Skip to main content Skip to main navigation menu Skip to site footer
Responsive image
Issue: Vol. 5301 No. 4: 13 Jun. 2023
Type: Article
Published: 2023-06-13
DOI: 10.11646/zootaxa.5301.4.2
Page range: 427-446
Abstract views: 323
PDF downloaded: 28

On the types of Bolboceratidae (Coleoptera: Scarabaeoidea) in the Swedish Museum of Natural History in Stockholm

Department of Agronomy; Food; Natural resources; Animals and Environment (DAFNAE); University of Padua; Via dell’Università 16; 35020 Legnaro (PD); Italy
Swedish Museum of Natural History; PO Box 50007; SE-104 05 Stockholm; Sweden
Coleoptera Natural history collections Nomenclature Taxonomy Stockholm museum Boheman Péringuey Wahlberg South Africa

Abstract

The investigation of the Bolboceratidae collection of the Stockholm Museum of Natural History (NHRS) yielded 12 type specimens of eight Afrotropical and one Australian species. The majority of these types belongs to species described by Carl Henrik Boheman (1796–1868) based on specimens collected by Johan August Wahlberg (1810–1856) in southern Africa. In two of these species, we found ambiguity among potential syntypes. Therefore, based on comparison with original descriptions, we designate lectotypes for Bolboceras caffrum Boheman, 1857 and Bolboceras exasperans Péringuey, 1908, both of which are today classified in the genus Bolbocaffer Vulcano, Martínez & Pereira, 1969. Finally, we establish Bolboceras matabele Péringuey, 1908 (syn. nov.) as a junior subjective synonym of B. exasperans, we confirm a few synonymies that were historically established without study of type specimens, and we resurrect Bolboceras dorsuale Boheman, 1857 from previous synonymy under B. maculicolle Boheman, 1857 (Mimobolbus dorsualis (Boheman, 1857) n. comb. and stat. rev.).

 

References

  1. Agoglitta, R., Barbero, E., Ragusa, E. & Zunino, M. (2006) Catalogo sistematico e topografico dei Geotrupidae e Scarabaeidae degradatori della Sicilia e isole circumsiciliane (Coleoptera: Scarabaeoidea). Boletín de la Sociedad Entomológica Aragonesa, 39, 181–204.
  2. Ahrens, D., Schwarzer, J. & Vogler, A.P. (2014) The evolution of scarab beetles tracks the sequential rise of angiosperms and mammals. Proceedings of the Royal Society B: Biological Sciences, 281 (1791), 1–10. https://doi.org/10.1098/rspb.2014.1470 DOI: https://doi.org/10.1098/rspb.2014.1470
  3. Baguena Corella, L. (1967) Scarabaeoidea de la fauna Ibero-Balear y Pirenaica. Consejo superior de investigaciones cientificas, Instituto Español de Entomologia, Madrid, 576 pp.
  4. Baraud, J. (1977) Coléoptères Scarabaeoidea. Faune de l’Europe occidentale, Belgique, France Grande-Bretagne, Italie, Péninsule Ibérique. Nouvelle Revue d’Entomologie, 4, 1–352.
  5. Baraud, J. (1992) Coléoptères Scarabaeoidea d’Europe. Faune de France 78. Fédération française des Sociétés de Sciences naturelles et Société Linnéenne de Lyon, Paris, 856 pp. DOI: https://doi.org/10.3406/linly.1992.11007
  6. Blackburn, T. (1904) A revision of the Australian species of Bolboceras, with descriptions of new species. Proceedings of the Linnean Society of New South Wales, 29, 481–526. https://doi.org/10.5962/bhl.part.20169 DOI: https://doi.org/10.5962/bhl.part.20169
  7. Bocak, L., Barton, C., Crampton-Platt, A., Chesters, D., Ahrens, D. & Vogler, A.P. (2014) Building the Coleoptera tree-of-life for >8000 species: Composition of public DNA data and fit with Linnaean classification. Systematic Entomology, 39, 97–110. https://doi.org/10.1111/syen.12037 DOI: https://doi.org/10.1111/syen.12037
  8. Boheman, C.H. (1848) Insecta Caffrariae annis 1838–1845 a J. A. Wahlberg collecta. I. 1. Fritze, Stockholm, 297 pp. https://doi.org/10.5962/bhl.title.9375 DOI: https://doi.org/10.5962/bhl.title.9375
