Overview
As the name implies, the geotrupids are burrowers in the the soil ("geos" from Greek meaning earth and "trypetes" from Greek meaning borer). In Europe, geotrupids are referred to as dor beetles. Adults of most species provision larvae in earthen burrows with dead leaves, cow dung, horse dung, or humus.

Description
Length 5.0-45.0 mm. Shape oval or round. Color yellowish, brown, orange-brown, reddish-brown, purple, brown, or black (with or without metallic reflections). Head not deflexed. Antennae 11-segmented with 3-segmented, opposable club (all segments tomentose). Eyes with eucone or exocone ommatidia, completely or partially divided by canthus. Clypeus often with tubercle or horn. Labrum truncate, prominent, produced beyond apex of clypeus. Mandibles produced beyond apex of labrum, prominent. Maxillae with 4-segmented palpi; labium with 3-segmented (Lethrinae) or 4-segmented (remaining taxa) palpi. Pronotum convex with base wider than or subequal to elytral base and with or without tubercles, ridges, horns, or sulci. Elytra convex, with or without striae. Pygidium concealed by elytra. Scutellum exposed, triangular. Legs with coxae transverse, mesocoxae separated or contiguous; protibiae serrate on outer margin, apex with 1 spur; meso- and metatibia with ridges, apex with 2 spurs; spurs mesad, adjacent (not separated by basal metatarsal segment); tarsi 5-5-5; claws equal in size, simple; empodium present, extending beyond fifth tarsal segment, with 2 setae. Abdomen with 6 free sternites; 7 functional abdominal spiracles situated in pleural membrane (spiracles 1-7) and vestigial spiracle in pleural membrane (spiracle 8) [Bolboceratinae] or with 8 functional spiracles situated in the pleural membrane (spiracles 1-7) and the 8th pair in the tergite [Geotrupinae]. Wings well developed, M-Cu loop and two apical detached veins present. Male genitalia variable. References: Howden 1955; Scholtz 1990; Scholtz and Browne 1996.

Classification Status
There is considerable debate concerning the classification of the Geotrupidae. The diversity in structure in both adults and larvae has led to differences of opinion regarding classification, evolution, and monophyly of the group and the genera assigned to it. There is evidence that the group (as defined in this work) includes two distinct lineages: the Bolboceratinae and Athyreinae forming one lineage, and the Geotrupinae, Taurocerastinae, and Lethrinae forming the other. Scholtz and Browne (1996) recently proposed the family Bolboceratidae for the former lineage. In addition to the possible division of the group into two families, the geotrupids are often considered as a subfamily of the family Scarabaeidae. In this volume, we follow Lawrence and Newton (1995) and consider the group a family. There is ample evidence that the group is not monophyletic, but the taxa that should be included, characters that support the groupings, and the ranking of the groupings are still debated. Past workers (e.g., Davis 1935; Ritcher 1947) have also included the genus Pleocoma (treated in the family Pleocomidae in this work) in the Geotrupidae based on the 11-segmented antenna. However, Pleocoma species differ from the geotrupids based on the open procoxal cavities (closed in geotrupids) and 4 to 7-segmented club (3-segmented in geotrupids). Howden (1982) hypothesized that the Geotrupinae (Geotrupini, Athyreini, Bolboceratini, and Lethrini) form a monophyletic lineage that is most closely related to the Pleocominae (=Pleocomidae) based on characters such as the form of the antennal club, adult provisioning, and diet. Alternatively, it was hypothesized (Browne and Scholtz 1995; Scholtz and Browne 1996; Scholtz and Chown 1995) that the geotrupids are a polyphyletic group: the Bolboceratidae (including Athyreinae) is part of a clade that includes the Glaphyridae, Trogidae, and Pleocomidae while the Geotrupidae (including Geotrupinae, Taurocerastinae, and Lethrinae) is part of a separate clade that includes the Ochodaeidae, Hybosoridae, and Ceratocanthidae. The placement of the unusual South American genera Taurocerastes and Frickius has also been a source of much debate, being placed with genera in the Geotrupini (e.g., Howden 1982; Howden and Peck 1987) or in its own group, the Taurocerastinae (e.g., Zunino 1984a; Browne and Scholtz 1995). Taxonomy of the world Geotrupidae is well established, primarily due to the prolific work of Howden (e.g., Howden 1955, 1964, 1974, 1980; Howden and Gill 1984).

