Abstract

This paper presents a taxonomic review of the pilularius species group of the genus Canthon Hoffmannsegg, 1817, a New World group of ball-rolling (telocoprid) dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). This species group comprises five North American taxa: Canthon vigilans LeConte; C. chalcites (Haldeman); C. pilularius (Linnaeus); C. imitator Brown; and C. floridanus Brown. Each species is keyed, diagnosed, illustrated and presented with information on distribution, relationships, and biology. Nomenclatural actions taken include (a) designation of lectotype for Scarabaeus pilularius Linnaeus, 1758; (b) designation of lectotype for Scarabaeus hudsonias Forster, 1771; (c) designation of lectotype for Scarabaeus laevis Drury, 1773; (d) designation of lectotype and correction of type locality for Scarabaeus volvens Fabricius, 1792; and (e) correction of type locality for Canthon floridanus Brown. In addition, a new synonymy is designated: Scarabaeus hudsonias Forster, 1771, as senior objective synonym of S. laevis Drury, 1773.

References

• Adler K. 2015. Catesby’s fundamental contributions to Linnaeus’ binomial catalog of North American animals. p. 251–264. In: Nelson EC, Elliott DJ (eds.). The curious Mister Catesby: a “truly ingenious” naturalist explores new worlds. University of Georgia Press; Athens, GA. 456 p 
• Balthasar V. 1939. Eine Vorstudie zur Monographie der Gattung Canthon Hffsg. 10. Beitrag zur Kenntnis der Scarabaeiden der neotropische Region. Folia Zoologica et Hydrobiologica 9: 179–238. 
• Bauer AM. 2015. Catesby’s animals (other than birds) in The Natural History of Carolina, Florida and Bahama Islands. p. 231–250. In: Nelson EC, Elliott DJ (eds.). The curious Mister Catesby: a “truly ingenious” naturalist explores new worlds. University of Georgia Press; Athens, GA. 456 p 
• Blatchley WS. 1927. The Scarabaeidae of Florida. The Florida Entomologist 11: 55–62. 
• Blatchley WS. 1928. Notes on some Florida Coleoptera with descriptions of new species. The Canadian Entomologist 60: 60–73. 
• Blanchard F. 1885. On the species of Canthon and Phanaeus of the United States with notes on other genera. Transactions of the American Entomological Society 12: 163–169. 
• Bouchard P, Bousquet Y, Davies AE, Cai C. 2024. On the nomenclatural status of type genera in Coleoptera (Insecta). ZooKeys 1194: 1–981. 
• 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. 
• Bragg AN. 1957. Use of carrion by the beetle, Canthon laevis (Coleoptera: Scarabaeidae). The Southwestern Naturalist 2: 173. 
• Brown WJ. 1927. An annotated list of the coprophagous Scarabaeidae known to occur in Oklahoma. Proceedings of the Oklahoma Academy of Science 7: 24–28. 
• Brown WJ. 1946. Notes on some species of Canthon and Dichelonyx (Coleoptera, Scarabaeidae). The Canadian Entomologist 78: 104–109. 
• Calhoun JV. 2008. A glimpse into a “flora et entomologia”: The natural history of the rarer lepidopterous insects of Georgia” by J. E. Smith and J. Abbot (1797). Journal of the Lepidopterists Society 60: 1–37. 
• Calvert PP. 1930. Dynastes tityus (scarabaeid) in Pennsylvania and the Rathvon and Auxer collections of Coleoptera. Entomological News 41: 195–201. 
