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Superregnum: Eukaryota
Cladus: Unikonta
Cladus: Opisthokonta
Cladus: Holozoa
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Cladus: Metapterygota
Infraclassis: Neoptera
Cladus: Eumetabola
Cladus: Endopterygota
Superordo: Coleopterida
Ordo: Coleoptera
Subordo: Polyphaga
Infraordo: Scarabaeiformia
Superfamilia: Scarabaeoidea

Subfamilia: Ceratocanthinae
Tribus: Ceratocanthini - Ivieolini - Scarabatermitini
Overview of genera

Acanthocerodes – Afrocloetus – Aneilobolus – Anopsiostes – Astaenomoechus – Aulisostes – Balleriodes – Baloghianestes – Belloanopsiostes – Besuchetostes – Callophilharmostes – Carinophilharmostes – Ceratocanthoides – Ceratocanthopsis – Ceratocanthus – Chaetophilharmostes – Cloeotus – Congomostes – Cryptophilharmostes – Cryptosphaeroides – Cyphopisthes – Ebbrittoniella – Eusphaeropeltis – Germarostes – Glyptopterus – Goudotostes – Hodotermopsis – Ivieolus – Madrasostes – Martinezostes – Melanophilharmostes – Nesopalla – Oxymorostes – Paulianostes – Perignamptus – Petrovitzostes – Philharmostes – Pseudopedaria – Pseudopterorthochaetes – Pseudosynarmostes – Pterorthochaetes – Scarabaeinus – Scarabatermes – Synarmostes – Termitariophilous – Trachycrusus – Xenocanthus

Ceratocanthidae A. Martínez, 1968

Acanthoceridae Lacordaire, 1856


Template:Martinez, 1968

Ballerio, A. 2000: A new genus and species of Ceratocanthidae from Tanzania (Coleoptera Scarabaeoidea). African zoology, 35: 131–137.
Ballerio, A. 2000: Revision of the genus Ebbrittoniella Martinez (Coleoptera: Scarabaeoidea: Ceratocanthidae). Revue suisse de zoologie, 107: 259–275.
Ballerio, A. 2001: Description of Philharmostes werneri n. sp. from Tanzania with notes on the “Philharmostes” generic group (Coleoptera, Ceratocanthidae). Fragmenta entomologica, 33: 147–157.
Ballerio, A. 2005: The discovery of a second species of Cryptophilharmostes Ballerio from Tanzania. Zootaxa, 1004: 29–36.
Ballerio, A. 2006: The enigmatic Afrotropical genus Afrocloetus Petrovitz, 1968, with description of a second species from Kenia. Giornale italiano di entomologia, 11: 307–313.
Ballerio, A. 2008: Descriptions of two new endemic genera and four new species of Ceratocanthinae (Insecta, Coleoptera, Scarabaeoidea, Hybosoridae) from Madagascar. Zoosystema, 30(3): 605–628.
Ballerio, A. 2009: Unusual morphology in a new genus and species of Ceratocanthinae from New Guinea (Coleoptera: Scarabaeoidea: Hybosoridae). Coleopterists bulletin, 63: 44–53. DOI: 10.1649/0010-065X-63.1.44
Ballerio, A. 2014: New data on Palaearctic Ceratocanthinae with description of a new species from China. Koleopterologische rundschau, 84: 277–280. [Not seen] Reference page.
Ballerio, A.; Gill, B.D. 2008: Notes on some Germarostes s.str. Paulian, 1982 from the cloud forests of Ecuadorian Andes with remarks on allied Ceratocanthinae genera (Coleoptera Scarabaeoidea Hybosoridae). Pp. 407-416 in: Giachino, P.M. (ed) Biodiversity of South America. I. Memoirs on biodiversity, 1 [seen in part]
Ballerio, A.; Maruyama, M. 2010: The Ceratocanthinae of Ulu Gombak: high species richness at a single site, with descriptions of three new species and an annotated checklist of the Ceratocanthinae of Western Malaysia and Singapore (Coleoptera, Scarabaeoidea, Hybosoridae). In: Ratcliffe, B.; Krell, F.-T. (eds) Current advances in Scarabaeoidea research. ZooKeys, 34: 77–104. DOI: 10.3897/zookeys.34.268
Gao, L. 2010: A new species of Ceratocanthidae (Coleoptera) from China. Oriental insects, 43: 155–158. ISSN: 0030-5316 [not seen]
Grebennikov, V.V. 2021a. Phylogenetic placement of a new paleoendemic pill scarab from the Udzungwa Mountains, Tanzania, triggers biogeographic interpretations (Coleoptera: Hybosoridae, Ceratocanthinae). Fragmenta entomologica 53(2): 283–298. DOI: 10.13133/2284-4880/547. Open access. Reference page.
Howden, H.F.; Gill, B.D. 1988: A fourth genus of unusually modified Ceratocanthinae (Coleoptera: Scarabaeidae) from South America. Canadian journal of zoology, 66: 2077–2079. ISSN: 1480-3283 DOI: 10.1139/z88-306
Howden, H.F.; Gill, B.D. 2000: Tribes of New World Ceratocanthinae, with keys to genera and descriptions of new species (Coleoptera: Scarabaeidae). Sociobiology, 35: 281–329. ISSN: 0361-6525 [not seen]
Paulian, R., 1978: Révision des Ceratocanthidae [Col. Scarabaeoidea] II - Les espèces orientales et australiennes. Annales de la Societé Entomologique de France (N.S.) 14: 479–514.
Paulian, R., 1981. Un nouveau Madrasostes des Iles Bismarck (Coleoptera, Ceratocanthidae). Revue Suisse de Zoologie 88: 343–344.
Poinar, G.O. & Ballerio, A. 2017. Remarks on some Ceratocanthinae (Coleoptera: Hybosoridae) in Dominican amber. Zootaxa 4286(1): 125–128. DOI: 10.11646/zootaxa.4286.1.9. Reference page.


