819, 1820, 1821, et 1822, par M. Frédéric Cailliaud de Nantes, 4. Imprimerie Royale, Deleure, Paris, France, pp. 271–292.
LePeletier de Saint Fargeau, A.L.M. & Audinet-Serville, J.G. de (1828) Scarabé, Scarabaeus. In: Encyclopédie méthodique, ou par ordre de matières; par une société de gens de lettres, de savant et d’artistes; précédé d’un vocabulaire universel, servans de table pour tout l’ouvrage, ornée des portraits de Mm. Diderot & d’Alembert, premiers éditeurs de l’Encyclopédie. Tome dixième. Pars 2. Mme. veuve Agasse, Paris, France, pp. 345–832.
MacRae, T.C. & Penn, S.R. (2001) Additional records of adventive Onthophagus Latreille (Coleoptera: Scarabaeidae) in North America. The Coleopterists Bulletin, 55, 49–50.
https://doi.org/10.1649/0010-065X(2001)055[0049:AROAOL]2.0.CO;2
Maddison, W.P. & Maddison, D.R. (2009) Mesquite: a modular system for evolutionary analysis. Version 3.10. Available from: mesquiteproject.org (accessed 1 August 2016)
Masumoto, K. (1988) Coprophagid-beetles from Northwest Thailand (II). The Entomological Review of Japan, 43, 135–143.
Mlambo, S., Sole, C.L. & Scholtz, C.H. (2015) A molecular phylogeny of the African Scarabaeinae (Coleoptera: Scarabaeidae). Arthropod Systematics & Phylogeny, 73, 303–321.
Monaghan, M.T., Inward, D.J.G., Hunt, T. & Vogler, A. (2007) A molecular phylogenetic analysis of the Scarabaeinae (dung beetles). Molecular Phylogenetics and Evolution, 45, 674–692.
https://doi.org/10.1016/j.ympev.2007.06.009
Moretto, P. & Bordat, P. (2007) Les scarabéides coprophages du Parc National du Niokolo-Koba au Sénégal (Coleoptera, Scarabaeoidea). Nouvelle Revue d’Entomologie (N.S.), 23, 115–145.
Motschulsky, V. de (1860) Sur les Collections Coléoptèrologiques de Linné et de Fabricius (Continuation.). Études Entomologiques, rédigées par Victor de Motschulsky, 8, 147–168.
Motschulsky, V. de (1863) Essaie d’un catalogue des insectes de l’Ile Ceylan. Bulletin de la Société Impériale des Naturalistes de Moscou, 36, 421–532.
Müller, G. (1942) Coleotterie del Benadir (Somalia) raccolti dal Prof. Giuseppe Russo. Bollettino del Laboratorio di Zoologia generale e agraria della Facoltà Agraria in Portici, 32, 70–103.
Naskrecki, P. (2008) Mantis v. 2.0 - A Manager of Taxonomic Information and Specimens. Available from: http://140.247.119.225/Mantis/index.htm (accessed 14 January 2015)
Nixon, K.C. (2002) WinClada, Version 1.00.08. Published by the author, Ithaca, New York, United States of America.
Noriega, J.A. (2002) First report of the presence of the genus Digitonthophagus (Coleoptera: Scarabaeidae) in Colombia. Primera observación sobre la presencia del género Digitonthophagus (Coleoptera: Scarabaeidae) en Colombia. Caldasia, 24, 213–215.
Noriega J.A., Solis C., Quintero I., Perez L.G., Garcia H.G. & Ospino D.A. (2006) Registro continental de Digitonthophagus gazella (Coleoptera: Scarabaeidae) en Colombia. First report of Digitonthophagus gazella (Coleoptera: Scarabaeidae) in the continental area of Colombia. Caldasia, 28, 379–381.
Noriega, J.A., Horgan, F.G., Larsen, T.H. & Valencia, G. (2010) Records of an invasive dung beetle species, Digitonthophagus gazella (Fabricius, 1787) (Coleoptera: Scarabaeidae), in Peru. Acta Zoologica Mexicana (n.s.), 26, 451–456.
Noriega J.A., Moreno J. & Otavo S. (2011) Quince años del arribo del escarabajo coprófago Digitonthophagus gazella (Fabricius, 1787) (Coleoptera: Scarabaeidae) a Colombia: proceso de invasión y posibles efectos de su establecimiento. Biota Colombiana, 12, 35–43.
Ochi, T. (2003) Studies on the coprophagous scarab beetles from East Asia. VII Descriptions of two new subgenera of the genus Onthophagus (Co/eoptera, Scarabaeidae). Giornale Italiano di Entomologia, 10, 259–274.
