1 Phylogeny of Apioceridae and Mydidae based on morphological characters of imagines Apioceridae: Apiocera sp. © J. Hort Mydidae: Miltinus minutus © J. Hort Torsten Dikow Smithsonian @TDikow #asiloidflies National Museum of Natural History Apioceridae – “flower-loving flies” 2 ◊ imagines flower-feeders ◊ size = 15 – 30 mm ◊ 138 species in 1 genus (4 subgenera) ◊ occur only in › Argentina and Chile –> 4 species › Australia –> 73 species Apiocera (Apiocera) pica © J. Hort › South Africa –> 3 species › western North America (Canada, Mexico, USA) –> 64 species Apiocera (Apiocera) sp. © J. Hort Mydidae – “mydas flies” 3 ◊ imagines flower-feeders › some adults do not feed at all ◊ size = 9 – 60 mm (individual wing length up to 52 mm) ◊ 473 species in 66 genera ◊ world-wide primarily in arid regions › southern Africa most speciose Opomydas townsendi ◊ rarely collected ◊ mimetic coloration, morphology, and behavior present in several clades › aculeate Hymenoptera ◊ Cretaceous fossil Eremomidas arabicus © D. Gardner taxon sampling 4 ◊ 30 Apiocera species (= 21 %) › 14 Australian species › 12 North American species › 2 Chilean species › 2 South African species ◊ 138 Mydidae species (= 29.1 %) Apioceridae › 46 genera (= 69.7 %) › 7 tribal taxa (= 87.5 %) › 11 subfamily taxa (= 100.0 %) ◊ Outgroup taxa › 19 Asilidae species › 10 Asiloidea species › 3 Nemestrinoidea › 1 Tabanomorpha Mydidae collecting localities of specimens included characters 5 ◊ 214 discrete, parsimony informative characters ◊ 36 non-genitalic features sexually dimorphic tagma # characters remarks Head 44 10 sexually dimorphic Thorax 101 22 sexually dimorphic 45 chaetotaxy, sclerites etc. 22 wing 34 leg Abdomen 67 4 sexually dimorphic 11 chaetotaxy, sclerites etc. 30 ? terminalia 26 / terminalia characters – sexual dimorphism 6 Syllegomydas vittatus ? 36 characters / › 30 char. ? terminalia › 19 char. / terminalia phylogenetic analysis 7 ◊ TNT 1.1 (February 2014) Goloboff et al. 2008 ◊ exhaustive search of tree space (MacPro 3.5 GHz 6-core Intel Xeon E5, 64 GB RAM) command(s) time (min) # re-arrangements examined mult1000 02:19 38,676,135,139 xmult=rss hits 20 fuse 2 drift 3 ratchet 10 multiply consense 10 15:54 196,237,489,228 random sectorial searches after hitting best length, find additional trees by fusing suboptimal with optimal trees hit best length 20x consense until strict consensus is stabilized 10x ◊ Bremer support Bremer 1988 ◊ character analysis Mesquite (2.75) and WinClada/Asado (1.08) monophyly + sister taxon 8 outgroups Apiocera An obscura Apiocera An philippii Apiocera Ap helenae ◊ (233 most parsimonious trees) Apiocera Ap imminuta tree length 1870 steps Apiocera Ap tonnoiri Apiocera R alastor M Apiocera R braunsi Apiocera Ap goerlingi Apiocera Ap asilica Apiocera Ap brevicornis Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Apioceridae 30 ◊ Apiocera P bilineata Asilidae + (Apioceridae + Mydidae) Apiocera P mexicana M Apiocera P acuticauda Apiocera P barri Apiocera P martinorum Apiocera P parahydra Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett M Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Tabanidae 1 Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Midacritus stuardoanus M Midacritus wagenknechti Nemestrinidae 2 Miltinus sp. (Northern Territory) 15 Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Ectyphus amboseli M Ectyphus armipes Ectyphus pinguis Parectyphus namibiensis Heteromydas bicolor Opomydas townsendi Acroceridae 1 Paramydas igniticornis Messiasia californica ≥21 Messiasia painteri Mydas xanthopterus Stratiomydas lividus Mydas clavatus Mydas tricinctus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Plyomydas sp nov Tandil Plyomydas peruviensis M Cacatuopyga basifascia Bombyliidae 2 Cacatuopyga ruficornis M 4 Anomalomydas australicus F Anomalomydas mackerrasi 1 Anomalomydas sp nov Simpson F Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Mydidae 138 Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Halterorchis sp nov Cape Recife M Bombyliidae 1 Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M 2 Leptomydas notos Nemomydas pantherinus Nemomydas