Phylogeny and evolution of wasps, ants and bees (Hymenoptera, Chrysidoidea, Vespoidea and Apoidea)

The comprehensive cladistic study of family-level phylogeny in the Aculeata (sensu lato) by Brothers and Carpenter, published in 1993, is briefly reviewed and re-evaluated, particularly with respect to the sections dealing with Vespoidea and Apoidea. This remains the most recent general treatment of the subject, but several of the relationships indicated are only weakly supported, notably those of Pompilidae and Rhopalosomatidae. Characters used were almost entirely morphological, and re-evaluation of ground-plan states and hypotheses of character-state changes, specially from examination of different exemplars, is likely to lead to slightly different conclusions for some taxa, as is the use of additional or new characters, including molecular ones. The relationships of taxa within the Vespoidea are much better known than for those in the Apoidea, but recent work on the two major groups of bees (by Michener and colleagues) and various groups of sphecoid wasps (by Alexander and Melo) have provided greater clarity, for some families at least. A single cladogram showing the putative relationships of those taxa which should be recognized at the family level for the entire Aculeata is presented. These are, for the Chrysidoidea, Apoidea and Vespoidea, respectively (limits indicated by curly brackets): {Phimariidae + (Scolebythidae + ((Bethylidae + Chrysididae) + (Sclerogibbidae + (Dryinidae + Embolemidae))))} + ({Heterogynaidae + (Ampulicidae + (Sphecidae + (Crabronidae + Apidae)))} + {Sierolomorphidae + ((Tiphiidae + (Sapygidae + Mutillidae))+ ((Pompilidae + Rhopalosomatidae) + (Bradynobaenidae + (Formicidae + (Vespidae + Scoliidae)))))}). Current knowledge of the relationships within the families of Vespoidea is reviewed. A new analysis of the subtaxa of Mutillidae, based on that of Brothers published in 1975, is presented; the subfamilies to be recognized are: (Myrmosinae [ = Myrmosini + Kudakrumiini] + (Pseudophotopsidinae + (Ticoplinae + (Rhopalomutillinae + (Sphaeropthalminae [ = Dasylabrini + (Sphaeropthalmina + Pseudomethocina)] + (Myrmillinae + Mutillinae [ = Ephutini + (Mutillina + Smicromyrmina)])))))).