Carabidae

Introduction

Carabids (as here treated) include all of the terrestrial adephagans other than trachypachids. This is by far the largest family of Adephaga, with over 30,000 described species. Among the more well-known members of the family are the genus Carabus (Carabini), bombardier beetles (Brachinini), and tiger beetles (Cicindelitae).

The vast majority of carabids are predacious. Most of these are generalist predators, but there are a number of groups that have become specialized (e.g., Peleciini and Promecognathini on millipedes, Cychrini and Licinini on snails). A few clades have larvae that are ectoparasitoids on other arthropods (e.g., Lebiini, Brachinitae, and Peleciini). Others are seed-eaters (e.g., Harpalini).

Characteristics

There are very few derived features that delimit carabids. In adults, the metacoxae are narrower than other adephagans, with the metapleuron extending posteriorly to contact the second abdominal sternite. There are also a few minor features in the head structure and musculature of larvae (see Beutel, 1995, for a summary).

Discussion of Phylogenetic Relationships

While carabid phylogeny has been extensively studied, the convergences and reversals present in morphological traits has lead to a great deal of controversy about many groups. Two of these groups, the tiger beetles (Cicindelitae) and wrinkled bark beetles (Rhysodini) are often considered outside the carabid clade. The phylogeny shown of carabid tribes on this and other pages is a conservative consensus view, in which a large number of "basal" groups give rise to a middle and upper grade of carabids. Within this latter group is a large, relatively uniform clade, the Harpalinae, which includes many of the larger, more common carabids.

Included below the tree are a number of especially enigmatic groups, including Gehringiini and Rhysodini, which may be older lineages, related to groups in this page, or they may instead be related to groups within the Carabidae Conjunctae. Their placement, along with the resolution of other aspects of carabid phylogeny, awaits numerical analysis of available morphological and molecular data.

Other Names for Carabidae

• carabids
• Ground beetles and tiger beetles

References

Arndt, E. 1993. Phylogenetische Untersuchungen larvalmorphologischer Merkmale der Carabidae (Insecta: Coleoptera). Stuttgarter Beiträge zur Naturkunde Serie A 488: 1-56.

Baehr, M. 1979. Vergleichende Untersuchungen am Skelett und an der Coxalmuskulatur des Prothorax der Coleoptera, ein Beitrag zur Klärung der phylogenetischen Beziehungen der Adephaga (Coleoptera, Insecta). Zoologica 44(4): 1–76.

Ball, G. E. 1979. Conspectus of carabid classification: history, holomorphology, and higher taxa. In T. L. Erwin, G. E. Ball, D. R. Whitehead and A. L. Halpern (ed.), Carabid Beetles: Their Evolution, Natural History, and Classification. The Hague, Dr. W. Junk bv. pp. 63–111.

Bell, R. T. 1964. Does Gehringia belong to the Isochaeta? (Coleoptera: Carabidae). The Coleopterists Bulletin 18: 59–61.

Bell, R. T. 1967. Coxal cavities and the classification of the Adephaga (Coleoptera). Annals of the Entomological Society of America 60(1): 101–107.

Beutel, R. G. 1990. Metathoracic features of Omoglymmius hamatus and their significance for classification of Rhysodini (Coleoptera: Adephaga). Entomologia Generalis 15(3): 185-201.

Beutel, R. G. 1991b. Larval head structures of Omophron and their implications for the relationships of Omophronini (Coleoptera: Carabidae). Entomologica Scandinavica 22(1): 55-67.

Beutel, R. G. 1992a. Larval head structures of Omoglymmius hamatus and their implications for the relationships of Rhysodidae (Coleoptera: Adephaga). Entomologica Scandinavica 23(2): 169-184.

Beutel, R. G. 1992b. Phylogenetic analysis of thoracic structures of Carabidae (Coleoptera: Adephaga). Zeitschrift für Zoologische Systematik und Evolutionsforschung 30(1): 53-74.

Beutel, R. G. 1992c. Study on the systematic position of Metriini based on characters of the larval head (Coleoptera: Carabidae). Systematic Entomology 17(3): 207-218.

Beutel, R. G. 1993. Phylogenetic analysis of Adephaga (Coleoptera) based on characters of the larval head. Systematic Entomology 18: 127-147.