Distribution
The family Geotrupidae includes 68 genera and about 620 species worldwide (Scholtz and Browne 1996). The subfamily Geotrupinae (including Geotrupini, Taurocerastini, and Lethrini) is distributed in the Holarctic region. In the New World, the Geotrupinae occurs from Canada to El Salvador. The subfamily Bolboceratinae (including Bolboceratini and Athyreini) is best represented in Australia, Africa, and South America. In the New World, the Bolboceratinae is distributed from Canada to Central America, and the Athyreinae is distributed from Mexico to South America. The subfamily Taurocerastinae is distributed in southern South America. Sixteen genera of geotrupids occur in the New World. Keys to genera and species: Howden 1955; Olson et al. 1954; Howden 1964. Reference: Scholtz and Browne 1996; Zunino 1984b. Biology: Howden 1955. Larvae: Ritcher 1966.

New World Subfamilies and Genera

Subfamilies	Tribes	Genera
BOLBOCERATINAE	BOLBOCERATINI	Bolbapium Bolbelasmus Bolboceras Bolbocerastes Bolbocerosoma Bolborhinum Bolborhombus Bolbothyreus Bradycinetulus Eucanthus Halffterobolbus Parabolbapium
	ATHYREINI	Athyreus Neothyreus Parathyreus
GEOTRUPINAE	GEOTRUPINI	Ceratophyus Geotrupes Mycotrupes Peltotrupes
	TAUROCERASTINI	Frickius Taurocerastes

Ecology
Life histories of the geotrupids are diverse, and food habits vary from saprophagous to coprophagous and mycetophagous, and some adults apparently do not feed. Adults of most species are secretive, living most of their life in burrows. Although adults do not tend larvae, adults provision food for larvae in brood burrows. There is overlapping of generations in some species. For example, in the genus Bolboceras, eggs, larvae, pupae, and adults have been observed together in a single branching burrow. Adults dig vertical burrows (15-200 cm in depth) and provision larval cells with dead leaves, cow dung, horse dung, or humus. Burrows of some species extend to a depth of 3.0 meters. In restricted habitats, some species are semi-colonial. Geotrupids are not of economic importance, although their burrowing has occasionally caused damage in lawns. Adults of many geotrupids are nocturnal and are frequently attracted to lights at night. Some species are attracted to fermenting malt and molasses baits. Most adults and larvae stridulate. The biology and behavior of many species, especially the Bolboceratinae, are poorly known. References: Howden 1955; Woodruff 1973.

Larvae
Form scarabaeiform (C-shaped, cylindrical). Color creamy-white or yellow (except at caudal end which may be darkened by accumulated feces). Cranium heavily sclerotized, brown to dark brown. Antennae 3-segmented, penultimate segment bearing 1 or more distal sense organs, last segment reduced in diameter. Lateral ocelli absent. Frontoclypeal suture absent (Geotrupinae and Bolboceratinae) or present (Taurocerastinae). Labrum at apex with 3 weak lobes or rounded. Epipharynx usually trilobed with symmetrical tormae. Maxilla with galea and lacinia distinctly separate; maxillary stridulatory area with teeth; maxillary palpi 4-segmented. Abdominal segments 3 to 7 with 2 annuli, each with one or more transverse rows of short setae. Spiracles cribriform (Geotrupinae and Lethrinae) or biforous (Bolboceratinae and Taurocerastinae). Venter of last abdominal segment V-shaped or Y-shaped, surrounded by fleshy lobes in some taxa. Legs 4-segmented (some Bolboceratinae) or pro- and mesothoracic legs 3-segmented and metathoracic leg reduced in size and 2-segmented (Geotrupinae and Taurocerastinae); stridulatory apparatus on meso- and metathoracic legs present (some Geotrupinae, some Bolboceratinae, some Taurocerastinae) or absent (some Geotrupinae, some Bolboceratinae, and Lethrinae); claws absent (Geotrupinae, Taurocerastinae, and some Bolboceratinae) or present (Eucanthus and Bolbocerosoma). References: Ritcher 1966; Scholtz 1990; Scholtz and Browne 1996.

References Cited
BROWNE, D. J. and C. H. SCHOLTZ. 1995. Phylogeny of the families of the Scarabaeiodea (Coleoptera) based on characters of the hindwing articulation, hindwing base and wing venation. Systematic Entomology 21: 145-173.

DAVIS, A. C. 1935. A revision of the genus Pleocoma. Bulletin of the Southern California Academy of Science 33: 123-130.

