Are Miquihuana rhadiniformis Barr, 1982 and Pseudamara arenaria (LeConte, 1847) (Coleoptera, Carabidae) sphodrines? Phylogenetic analysis of data from next-generation sequencing of museum specimens resolves the tribal-group relationships of these enigmatic taxa

Miquihuana rhadiniformis Barr, 1982 and Pseudamara arenaria (LeConte, 1847) are enigmatic North American carabid beetles whose tribal-group placement has been in doubt. Both are currently placed in Sphodrini, although Pseudamara has typically been placed in Zabrini by most authors. As these species are relatively rarely collected, neither Miquihuana Barr, 1982 nor Pseudamara Lindroth, 1968 have been sampled in previous phylogenetic studies and attempts to obtain DNA sequences through standard PCR of available specimens failed. We used Illumina sequencing to recover nuclear gene fragments (18S rDNA, 28S rDNA, wingless, and CAD) from pinned M. rhadiniformis and 70%-ethanol-preserved P. arenaria. Combining these data with data from exemplar taxa from Sphodrini, Zabrini, Pterostichini, Platynini, and other, putatively related tribes, we examine the tribal relationships of Miquihuana and Pseudamara. The phylogenetic reconstructions consistently and with high support place M. rhadiniformis wi...