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MacLeay, 1819
Glaphyrid scarab beetles

by David C. Carlson

Glaphyrid beetles are active fliers during the day. Adults of many species are brightly colored and hairy and often possess markings and coloration resembling bees and bumblebees. They are strong fliers and are often observed hovering near flowers or foliage or flying over sandy areas.

Length 6.0-20.0 mm. Shape elongate. Color testaceous to black, often with metallic reflections; setae dense, moderately long, color variable (white, yellow, orange, red, brown, or black). Head deflexed. Antennae 9 or 10-segmented with 3-segmented, opposable club (all segments tomentose). Eyes with eucone ommatidia, completely or partially divided by a canthus. Clypeus usually simple, anterior margin with or without teeth. Labrum emarginate, truncate or rounded, produced beyond apex of clypeus, prominent. Mandibles produced beyond apex of labrum, prominent. Maxillae filiform, plumose or truncate, with 4 or 5-segmented palpi. Labium with 4-segmented palpi. Pronotum convex, usually subquadrate, often densely punctate and setose, without tubercles, ridges, horns, or sulci. Elytra elongate, often thin and dehiscent at apex, without striae, often setose. Pygidium usually visible beyond elytra. Scutellum exposed, U-shaped or triangular. Legs with procoxae conical or transverse, meso- and metacoxae transverse; mesocoxae separated or contiguous; protibiae dentate on outer margin, apex with one spur; meso- and metatibia generally simple but some with apical modifications (spines or emarginations), apex with 2 spurs; spurs mesad, adjacent (e.g., Lichnanthe) or separated by basal metatarsal segment (e.g., Arctodium ) ; tarsi 5-5-5, foretarsi modified medially (pectinate) in some Old World genera; claws equal in size with 1 tooth; empodium exposed beyond fifth tarsal segment, dorso-ventrally flattened, with 2 setae. Abdomen with 6 free sternites, usually with 8 pairs of spiracles; spiracles 1-6 or 1-7 situated in pleural membrane, spiracles 7 or 8 in tergites (Glayphyrinae and some Lichniinae) or spiracles 1-8 in pleural membrane (some Lichniinae). Wings well developed, M-Cu loop present with 1 apical, detached vein. Male genitalia with well sclerotized, strongly arched basal piece, basal piece large relative to parmeres; internal sac variable. Ovary with 6 ovarioles. References: Carlson 1980; Chapin 1938; d'Hotman and Scholtz 1990; Ritcher 1969; Ritcher and Baker 1974; Scholtz 1990.

Classification Status
The uniqueness of the glaphyrids has been recognized for a very long time, and the genera included in the group have changed little. However, the status of the group has been the subject of debate. Superfamily status was proposed by Machatschke (1959) but was not generally accepted. Workers have vacillated between using subfamily or family status, and familial status is now generally accepted (Browne and Scholtz 1995; d'Hotman and Scholtz 1990; Scholtz 1990).

The phylogenetic position of the Glaphyridae within the Scarabaeoidea has been discussed by numerous authors and is currently considered by most to be among the intermediate scarabaeoid families (Browne and Scholtz 1995; d'Hotman and Scholtz 1990; Scholtz 1990). Browne and Scholtz (1995) consider the Glaphyridae to be a monophyletic sister group of the trogid subgroup (Trogidae, Bolboceratinae [Geotrupidae] and Pleocomidae) based on characters of wing articulation.

Taxonomy of the world Glaphyridae is not well established. Comprehensive taxonomic treatments are available for Lichnanthe Burmeister (Carlson 1980), Anthypna Latreille (Endrödi 1952), and Pygopleurus Motschulsky (Petrovitz 1958). Most other genera have not been reviewed comprehensively. The taxonomy and nomenclature of the group were discussed by Chapin (1938) and Machatschke (1959).

The extreme color polymorphism exhibited by many species has resulted in a proliferation of form, variety, or color morph names for some genera, many of which are synonyms. The most recent world catalog for the family was Arrow (1912).