  9. Boheman, C.H. (1851) Insecta Caffrariae annis 1838–1845 a J. A. Wahlberg collecta. I. 2. Fritze, Stockholm, 328 pp.
  10. Boheman, C.H. (1857) Insecta Caffraria annis 1838–1845 a J A. Wahlberg collecta. II. Fritze, Stockholm, 395 pp.
  11. Boheman, C.H. (1860) Coleoptera samlade af J. A. Wahlberg i sydvestra Afrika. Öfversigt af Kongl. Svenska Vetenskapsakademiens Förhandlingar, 17, 107–120.
  12. Bouchard, P., Bousquet, 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, A. & Smith, A.B.T. (2011) Family-group names in Coleoptera (Insecta). ZooKeys, 88, 1–972. https://doi.org/10.3897/zookeys.88.807 DOI: https://doi.org/10.3897/zookeys.88.807
  13. Boucomont, A. (1902) Coleoptera Lamellicornia Fam. Geotrupidae. Genera Insectorum. Proceedings of the United States National Museum, 7, 1–20.
  14. Boucomont, A. (1910) Contribution à la classification des Geotrupidae. Annales De La Société Entomologique De France, 79, 333–350.
  15. Boucomont, A. (1912) Scarabaeidae: Taurocerastinae, Geotrupinae. In: Junk, W. & Schenkling, S. (Eds.), Coleopterum Catalogus. Pars 46. Junk, W., Berlin, pp. 1–47.
  16. Browne, D.J. & Scholtz, C.H. (1995) Phylogeny of the families of Scarabaeoidea (Coleoptera) based on characters of the hindwing articulation, hindwing base and wing venation. Systematic Entomology, 20, 145–173. https://doi.org/10.1111/j.1365-3113.1995.tb00089.x DOI: https://doi.org/10.1111/j.1365-3113.1995.tb00089.x
  17. Cambefort, Y. (1991) Biogeography and Evolution. In: Hanski, I. & Cambefort, Y. (Eds.), Dung Beetle Ecology. Princeton University Press, Princeton, pp. 51–67. https://doi.org/10.1515/9781400862092.51 DOI: https://doi.org/10.1515/9781400862092.51
  18. Carpaneto, G.M., Mignani, R. & Piattella, E. (1993) A revision of the Afro-Indian genus Bolboceratops. Journal of African Zoology, 107, 329–353.
  19. Cartwright, O.L. (1953) Scarabaeid beetles of the genus Bradycinetulus and closely related genera in the United States. Proceedings of the United States National Museum, 103, 95–120. https://doi.org/10.5479/si.00963801.103-3318.95 DOI: https://doi.org/10.5479/si.00963801.103-3318.95
  20. Crowson, R.A. (1981) The biology of the Coleoptera. Harcourt Brace Jovanovich Publisher (Academic Press), London, 802 pp. https://doi.org/10.1016/c2013-0-07304-5 DOI: https://doi.org/10.1016/C2013-0-07304-5
  21. Dean, W.R.J., Hlander, E. & Johansson, U.S. (2022) Avian type localities and the type specimens collected by Johan August Wahlberg in southern Africa. Zootaxa, 5134 (4), 521–560. https://doi.org/10.11646/zootaxa.5134.4.3 DOI: https://doi.org/10.11646/zootaxa.5134.4.3
  22. Dietz, L., Seidel, M., Eberle, J., Misof, B., Pacheco, T.L., Podsiadlowski, L., Ranasinghe, S., Gunter, N.L., Niehuis, O., Mayer, C. & Ahrens, D. (2023) A transcriptome-based phylogeny of Scarabaeoidea confirms the sister group relationship of dung beetles and phytophagous pleurostict scarabs (Coleoptera). bioRxiv. [preprint] https://doi.org/10.1101/2023.03.11.532172 DOI: https://doi.org/10.1101/2023.03.11.532172
  23. Engström, K. (1989) Naturen berättar. Utveckling och forskning vid Naturhistoriska Riksmuseet. Naturhistoriska Riksmuseet, Stockholm, 184 pp.