melanopogon Namadytes cimbebasiensis Namadytes vansoni Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Apsilocephalidae 1 Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Nemomydas lamia 1 Agaperemius sp Schlafkuppe M Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata M Scenopinidae 3 Cephalocera sp Ratelfontein 2 Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F 2 Mydaselpis ngurumani Neolaparopsis puncturatus Mydaselpis sp nov Daan Therevidae 3 Mydaselpis sp nov Gamsberg M 1 Vespiodes leopolditertii Vespiodes phaios Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Afroleptomydas A suffusipennis Afroleptomydas A paganus Afroleptomydas A inhacae Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Asilidae 19 Afroleptomydas C sp Koeroegab 5 Afroleptomydas A omeri Afroleptomydas A rufithorax Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Afroleptomydas C sp Gobabeb Stenomydas gen nov sp nov Panne M Syllegomydas S lineatus Syllegomydas sp nov Ruacana Mimadelphus sp Gobabis M Hispanomydas hispanicus 1 Syllegomydas S vittatus Syllegomydas S palestinensis Apioceridae 30 Syllegomydas S algiricus 13 Syllegomydas S proximus Syllegomydas S elachys Syllegomydas N astrictus Namibimydas psamminos Namibimydas stuckenbergi Nothomydas picketti Cleamydas gen nov sp nov Ugab F 5 Syll gen nov sp nov minute proboscis F Syllegomydas N arnoldi M Syllegomydas N dispar Syllegomydas N heothinos Diochlistinae Argentina F Mydidae 138 Mitrodetus nanoglossa M Mitrodetus microglossa 5 Mitrodetus dentitarsis strict consensus topology Mitrodetus dimidiatus relationships within Apioceridae 9 ◊ Afrotropical + North American subgenera monophyletic ◊ Australian Apiocera non-monophyletic Apiocera An obscura Apiocera An philippii Apiocera (Anypenus) Apiocera Ap helenae Apiocera Ap imminuta Apiocera (Apiocera) Apiocera Ap tonnoiri 11 32 35 37 59 60 64 7484 85131 4 63 64 97 121203 213 Apiocera R alastor M Apiocera (Ripidosyrma) 1 0 5 0 0 2 1 1 3 3 0 1 0 0 1 1 1 1 Apiocera R braunsi 13 7 3 4 64 75 Apiocera Ap goerlingi Apiocera Ap asilica 3 3 1 1 Apiocera Ap brevicornis 3 5 Apiocera Ap sp Stuart M 2 Apiocera (Apiocera) Apiocera Ap hardyi 4 Apiocera Ap pica 2 Apiocera Ap sp Lloyd 2 Apiocera Ap similis 9 Apiocera Ap maritima 9 Apiocera Ap minor ≥21 Apiocera Ap newmani Apiocera P bilineata 6 Apiocera P mexicana M 3 38 74 97 130194 199203215 Apiocera P acuticauda 1 2 0 1 0 1 1 1 1 Apiocera P barri 2 Apiocera P martinorum Apiocera (Pyrocera) 2 Apiocera P parahydra Apiocera P haruspex Apiocera P aldrichi strict consensus topology 2 5 Apiocera P painteri 2 Apiocera P arena unambiguous optimization Apiocera P parkeri 7 Bremer below branches Apiocera P sp Colnett M relationships within Mydidae 10 Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri √ Rhaphiomidinae Tongamya miranda ◊ Rhaphiomidinae and Megascelinae earliest branches Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M √ Megascelinae Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistinae Diochlistus melleipennis Midacritus stuardoanus M ◊ Mydinae to include Neotropical genera Paramydas and Midacritus wagenknechti Miltinus sp. (Northern Territory) Apiophora paulseni Apiophora quadricinctata Apiophorinae Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Ectyphus amboseli M Plyomydas and Oriental genus Cacatuopyga Ectyphus armipes Ectyphus pinguis Parectyphus namibiensis √ Ectyphinae Heteromydas bicolor Opomydas townsendi Paramydas igniticornis Apiophorinae Messiasia californica Messiasia painteri Mydas xanthopterus Stratiomydas lividus Mydas clavatus Mydinae Mydas tricinctus ◊ Australian Anomalomydas distinct clade Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Plyomydas sp nov Tandil Plyomydas peruviensis M Cacatuopyga basifascia Cacatuopyga ruficornis M Anomalomydas australicus F √ Cacatuopyginae Anomalomydas mackerrasi Anomalomydas sp nov Simpson F √ Anomalomydinae Pseudorhopalia mirandai Rhopalia annulata ◊ Rhopalia cincta speciose Leptomydinae and Syllegomydinae Rhopalia olivieri Rhopaliinae Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Halterorchis sp nov Cape Recife M Haplomydas crassipes non-monophyletic