Beutel, R.G. 1995. The Adephaga (Coleoptera): phylogeny and evolutionary history. Pp. 173-217 in Pakaluk and Slipinski (1995).

Bils, W. 1976. Das Abdomende weiblicher, terrestrisch lebender Adephaga (Coleoptera) und seine Bedeutung für die Phylogenie. Zoomorphologie 84(2): 113–193.

Bousquet, Y. and A. Smetana. 1991. The tribe Opisthiini (Coleoptera: Carabidae): Description of the larvae, note on habitat, and brief discussion on its relationships. Journal of the New York Entomological Society 99(1): 104-114.

Burmeister, E.G. 1980. Funktionsmorphologie und Evolution des Ovipositor der Adephaga (Coleoptera). Verhandlungen der naturwissenschaftlicher Verein, Hamburg (NF) 24(1): 89–184.

Deuve, T. 1988. Étude phylogénétique des Coléoptères Adephaga: redéfinition de la famille des Harpalidae, sensu novo, et position systématique des Pseudomorphinae et Brachinidae. Bull. Soc. Entomol. France 92(5-6): 161-182.

Deuve, T. 1993. L'abdomen et les genitalia des femelles de Coléoptères Adephaga. Mémoires du Muséum National D'Histoire Naturelle 155: 1-184.

Erwin, T. L. 1979. Thoughts on the evolutionary history of ground beetles: hypotheses generated from comparative faunal analyses of lowland forest sites in temperate and tropical regions. In T. L. Erwin, G. E. Ball, D. R. Whitehead and A. L. Halpern (ed.), Carabid Beetles: Their Evolution, Natural History, and Classification. The Hague, Dr. W. Junk bv. pp. 539-592.

Erwin, T. L. 1981. Taxon pulses, vicariance, and dispersal: An evolutionary synthesis illustrated by carabid beetles. In G. Nelson and D. E. Rosen (ed.), Vicariance Biogeography: A Critique. New York, Columbia University Press. 159-196.

Erwin, T. L. 1985. The taxon pulse: A general pattern of lineage radiation and extinction among carabid beetles. In G. E. Ball (ed.), Taxonomy, Phylogeny and Zoogeography of Beetles and Ants. A volume dedicated to the Memory of Philip Jackson Darlingon, Jr. (1904-1983). Dordrecht, Dr W. Junk. 437-472.

Erwin, T. L. and N. E. Stork. 1985. The Hiletini, an ancient and enigmatic tribe of Carabidae with a pantropical distribution (Coleoptera). Systematic Entomology 10(4): 405-451.

Evans, M. E. G. 1982. Early evolution of the Adephaga—some locomotor speculations. The Coleopterists Bulletin 36(4): 597–607.

Forsyth, D. J. 1972. The structure of the pygidial defence glands of Carabidae (Coleoptera). Trans. Zool. Soc. London 32: 249–309.

Hammond, P. M. 1979. Wing-folding mechanisms in beetles with special reference to investigations of adephagan phylogeny (Coleoptera). In T. L. Erwin, G. E. Ball, D. R. Whitehead and A. L. Halpern (ed.), Carabid Beetles: Their Evolution, Natural History, and Classification. The Hague, Dr. W. Junk bv. pp. 113–180.

Heath, R. V. and M. E. G. Evans. 1990. The relationship between the ventral nerve cord, body size and phylogeny in ground beetles (Coleoptera: Carabidae). Zoological Journal of the Linnean Society 98(3): 259-293.

Hlavac, T. F. 1971. Differentiation of the carabid antenna cleaner. Psyche 78(1–2): 51–66.

Jeannel, R. 1941. Coléoptères Carabiques, Première Partie. Faune de France, 39. Paris, Lechevalier, 571 pp.

Kanehisa, K. and K. Kawazu. 1985. Differences in neutral components of the defensive secretion in formic acid-secreting carabid beetles. Applied Entomology and Zoology 20(3): 299-304.

Kanehisa, K. and M. Murase. 1977. Comparative study of the pygidial defensive systems of carabid beetles. Applied Entomology and Zoology 12: 225-235.

