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Bombyliidae is a very species‐rich and widespread family of parasitoid flies with more than 250 genera classified into 17 extant subfamilies. However, little is known about their evolutionary history or how their present‐day diversity was shaped. Transcriptomes of 15 species and anchored hybrid enrichment (AHE) sequence captures of 86 species, representing 94 bee fly species and 14 subfamilies, were used to reconstruct the phylogeny of Bombyliidae. We integrated data from transcriptomes across each of the main lineages in our AHE tree to build a data set with more genes (550 loci versus 216 loci) and higher support levels. Our overall results show strong congruence with the current classification of the family, with 11 out of 14 included subfamilies recovered as monophyletic. Heterotropinae and Mythicomyiinae are successive sister groups to the remainder of the family. We examined the evolution of key morphological characters through our phylogenetic hypotheses and show that neither the "sand chamber subfamilies" nor the "Tomophthalmae" are monophyletic in our phylogenomic analyses. Based on our results, we reinstate two tribes at the subfamily level (Phthiriinae stat. rev. and Ecliminae stat. rev.) and we include the genus Sericosoma Macquart (previously incertae sedis) in the subfamily Oniromyiinae, bringing the total number of bee fly subfamilies to 19. Our dating analyses indicate a Jurassic origin of the family (165–194 Ma), with the sand chamber evolving early in bee fly evolution, in the late Jurassic or mid‐Cretaceous (100–165 Ma). We hypothesize that the angiosperm radiation and the hothouse climate established during the late Cretaceous accelerated the diversification of bee flies, by providing an expanded range of resources for the parasitoid larvae and nectarivorous adults. [ABSTRACT FROM AUTHOR]

Assassin or robber flies (Asilidae) and bee flies (Bombyliidae) are two of the largest and most diverse fly families with over 7400 and 4700 described species worldwide, respectively. Adult assassin flies are predatory, feeding on smaller insects captured on the wing, and larvae predate on soft‐bodied arthropods in the soil or rotting wood. Adult bee flies feed on nectar and pollen. Almost all bee fly larvae are external or internal parasitoids of other insect larvae, usually in the soil. Although the interaction between bee fly larvae and their hosts has been rarely observed, we recently discovered assassin fly larvae hosting bee fly Lepidanthrax linguatus (Anthracinae: Villini) endoparasitoids in riverine sandy soil in southern NSW, Australia. While bee fly larvae have been known to attack other fly larvae, Asilidae is an uncommon host group. Furthermore, to find bee fly larvae inside the host is even less common because most bee fly larvae are ectoparasitoids. Here we describe and illustrate the host assassin fly (Bathypogon nigrinus: Bathypogoninae) larva and pupa, as well as the first instar bee fly larva (planidium) though careful external examination of the surface of the asilid larva, which is translucent, allowing us to see the internal parasite. This is the first larva and pupa from Bathypogoninae to be described. We also illustrate the bee fly larva in detail based on scanning electron microscopy of a larva in situ, after dissection from the host. Finally, we describe the late instar larval and pupal exuvia of the endoparasitoid. Like other endoparasitoid bombyliid larvae in the Villini, the first instar larva causes the host to produce a breathing tube that surrounds the posterior half of the parasitoid, allowing the parasitoid to maintain contact with atmospheric oxygen. [ABSTRACT FROM AUTHOR]

The Bombyliinae comprises over 1100 described species in 73 known genera distributed worldwide. It is one of the largest subfamilies of bee flies (Diptera: Bombyliidae). We present the first phylogenetic hypothesis for this subfamily, based on 157 adult morphological characters scored for 123 species representing 60 genera, including all the tribes of Bombyliinae, and the related subfamilies Lordotinae and Toxophorinae. Four most parsimonious trees were generated from our analysis under equal weighting schemes. The monophyly of Bombyliinae is supported, and Lordotinae is sister to the Bombyliinae. Within Bombyliinae, Conophorini is sister to the remaining tribes. Five previously recognized tribes are revised and four new tribes are erected. We placed almost all genera in our tribal classification, based on our phylogenetic results and available character evidence. The genus Parabombylius is proposed as a synonym of Bombylius. The Gondwanan origin for the major lineages of Bombyliinae is strongly indicated by our biogeographic analysis which reconstructs ancestral areas. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:1EC5C827‐34D5‐4A95‐BA78‐4ACF457F6D40. [ABSTRACT FROM AUTHOR]