| A comprehensive phylogeny of the Figitidae (Hymenoptera: Cynipoidea) is presented based on simultaneous analysis of molecular (28S-D2 and D3, COI and 18S-E23), morphological data and biological data. Data are analyzed by parsimony and Bayesian inference methods. A total of 94 genera and 168 species were sampled. The alignment of the 28S D2+D3 gene fragment based on a structural model resulted in the most defendable and least conflicting alignment tested. Partitioned Bremer support, sensitivity analysis and bootstrap support provide good measures of robustness for all major clades, though the morphological/biological data partition was required to recover basal lineage monophyly. All subfamilies of Figitidae were found to be monophyletic except for Figitinae, which is rendered paraphyletic by Lonchidia. Melanips (Figitinae) was consistently found to be the sister group of the Aspicerinae. The general evolutionary trend of early figitids being parasitoids of gall inducing insects, with later host shifts to exposed hosts associated with the aphid community, was supported. Non clock-like estimates of divergence times of all major lineages suggests that most figitid groups diverged in the mid to late Cretaceous except for Eucoilinae which diverged in the Eocene (44 mya). The divergence of major lineages of Eucoilinae, the most diverse and successful of all figitid groups, has been estimated to follow the divergence estimates of their hosts, Muscomorpha Diptera (48--51 mya).; The presence of the ovipositor clip is surveyed throughout Figitidae. This morphological structure is postulated to restrain the parasitoid host during oviposition. All Figitinae and Eucoilinae that attack semi-concealed dipterous hosts were found to possess the clip. Figitids that attack fully concealed hosts all lacked the ovipositor clip. It is hypothesized here that two subfamilies, the Anacharitinae and Aspicerinae, attack fully exposed hosts but lack the clip in order to quickly oviposit and prevent fighting with the host. Mapping of the presence/absence of the ovipositor clip onto two competing phylogenies suggests this structure evolved in parallel in Figitinae and Eucoilinae. The evolution of the ovipositor clip is likely responsible for the successful colonization exposed dipterous larvae by Eucoilinae.; The descriptions of Preseucoela imallshookupis Buffington and Nordlandiella semirufa (Kieffer) are also included. |
