Overview
The Ochodaeidae is a relatively small, widely distributed family. Adults are small, mostly reddish-brown, non-metallic beetles that are predominately active at night. They are most often collected at lights, sometimes in large numbers. Adults of a few species are active during the day. Most species prefer sandy areas, and many stridulate. Woodruff (1973) suggested that adults may spend the daylight hours in subterranean burrows, and that they might feed on fungi. Little else is known about the habits of adult or immature stages.

Description
Length 3.0-10.0 mm. Shape elongate and convex. Color yellowish, brown, reddish-brown, brown, or black; infrequently bicolorous. Head not deflexed. Antennae 9 or 10-segmented, with 3-segmented, opposable, club (all segments tomentose). Eyes with eucone ommatidia, not divided by canthus. Clypeus simple or with tubercle(s) on anterior margin. Labrum produced beyond apex of clypeus, often bilobed and emarginate, prominent. Mandibles produced beyond apex of labrum, prominent. Maxillae with 4 or 5-segmented palpi; labium with 3 or 4-segmented palpi. Pronotum convex, subquadrate; usually punctate and setose; without tubercles, ridges, horns, or sulci. Elytra convex, with or without striae, often punctate or granulate and setose, sometimes smooth. Scutellum exposed, triangular. Pygidium exposed or concealed by elytra. Legs with procoxae conical or transverse; meso- and metacoxae transverse, mesocoxae separated or contiguous; protibia dentate on outer margin, apex with 1 spur; meso- and metatibia with 2 apical spurs; 1 mesotibial spur pectinate/crenulate, pro- and metatibial spur crenulate/pectinate in some; spurs mesad, adjacent (not separated by basal metatarsal segment); tarsi 5-5-5; claws equal in size, simple; empodium absent. Abdomen with 6 visible sternites; stridulatory peg present in some; 8 functional abdominal spiracles with spiracles 1-6 situated in pleural membrane and spiracles 7-8 situated in tergites; tergite, pleurite, and sternite of female 9th abdominal segment visible as distinct sclerites. Wings well developed, M-Cu loop and two apical detached veins present. Male genitalia with divided basal piece, symmetrical parameres, partially sclerotized membranous median lobe and large internal sac; internal sac armed with spines, hooks, and toothed sclerites in many. Female genitalic hemisternites with styli present. Six ovarioles per ovary. References: Browne and Scholtz 1995; Carlson 1975; Carlson and Ritcher 1974; Ritcher 1969a, b; Ritcher and Baker 1974; Scholtz 1990; Scholtz et al. 1988.

Classification Status
The ochodaeids have long been recognized as a distinct group. Their status has vacillated between the subfamily and family levels, but more recent works have tended to favor familial status (Scholtz 1990; Scholtz and Evans 1987; Scholtz and Chown 1995; Scholtz et al. 1988). Two subfamilies, Chaetocanthinae Scholtz and Ochodaeinae Mulsant and Rey are currently recognized (Lawrence and Newton 1995; Scholtz et al. 1988).

The phylogentic position of the Ochodaeidae within the Scarabaeoidea has been discussed by numerous authors. Current views consider the ochodaeids to be an "intermediate" scarabaeoid family (Browne and Scholtz 1995; Scholtz 1990). A close relationship with the Hybosoridae was suggested (Carlson and Ritcher 1974). Recent studies suggest a close relationship between the Ochodaeidae and Hybosoridae, and consider the Ochodaeidae to be the sister group to the Hybosoridae (d'Hotman and Scholtz 1990; Scholtz et al. 1988; Browne and Scholtz 1995). The presence of a pectinate/crenulate mesotibial spur in the Ochodaeidae is unique among the Scarabaeoidea and, according to Scholtz (1990), establishes the monophyly of the group. Within the Ochodaeidae, d'Hotman and Scholtz (1990) consider Ochodaeus Serville and Codocera Eschscholtz to be the most primitive based on the structure of the male genitalia, and Chaetocanthus Peringuey and Namibiotalpa Scholtz and Evans are considered the most derived.

Taxonomy of the world Ochodaeidae is fairly well established for the Chaetocanthinae Scholtz and the tribe Enodognathini Scholtz (Ochodaeinae) (Scholtz and Evans 1987; Scholtz et al. 1988). The tribe Ochodaeini, however, has recently undergone the creation of new genera. Nikolajev (1995) created two new Nearctic genera, but did not fully assign species to the new genera. Paulsen (2007) created two additional genera for the Nearctic fauna and assigned all Nearctic species to the appropriate genera. Nikolajev (1995) and Paulsen (2007) based these genera on characters that are well-recognized as separating groups within Ochodaeidae (Carlson 1975; Fall 1909; Horn 1876). The Palearctic and Neotropical members of the Ochodaeini are currently not placed into the appropriate genera, with many species remaining in Ochodaeus. All described Neotropical species of Ochodaeinae belong in either Neochodaeus or Parochodaeus, thus the genus Ochodaeus is not listed here as occurring in the New World fauna. Paulsen created the chaetocanthine genus Gauchodaeus for the first member of that subfamily known from South America (Paulsen & Ocampo 2012).

Distribution
The family Ochodaeidae includes thirteen extant and two fossil genera and about 80 species worldwide (Arrow 1912; Nikolayev 1995; Scholtz and Evans 1987; Scholtz et al. 1988; Paulsen 2007; Paulsen & Ocampo 2012). In the New World, the family includes seven genera.

