Overview
Stag beetles range in size from less than 1 cm to 9 cm. Many adult males possess
greatly enlarged, curving mandibles that are sometimes used in combat with male
opponents during fights to establish dominance. In the males, development of
the mandibles is usually allometric, that is, the size of the mandibles is proportional
to the size of the body. Those males with the largest mandibles are referred
to as "male majors" and those with the smallest mandibles are called "male
minors."
Description
Length 8.0-90.0 mm. Shape usually weakly convex, subdepressed, or cylindrical,
elongate. Color usually black to reddish brown, sometimes testaceous or metallic.
Scales often present on dorsum. Head prognathus, not deflexed. Antennae geniculate
or straight, 10-segmented, with 3-7 segmented club; first segment elongate and
often subequal to remaining segments. Eyes with eucone or acone ommatidia; eye
canthus present or absent. Clypeus and labrum fused to frons. Mandibles produced
beyond apex of labrum, prominent (males often with large, curved, elongate mandibles).
Maxillae with 4-segmented palpi; labium with 3-segmented palpi. Pronotum variably
convex, with or without tubercles. Elytra weakly convex, with or without impressed
striae. Scutellum exposed, triangular or parabolic. Pygidium concealed by elytra
or only weakly exposed. Legs with coxae transverse, mesocoxae separated; protibiae
dentate on outer margin, apex with one 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 weakly beyond
fifth tarsal segment or extending nearly one half claw length, with 2 to several
setae. Abdomen with 5 or 6 visible sternites; 8 functional abdominal spiracles
situated in pleural membrane. Wings usually well developed, with M-Cu loop and
two, apical, detached veins. Male genitalia with paired symmetrical parameres
and a median lobe. Median lobe often terminating in a ribbon-like permanently
everted internal sac. References: Didier and Seguy 1953; Scholtz 1990.
Classification Status
The family Lucanidae has long been considered one of the most primitive groups
in the Scarabaeoidea (Crowson 1967; Howden 1982; Ritcher 1966), and scarabaeoid
classifications and evolutionary hypotheses have generally regarded the Lucanidae
as basal to all scarabaeoids (Howden 1982; Iablokoff-Khnzorian 1977; Lawrence
and Newton 1995). However, based on comparison with "primitive" scarabaeoid
groups, Scholtz et al. (1994) hypothesized that the scarabaeoid family Glaresidae,
rather than the Lucanidae, is the most primitive scarabaeoid. According to this
hypothesis, the Lucanidae is a member of a clade that includes the Passalidae,
Diphyllostomatidae, Glaphyridae, Trogidae, Pleocomidae, and Bolboceratinae (Geotrupidae).
Prior to the taxonomic elevation of the genus Diphyllostoma to the family
Diphyllostomatidae (Holloway 1972), the Lucanidae was hypothesized to be most
closely related to the Passalidae (Howden 1982). Based on shared characters, it
is now thought that the Lucanidae is most closely related to the Diphyllostomatidae
(Caveney 1986; Browne and Scholtz 1995).
The world Lucanidae (about 800 species) have been treated in checklists by Benesh
(1960) and Maes (1992) and in illustrated catalogs by Didier and Seguy (1953)
and Mizunuma and Nagai (1994). The latter is spectacular for its colored plates
of the world fauna. Benesh (1960) recognized eight subfamilies, four of which
occurred in the United States. Howden and Lawrence (1974) and Kikuta (1986) proposed
significant changes of genera within subfamilies. Based on the work of Holloway
(1960, 1968, 1969), the most recent subfamilial classification is followed here,
with four subfamilies now recognized as occurring in North America: Aesalinae,
Syndesinae, Platycerinae, and Lucaninae. One member of the remaining lucanid subfamily,
Lampriminae, occurs in South America. The subfamily Penichrolucaninae was formerly
used for two genera of aberrant termitophilous lucanids, but these are now considered
to be derived members of Lucaninae (Bartolozzi 1989).
New World subfamilies and genera
(click on generic names for information)
Distribution The world
fauna consists of about 800 species (Mizunuma and Nagai 1994) with more species
found in Asia than in other areas. Keys to adults: Benesh 1946; Howden and Lawrence
1974; Ratcliffe 1991. Keys to larvae: Ritcher 1966. Catalogs of North America
species: Benesh 1960; Blackwelder and Arnett 1974. Biology: Milne 1933; Hoffman
1937, Paulsen 2005.