The family Glaphyridae includes eight genera and about 80 species worldwide. The subfamily Lichniinae (including three genera) is limited to western South America, and the subfamily Glaphyrinae (including five genera) is widely distributed in the Holarctic region. Four genera are found in the New World. Key to genera and subgenera: Chapin 1938. Key to U.S. species: Carlson 1980. Larvae: Ritcher 1966.

New World Subfamilies and Genera of Glaphyridae

Lichnanthe Burmeister 1844


Dasychaeta Erichson 1847
Lichnia Erichson 1847
Arctodium Burmeister 1844

Except for a few species, life histories of the glaphyrids are poorly documented. Adults are often brightly colored, densely setose, active diurnally, and strong fliers. Many species have colored setal bands on the abdomen and resemble various Hymenoptera (bumble bees and metallic bees). They have been observed frequenting flowers and foliage. Larval are free living in sandy areas (riparian and coastal dunes) where they feed on decaying leaf litter and detritis that is layered in the sand. Larvae of Lichnanthe vulpina (Hentz) may be a pest of cranberry bogs in the northeastern United States. References: Carlson 1977, 1980; Ritcher 1966; Westcott 1976.

Form scarabaeiform (C-shaped, cylindrical). Color bluish-white to yellow (prepupae) (except at caudal end which maybe darkened by accumulated feces). Head capsule heavily sclerotized, reddish-brown (Lichnanthe with conspicuous median, circular depression on frons). Antennae 4-segmented, third segment with small sensory pits. Ocelli present or absent. Frontoclypeal suture present. Labrum trilobed. Epipharynx with asymmetrical tormae not fused. Maxilla with galea and lacinia separate; maxillary palpi 4-segmented; labial palpi 2-segmented; maxillary and mandibular stridulatory areas present. Abdominal segments 1 to 8 with 3 dorsal annuli. Spiracles cribriform. Anal slit transverse, located caudally on dorsum of last abdominal segment. Legs well developed, 4-segmented, lacking stridulatory organs; claws present. References: Ritcher 1966; Scholtz 1990.

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

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

CARLSON, D. C. 1977.
Taxonomic revision of Lichnanthe Burmeister with studies on the biology of L. rathvoni (LeConte) (Coleoptera: Scarabaeidae). Ph.D. Thesis, Oregon State University, Corvallis.

CARLSON, D. C. 1980. Taxonomic revision of Lichnanthe Burmeister (Coleoptera: Scarabaeidae). Coleopterists Bulletin 34: 177-208.

CHAPIN, E. A. 1938. The nomenclature and taxonomy of the genera of the scarabaeid subfamily Glaphyrinae. Proceedings of the Biological Society of Washington 51: 79-86.

d'HOTMAN, D. and C. H. SCHOLTZ. 1990.
Comparative morphology of the male genitalia of derived groups of Scarabaeoidea (Coleoptera). Elytron 4: 3-39.

ENDRODI, S. 1952.
Monographie der Gattung Anthypna Latr. Folia Entomologica Hungarica (New Series) 5: 1-40.

MACHATSCHE, J. W. 1959. Untersuchungen uber die verwantschaftlichen Beziehungen der Gattungen der bisherigen Glaphyrinae (Coleoptera: Lamellicornia). Beitrage zur Entomologie 9: 528-545.

PETROVITZ, R. 1958. Das Subgenus Pygopleurus Motschulsky der Gattung Amphicoma Latreille (Col., Scarabaeidae, Glaphyrinae). Entomologisk Tidskrift 78: 38-68.

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

RITCHER, P. O. 1969. 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. 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: 1,027-1,066.

WESTCOTT, R. L. 1976. Observations on the biology and ethology of Lichnante rathvoni LeConte (Coleoptera: Scarabaeidae) with emphasis on mating. University of Idaho, Department of Entomology Anniversary Publication, No. 11: 85-90.