Phylogeny of psychodid subfamilies (Diptera: Psychodidae) inferred from nuclear DNA sequences with a review of morphological evidence for relationships

Psychodidae is a diverse family of flies with approximately 3000 described species in six subfamilies, including Phlebotominae vectors of human disease. Psychodidae has been the subject of few phylogenetic investigations and development of a stable classification has been hampered by poor understanding of the morphology of larvae, pupae and adults. Specimens were collected, and we analysed DNA sequence data from two nuclear genes for one or more representatives of all subfamilies. The subfamilies with multiple representatives included were resolved as monophyletic with good support. Placement of Horaiellinae, Sycoracinae and Trichomyiinae remains unclear, whereas Bruchomyiinae is hypothesized as the sister group to (Phlebotominae + Psychodinae). Representatives of some psychodine tribes were resolved in agreement with previous hypotheses. Relationships among and within subfamilies are discussed, and morphological characters supporting these relationships are reviewed. One compelling synapomorphy of the male genitalia supporting a relationship between Phlebotominae and Psychodinae is the presence of articulated surstyli with apical retinacula. Only cerci are present and sometimes developed into clasping structures in males of other subfamilies.