  24. Fåhraeus, O.I. (1870) Coleoptera Caffrariae. Öfversikt af Kongl. Vetenskapsakademiens Förhandlingar, 27 & 28, 243–357 & 3–69 + 197–291 + 433–452 + 661–671.
  25. Felsche, C. (1910) Ateuchini und Coprini. In: Sjöstedt, Y. (Ed.), Wissenschaftliche Ergebnisse der schwedischen zoologischen Expedition nach dem Kilimandjaro, dem Meru und den umgebenden Massaisteppen Deutsch-Ostafrikas 1905–1906 unter Leitung von Prof. Dr. Yngve Sjöstedt. P. Palmquist Aktiebolag, Stockholm, pp. 39–44.
  26. Forshage, M. (2020) Old entomology collections kept separate in the Stockholm Museum of Natural History, occasionally referred to as “historical insect collections.” Compendium and pdf, Swedish Museum of Natural History, 34. [unpaginated]
  27. Forshage, M. & Vårdal, H. (2019) Den entomologiska verksamheten på Riksmuseet 1915–2013. Entomologisk tidskrift, 140 (2), 89–106.
  28. Gemminger, M. & Harold, E. von (1869) In: Gummi, E. (Ed.), Catalogous Coleopterorum hucusque descriptorum synonymicus et systematicus. Tomus IV. Scarabaeidae. sumptu E.H. Gummi, Monachii, pp. 979–1346. https://doi.org/10.5962/bhl.title.9089 DOI: https://doi.org/10.5962/bhl.title.9089
  29. Gillet, J.J.E. (1925) Results of Dr. E. Mjöberg’s Swedish Scientific Expeditions to Australia, 1910–1913. 40. Scarabaeidae: Geotrupinae et Coprinae. Arkiv För Zoologi, Stockholm, 17 (7), 1–16.
  30. Grebennikov, V.V. & Scholtz, C.H. (2004) The basal phylogeny of Scarabaeoidea (Insecta: Coleoptera) inferred from larval morphology. Invertebrate Systematics, 18 (3), 321–348. https://doi.org/10.1071/is03013 DOI: https://doi.org/10.1071/IS03013
  31. Gussmann, S.M.V. & Scholtz, C.H. (2000) Systematic revision of endemic southern African genera of Bolboceratinae. Journal of Natural History, 34 (7), 1045–1123. https://doi.org/10.1080/00222930050020122 DOI: https://doi.org/10.1080/00222930050020122
  32. Gussmann, S.M.V. & Scholtz, C.H. (2001) Systematic revision of the Afrotropical genus Bolbaffer, with descriptions of eight new species. Journal of Natural History, 35 (7), 1013–1084. https://doi.org/10.1080/002229301300323910 DOI: https://doi.org/10.1080/002229301300323910
  33. Hillert, O., Arnone, M., Král, D. & Massa, B. (2016) The genus Bolbelasmus in the western and southern regions of the Mediterranean Basin. Acta Entomologica Musei Nationalis Pragae, 56 (1), 211–254.