Olivier, A.G. (1789) Entomologie, ou histoire naturelle des insectes, avec leurs caractères génériques et spécifiques, leur descriptions, leur synonymie, et leur figure enluminée. Coléoptères. Tome premier. Beaudoin, Paris, France, xx + 497 pp.
https://doi.org/10.5962/bhl.title.61905
Pallas, P.S. (1781) Icones Insectorum praesertim Rossiae Sibiriaeque peculiarium quae collegit et descriptionibus illustravit. Wolfgangi Waltheri, Erlangen, Germany, [6] + 104 pp, 8 plates.
Paulian, R. (1936) Faune entomologique de Madagascar. Coleoptera Lamellicornia, Scarabaeitae, Scarabaeini et Aphodiini. Bulletin de l’Académie Malgache, Nouvelle série, 18, 121–144.
Paulian, R. (1937) Contribution à l’étude de la faune entomologique de L’Angola, Mission scientifique suisse en Angola. Coléoptères Lamellicornes Laparoscticti. Memórias e Estudos do Museu Zoológico da Universidade de Coimbra, 1 (105), 1–41.
Paulian, R. (1945) Coléoptères Scarabéides de l’Indochine. Faune de l’empire Français, 3. Librairie Larose, Paris, France, 227 pp.
Paulian, R. (1948) 12. Coleoptera, Scarabaeidae: Troginae, Geotrupinae, Dynamopinae, Hybosoridae, Coprinae, Aphodiinae. British Museum (Natural History) Expedition to South-Est Arabia 1937-38, Volume 1 (9–15). British Museum, London, United Kingdom, pp. 141–155.
Paulian, R. (1986) Catalogue des coléoptères Scarabaeidae de Madagascar. Bulletin de l’Académie Malgache, 62, 89–111.
Péringuey, L. (1892) Third contribution to the South-African coleopterous fauna. The Transactions of the South African Philosophical Society, 6 (2), 1–94.
Péringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa (Lucanidae and Scarabaeidae). Transactions of the South African Philosophical Society, 12, 1–563, 9 plates.
Petrovitz, R. (1955) The 3rd Danish expedition to central Asia. Zoological resuts 18. Scarabaeidae (Insecta) aus Afganistan. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening i København, 117, 275–287.
Petrovitz, R. (1961) Zoologische Ergebnisse der Österreichischen Karakorum-Expedition 1958. II. Teil. Coleoptera: Scarabaeidae. Sitzungsberichte der Österreichischen Akademie der Wissenschaften. Mathematisch-naturwissenschaftliche Klasse, Abtelung 1, 170, 101–118.
Quedenfeldt, G. (1884) Verzeichniss der von Herrn Major a. D. von Mechow in Angola und am Quango-strom 1878–1881 gesammelten Pectinicornen und Lamellicornen. Berliner Entomologische Zeitschrift, 28, 265–340.
https://doi.org/10.1002/mmnd.18840280207
Reiche, L.J. (1840) Notes sur deux espèces d’Ontophages. Revue Zoologique, 1840, 243–244.
Reitter, E. (1892) Bestimmungs-Tabelle der Lucaniden und coprophagen Lamellicornen des palaeactischen Faunengebietes. Edmund Reitter, Brno, Czech Republic, 230 pp.
Reitter, E. (1893) Bestimmungs-Tabelle der Lucaniden und coprophagen Lamellicornen des palaeactischen Faunengebietes. Verhandlungen des Naturforschenden Vereins, 31, 1–230.
Ripa, S.R., Rojas, P.S. & Velasco, G. (1995) Releases of biological agents of insect pests on Easter Island (Pacific Ocean). Entomophaga, 40, 427–440.
https://doi.org/10.1007/BF02373730
Roggero, A., Barbero, E. & Palestrini, C. (2016a) Revised classification and phylogeny of an Afrotropical species group based on molecular and morphological data, with the description of a new genus (Coleoptera: Scarabaeidae: Onthophagini). Organisms Diversity & Evolution, 2016, [1–18].
https://doi.org/10.1007/s13127-016-0297-z
Roggero, A., Dierkens, M., Barbero, E. & Palestrini, C. (2016b) Combined phylogenetic analysis of two new Afrotropical genera of Onthophagini (Coleoptera, Scarabaeidae). Zoological Journal of the Linnean Society, 2016, 1–23.
https://doi.org/10.1111/zoj.12498
Rougon, C. & Rougon, D. (1980) Contibution à la biologie des Coléoptères coprophages en région sahélienne. Étude du développement d’Onthophagus gazella (Coleoptera : Scarabaeidae). Revue d’Écologie et de Biologie du Sol, 17, 379–392.
