Cladistics
Cladistics 27 (2011) 1–25 10.1111/j.1096-0031.2011.00358.x
Combined molecular and morphological phylogeny of Eulophidae (Hymenoptera: Chalcidoidea), with focus on the subfamily Entedoninae
Roger A. Burksa,*, John M. Heratya, Marco Gebiolab and Christer Hanssonc aDepartment of Entomology, University of California, Riverside, CA, USA; bDipartimento di Entomologia e Zoologia agraria ‘‘F. Silvestri’’ Universita` degli Studi di Napoli ‘‘Federico II’’, Italy; cDepartment of Biology, Zoology, Lund University, Lund, Sweden Accepted 9 March 2011
Abstract
A new combined molecular and morphological phylogeny of the Eulophidae is presented with special reference to the subfamily Entedoninae. We examined 28S D2–D5 and CO1 gene regions with parsimony and partitioned Bayesian analyses, and examined the impact of a small set of historically recognized morphological characters on combined analyses. Eulophidae was strongly supported as monophyletic only after exclusion of the enigmatic genus Trisecodes. The subfamilies Eulophinae, Entiinae (=Euderinae) and Tetrastichinae were consistently supported as monophyletic, but Entedoninae was monophyletic only in combined analyses. Six contiguous bases in the 3e¢ subregion of the 28S D2 rDNA contributed to placement of nominal subgenus of Closterocerus outside Entedoninae. In all cases, Euderomphalini was excluded from Entiinae, and we suggest that it be retained in Entedoninae. Opheliminae n. stat. is raised from tribe to subfamily status. Trisecodes is removed from Entedoninae but retained as incertae sedis in Eulophidae until its family placement can be determined new placement. The genera Neochrysocharis stat. rev. and Asecodes stat. rev. are removed from synonymy with Closterocerus because strong molecular differences corroborate their morphological differences. Closterocerus (Achrysocharis) germanicus is transferred to the genus Chrysonotomyia n. comb. based on molecular and morphological characters. The Willi Hennig Society 2011.
Eulophidae is one of the largest families of Chalci- subgroups of eulophids, including many genera, have doidea, with over 4400 described species in four very broad host ranges that may include several orders subfamilies (Noyes, 2003). It is also one of the most of insects or other taxa as well. A notable exception is diverse and economically important chalcidoid families, the tribe Euderomphalini, which apparently contains with many species important for biological control of only parasitoids of whiteflies (Hemiptera: Aleyrodidae). agricultural pests, especially of leaf-mining Diptera Geographic distribution is similarly broad for most (Clausen, 1978). Other species are gall-formers on a eulophid groups, with new continental distribution variety of plants, including Eucalyptus (Boucˇ ek, 1988; records of genera being discovered on a regular basis La Salle, 2005), but the range of diversity in life-history (Boucˇ ek, 1988; LaSalle, 1994; Schauff et al., 1997; traits of eulophids is comparable with that of Chalci- Burks, 2003). It seems likely that most genera with a doidea itself, with several unique examples. Hosts significant number of species occur in most continents. include species from most insect orders, and some However, there is another notable exception in this case eulophids are even predators in spider egg sacs or in as well, as the unplaced tribe Anselmellini is strictly galls of mites or nematodes (LaSalle, 1994). Most Australasian (Boucˇ ek, 1988). Although Eulophidae and most of its major sub- *Corresponding author: groups cannot be characterized succinctly in terms of life E-mail address: [email protected] history or distribution, the family can be defined by a