Ecology
Lucanids are usually associated with decaying wood and logs in coniferous and
deciduous forest habitats. Adults of some species are attracted to lights at night
and some feed at sap flows from fluxing trees. Adults of some of the smaller species
have been observed feeding on flowers. The eggs are customarily laid in crevices
in bark or logs, and the larvae feed on decaying wood and are not economically
injurious. The larvae resemble those of Scarabaeidae, but in lucanids the anal
opening is longitudinal or Y-shaped, whereas in scarabs it is usually transverse
or occasionally Y-shaped. References: Ratcliffe 1991.
Larvae
Form scarabaeiform (c-shaped, subcylindrical). Color creamy-white or yellowish
(except at caudal end which may be darkened by accumulated feces). Cranium heavily
sclerotized, lightly pigmented. Antennae 3-4 segmented, last segment greatly reduced
in size. Ocelli absent (present in Platycerus). Frontoclypeal suture
present. Labrum at apex rounded or weakly lobed. Epipharynx rounded or lobed,
with symmetrical tormae. Maxilla with galea and lacinia distinctly separate; maxillary
stridulatory teeth absent (present in Platycerus); maxillary palpus 4-segmented.
Mandibles elongate, asymmetrical. Abdominal segments 3-7 with 2 annuli, each with
1 or more transverse rows of short setae. Spiracles cribriform. Anal opening Y-shaped
or longitudinal, surrounded by 2 fleshy lobes. Legs 4-segmented. Stridulatory
apparatus on meso- and metathoracic legs present; claws present. References: Ritcher
1966; Scholtz 1990.
References Cited BARTOLOZZI, L. 1989. Taxonomic revue of the genus Penichrolucanus
Deyrolle 1863 (Coleoptera Lucanidae) with notes on its biology. Tropical Zoology
2: 37-44.
BENESH, B. 1946. A systematic revision of the Holarctic genus Platycerus
Geoffroy. Transactions of the American Entomological Society 72: 139-202.
BENESH, B. 1960. Coleopterorum Catalogus Supplementa, pars 8:
Lucanidae. W. Junk, Gravenhage, Netherlands.
BLACKWELDER, R. E. and R. H. ARNETT, JR. 1974. Checklist of the
beetles of Canada, United States, Mexico, Central America and the West Indies.
Volume 1, Part 3. The scarab beetles, ant-loving beetles, clown beetles, and related
groups (red version). The Biological Research Institute of America, Inc., Latham,
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BROWNE, D. J. and C. H. SCHOLTZ. 1995. Phylogeny of the families of the
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base and wing venation. Systematic Entomology 21: 145-173.
CAVENEY, S. 1986. The phylogenetic significance of ommatidium
structure in the compound eyes of polyphagan beetles. Canadian Journal of Zoology
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CROWSON, R. A. 1967. The natural classification of the families of Coleoptera.
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1-223.
HOFFMAN, C. H. 1937. Biological notes on Pseudolucanus placidus
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HOLLOWAY, B. A. 1969. Further studies on generic relationships
in Lucanidae (Insecta: Coleoptera) with special reference to the ocular canthus.
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IABLOKOFF-KHNZORIAN, S. M. 1977. Über die Phylogenie der
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KIKUTA, T. 1986. On the higher taxa of the stag beetle family Lucanidae,
pp. 131-138. In: J. Aoki (editor). Papers on Entomology Presented to Professor
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and Classification of Coleoptera. Papers Celebrating the 80th Birthday of Roy
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MAES, J.-M. 1992. Lista de los Lucanidae (Coleoptera) del mundo.
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MIZUNUMA, T. and S. NAGAI. 1994. The Lucanid Beetles of the World. Mushi-sha,
Tokyo. 337 pp.
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Online publication Accessed here.
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Coleoptera) of Nebraska. Great Plains Research 1: 249-282.
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SCHOLTZ, C. H., D. J. BROWNE and J. KUKALOVA-PECK. 1994. Glaresidae,
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