HarpalinaeKipling Will and David R. Maddison
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IntroductionThis large clade contains more than half of carabid species. It is supported as monophyletic by both morphological and molecular data (Maddison et al., 1999; Ober, 2002).
CharacteristicsThe Harpalinae comprise those carabids with conjunct mesocoxae (as explained on the page for the Carabidae Conjunctae, and which also have the following features:
- male parameres conchoid and asetose (in rare cases, e.g. Pseudamara arenaria, parameres have some apical setae)
- mandible lacking scrobal seta (various taxa may have numerous short fine setae in the scrobe and in rare cases, e.g. Metius robustus, there may be a single more substantial seta. However, this condition is recognizably different than the single robust fixed seta in the scrobe of non-harpalinae taxa)
- Configuration of abdomen:Tergum VIII partially or totally invaginated with anterolateral apophyses present. Pygidial gland opening near the posterior margin of tergum VIII.
- most species spray hydrocarbons and formic acid in defense secretions (Moore, 1979)
- male chromosome number in most species 2n=36+x
- high-grade wedge-pushing ability
Maddison, D.R., M.D. Baker, and K.A. Ober. 1999. Phylogeny of carabid beetles as inferred from 18S ribosomal DNA (Coleoptera: Carabidae). Systematic Entomology, 24:103-138.
Ober, K.A. 2002. Phylogenetic relationships of the carabid subfamily Harpalinae (Coleoptera) based on molecular sequence data. Molecular Phylogenetics and Evolution, 24: 228-248.
About This Page
University of California, Berkeley, California, USA
David R. Maddison
Oregon State University
Correspondence regarding this page should be directed to Kipling Will at and David R. Maddison at
Page copyright © 2012 Kipling Will and David R. Maddison
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- Content changed 07 July 2006
Citing this page:
Will, Kipling and David R. Maddison. 2006. Harpalinae. Version 07 July 2006 (under construction). http://tolweb.org/Harpalinae/100/2006.07.07 in The Tree of Life Web Project, http://tolweb.org/