• Catesby M. 1747. Appendix to Catesby M. 1729–1743. The natural history of Carolina, Florida and the Bahama islands: containing the figures of birds, beasts, fishes, serpents, insects, and plants: particularly the forest-trees, shrubs, and other plants, not hitherto described, or very incorrectly figured by authors. Together with their descriptions in English and French. To which are added, observations on the air, soil, and waters: with remarks upon agriculture, grain, pulse, roots, &c. To the whole is prefixed a new and correct map of the countries treated of. Privately published; London, UK. 20 p  (Volume I was published in five parts dated 1729–1732; Volume II was published in five parts dated 1734–1743. The Appendix was issued in 1747 and comprised of 20 printed pages of animal and plant descriptions accompanied by 20 plates; it is sometimes inappropriately referred to as “Vol. 3” of Catesby’s classic and is often bound together with Volume 2.)
• Cupello M, Silva SAB, Vaz-de-Mello FZ. 2023. The taxonomic revolution of New World dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae). Frontiers in Ecology and Evolution 11: 1–42. 
• Cupello M, Vaz de Mello FZ. 2018. A monographic revision of the Neotropical dung beetle genus Sylvicanthon Halffter & Martínez, 1977 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini), including a reappraisal of the taxonomic history of ‘Canthon sensu lato’. European Journal of Taxonomy 467: 1–205. 
• Daniel GM, Davis ALV. 2023. Dung beetle tribal classification (Coleoptera: Scarabaeidae: Scarabaeinae): progress, problems and prospects. Annals of the Entomological Society of America 117: 1–13. 
• Dimmock G. 1884. Order VII. Coleoptera. p. 297–402. In: Kingsley JS (ed.). The standard natural history. Edited by John Sterling Kingsley. Vol. II. Crustacea and insects. Illustrated by six hundred and sixty-six woodcuts and twenty full-page plates. S.E. Cassino & Co.; Boston. 662 p 
• Drury D. 1770. Illustrations of natural history. Wherein are exhibited upwards of two hundred and forty figures of exotic insects, according to their different genera; very few of which have hitherto been figured by any author, being engraved and coloured from nature, with the greatest accuracy, and under the author’s own inspection, on fifty copper-plates. With a particular description of each insect: interspersed with remarks and reflections on the nature and properties of many of them. Vol. 1. B. White; London. 130 p  (See Bousquet 2016 for collation details and comments.)
• Drury D. 1773. Illustrations of natural history. Wherein are exhibited upwards of two hundred and twenty figures of exotic insects, according to their different genera; very few of which have hitherto been figured by any author, being engraved and coloured from nature, with the greatest accuracy, and under the author’s own inspection, on fifty copper-plates. With a particular description of each insect: interspersed with remarks and reflections on the nature and properties of many of them. Vol. 2. B. White; London. 90 p. [+ indices to volumes 1 and 2]  (See Bousquet 2016 for collation details and comments.)
• Dubois A, Nemésio A, Bour R. 2014. Primary, secondary and tertiary syntypes and virtual lectotype designation in zoological nomenclature, with comments on the recent designation of lectotype for Elephas maximus Linnaeus, 1758. Bionomina 7: 45–64. 
• Duponchel P. 1842. Canthon. Capnodis. Cardiorhinus. Ceraspis. In: Orbigny C. d’ (ed.). Dictionnaire universel d’histoire naturelle... Tome troisième. [Livraisons 27, 28, 29]. MM. Renard, Martinet et Cie., Paris. p. 131 
• Edmonds WD. 2018. The dung beetle fauna of the Big Bend region of Texas (Coleoptera: Scarabaeidae). Insecta Mundi 0642: 1–30. 
• Edmonds WD. 2022. Taxonomic review of the North American dung beetle genus Boreocanthon Halffter, 1958 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini). Insecta Mundi 0952: 1–65. 
• Edmonds WD. 2023. Taxonomic review of the North American dung beetle genus Melanoocanthon Halffter, 1958 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini). Insecta Mundi 1014: 1–28. 
