Characteristics
Clypeus expanded, covering mouthparts. Mandibles lamelliform, mostly membranous,
with only outer margin hardened. Antennae with 8 or 9 segments, club with
3. Middle coxae widely separated. Posterior tibiae nearly always with single
apical spur; if two spurs present (i.e., Melanocanthon), spurs mesad,
adjacent (not separated by basal metatarsal segment). Elytra exposing pygidium,
6 visible, fused abdominal sternites. Anterior tarsi sometimes absent either
in female or in both sexes.
This subfamily was divided by Cambefort (1991) into 12 tribes, ten of which
are represented in the New World and two of which are limited to the Old World
(Gymnopleurini and Scarabaeini). Members of this subfamily are commonly called
dung beetles. Although many species feed on mammalian dung, others specialize
to varying degrees upon the dung of other vertebrates and invertebrates, as
well as on carrion, mushrooms, rotting fruit, and other decomposing plant
material. The World fauna includes slightly over 5,000 described species in
234 genera, with close to 1,800 of these species belonging to the genus Onthophagus.
Biology: Halffter and Matthews 1966, Halffter and Edmonds 1982, Hanski and
Cambefort 1991. Keys to larvae: Ritcher 1966; Edmonds and Halffter 1978.
New World Tribes
Canthonini
Coprini
Dichotomini
Eucraniini
Eurysternini
Oniticellini
Onitini
Onthophagini
Phanaeini
Sisyphini
TRIBE CANTHONINI
Characteristics
Body size small to large (length 2-25 mm), shape round or oval. Head and pronotum
lacking horns or carinae. Antenna with 9 segments. Middle and posterior tibiae
slender, curved, only slightly expanded apically. Sexual dimorphism minimal.
The New World has 340
species in 27 genera, which equates to a third of the World fauna. Adults
of most species are ball-rollers, and they shape carrion or dung into balls
that are rolled away and buried at a distance from the food source. Key to
genera: Halffter and Martínez 1977. Biology: Gordon and Cartwright
1974; Halffter and Halffter 1989.
TRIBE COPRINI
Characteristics: Body 8-20 mm in length, elongate, robust,
convex, generally black in color. Head or pronotum often with horns or tubercles,
especially in males. Elytra with 9 discal striae. Scutellum hidden. Antenna
with 9 segments. Middle and posterior tibia strongly expanded apically.
The Coprini is a relatively small tribe with ten genera and slightly under
400 species worldwide. Copris is the only representative of the tribe
in the New World.
TRIBE DICHOTOMINI
Characteristics: Body small and compact or large and robust
(length 4-30 mm). Elytra with 7 or 8 discal striae. Scutellum hidden. Antenna
with 9 segments. Middle and posterior tibia strongly expanded apically.
The Dichotomini is a diverse tribe with approximately 30 genera and 750 species
worldwide. The New World has a large proportion of this diversity with close
to 600 species in 23 genera.
TRIBE ONITICELLINI
Characteristics: Body 7-14 mm in length, elongate, usually
flattened. Pronotum without basal pits near midline. Scutellum visible. Antenna
with 8 segments.
Most of the 14 genera and 165 species in the tribe are found in Africa and
Asia although Anoplodrepanus is restricted to the West Indies (Simonis
1981). Key to adults: Janssens 1953. Biogeography: Zunino 1982.
TRIBE ONITINI
Characteristics: Body 10-20 mm in length, oblong, robust.
Pronotum with pit on each side of midline near base. Scutellum small but visible.
Elytron with prominent lateral carina. Antenna with 9 segments. Anterior tarsi
lacking in both sexes. Middle and posterior tibiae strongly expanded at apex.
The Onitini is an Old World group with 18 genera and about 200 species.
TRIBE ONTHOPHAGINI
Characteristics: Body 2-12 mm in length, oval, convex. Head
and/or pronotum often with horns or carinae. Scutellum hidden. Elytra with
7 discal striae. Third segment of labial palpus small or inconspicuous. Antenna
with 9 segments. Tarsi and tarsal claws present on all legs. Middle and posterior
tibiae broadly expanded at apex.
Sexual dimorphism is
usually well developed in Onthophagus species. Males typically have
large horns on the head and/or pronotum (females with only carinae or rudimentary
horns) or males have elongated foretibiae with a long setal brush at the apex.
The world fauna includes over 2,200 species in 34 genera.