HOWDEN, H. F. 1955. Biology and taxonomy of North American beetles of the subfamily Geotrupinae with revisions of the genera Bolbocerosoma, Eucanthus, Geotrupes and Peltotrupes (Scarabaeidae). Proceedings of the United States National Museum 104: 151-319.

HOWDEN, H. F. 1964. The Geotrupinae of North America and Central America. Memoires of the Entomological Society of Canada 39: 1- 91.

HOWDEN, H. F. 1974. Additional records and descriptions of North and Central American Geotrupinae (Coleoptera, Scarabaeidae). Canadian Journal of Zoology 52: 567-573.

HOWDEN, H. F. 1980. Key to the Geotrupini of Mexico and Central America, with the description of a new species (Scarabaeidae, Geotrupinae). Canadian Journal of Zoology 58: 1959-1963.

HOWDEN, H. F. 1982. Larval and adult characters of Frickius Germain, its relationship to the Geotrupini, and a phylogeny of some major taxa in the Scarabaeoidea (Insecta: Coleoptera). Canadian Journal of Zoology 60: 2713-2724.

HOWDEN, H. F. 1984. A Catalog of the Coleoptera of America North of Mexico. Family Scarabaeidae. Subfamily: Geotrupinae. United States Department of Agriculture, Agriculture Handbook 529-34a. 17 pp.

HOWDEN, H. F. and B. GILL. 1984. Two new species of Neoathyreus Howden and Martínez from Costa Rica with distribution notes on other Athyreini from Mexico and Central America (Coleoptera: Scarabaeidae). Canadian Entomologist 116: 1637-1641.

HOWDEN, H. F and S. B. PECK. 1987. Adult habits, larval morphology, and phylogenetic placement of Taurocerastes patagonicus Philippi (Scarabaeidae: Geotrupinae). Canadian Journal of Zoology 65: 329-332.

LAWRENCE, J. F. and A. F. NEWTON, JR. 1995. Families and subfamilies of Coleoptera (with selected genera, notes, and references and data on family-group names), pp. 779-1006. In J. Pakaluk and S. A. Slipinski (eds.), Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warszawa, Poland.

OLSON, A. L., T. H. HUBBELL and H. F. HOWDEN. 1954. The beetles of the genus Mycotrupes (Coleoptera: Scarabaeidae: Geotrupidae). Miscellaneous Publications of the Museum of Zoology, University of Michigan 84: 1-59.

RITCHER, P. O. 1947. Larvae of Geotrupinae with keys to tribes and genera (Coleoptera: Scarabaeidae). Bulletin 506 of the Kentucky Agricultural Experiment Station, University of Kentucky, Lexington. 27 pp.

RITCHER, P. O. 1966. White Grubs and Their Allies: A Study of North American Scarabaeioid Larvae. Oregon State University Press, Corvallis. 219 pp.

SCHOLTZ, C. H. 1990. Phylogenetic trends in the Scarabaeoidea (Coleoptera). Journal of Natural History 24: 1027-1066.

SCHOLTZ, C. H. and D. J. BROWNE. 1996. Polyphyly in the Geotrupidae (Coleoptera: Scarabaeoidea): a case for a new family. Journal of Natural History 30: 597-614.

SCHOLTZ, C. H. and S. L. CHOWN. 1995. The evolution of habitat use and diet in the Scarabaeoidea: a phylogenetic approach, pp. 355-374. In J. Pakaluk and S. A. Slipinski (eds.), Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warszawa, Poland.

WOODRUFF, R. E. 1973. Arthropods of Florida and Neighboring Land Areas. Volume 8. The Scarab Beetles of Florida (Coleoptera: Scarabaeidae). Part 1. The Laparosticti (Subfamilies: Scarabaeinae, Aphodiinae, Hybosorinae, Ochodaeinae, Geotrupinae, Acanthocerinae). Florida Department of Agriculture and Consumer Services, Contribution No. 260 Bureau of Entomology, Gainesville, Florida. 220 pp.

ZUNINO, M. 1984a. Analisi sistematica e zoogeografica della sottofamiglia Taurocerastinae Germain (Coleoptera, Scarabaeoidea: Geotrupidae). Bollettino del Museo Regionale di Scienze Naturali, Torino 2: 445-464.

ZUNINO, M. 1984b. Sistematica generica dei Geotrupinae (Coleoptera, Scarabaeoidea: Geotrupidae), filogenesi della sottofamiglia e considerazioni biogeografiche. Bollettino del Museo Regionale di Scienze Naturali, Torino 2: 9-162.