• Erichson WF. 1847. Naturgeschichte der Insecten Deutschlands. Erste Abtheilung. Coleoptera. Dritter Band. Fünfte Lieferung. p. 641–800  Nicolaische Buchhandlung, Berlin
• Fabricius JC. 1792. Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species adjectis synonimis, locis, observationibus, descriptionibus. Tom. I, Pars I. Christ. Gottl. Proft; Copenhagen, Denmark. 330 p 
• Fabricius JC. 1801. Systema eleutheratorvm secvndvm ordines, genera, species: adiectis synonymis, locis, observationibvs, descriptionibvs. Tomvs I. Bibliopolii Academici Novi; Kiel, Germany. 506 p 
• Fincher GT. 1986. Importation, colonization, and release of dung-burying scarabs. Entomological Society of America, Miscellaneous Publication No. 61: 69–76. 
• Fincher GT, Blume RR, Hunter III JS, Beerwinkle KR. 1986. Seasonal distribution and diel flight activity of dung-feeding scarabs in open and wooded pasture in east-central Texas. Southwestern Entomologist, Supplement 10: 1–35. 
• Forster JR. 1771. Novae species insectorum. Centuria I. T. Davies et B. White; London. 100 p 
• Freese EL, Veal DA, Lago PK. 2020. The Scarabaeoidea (Coleoptera) of Iowa: An annotated checklist. Insecta Mundi 0787: 1–83. 
• Génier F. 2019. On the identity of Canthon imitator floridanus Brown, 1946 (Coleoptera: Scarabaeidae: Scarabaeinae). The Coleopterists Bulletin 73: 300–306. 
• Goldstein PZ, Simmons T. 2002. A checklist and commentary on the scarabaeid fauna of the Massachusetts offshore islands (Coleoptera: Scarabaeidae). Journal of the New York Entomological Society 110: 389–401. 
• Gordon RD, Cartwright OL. 1974. Survey of food preferences of some North American Canthonini. Entomological News 85: 181–185. 
• Haldeman SS. 1843. Descriptions of North American species of Coleoptera, presumed to be undescribed. Proceedings of the Academy of Natural Sciences of Philadelphia 1: 298–304. 
• Halffter G. 1958. Dos nuevos géneros de Canthonini (Col. Scarabaeidae). Ciencia (Mexico) 17: 207–212. 
• Halffter G. 1961. Monografía de las especies norteamericanas del género Canthon Hoffsg. (Coleopt. Scarab.) Ciencia (Mexico) 20: 225–320. 
• Halffter G, Edmonds WD. 1981. The nesting behavior of dung beetles (Scarabaeinae), an ecological and evolutive approach. Instituto de Ecología; Mexico City. 176 p 
• Halffter G, Espinoza de los Monteros A, Nolasco-Soto J, Arriaga-Jiménez A, Rivera-Gasperín S. 2022. Bajacanthon, a new subgenus for the Mexican Deltochilini (Coleoptera: Scarabaeidae: Scarabaeinae) fauna. Diversity 14 (109): 1–13. 
• Halffter G, Matthews EG. 1966. The natural history of dung beetles of the subfamily Scarabaeinae. Folia Entomológica Mexicana, Números 12–14: 1–312. 
• Halffter G, Morrone JJ. 2017. An analytical review of Halffter’s Mexican transition zone, and its relevance for evolutionary biogeography, ecology and biogeographical regionalization. Zootaxa 4226: 1–46. 
• Halffter G, Rivera Cervantes LE, Halffter V. 2015. Diversificación del grupo humectus del género Canthon (Coleoptera: Scarabaeidae: Scarabaeinae) en el occidente de México. Acta Zoológica Mexicana 31: 208–220. 
• Harold E von. 1868. Monographie der Gattung Canthon. Berliner Entomologische Zeitschrift 12: 1–144. 
• Harold E von. 1869. Vol. 4, Scarabaeidae. In: Gemminger M, Harold E von. Catalogus coleopterorum hucusque descriptorum synonymicus et systematicus. Gummi; Munich. 368 p. [numbered 979–1346] 
• Harold E von. 1870. Berichtigungen und Zusätze zum Catalogus Coleopterorum synonymicus et systematicus. Coleopterologische Hefte 7: 94–111. 