  34. Houston, T.F. (2011) Egg gigantism in some Australian earth-borer beetles (Coleoptera: Geotrupidae: Bolboceratinae) and its apparent association with reduction or elimination of larval feeding. Australian Journal of Entomology, 50 (2), 164–173. https://doi.org/10.1111/j.1440-6055.2010.00794.x DOI: https://doi.org/10.1111/j.1440-6055.2010.00794.x
  35. Houston, T.F. & Bougher, N.L. (2010) Records of hypogeous mycorrhizal fungi in the diet of some Western Australian bolboceratine beetles (Coleoptera: Geotrupidae, Bolboceratinae). Australian Journal of Entomology, 49 (1), 49–55. https://doi.org/10.1111/j.1440-6055.2009.00720.x DOI: https://doi.org/10.1111/j.1440-6055.2009.00720.x
  36. Howden, H. (1975) A sixth species of Stenaspidius with a note on adult behaviour. Journal of the Australian Entomological Society, 14 (4), 439–441. https://doi.org/10.1111/j.1440-6055.1975.tb02062.x DOI: https://doi.org/10.1111/j.1440-6055.1975.tb02062.x
  37. Howden, H. (1985) A revision of the Australian genera Bolboleaus, Blackbolbus and Bolborhachium. Australian Journal of Zoology, Supplementary Series, 33 (111), 1–179. https://doi.org/10.1071/ajzs111 DOI: https://doi.org/10.1071/AJZS111
  38. Howden, H. & Cooper, J.B. (1977) The generic classification of the Bolboceratini of the Australian region, with descriptions of four new genera. Australian Journal of Zoology, Supplementary Series, 25 (50), 1–50. https://doi.org/10.1071/ajzs050 DOI: https://doi.org/10.1071/AJZS050
  39. Howden, H., Howden, A. & Holloway, G. (2007) Digging down under: Australian Bolboceratini, their habits and a list of species (Coleoptera: Scarabaeoidea: Geotrupidae). Zootaxa, 1499 (1), 47–59. https://doi.org/10.11646/zootaxa.1499.1.2 DOI: https://doi.org/10.11646/zootaxa.1499.1.2
  40. Howden, H.F. (1955) Biology and taxonomy of North American Beetles of the subfamily Geotrupinae with revision of the genera Bolbocerosomas, Eucanthus, Geotrupes and Peltotrupes. Proceedings of the United States National Museum, 104 (3342), 151–319. https://doi.org/10.5479/si.00963801.104-3342.151 DOI: https://doi.org/10.5479/si.00963801.104-3342.151
  41. Howden, H.F. (1964) The Geotrupinae of North and Central America. The Memoirs of the Entomological Society of Canada, 96, 5–91. https://doi.org/10.4039/entm9639fv DOI: https://doi.org/10.4039/entm9639fv
  42. Hunt, T., Bergsten, J., Levkanicova, Z., Papadopoulou, A., John, O.S., Wild, R., Hammond, P.M., Ahrens, D., Balke, M., Caterino, M.S., Gómez-Zurita, J.,Ribera, I., Barraclough, T.G., Bocakova, M., Bocak, L. & Vogler, A.P. (2007) A comprehensive phylogeny of beetles reveals the evolutionary origins of a superradiation. Science, 318 (5858), 1913–1916. https://doi.org/10.1126/science.1146954 DOI: https://doi.org/10.1126/science.1146954
  43. Jaquelin du Val, P.N.C. (1863) Manuel entomologique. Genera des Coléoptères d’Europe comprenant leur classifi cation en familles naturelles, la description de tous les genres, des Tableaux synoptiques destinés à faciliter l’étude, le Catalogue de toutes les espèces, de nombreux dessins au trait de caractères. Tome troisiéme (1859–1863). A. Deyrolle, Paris, 463 pp.
  44. Kolbe, H.J. (1907) Neue Coprophagen aus Afrika I. Annales de la Société Entomologique de Belgique, 51, 25–31.
  45. Král, D., Löbl, I. & Nikolajev, G.V. (2006) Bolboceratidae. In: Löbl, I. & Smetana, A. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 3. Apollo Books, Stenstrup, pp. 82–84.
  46. Krikken, J. (1977) The genus Bolbelasmus Boucomont in Asia with notes on species occurring in other regions. Zoologische Mededelingen Leyden, 51 (17), 277–292.
  47. Krikken, J. (1984) A generic reclassification of the Afrotropical Bolboceratini. Zoologische Mededelingen Leyden, 58 (3), 23–45.