Catalogue of Palearctic Coleoptera. Vol. 3 ed. I. Lobl, & A. Smetana, Apollo Books, Stenstrup, Denmark, 2006
ISBN 87-88757-59-5, p.96

Vernacular names
English: Pill scarab beetle / Ball beetle / Contractile Scarab beetle

Ceratocanthinae is a subfamily of the Ceratocanthidae which is a sister group of the Hybosoridae in which they are sometimes placed in the superfamily Scarabaeoidea. It includes three tribes comprising 43 genera and 366 species.


Ceratocanthinae are small sized beetles from 2.0 to 10.0 millimeters in length. Adult beetles can be found on the bark and branches of dead trees and on fungus.

Ceratocanthinae are relatively widespread. They can be found in Australian, Afrotropical, Indomalaysian, Neotropical, Nearctic, and Palaearctic regions.

The adults have been found to associate with termites and ants. Larvae live under bark and in burrows of bessbugs (Passalidae).

The subfamily Ceratocanthinae contains 43 genera:[1]

Acanthocerodes Péringuey, 1901
Afrocloetus Petrovitz, 1968
Aneilobolus Hesse, 1948
Anopsiostes Paulian, 1982
Astaenomoechus Martínez & Pereira, 1959
Aulisostes Howden & Gill, 2000
Baloghianestes Paulian, 1968
Besuchetostes Paulian, 1972
Callophilharmostes Paulian, 1968
Carinophilharmostes Paulian, 1968
Ceratocanthoides Paulian, 1982
Ceratocanthopsis Paulian, 1982
Ceratocanthus White, 1842
Chaetophilharmostes Paulian, 1977
Cloeotus Germar, 1843
Congomostes Paulian, 1968
Cryptophilharmostes Ballerio, 2000
Cryptosphaeroides Ballerio, 2009
Cyphopisthes Gestro, 1898
Ebbrittoniella Martínez, 1962
Eusphaeropeltis Gestro, 1898
Germarostes Paulian, 1982
Glyptogermarostes Ocampo & Ballerio, 2006
Goudotostes Paulian, 1979
Ivieolus Howden & Gill, 1988
Macrophilharmostes Paulian, 1978
Madrasostes Paulian, 1975
Martinezostes Paulian, 1982
Melanophilharmostes Paulian, 1968
Nesopalla Paulian & Howden, 1982
Oxymorostes Ballerio, 2009
Paulianostes Ballerio, 2000
Perignamptus Harold, 1877
Petrovitzostes Paulian, 1977
Philharmostes Kolbe, 1895
Pseudopterorthochaetes Paulian, 1977
Pseudosynarmostes Ballerio, 2009
Pterorthochaetes Gestro, 1898
Scarabaeinus Silvestri, 1940
Scarabatermes Howden, 1973
Synarmostes Germar, 1843
Trachycrusus Howden & Gill, 1995
Xenocanthus Howden & Gill, 1988


Ballerio, Alberto; Grebennikov, Vasily (14 June 2016). "Rolling into a ball: phylogeny of the Ceratocanthinae (Coleoptera: Hybosoridae) inferred from adult morphology and origin of a unique body enrollment coaptation in terrestrial arthropods" (PDF). Arthropod Systematics & Phylogeny. 74 (1): 23–52. eISSN 1864-8312. ISSN 1863-7221.

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