• Hayek CMF von. 1985. On the type material of the species of Coleoptera described from the Drury collection by D. Drury and J. C. Fabricius with notes on some Coleoptera from the Milne collection preserved in the British Museum (Natural History). Archives of Natural History 12: 143–152. 
• Hoffmannsegg JC von. 1817. Entomologische Bemerkungen bei Gelegenheit der Abhandlungen über amerikanischen Insecten, in der vierten nach sechten Lieferung von den Recueils d’observations de Zoologie et d’Anatomie comparée, oder dem zten Theile der Reise, der Herren A. v. Humboldt und A. Bonpland, nemlich: No. IX. In Livr. 4. p. 197–283. und No. XI. XII. In Livr. 5. 6. p. 294–397. Zoologische Magazin 1: 8–56. 
• Hope FW. 1837. The coleopterist’s manual, containing the lamellicorn insects of Linneus [sic] and Fabricius. Henry G. Bond; London. 121 p 
• Hope FW. 1838. Observations on the lamellicorns of Olivier. The Entomological Magazine 5: 312–326. 
• Horn GH. 1870. Notes on some genera of coprophagous Scarabaeidae of the United States. Transactions of the American Entomological Society 3: 42–51. 
• Gaedike R. 1990. Collectiones entomologicae. Ein Kompendius über den Verbleib entomologischer Sammlungen der Welt bis 1960. Teil I: A bis K. Berlin; Akademie der Landwirtschaftswissenschaften der Deutschen Demokratischen Republik. 220 p 
• ICZN [International Commission on Zoological Nomenclature]. 1957. Opinion 474 (Case 930) Determination of the dates to be assigned for the purposes of the Law of Priority to the names published in Dru Drury’s Illustrations of Natural History in the period 1770–1782. Opinions and Declarations rendered by the ICZN, 16(16): 297–306. 
• ICZN [International Commission on Zoological Nomenclature]. 1999. International Code of Zoological Nomenclature. Fourth Edition. The International Trust for Zoological Nomenclature; London. 306 p 
• Illiger JKW. 1803. Verzeichniss der in Portugall einheimischen Käfer. Erste Lieferung. Magazin für Insektekunde 2: 168–247. 
• Kadiri N, Lumaret J-P, Floate KD. 2014. Functional diversity and seasonal activity of dung beetles (Coleoptera: Scarabaeoidea) on native grasslands in southern Alberta, Canada. Canadian Entomologist 146: 291–305. 
• Kohlmann B, Halffter G. 1990. Reconstruction of a specific example of insect invasion waves: The cladistic analysis of Canthon (Coleoptera: Scarabaeidae) and related genera in North America. Quaestiones Entomologicae 26: 1–20. 
• Krell F-T. 2010. Catalogue of Colorado scarab and stag beetles (Coleoptera: Scarabaeoidea), based on literature records. DMNS Technical Report 2010–4. Denver; Denver Museum of Nature and Science. 84 p. Kriska NL, Young DK 2002. An annotated checklist of Wisconsin Scarabaeoidea (Coleoptera). Insecta Mundi 16: 31–48. 
• Landin B-O. 1956. The Linnean species of Lamellicornia described in “Systema Naturae”, Ed. X (1758). (Col.). Entomologisk Tidskrift 77: 1–18. 
• Lane F. 1947. Sobre os tipos e a sinonimia de alguns Canthonini (Col. Scarabaeidae). II. Papéis Avulsos do Departamento de Zoologia 9: 109–121. 
• Lane F. 1950. Sobre os tipos e a sinonimia de alguns Canthonini (Col. Scarabaeidae). III. Nota sobre a data certa de Canthon laevis Drury. Papéis Avulsos do Departamento de Zoologia 9: 79–82. 
