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Diversification of subgenus Calathus (Coleoptera: Carabidae) in the Mediterranean region – glacial refugia and taxon pulses
Authors:Carlos Ruiz  Bjarte H Jordal  José Serrano
Institution:1. Departamento de Zoología y Antropología Física, Facultad de Veterinaria, Universidad de Murcia, 30071 Murcia, Spain;2. Museum of Natural History, University of Bergen, NO‐5007 Bergen, Norway
Abstract:Aim To investigate the effects of Pleistocene climatic variations on the diversification rate of the subgenus Calathus (Coleoptera: Carabidae), and to estimate the role of vicariance and dispersal for explaining current distributional patterns. Location Western Palaearctic Region, particularly the Mediterranean Basin. Methods Fragments of the mitochondrial cox1–cox2 and the nuclear 28S and EF1α genes were analysed by Bayesian inference. Lineage divergence times were estimated using a Bayesian relaxed molecular clock. Three diversification rate analyses were conducted, namely gamma (γ)‐statistic, birth–death likelihood (BDL) test and survival analyses, in order to test departures from a constant rate model of diversification. A Bayesian approach to dispersal–vicariance analysis was developed to reconstruct the most probable ancestral area of subgenus Calathus and subsequent events of dispersal and colonization. Results A constant rate of speciation events from the late Miocene onwards was found for the subgenus Calathus, whereas recent Pleistocene climatic oscillations played an important role only in shaping intraspecific diversity. Overall diversification patterns for the subgenus are best explained by at least four westward dispersal events from the eastern Mediterranean Basin. Three distinct phylogroups were found for the widely distributed Calathus fuscipes. Incongruence between mitochondrial and nuclear loci was found for a number of species. Main conclusions Diversification analyses suggest either a constant rate of diversification (BDL analysis) or a decrease in diversification rates for the subgenus (survival or γ‐statistics analyses), but not an increase related to the effects of glaciation cycles. Diversification patterns in the subgenus Calathus agree with predictions of the taxon pulse model. From the middle Miocene onwards the Anatolian Peninsula was possibly the main centre of diversification, with successive dispersal events towards the western Mediterranean Basin. Range expansion and secondary contact zones are postulated between members of different phylogroups in C. fuscipes.
Keywords:BDL test  dispersal–  vicariance analysis  divergence time  diversification analyses  Pleistocene climatic oscillations  subgenus Calathus  taxon pulse model  Western Palaearctic Region
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