  48. Krikken, J. & Li, C.L. (2013) Taxonomy of the Oriental genus Bolbochromus: a generic overview and descriptions of four new species. Zootaxa, 3731 (4), 495–519. https://doi.org/10.11646/zootaxa.373 1.4.4 DOI: https://doi.org/10.11646/zootaxa.3731.4.4
  49. Lawrence, J.F. & Newton, A.F. (1995) Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names. In: Pakaluk, J. & Ślipiński, A. (Eds.), Biology, Phylogeny, and Classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii Polska Akademia Nauk, Warszawa, pp. 779–1006.
  50. Lawrence, J.F., Ślipiński, A., Seago, A.E., Thayer, M.K., Newton, A.F. & Marvaldi, A.E. (2011) Phylogeny of the Coleoptera based on morphological characters of adults and larvae. Annales Zoologici, 61 (1), 1–217. https://doi.org/10.3161/000345411X576725 DOI: https://doi.org/10.3161/000345411X576725
  51. Lea, A.M. (1916) Notes on some miscellaneous Coleoptera with descriptions of new species. Part II. Transactions of the Royal Society of South Australia, 40, 272–436. DOI: https://doi.org/10.5962/bhl.part.18882
  52. Li, C.L., Krikken, J. & Wang, C.C. (2019) Review of the genus Bolbochromus in the Philippines. ZooKeys, 842, 135–152. https://doi.org/10.3897/zookeys.842.32315 DOI: https://doi.org/10.3897/zookeys.842.32315
  53. Li, C.L., Wang, C.C., Masumoto, K., Ochi, T. & Yang, P.S. (2008) Review of the Tribe Bolboceratini s.l. from Taiwan with a Key to the Eurasian Genera. Annals of the Entomological Society of America, 101 (3), 474–490. https://doi.org/10.1603/0013-8746(2008)101[474:rottbs]2.0.co;2 DOI: https://doi.org/10.1603/0013-8746(2008)101[474:ROTTBS]2.0.CO;2
  54. Lönnberg, E. (1916) Naturhistoriska Riksmuseets historia. Dess uppkomst och utveckling. Kungl. Vetenskapsakademien, Stockholm, 290 pp.
  55. López-Colón, J.I. (2003) Lista preliminar del los Scarabaeoidea (Coleoptera) de la fauna Europea (Parte I). Boletín de la Sociedad Entomológica Aragonesa, 33, 135–144.
  56. Marseul, S.A. (1857) Catalogue des coléoptères d’Europe. A. Duménil, Paris, 200 pp. DOI: https://doi.org/10.5962/bhl.title.66031
  57. Martín-Piera, F. & López-Colón, J.I. (2000) Fauna Iberica. Vol. 14. Coleoptera, Scarabaeoidea I. Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Científicas (CSIC), Madrid, 526 pp.
  58. McKenna, D.D., Shin, S., Ahrens, D., Balke, M., Beza-Beza, C., Clarke, D.J., Donath, A., Escalona, H.E., Friedrich, F., Letsch, H., Liu, S., Maddison, D., Mayer, C., Misof, B., Murin, P.J., Niehuis, O., Peters, R.S., Podsiadlowski, L., Pohl, H., Scully, E.D., Yan, E.V., Zhou, X., Ślipiński, A. & Beutel, R.G. (2019) The evolution and genomic basis of beetle diversity. Proceedings of the National Academy of Sciences, 116 (49), 24729–24737. https://doi.org/10.1073/pnas.1909655116 DOI: https://doi.org/10.1073/pnas.1909655116
  59. Mckenna, D.D., Wild, A.L., Kanda, K., Bellamy, C.L., Beutel, R.G., Caterino, M.S., Farnum, C.W., Hawks, D.C., Ivie, M.A., Jameson, M.L., Leschen, R.A.B., Marvaldi, A.E., Mchugh, J.V., Newton, A.F., Robertson, J.A., Thayer, M.K., Whiting, M.F., Lawrence, J.F., Ślipiński, A., Maddison, D.R. & Farrell, B.D. (2015) The beetle tree of life reveals that Coleoptera survived end-Permian mass extinction to diversify during the Cretaceous terrestrial revolution. Systematic Entomology, 40 (4), 835–880. https://doi.org/10.1111/syen.12132 DOI: https://doi.org/10.1111/syen.12132
