An evolutionary radiation of beeflies in semi-arid Australia: systematics of the Exoprosopini (Diptera : Bombyliidae)

Almost half of the 4822 described beeflies in the world belong to the subfamily Anthracinae, with most of the diversity found in three cosmopolitan tribes: Villini, Anthracini, and Exoprosopini. The Australian Exoprosopini previously contained three genera, Ligyra Newman, Pseudopenthes Roberts and Exoprosopa Macquart. Pseudopenthes is an Australian endemic, with two species including Ps. hesperis, sp. nov. from Western Australia. Two new species of the exoprosopine Atrichochira Hesse, Atr. commoni, sp. nov. and Atr. paramonovi, sp. nov., are also described from Australia, extending the generic distribution from Africa. Cladistic analysis clarified the phylogenetic relationships between the recognised groups of the Exoprosopini and determined generic limits on a world scale. Inclusion of 18 Australian exoprosopines placed the Australian species in the context of the world fauna. The Exoprosopini contains six large groups. The basal group I contains species previously included in Exoprosopa to which the name Defilippia Lioy is applied. Group II contains Heteralonia Rondani, Atrichochira, Micomitra Bowden, Pseudopenthes, and Diatropomma Bowden. Colossoptera Hull is newly synonymised with Heteralonia. Group III is a paraphyletic assemblage of Pterobates Bezzi and Exoprosopa including the Australian Ex. sylvana ( Fabricius). Ligyra is paraphyletic, forming two well-separated clades. The African clade is described as Euligyra Lambkin, gen. nov., which, together with Litorhina Bezzi and Hyperalonia Rondani, form group IV. The Australian group V is true Ligyra. The remaining monophyletic lineage of exoprosopines, group VI, the Balaana-group of genera, shows evidence of an evolutionary radiation of beeflies in semi-arid Australia. Phylogenetic analysis of all 42 species of the Balaana-group of genera formed a basis for delimiting genera. Seven new genera are described by Lambkin & Yeates: Balaana, Kapua, Larrpana, Munjua, Muwarna, Palirika and Wurda. Four non-Australian species belong to Balaana. Thirty two new Australian species are described: Bal. abscondita, Bal. bicuspis, Bal. centrosa, Bal. gigantea, Bal. kingcascadensis, K. corusca, K. irwini, K. westralica, Lar. collessi, Lar. zwicki, Mun. erugata, Mun. lepidokingi, Mun. paralutea, Mun. trigona, Muw. vitreilinearis, Pa. anaxios, Pa. basilikos, Pa. blackdownensis, Pa. bouchardi, Pa. cyanea, Pa. danielsi, Pa. decora, Pa. viridula, Pa. whyalla, W. emu, W. impatientis, W. montebelloensis, W. norrisi, W. patrellia, W. skevingtoni, W. windorah, and W. wyperfeldensis. The following new combinations are proposed: from Colossoptera: Heteralonia latipennis (Brunetti); from Exoprosopa: Bal. grandis (Pallas), Bal. efflatounbeyi (Paramonov), Bal. latelimbata ( Bigot), Bal. obliquebifasciata ( Macquart), Bal. tamerlan (Portschinsky), Bal. onusta ( Walker), Def. busiris (Jaennicke), Def. efflatouni ( Bezzi), Def. eritreae (Greathead), Def. gentilis ( Bezzi), Def. luteicosta ( Bezzi), Def. minos (Meigen), Def. nigrifimbriata ( Hesse), Def. rubescens ( Bezzi), K. adelaidica ( Macquart), Lar. dimidiatipennis ( Bowden), Muw. stellifera ( Walker), and Pa. marginicollis ( Gray); from Ligyra: Eu. enderleini ( Paramonov), Eu. mars ( Bezzi), Eu. monacha (Klug), Eu. paris ( Bezzi), Eu. sisyphus ( Fabricius), and Eu. venus (Karsch).