• Latreille PA. 1812. Insectes de l’Amérique équinoxiale, recueillis pendant le voyage de MM. de Humboldt et Bonpland. In: Recueil d’observations de zoologie et d’anatomie comparée; faites dans l’océan Atlantique, dans l’intérieur du nouveau continent et dans la mer du sud, pendant les années 1799, 1800, 1801, 1802 et 1803, par Al. de Humboldt et A. Bonpland. Premier Volume (Second edition). F. Schell, G. Dufour; Paris. 368 p  (The second edition carries the publication date 1811 on the title page but was issued in 1812. See Bousquet 2016.)
• Latreille PA. 1829. Le règne animal distribué d’après son organisation, pour servir de base à l’histoire naturelle des animaux et d’introduction à l’anatomie comparée. Par M. le Baron Cuvier. Avec figures, dessinées d’après nature. Nouvelle édition, revue et augmentée. Tome IV. Crustacés, arachnides et partie des insectes. Déterville; Paris. 584 p 
• LeConte JL. 1858. Catalogue of Coleoptera of the regions adjacent to the boundary line between the United States and Mexico. Journal of the Academy of Natural Sciences of Philadelphia (Series 2) 4: 9–42. 
• LeConte JL. 1859. The Coleoptera of Kansas and eastern New Mexico. Smithsonian Contributions to Knowledge, Volume 2, Article 6. 58 p 
• LeConte JL. 1863. List of the Coleoptera of North America, prepared for the Smithsonian Institution. Part 1. Smithsonian Miscellaneous Collections No. 140. Smithsonian Institution: Washington D.C.. 50 p 
• Legner EF. 1978. Part I. Parasites and predators introduced against arthropod pests. Diptera. p. 346–355. In: Bartlett BR, Clausen CP (eds.). Introduced parasites and predators of arthropod pests and weeds: A world review. United States Department of Agriculture, Agricultural Handbook No. 48. USDA-ARS; Washington, DC. 545 p 
• Leng CW. 1920. Catalogue of the Coleoptera of America, north of Mexico. John D. Sherman: Mount Vernon, NY. 470 p 
• Lindquist AW. 1935. Notes on the habits of certain coprophagous beetles and methods of rearing them. United States Department of Agriculture Circular No. 351. 9 p 
• Linnaeus C. 1758. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tomus I. Editio decima, reformata. Laurentii Salvii; Stockholm. 823 p 
• Linnaeus C. 1764. Museum S:ae R:ae M:tis Ludovicae Ulricae Reginae Svecorum, Gothorum, Vandalorumque &c. &c. &c. In quo animalia rariora, exotica, imprimis insecta & conchilia describuntur & determinantur. Prodromi instar editum. Laurentii Salvii; Stockholm. 720 p 
• Linnaeus C. 1767. Systema naturae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Editio duodecima, reformata. Tom. I. Pars II. Laurentii Salvii; Stockholm. 795 p. [numbered 533–1327] 
• Majka CG, Chandler DS, Donahue CP. 2011. Checklist of the beetles of Maine, USA. Empty Mirrors Press; Halifax, NS. 328 p 
• Mamantov MA, Sheldon KS. 2023. Seasonality, distribution and diversity of dung beetles (Coleoptera: Scarabaeidae: Scarabaeinae, Aphodiinae and Geotrupidae: Geotrupinae) in Great Smoky Mountains National Park. The Coleopterists Bulletin 77: 285–295. 
• Matthews EG. 1963. Observations on the ball-rolling behavior of Canthon pilularius (L.) (Coleoptera, Scarabaeidae). Psyche 70: 75–93. 
• Medina CA, Scholtz CH, Gill BD. 2003. Morphological variation and systematics of Canthon Hoffmannsegg 1817, and related genera of New World Cathonini dung beetles (Coleoptera, Scarabaeinae). Deutsche Entomologische Zeitschrift 50: 23–68. 
