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
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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,
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CARLSON, D. C. and P. O. RITCHER. 1974. A new genus of Ochodaeinae
and a description of the larva of Pseudochodaeus estriatus
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CARLSON, D. C. 1975. Taxonomic characters of the genus Ochodaeus
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DELOYA, C. 1988. Coleopteros lamelicornios asociados a depositos
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