Cretaceous Promecognathinae (Goleoptera: Garabidae): a new genus, phylogenetic reconstruction and zoogeography

A fossil carabid (Coleoptera) belonging to the subfamily Promecognathinae is described from Cretaceous crater lake deposits at Orapa, Botswana. The specimen, which is the first ever fossil promecognathine, is placed in a new genus Palaeoxinidium. The fossil supports a Cretaceous age determination for the sediments, and it indicates that the Orapa crater was, at least in part, heavily vegetated at the time of deposition. Cladistic analysis of the fossil and extant Promecognathinae indicates that the fossil represents the sister group of the extant taxa and that the latter, in turn, comprise a pair of sister groups. It is, therefore, proposed that the Promecognathinae should be divided into two tribes: the Palaeoaxinidiini, represented by the fossil, and the Promecognathini, represented by the extant groups. The Promecognathini should be subdivided further into the subtribes Promecognathina and Axinidiina, represented by the American and African species, respectively. Three alternative zoogeographic hypotheses that explain the distributions of the fossil and extant promecognathines are discussed and tests for the hypotheses are proposed.