• Melsheimer FE. 1853. Catalogue of the described Coleoptera of the United States. Smithsonian Institution; Washington D. C.. 174 p  (revised by S. S. Haldeman and J. L. LeConte)
• Miller A. 1954. Dung beetles (Coleoptera: Scarabaeidae) and other insects in relation to human feces in a hookworm area of southern Georgia. The American Journal of Tropical Medicine and Hygiene 3: 372–389. 
• Mulsant ME. 1842. Histoire naturelle des coléoptères de France. Maison, Libraire ; Paris. 304 p 
• Nemes SN, Price DL. 2015. Illustrated keys to the Scarabaeinae (Coleoptera: Scarabaeidae) of Maryland. Northeastern Naturalist 22: 318–344. 
• Nishida GM. 2002. Hawaiian terrestrial arthropod checklist. Fourth edition. Bishop Museum Technical Report No. 22. Hawaii Biological Survey; Honolulu, HI. 313 p 
• Nolasco-Soto J, González-Astorga J, Espinosa de los Monteros A, Favila ME. 2023. Evolutionary history and diversity in the ball roller beetle Canthon cyanellus. Frontiers in Ecology and Evolution 10: 1066439. 
• Nunes LG de OA, Nunes RV, Vaz de Mello FZ. 2018. Taxonomic revision of the South American subgenus Canthon (Goniocanthon) Pereira & Martínez, 1956 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini). European Journal of Taxonomy 437: 1–31. 
• Nunes LG de OA, Nunes RV, Vaz de Mello FZ. 2020. Taxonomic revision of the South American subgenus Canthon (Peltecanthon) Pereira, 1953 (Coleoptera: Scarabaeidae: Scarabaeinae: Deltochilini). European Journal of Taxonomy 594: 1. 
• Olivier G-A. 1789. No. 3. Scarabé. Scarabaeus. In: Entomologie, ou histoire naturelle des insectes, avec leurs caractères génériques et spécifiques, leur description, leur synonymie, et leur figure enluminée. Coléoptères. Tome premier. Baudouin; Paris, France. 433 p  (This volume treats eight genera, each paginated separately; No. 3 is Scarabaeus, paginated 1–190. See Bousquet 2016)
• Olivier G-A. 1790. Encyclopedie méthodique. Histoire naturelle. Insectes. Vol. 5. Panckoucke Librarie; Paris, France. 793 p 
• Paulian R. 1939. Contribution a l’etude des Canthonides américains [Coleopt. Lamellic.]. Annales de la Société Entomologique de France 108: 1–48. 
• Peck SB, Howden HF. 1985. Biogeography of scavenging scarab beetles in the Florida Keys: post-Pleistocene land-bridge islands. Canadian Journal of Zoology 63: 2730–2737. 
• Pokhrel MR, Cairns SC, Hemmings Z, Floate KD, Andrew NR. 2021. A review of dung beetle introductions in the Antipodes and North America: Status, opportunities, and challenges. Environmental Entomology 50: 762–780. 
• Price DL, Ratcliffe B. 2023. The scarabaeoid beetles of Maryland (Coleoptera). Bulletin of the University of Nebraska State Museum 33: 1–330. 
• Ray J. 1710. Historia insectorum. A.&J. Churchill; London. 400 p 
• Reiche L. 1841. Tableau d’une division systématique de la tribu des coprophages, dans la famille des lamellicornes. Revue Zoologique 1841: 211–213. 
• Reiche L. 1842. D’une classification methodique de la tribu des coprophages, famille des Lamellicornes, division des Scaraboeides [sic], Coléoptères, Pentamères. Annales de la Société Entomologique de France 11: 59–94. 
• Reveal JL. 2015. Identification of the plants and animals illustrated by Mark Catesby for The Natural History of Carolina, Florida and the Bahama Islands. p. 331–350. In: Nelson EC, Elliott DJ (eds.). The curious Mister Catesby: a “truly ingenious” naturalist explores new worlds. University of Georgia Press; Athens, GA. 456 p 
• Robinson M. 1948. A review of the species of Canthon inhabiting the United States. Transactions of the American Entomological Society 74: 83–99. 