  60. Mulsant, E. (1843) Description de quelques Coléoptères inédits. Annales De La Société Agricole De Lyon, 6, 276–286.
  61. Nikolajev, G. (1970) The taxonomic rank of the groups within the subfamily Geotrupinae (Coleoptera, Scarabaeidae). In: Proceedings of the II Scientific Conference of Young Researchers and Postgraduates, dedicated to the 100th Anniversary of the Birth of V. I. Lenin and to the 50th Anniversary of Kazakhstan, Almaty, Kazakhstan, 1970, pp. 31–34.
  62. Nikolajev, G.V., Král, D. & Bezděk, A. (2016) Geotrupidae. In: Löbl, I. & Löbl, D. (Eds.), Catalogue of Palaearctic Coleoptera. Vol. 3. Scarabaeoidea, Scirtoidea, Dascilloidea, Buprestoidea, Byrrhoidea. Revised and updated edition. Brill NV, Leiden, pp. 33–52. DOI: https://doi.org/10.1163/9789004309142_003
  63. Niogret, J., Felix, A.E., Nicot, A. & Lumaret, J.P. (2019) Chemosystematics using cuticular compounds: A powerful tool to separate species in Mediterranean dung beetles (Coleoptera: Geotrupidae). Journal of Insect Science, 19 (2), 1–11. https://doi.org/10.1093/jisesa/iez026 DOI: https://doi.org/10.1093/jisesa/iez026
  64. Paulian, R. (1941) Coprophaga Africana: Deuxième note, Revision des Bolboceras Africains. Revue de Zoologie et de Botanique Africains, 35, 1–71.
  65. Paulian, R. & Baraud, J. (1982) Faune des Coléoptères de France. II. Lucanoidea et Scarabaeoidea, Encyclopédie Entomologique. Vol. 43. Lechevalier, Paris, 477 pp.
  66. Péringuey, L. (1901) Descriptive catalogue of the Coleoptera of South Africa. Transactions of the South African Philosophical Society, 12, 1–563. DOI: https://doi.org/10.1080/21560382.1901.9525976
  67. Péringuey, L. (1908) Descriptive catalogue of the Coleoptera of South Africa. Additions and corrections. Transactions of the South African philosophical Society, 13, 547–752.
  68. Petrovitz, R. (1975) Neue Aphodiinae, Hybosorinae, Bolbocerinae und Orphninae. Revue Suisse De Zoologie, 82 (3), 615–624. DOI: https://doi.org/10.5962/bhl.part.78280
  69. Scholtz, C.H. & Grebennikov, V. (2005) Scarabaeoidea. In: Beutel, R. & Leschen, R. (Eds.), Coleoptera, Beetles. Vol. 1. Morphology and Systematics (Archostermata, Adephaga, Myxophaga Polyphaga partim). Handbook of zoology, Arthropoda: Insecta. Walter de Gruyter, Berlin, pp. 443–525.
  70. Scholtz, C.H. & Browne, D. J. (1996) Polyphyly in the geotrupidae (Coleoptera: Scarabaeoidea): A case for a new family. Journal of Natural History, 30, 597–614. https://doi.org/10.1080/00222939600770311 DOI: https://doi.org/10.1080/00222939600770311
  71. Schoolmeesters, P. (2023) World Scarabaeidae Database. In: Bánki, O., Roskov, Y., Döring, M., Ower, G., Vandepitte, L., Hobern, D., Remsen, D., Schalk, P., DeWalt, R.E., Keping, M., Miller, J., Orrell, T., Aalbu, J. Abbott, R., Adlard, R., Adriaenssens, E.M., Aedo, C., Aescht, E. & Akkari, N. et al., Catalogue of Life Checklist. Version 9 February 2023). Available from: https://www.catalogueoflife.org/ (accessed 22 May 2023)
  72. Seabra, A.F. (1943) Contribuições para o inventario da fauna lusitânica. Insecta, Coleoptera. Memorias e Estudos do Museo Zoologico da Universidade de Coimbra, 142, 47–56.
