Invertebrate Systematics Invertebrate Systematics Society
Systematics, phylogeny and biogeography

The evolutionary relationships of North American Diplous Motschulsky (Coleoptera : Carabidae : Patrobini) inferred from morphological and molecular evidence

Paul E. Marek A B C D and David H. Kavanaugh A
+ Author Affiliations
- Author Affiliations

A Department of Entomology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118, USA.

B Department of Biology, San Francisco State University, 1600 Holloway Avenue, San Francisco, California 94132, USA.

C Current address: Department of Biology, East Carolina University, Howell Science Complex, Greenville, North Carolina 27858, USA.

D Corresponding author. Email:

Invertebrate Systematics 19(2) 145-168
Submitted: 13 April 2004  Accepted: 12 April 2005   Published: 29 June 2005


Individuals of the ground beetle genus Diplous Motschulsky, 1850 occur in riparian areas predominately throughout boreal North America and Asia. In order to infer the species phylogeny of the North American Diplous, we examined 97 morphological characters (56 quantitative characters and 41 qualitative characters) and 458 bp of the mitochondrial cytochrome oxidase subunit I region. We used the four North American species, four Palearctic species, and one undescribed species of a closely related genus to test the monophyly and the direction of character state change in North American Diplous. Overall, we found that North American Diplous appear to represent a monophyletic group, but that the morphological and molecular evidence did not support the same relationships in the placement of one of the species. We found that the total evidence trees agreed most with biogeography and considerations of accelerated morphological evolution. In this paper, we present a morphological phylogenetic tree, a molecular phylogenetic tree, a total evidence phylogenetic tree, a species key, species diagnoses, and a distribution map of Nearctic Diplous.

Additional keywords: COI, cytochrome oxidase subunit I, Diplous, glacier, ground beetle, Platidius, Pleistocene, refugia, riparian, total evidence, vicariance.


This project was supported by a Graduate Assistantship from the California Academy of Sciences. To the Entomology Department staff at California Academy of Sciences, the first author is grateful for accommodating me as a graduate student and for providing a great research atmosphere. Thanks are also due to lenders of museum specimens (Liang Hongbin for specimens from IOZB and Stuart Hine for specimens from BMNH), and the collectors who obtained fresh specimens on my behalf (Robert Nelson for specimens from Maine and Tracy Mastranicola for specimens from Washington). We would also like to thank Greg Spicer, John Hafernik, Corrie Saux, and Alexander Zamotajlov for their help on technical and theoretical issues, as well as Jason Bond, Petra Sierwald, and anonymous reviewers for providing very helpful suggestions on the final versions of the manuscript.


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Appendix 1.  Median values for morphometric ratios by species for males (characters: 1–31) and females (characters: 32–56)
n, species sample size; values rounded to the thousandth; numbers refer to characters listed in Table 2
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Appendix 1a. continued
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Appendix 2.  Morphology character state matrix (TreeBASE Acc. M2288)
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Appendix 2a. continued
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