• Say T. 1824. American entomology, or descriptions of the insects of North America. Illustrated by coloured figures from original drawings executed from nature. [Vol. 1] Samuel Augustus Mitchell; Philadelphia. 105 p. [unnumbered], 18 plates [I–XVIII] 
• Schmidt A. 1920. Beitrag zur Kenntnis der Gattungen Canthon Hffsg., Sybax Boh., Aphodius Ill., Simogenius Har., Ataenius Har. Archiv für Naturgeschichte 86: 114–147. 
• Schmidt A. 1922. 1. Bestimmungstabelle der mir bekannten Canthon-Arten. 2. Verbreitungsgebiete der Canthon-Arten. 3. Neubeschreibungen von Canthon, Saprositis, Mendidius, Euparia, und Ataenius. Archiv für Naturgeschichte 88: 61–103. 
• Smith JE, Abbot J. 1797. The natural history of the rarer lepidopterous insects of Georgia: including their systematic characters, the particulars of their several metamorphoses, and the plants on which they feed. Collected from the observation of Mr. John Abbot, many years resident in that country. Vol.1, 2. J. Edwards, Cadell et Davies, J. White; London. 314 p 
• Solís A, Kohlmann B. 2002. El género Canthon (Coleoptera: Scarabaeidae) en Costa Rica. Gironale Italiano di Entomologia 10: 1–68. 
• Stanbrook R, King JR. 2021. Dung beetle community composition affects dung turnover in subtropical US grasslands. Ecology and Evolution 12: e8660.https://doi.org/10.1002/ece3.8660 
• Viera MK, Vaz de Mello FZ, Silva FAB. 2019. A taxonomic revision of Pseudepilissus Martínez, 1954 (Coleoptera: Scarabaeidae: Scarabaeinae). Insect Systematics & Evolution 51: 696–752. 
• Vulcano MA, Pereira FS. 1964. Catalogue of the Canthonini (Col. Scarab.). Entomologischen Arbeiten aus dem Museum G. Frey, Tutzing 15: 570–685. 
• Wagner PM, Abagandura GO, Mamo M, Weissling T, Wingeyer A, Bradshaw JD. 2020. Abundance and diversity of dung beetles (Coleoptera: Scarabaeoidea) as affected by grazing management in Nebraska sandhills ecosystem. Environmental Entomology 50: 222–231. 
• Wallace FL, Drew WA. 1965. The Scarabaeinae of Oklahoma. Proceedings of the Oklahoma Academy of Science 45: 82–93. 
• Wallin L. 2001. Catalogue of type specimens. Part 4. Linnean specimens. Version 6. Uppsala University Museum of Evolution, Zoology Section. 128 p 
• Wheeler AG Jr., Miller GL. 2006. Simon Snyder Rathvon: Popularizer of agricultural entomology in mid-19th century America. American Entomologist 52: 36–47. 
• Whipple SD, Hoback WW. 2012. A comparison of dung beetle (Coleoptera: Scarabaeidae) attraction to native and exotic mammal dung. Environmental Entomology 41: 238–244. 
• Woodruff RE. 1973. The scarab beetles of Florida (Coleoptera: Scarabaeidae). Part I. The Laparosticti (Subfamilies: Scarabaeinae, Aphodiinae, Hybosorinae, Ochodaeinae, Geotrupinae, Acanthocerinae). Arthropods of Florida and neighboring land areas, Vol. 8. Florida Department of Agriculture and Consumer Services; Gainesville, Florida. 220 p 
• Ziegler D. 1846. Descriptions of new North American Coleoptera. Proceedings of the Academy of Natural Sciences of Philadelphia 2: 43–47.