Molecular systematics and evolution of the subgenus Mesocarabus Thomson, 1875 (Coleoptera: Carabidae: Carabus), based on mitochondrial and nuclear DNA

Abstract

The subgenus Mesocarabus Thomson, 1875 is a western Palaearctic group that currently includes five species: four of them inhabiting western Europe (Carabus lusitanicus Fabricius, 1801, Carabus problematicus Herbst, 1786, Carabus dufourii Dejean & Boisduval, 1829, and Carabus macrocephalus Dejean, 1826) and one found in the Rif Mountains in northern Morocco (Carabus riffensis Fairmaire, 1872). Representatives of Mesocarabus have been included in previous molecular phylogenetic studies, but taxon- or gene-sampling limitations yielded inconclusive results regarding its monophyly and sister relationship. Here we perform molecular phylogenetic analyses based on five mitochondrial (3625 nt) and eight nuclear (5970 nt) genes sequenced in many Mesocarabus populations, and in related western Palaearctic Carabus Linnaeus, 1758. We conducted parsimony, maximum-likelihood, and Bayesian analyses and found a well-supported sister relationship between a monophyletic Mesocarabus with Iberian species of the subgenus Oreocarabus Géhin, 1876. Within Mesocarabus, the European species form a monophyletic lineage sister to Moroccan C. riffensis. A time-calibrated phylogeny suggests the split between Mesocarabus and Oreocarabus occurred at 11.8 Mya (95% highest posterior density, HPD, 8.7–15.3 Mya), and the divergence between C. riffensis and European Mesocarabus at 9.5 Mya (95% HPD 7.0–12.5 Mya). The early diversification of Mesocarabus and related subgenera during the Miocene, and alternative hypotheses concerning the origin of Mesocarabus in the Iberian Peninsula and the Betic-Riffian plate are discussed using calibration data and dispersal–vicariance biogeographic analyses. Finally, we found instances of incongruence between mitochondrial DNA and nuclear-based phylogenies of Mesocarabus, which are hypothesized to be the result of introgressive hybridization.