Rougon, C. & Rougon, D. (1981) Description de la larve d'Onthophagus gazella Fabricius (Col. Scarabaeidae). Bulletin de la Société entomologique de France, 86, 6–13.
Rougon, D. & Rougon, C. (1983) Nidification des Scarabaeinae et cleptoparasitisme des Aphodiidae en zone sahélienne (Niger). Bulletin de la Société entomologique de France, 88, 496–513.
Roy L., Bon M.-C., Cesarini C., Srin, J. & Bonato, O. (2015) Pinpointing the level of isolation between two cryptic species sharing the same microhabitat: a case study with a scarabeid species complex. Zoologica Scripta, 45, 407–420.
https://doi.org/10.1111/zsc.12166
Schönherr, C.J. (1806) Synonymia insectorum, ode: Versuch einer Synonymie aller bisher bekannten Insecten; nach Fabricii Systema Eleutheratorum geordnet, mit Berichtigungen und Anmerkungen, wie auch Beschreibungen neuer Arten und illuminirten Kupfern. Erter Band. Eleutherata oder Käfer. Erster Theil. Lethrus ... Scolytes. A. Nordström, Stockholm, Sweden, xxii + 293 pp., 3 plates.
Schröter, J.S. (1776) Abhandlungen über verschiedene Gegenstände der Naturgeschichte. J. Just. Gebauers Witwe und J. Jac. Gebauer, Halle, Germany, 488 pp.
https://doi.org/10.5962/bhl.title.36902
Shorthouse, D.P. (2010) SimpleMappr, an online tool to produce publication-quality point maps. Avaliable from: http:// www.simplemappr.net (Accessed 5 Oct. 2016)
Sturm, J. (1826) Catalog meiner Insecten-Sammlung. Erster Theil. Käfer. J. Sturm, Nuremberg, Germany, viii + 207 pp., 4 plates.
Sturm, J. (1843) Catalog der Kaefer-Sammlung von Jacob Sturm. Jacob Sturm, Nurenberg, Germany, xii + 386 pp., 6 plates.
Tarasov, S. & Génier, F. (2015) Innovative Bayesian and parsimony phylogeny of dung beetles (Coleoptera, Scarabaeidae, Scarabaeinae) enhanced by ontology-based partitioning of morphological characters. PLoS ONE, 10 (3), e0116671.
https://doi.org/10.1371/journal.pone.0116671
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 beetles (Coleoptera: Scarabaeidae: Scarabaeinae). Cladistics, 27, 1–39.
https://doi.org/10.1111/j.1096-0031.2011.00351.x
Veenakumari, K. & Veeresh, G.K. (1996) Some aspects of the reproductive biology of Onthophagus gazella (F) and Onthophagus rectecornutus (Lansb) (Coleoptera, Scarabaeidae). Journal of Bombay Natural History Society, 93, 252–256.
Wallengren, H.D.J. (1881) Coleoptera Transvaaliensia. Bidrag till kännedora om Transvaal-Landets I S. Africa Coleopter-fauna. Entomologisk Tidskrift, 2, 9–22.
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.
https://doi.org/10.1111/j.1095-8312.2007.00924.x
Whipple, S.D., Lindroth, E.J., Hoback, W.W., Albrecht, M.C. & Foster, J.E. (2012) Genetic variability of Digitonthophagus gazella (F.) (Coleoptera: Scarabaeidae) from Vieques, Puerto Rico and South Africa. The Coleopterists Bulletin, 66, 45–50.
https://doi.org/10.1649/072.066.0112
Wirta, H., Orisini, L. & Hanski, I. (2008) An old adaptative radiation of forest dung beetles in Madagascar. Molecular Phylogenetics and Evolution, 47, 1076–1089.
https://doi.org/10.1016/j.ympev.2008.03.010
Zimsen, E. (1964) The Type Material of I. C. Fabricius. Munksgaard, Copenhagen, Denmark, 280 pp.
Zunino, M. (1981) Insect of Saudi Arabia, Coleoptera: Fam. Scarabaeidae, Tribe Onthophagini. Fauna of Saudi Arabia, 3, 408–416.
Zunino, M. & Halffter, G. (1988) Analísis taxonómico, ecológico y biogeográfico de un grupo americana de Onthophagus (Coleoptera, Scarabaeidae). Monographie, Museo Regionale di Scienze Naturali, Torino, 9, 1–211.