New World Subfamilies and Genera

Ochodaeinae
Codocera Eschscholtz
Cucochodaeus Paulsen
Neochodaeus Nikolajev
Parochodaeus Nikolajev
Xenochodaeus Paulsen

Chaetocanthinae
Pseudochodaeus Carlson & Ritcher
Gauchochodaeus Paulsen

Ecology
Little is known about the biology of Ochodaeidae. There are few recorded observations of adult or larval habits except that adults of most species are nocturnal and are attracted to light, sometimes in large numbers. Adults of a few species are diurnally active and have been collected infrequently with sweep nets, malaise traps, or seining flumes. Adults of one species have been found associated with detritus deposits of harvester ants. Adults of another species were found to have basidiomycete spores in the midgut and hindgut. References: Arrow 1912; Carlson 1975; Carlson and Ritcher 1974; Deloya 1988.

Larvae
Form scarabaeiform (C-shaped, cylindrical). Color whitish (except at caudal end which may be darkened by accumulated feces). Cranium sclerotized, yellow-brown to red-brown. Antennae 3 or 4-segmented, penultimate and apical segment with sense organs. Ocelli absent. Frontoclypeal suture absent. Labrum trilobed. Epipharynx with complete, symmetrical zygum; tormae fused and symmetrical. Maxilla with galea and lacinia distinctly separate. Maxillary palpi 4-segmented; maxillary and mandibular stridulatory areas present. Abdominal segments 1-7 with 3 dorsal lobes, anterior 2 lobes with transverse row of setae. Spiracles cribriform, inconspicuous. Anal opening Y-shaped, surrounded by fleshy lobes. Legs well developed, 4-segmented, with well-developed claws, stridulatory apparatus lacking. References: Carlson and Ritcher 1974; Medvedev 1960; Scholtz 1990.

References Cited
ARROW, G. J. 1912. Scarabaeidae: Pachypodinae, Pleocominae, Aclopinae, Glaphyrinae, Ochodaeinae, Orphninae, Idiostominae, Hybosorinae, Dynamopinae, Acanthocerinae, Troginae. Coleoperorum Catalogus 19: 1-66.

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 20: 145-173.

CARLSON, D. C. and P. O. RITCHER. 1974. A new genus of Ochodaeinae and a description of the larva of Pseudochodaeus estriatus (Schaeffer) (Coleoptera: Scarabaeidae). Pan Pacific Entomologist 50: 99-110.

CARLSON, D. C. 1975. Taxonomic characters of the genus Ochodaeus Serville with descriptions of two new species in the O. pectoralis LeConte species complex (Coleoptera: Scarabaeidae). Bulletin of the Southern California Academy of Sciences 74: 49-65.

DELOYA, C. 1988. Coleopteros lamelicornios asociados a depositos de detritos de Atta mexicana (Smith) (Hymenoptera): Formicidae) en el sur del estado de Morelos, Mexico. Folia Entomologica Mexicana 75: 77-91.

d'HOTMAN, D. and C. H. SCHOLTZ. 1990. Phylogenetic significance of the structure of the external male genitalia in the Scarabaeoidea (Coleoptera). Republic of South Africa, Department of Agricultural Development, Entomology Memoir No. 77: 1-51.

FALL, H. F. 1909. A short synopsis ofthe species of Ochodaeus inhabiting the United States. Journal of the New York Entomological Society 17: 30-38.

HORN, G. H. 1876. Revision of the United States species of Ochodaeus and other genera of Scarabaeidae. Transactions of the American Entomological Society 5: 177-198.

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.

MEDVEDEV, S. I. 1960. Descriptions of the larva of eight species of lamelicorn beetles from the Ukraine and Central Asia. Zoologicheskiy Zhurnal 39: 381-393.

NIKOLAJEV, G. V. 1995. New data on the systematics of the subfamily Ochodaeinae (Coleoptera, Scarabaeidae). Zoologicheskiy Zhurnal 74: 72-82.

PAULSEN, M. J. 2007. Nomenclatural changes in the Nearctic Ochodaeinae and description of two new genera (Coleoptera: Scarabaeoidea: Ochodaeidae) Insecta Mundi 0021: 1-13. Download PDF

PAULSEN, M.J., and F.C. Ocampo. 2012. The Ochodaeidae of Argentina (Coleoptera, Scarabaeoidea). ZooKeys 174: 7-30. Download PDF

RITCHER, P. O. 1969a. Spiracles of adult Scarabaeoidea (Coleoptera) and their phylogenetic significance. I. The abdominal spiracles. Annals of the Entomological Society of America 62: 869-880.

RITCHER, P. O. 1969b. Spiracles of adult Scarabaeoidea (Coleoptera) and their phylogenetic significance. II. The thoracic spiracles and adjacent sclerites. Annals of the Entomological Society of America 62: 1388-1398.

RITCHER P. O. and C. W. BAKER. 1974. Ovariole numbers in Scarabaeoidea (Coleoptera: Lucanidae, Passalidae, Scarabaeidae). Proceedings of the Entomological Society of Washington 76: 480-494.

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

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.

SCHOLTZ, C. H. and A. V. EVANS. 1987. A revision of the African Ochodaeidae (Coleoptera: Scarabaeoidea). Journal of the Entomological Society of Southern Africa 50: 399-426.

SCHOLTZ, C. H., D. D'HOTMAN, A. V. EVANS and A. NEL. 1988. Phylogeny and systematics of the Ochodaeidae (Insecta: Coleoptera: Scarabaeoidea). Journal of the Entomological Society of South Africa 51: 207-240.

WOODRUFF, R. E. 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, 8: 1-220.