Kavanaugh, D. H. and T. L. Erwin. 1991. The tribe Cicindini Banninger (Coleoptera: Carabidae): Comparative morphology, classification, natural history, and evolution. Proceedings of the Entomological Society of Washington 93(2): 356-389.

Kavanaugh, D. H. and J. Négre. 1985. Notiokasiini—A new tribe of Carabidae (Coleoptera) from southeastern South America. Coleopterists Bulletin 36(4): 549–566.

Kryzhanovsky, O.L. 1976. Revised classification of the family Carabidae. Entomological Review, 1:80-91.

Kryzhanovskij, O.L., I.A. Belousov, I.I. Kabak, B.M. Kataev, K.V.Makarov, V.G.Shilenkov. 1995. A checklist of the ground-beetles of Russia and adjacent lands (Insecta, Coleoptera, Carabidae). 271 pp. Pensoft.

Liebherr, J. K. and G. E. Ball. 1990. The first instar larva of Eripus oaxacanus Straneo and Ball (Coleoptera: Carabidae: Peleciini): Indicator of affinity or convergence? Systematic Entomology 15(1): 69-79.

Lindroth, C.H. 1961. The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 2. Opuscula Entomologica, Supplementum XXX, pp. 1-200.

Lindroth, C.H. 1969. The ground-beetles (Carabidae, excl. Cicindelinae) of Canada and Alaska. Part 1. Opuscula Entomologica, Supplementum XXXV, pp. I-XLVIII.

Moore, B. P. 1979. Chemical defense in carabids and its bearing on phylogeny. In T. L. Erwin, G. E. Ball, D. R. Whitehead and A. L. Halpern (ed.), Carabid Beetles: Their Evolution, Natural History, and Classification. The Hague, Dr. W. Junk bv. 193–203.

Nagel, P. 1979. The classification of Carabidae. Misc. Pap. Agric. Univ. Wageningen 18: 7–14.

Nichols, S. W. 1985. Omophron and the origin of Hydradephaga (Insecta: Coleoptera: Adephaga). Proceedings of the Academy of Natural Sciences of Philadelphia 137: 182–201.

Noonan, G. R. 1973. The anisodactylines (Insecta: Coleoptera: Carabidae: Harpalini): Classification, evolution, and zoogeography. Quaestiones entomologicae 9: 266-480.

Pakaluk, J. and S.A. Slipinski (eds.) 1995. Biology, Phylogeny, and Classification of Coleoptera. Papers Celebrating the 80th Birthday of Roy A. Crowson. Muzeum i Instytut Zoologii PAN, Warszawa.

Ponomarenko, A. G. 1977. Suborder Adephaga, etc. In L. Arnoldy, V. V. Jerikin, L. M. Nikritin and A. G. Ponomarenko (ed.), Mesozoic Beetles. pp. 1–204.

Regenfuss, H. 1975. Die Antennen-Putzeinrichtung der Adephaga (Coleoptera), parallele evolutive Vervollkommnung einer komplexen Struktur. Zeitschrift für zoologische Systematik und Evolutionsforschung 13: 278–299.

Serrano, J. 1981b. Chromosome numbers and karyotypic evolution of Caraboidea. Genetica 55: 51-60.

Straneo, S. L. and G. E. Ball. 1989. Synopsis of the genera and subgenera of the tribe Peleciini, and revision of the Neotropical and Oriental species (Coleoptera: Carabidae). Insecta Mundi 3(2): 73-178.

Ward, R. D. 1979. Metathoracic wing structures as phylogenetic indicators in the Adephaga (Coleoptera). In T. L. Erwin, G. E. Ball, D. R. Whitehead and A. L. Halpern (ed.), Carabid Beetles: Their Evolution, Natural History, and Classification. The Hague, Dr. W. Junk bv. pp. 181–191.

Yahiro, K. 1990. A comparative morphology of the alimentary canal in the adults of ground-beetles (Coleoptera): I. Classification into the types. Esakia 0(SPEC. ISSUE 1): 35-44.

Information on the Internet

• Carabid Biologists
• Erwin's Illustrated Guide to Carabidae Genera
• Erwin's Western Hemisphere Carabidae database
• Ground Beetles of Ireland
• Familia Carabidae (INBIO, Costa Rica)
• Maine Carabid Survey
• The New Zealand Carabidae Web Site