  73. Sjöstedt, Y. (1910) Die Tierwelt der Steppen und Bergen. In: Sjöstedt, Y. (Ed.), Wissenschaftliche Ergebnisse der schwedischen zoologischen Expedition nach dem Kilimandjaro, dem Meru und den umgebenden Massaisteppen Deutsch-Ostafrikas 1905–1906 unter Leitung von Prof. Dr. Yngve Sjöstedt. Palmquist, Stockholm, pp. 1–80. DOI: https://doi.org/10.5962/bhl.title.1805
  74. Sjöstedt, Y. (1911) Bland storvildt i Ostafrika. Norstedts, Stockholm, 622 pp.
  75. Smith, A.B.T., Hawks, D.C. & Heraty, J.M. (2006) An overview of the classification and evolution of the major scarab beetle clades (Coleoptera: Scarabaeoidea) based on preliminary molecular analyses. The Coleopterists Bulletin, 60, 35–46. https://doi.org/10.1649/0010-065x(2006)60[35:aootca]2.0.co;2 DOI: https://doi.org/10.1649/0010-065X(2006)60[35:AOOTCA]2.0.CO;2
  76. TaxonWorks Community (2022) TaxonWorks [software and supporting resources], https://taxonworks.org.
  77. Verdú, J.R., Galante, E., Lumaret, J.P. & Cabrero-Sañudo, F.J. (2004) Phylogenetic analysis of Geotrupidae (Coleoptera, Scarabaeoidea) based on larvae. Systematic Entomology, 29, 509–523. https://doi.org/10.1111/j.0307-6970.2004.00256.x DOI: https://doi.org/10.1111/j.0307-6970.2004.00256.x
  78. Vulcano, M.A., Martínez, A. & Pereira, F.S. (1969) Notes on African Bolboceratini. Papéis Avulsos de Zoologia, 22, 159–174.
  79. de Vylder, G. (1998) The Journal of Gustaf de Vylder, naturalist in South-Western Africa 1873–1875. Van Riebeeck Society, Cape Town, 292 pp.
  80. Wahlberg, J.A. (1994) Johan August Wahlberg: travel Journals (and Some Letters) South Africa and Namibia/Botswana, 1838–1856. Translated from Swedish by Michael Roberts. Van Riebeeck Society, Cape Town, 249 pp.
  81. Wallengren, H.D.J. (1859) Kafferlandets Dag Fjärilar, insamlade åren 1838–1845 af J. A. Wahlberg. Kongl. Vetenskapsakademiens Handlingar, 2 (4), 1–55.
  82. Wallengren, H.D.J. (1865) Heterocer-fjärilar, samlade i Kafferlandet af J. A. Wahlberg. Kongl. Vetenskapsakademiens Handlingar, 5 (4), 1–83. DOI: https://doi.org/10.5962/bhl.title.13861
  83. Westwood, J.O. (1848) On the Australian species of the coleopterous genus Bolboceras Kirby. The Annals and Magazine of Natural History, 2, 11–18. DOI: https://doi.org/10.1111/j.1096-3642.1852.tb00495.x
  84. Winkler, A. (1929) Lamellicornia. In: Catalogus Coleopterorum regionis palaearcticae 2 pars 9. Winkler, A., Wien, pp. 1025–1134.
  85. Zhang, S.Q., Che, L.H., Li, Y., Dan, L., Pang, H., Ślipiński, A. & Zhang, P. (2018) Evolutionary history of Coleoptera revealed by extensive sampling of genes and species. Nature Communications, 9 (1), 1–11. https://doi.org/10.1038/s41467-017-02644-4 DOI: https://doi.org/10.1038/s41467-017-02644-4