A Combined Molecular and Morphological Approach to Explore the Higher Phylogeny of Entimine Weevils (Coleoptera: Curculionidae), with Special Reference to South American Taxa

The Entiminae are broad-nosed weevils constituting the most diverse subfamily of Curculionidae, with over 50 tribes. We performed Bayesian and Maximum Parsimony combined phylogenetic analyses with the main objective of testing higher-level relationships and the naturalness of the major Neotropical and Southern South American (Patagonia and Andes) tribes, including some members from other regions. We compiled a data matrix of 67 terminal units with 63 Entiminae species, as well as four outgroup taxa from Cyclominae, by 3522 molecular (from nuclear 18S rDNA and 28S rDNA, and mitochondrial 16S rDNA and COI gene sequences) and 70 morphological characters. The resulting trees recover a clade Entiminae with a monophyletic Cylydrorhinini and Premnotrypes branching off early. The tree resulting from parsimony analysis shows a clade of Leptopiini from the Australian region and another clade including taxa mainly distributed in the Palaearctic and Neotropical regions, but in the Bayesian tree the South American and Australian Leptopiini are grouped together. The mainly Palaearctic Entiminae (e.g., Brachyderini, Laparocerini, Otiorhynchini, Peritelini, Polydrusini, Phyllobiini and Sciaphylini) form a subclade separated from Southern Hemisphere taxa. Among the latter, the well-supported Naupactini are the sister group of the South American Tanymecini, excluding Platyaspistes, herein transferred to Leptopiini (new placement). Another well-justified clade is Eustylini-Geonemini, which also includes the enigmatic Galapagonotus, and the genus Artipus, thus corroborating its recent exclusion from Naupactini.