PHYLOGENY OF ASILIDAE INFERRED FROM MORPHOLOGICAL CHARACTERS OF IMAGINES (INSECTA: DIPTERA: BRACHYCERA: ASILOIDEA) TORSTEN DIKOW Division of Invertebrate Zoology American Museum of Natural History and Department of Entomology, Cornell University, Ithaca, NY 14853 ([email protected]) BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 319, 175 pp., 131 figures, 8 tables Issued March 17, 2009 Copyright E American Museum of Natural History 2009 ISSN 0003-0090 CONTENTS Abstract.......................................................... 4 Introduction . ...................................................... 4 Brief Historical Review of Classifications of Asilidae ........................ 11 MaterialsandMethods............................................... 12 Taxon Sampling .................................................. 13 PhylogeneticAnalysis.............................................. 18 Figure Abbreviations for Morphological Characters......................... 18 Character Analysis .................................................. 19 Head.......................................................... 20 Thorax. ..................................................... 36 Wing.......................................................... 51 Abdomen (Female and Male) ........................................ 58 FemaleAbdomen................................................. 61 MaleAbdomen................................................... 70 Characters Not Considered .......................................... 82 Results.......................................................... 83 A Revised, Higher-Level Classification of Asilidae.......................... 85 1.Laphriinae....................................................89 1.1. Laphystiini, new status ........................................ 89 1.2.Atomosiini................................................. 89 1.3.Hoplistomerini..............................................91 1.4. Laphriini .................................................. 91 1.5.Ctenotini.................................................. 98 1.6. Andrenosomatini ............................................ 98 2.Ommatiinae................................................... 98 3.Asilinae......................................................99 3.1.Asilini.................................................... 99 3.2.Apocleini,newstatus.......................................... 99 3.3.Lycomyini................................................. 99 3.4.Machimini................................................ 100 3.5. Neomochtherini ............................................ 100 3.6.Philonicini................................................ 100 4. Bathypogoninae, new status ....................................... 100 5. Phellinae, new status ............................................ 100 6. Tillobromatinae, new status ....................................... 101 7. Dasypogoninae ................................................ 101 7.1.Dasypogonini.............................................. 101 7.2.Saropogonini.............................................. 102 7.3.Thereutriini............................................... 102 7.4.Lastauracini............................................... 102 7.5. Blepharepiini .............................................. 102 7.6.Molobratiini............................................... 103 7.7. Megapodini ............................................... 103 7.7.1.Megapodina............................................ 104 7.7.2.Cyrtophryina........................................... 104 7.7.3.Lagodiina.............................................. 104 7.7.4. Senobasina ............................................. 104 8. Stenopogoninae. ............................................... 105 8.1. Enigmomorphini ............................................ 105 8.2. Plesiommatini .............................................. 105 8.3. Stenopogonini.............................................. 105 2 2009 DIKOW: PHYLOGENY OF ASILIDAE 3 9.Willistonininae,newstatus........................................ 106 10.Dioctriinae.................................................. 106 10.1.Dioctriini................................................ 106 10.2.Echthodopini............................................. 106 11. Leptogastrinae . ............................................ 107 11.1.Acronychini,newplacement................................... 107 11.2.Leptogastrini.............................................. 108 12.Trigonomiminae.............................................. 108 12.1.Trigonomimini............................................ 108 12.2.Xenomyzini.............................................. 108 13. Stichopogoninae . ............................................ 109 13.1.Stichopogonini............................................ 109 14. Brachyrhopalinae, new status ..................................... 109 14.1.Cyrtopogonini,newplacement................................. 110 14.2. Chrysopogonini, new placement ................................ 110 14.3.Brachyrhopalini,newplacement............................... 110 14.4. Ceraturgini, new placement ................................... 110 Incertaesedis................................................... 111 Phylogenetic Relationships of Higher-Level Taxa within Asilidae .............. 111 Discussion...................................................... 113 PreviousversusthePresentPhylogeneticHypothesisforAsilidae.............. 113 BiologyofAsilidae............................................... 115 Biogeographical Implications ........................................ 116 Some Morphological Characters Revisited .............................. 116 Transformation Series within Multistate Characters ........................ 121 Phylogenetic Placement of the Two Oldest Fossil Robber Flies................ 123 Sister Group to the Asilidae. ........................................ 124 Conclusions..................................................... 125 Acknowledgments................................................. 126 References...................................................... 126 Appendix1...................................................... 132 Appendix2...................................................... 174 Appendix3...................................................... 175 ABSTRACT A phylogenetic hypothesis is proposed for higher-level relationships within Asilidae, based on a sample of 158 species from 140 genera representing all 11 previously recognized subfamily taxa and 39 of the 42 tribal taxa and 220 discrete, parsimony informative, morphological characters from all tagmata of the imagines. Cladistic analysis results in 720 most parsimonious trees of 2760 steps in length, and a strict consensus topology of 2965 steps. The strict consensus cladogram is well resolved except for species of Apocleinae and Asilinae, which form a large polytomy. Monophyly of Asilidae is corroborated and supported by five autapomorphies: (1) labella of labium fused to prementum at least ventrally; (2) hypopharynx heavily sclerotized; (3) hypopharynx with dorsal seta-like spicules; (4) labrum short and at most half as long as labium; (5) cibarium trapezoidal. The clade Apioceridae + Mydidae is the sister group to Asilidae. The phylogenetic hypothesis indicates that five out of the 11 previously recognized subfamily taxa are non-monophyletic, i.e., Apocleinae, Asilinae, Dasypogoninae, Laphystiinae, and Stenopo- goninae. The present cladistic analysis forms the most comprehensive phylogenetic study on Asilidae to date and is used to revise the taxon’s phylogenetic classification in which 14 subfamily taxa are recognized. Ommatiinae, Trigonomiminae, and Stichopogoninae are recovered as monophyletic and contain the same genera as previously postulated. Dioctriinae and Leptogastrinae are also recovered as monophyletic, but the genera Myelaphus and Acronyches are transferred to them, respectively. Asilinae comprises all Apocleinae and Asilinae species and Laphriinae comprises all Laphriinae and Laphystiinae species sensu previous authors. Dasypogoninae and Stenopogoninae are divided into several taxa at phylogenetically unrelated positions in the cladogram. The Dasypogoninae comprises only Blepharepiini, Dasypogonini, Lastauracini, Megapodini (including Cyrtophryina, Lagodiina, Megapodina, and Senobasina), Molobratiini, Saropogonini, and Thereutriini as well as the unplaced genera Archilestris, Diogmites,andLestomyia. The remaining taxa possessing either a large prothoracic tibial spine, i.e., Brachyrhopalini and Chrysopogonini, or a small S-shaped spur, i.e., Cophura, Leptarthrus, and Nicocles, are part of the Brachyrhopalinae (new status). The Stenopogoninae comprises only Enigmomorphini, Plesiommatini, and Stenopogonini as well as the unplaced genera Ancylorhynchus and Scylaticus. Bathypogoninae (new status), Phellinae (new status), Tillobromatinae (new status), and Willistonininae (new status) are new subfamilial taxa previously assigned to Stenopogoninae. The remaining Stenopogoninae sensu previous authors represented here, i.e., Cyrtopogonini, Ceraturgini, Heteropogon, Holopogon, Metapogon,and Rhabdogaster, are assigned to the Brachyrhopalinae (new status). The genera Coleomyia and Oligopogon remain incertae sedis as neither genus groups with any other Asilidae, and are positioned as adelphotaxa to speciose clades. The higher-level relationships are: (Laphriinae ((Asilinae + Ommatiinae)