TRIBE PHANAEINI
Characteristics: Body length 12-25 mm, robust, often with
metallic coloration. Antenna with 9 segments, basal segment of club cup-shaped,
enclosing distal 2 segments. Tarsal claws absent. Anterior tarsi lacking in
males, present or absent in females.
This tribe is restricted
to the New World with most of the 9 genera and 150 species found in the Neotropics.
Many of the species exhibit strong sexual dimorphism with the males often
possessing long horns on the head and/or pronotum. Zunino (1985) considered
the Phanaeini to be a subtribe of the Onitini. Key to genera: Edmonds 1972,
1994.
TRIBE SISYPHINI
Characteristics: Body length 6-10 mm, oval, compressed laterally
with sides of pronotum and metasternum vertical. Color reddish brown to gray,
surface matte, with setae on dorsum. Scutellum hidden. Antenna with 8 segments.
Middle coxae very widely separated, forming lateral margin of body. Middle
and posterior tibiae elongate, curved, barely enlarged at apex.
This tribe includes three genera of ball-rolling dung beetles. Most of the
60 species in the tribe are found in Africa and Asia, with only a few representatives
in Europe and the New World.
References Cited
CAMBEFORT, Y. 1991. From saprophagy to coprophagy. In Hanski,
I. and Y. Cambefort (eds.), Dung Beetle Ecology, pages 22-35. Princeton University
Press, Princeton, NJ. 481 pp.
EDMONDS, W. D. 1972. Comparative skeletal morphology, systematics
and evolution of the Phanaeine dung beetles (Coleoptera: Scarabaeidae). University
of Kansas Science Bulletin 49: 731-874.
EDMONDS, W. D. 1994. Revision of Phanaeus Macleay, a New
World genus of Scarabaeine dung beetles (Coleoptera: Scarabaeidae, Scarabaeinae).
Natural History Museum of Los Angeles County Contributions in Science 443:
1-105.
EDMONDS, W. D. and G. HALFFTER. 1978. Taxonomic review of
immature dung beetles of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae).
Systematic Entomology 3: 307-331.
GORDON, R. D. and O. L. CARTWRIGHT. 1974. Survey of food preferences
of some North American Canthonini (Coleoptera: Scarabaeidae). Entomological
News 85: 181-185.
HALFFTER, G. and W. D. EDMONDS. 1982. The Nesting Behavior
of Dung Beetles (Scarabaeinae). An Ecological and Evolutive Approach. Instituto
de Ecología, México, D.F. 176 pp.
HALFFTER, G. and V. HALFFTER. 1989. Behavioral evolution
of the non-rolling roller beetles (Coleoptera: Scarabaeidae; Scarabaeinae).
Acta Zoologica Mexicana (N.S.) 32: 1-53.
HALFFTER, G. and A. MARTINEZ. 1977. Revision monographica de los
Canthonina americanos, IV parte. Clave para generos y subgeneros. Folia Entomologica
Mexicana 38: 29-107.
HALFFTER, G. and E. G. MATTHEWS. 1966. The natural history
of dung beetles of the subfamily Scarabaeinae (Coleoptera: Scarabaeidae).
Folia Entomologica Mexicana 12-14: 1-312.
HANSKI, I. and Y. CAMBEFORT (eds.). 1991. Dung Beetle Ecology.
Princeton University Press, Princeton, NJ. 481 pp.
JANSSENS, A. 1953. Oniticellini (Coleoptera Lamellicornia). In Exploration
du Parc National de l'Upemba, Mission G.F. De Witte. Institut des Parcs Nationaux
du Congo Belge. Fascicle 11: 1-118.
RITCHER, P. O. 1966. White Grubs and Their Allies. A Study
of North American Scarabaeoid Larvae. Oregon State University Press, Corvallis,
OR. 219 pp.
SIMONIS, A. 1981. Anoplodrepanus, nuovo genere di
Oniticellini (Coleoptera, Scarabaeidae). Bollettino del Museo di Zoologia
dell'Universiti di Torino 7: 87-94.
ZUNINO, M. 1982. Aspectos taxonomicos y biogeográficos
del poblamiento americano de Oniticellini (Coleoptera, Scarabaeidae). Acta
del VIII Congreso Latinoamericano de Zoología, Mérida, Venezuela
2: 1083-1089.
ZUNINO, M. 1985. Las relaciones taxonomicas de los Phanaeina
(Coleoptera, Scarabaeinae) y sus implicaciones biogeographicas. Folia Entomologica
Mexicana 64: 101-115.