Systematic Entomology (2014), 39, 486–499 DOI: 10.1111/syen.12065 Higher level molecular phylogeny of darkling beetles (Coleoptera: Tenebrionidae) GAEL J. KERGOAT 1 , LAURENT SOLDATI 1 , ANNE-LAURE CLAMENS 1 , HERVE JOURDAN 2 , ROULA JABBOUR-ZAHAB 1,3 , GWENAELLE GENSON 1 , PATRICE BOUCHARD 4 and F A B I E N L. CONDAMINE 1,5 1 INRA, UMR 1062 CBGP (INRA, IRD, CIRAD, Montpellier SupAgro), Campus de Baillarguet, Montferrier-sur-Lez, France, 2 Institut Méditerranéen de Biodiversité et d’Ecologie Marine et Continentale (IMBE), Aix-Marseille Université, UMR CNRS IRD Avignon Université, UMR 237 IRD, Centre IRD Nouméa, Nouméa Cedex, New Caledonia, 3 CNRS, UMR 5175 CEFE (CNRS, Université Montpellier 2), Montpellier, France, 4 Canadian National Collection of Insects, Arachnids and Nematodes, Agriculture and Agri-Food Canada, Ottawa, Canada and 5 CNRS, UMR 7641 CMAP (CNRS, École Polytechnique), Palaiseau, France Abstract. Insect diversity represents about 60% of the estimated million-and-a-half described eukaryotic species worldwide, yet comprehensive and well-resolved intra-ordinal phylogenies are still lacking for the majority of insect groups. This is the case especially for the most species-rich insect group, the beetles (Coleoptera), a group for which less than 4% of the known species have had their DNA sequenced. In this study, we reconstruct the frst higher level phylogeny based on DNA sequence data for the species-rich darkling beetles, a family comprising at least 20 000 species. Although amongst all families of beetles Tenebrionidae ranks seventh in terms of species diver- sity, the lack of knowledge on the phylogeny and systematics of the group is such that its monophyly has been questioned (not to mention those of the subfamilies and tribes con- tained within it). We investigate the evolutionary history of Tenebrionidae using multiple phylogenetic inference methods (Bayesian inference, maximum likelihood and parsi- mony) to analyse a dataset consisting of eight gene fragments across 404 taxa (including 250 tenebrionid species). Although the resulting phylogenetic framework only encom- passes a fraction of the known tenebrionid diversity, it provides important information on their systematics and evolution. Whatever the methods used, our results provide strong support for the monophyly of the family, and highlight the likely paraphyletic or polyphyletic nature of several important tenebrionid subfamilies and tribes, notably the polyphyletic subfamilies Diaperinae and Tenebrioninae that clearly require substantial revision in the future. Some interesting associations in several groups are also revealed by the phylogenetic analyses, such as the pairing of Aphtora Bates with Phrenapatinae. Furthermore this study advances our knowledge of the evolution of the group, providing novel insights into much-debated theories, such as the apparent relict distribution of the tribe Elenophorini. Introduction Despite the recent advances in DNA sequencing technologies and phylogenetic analyses (Delsuc et al., 2006; Edwards, 2009; Correspondence: Gael J. Kergoat, INRA, UMR 1062 CBGP (INRA, IRD, CIRAD, Montpellier SupAgro), Campus de Baillarguet, 34988 Montferrier-sur-Lez, France. E-mail: kergoat@supagro.inra.fr McCormack et al., 2013), the quest for a well-resolved and comprehensive tree of life remains ongoing (Drew, 2013). Glimpses of light may come from the recent assembly of complete or nearly complete trees for mammals (4510 and 5020 species out of 5488 species; Bininda-Emonds et al., 2007; Fritz et al., 2009, respectively), amphibians (6111 species out of about 7000 species; Fritz & Rahbek, 2012), squamata (4161 486 © 2014 The Royal Entomological Society