Phylogenetic position of the former subgenera Haptoderus (blue) and Pyreneorites (red), excerpt of the tree from combined analysis (Fig. 3) of CO1, 28S, 18S and 16S sequences based on maximum likelihood (ML) analysis. Numbers in nodes indicate ML bootstrap value / Bayesian posterior probability (only when >50), “x” node not recovered by Bayesian analysis. P. = Pterostichus, C. = Cryobius, M. = Morphnosoma, Ph. = Parahaptoderus. In brackets: specimen code (CrXX, this study) or GenBank accession number. P. (C.) riparius: chimera (JF888281 + EU142445).