TDikow Apioceridae + Mydidae morphological phylogeny ESA 2012
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Current (Nov 2012) phylogenetic hypothesis of relationships within Apioceridae and Mydidae (Diptera: Asiloidea)
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TDikow Apioceridae + Mydidae morphological phylogeny ESA 2012
- 1. 1 Phylogeny and historical biogeography of Apioceridae and Mydidae inferred from morphological characters of imagines (Diptera: Asiloidea) Apioceridae: Apiocera sp. © J. Hort Mydidae: Miltinus minutus © J. Hort Torsten Dikow Department of Entomology Smithsonian National Museum of Natural History National Museum of Natural History
- 2. . introduction 2 Diptera = 159,294 valid species (Pape et al. 2011) Deuterophlebiidae Nymphomyiidae Tipulomorpha Ptychopteridae Psychodomorpha Culicomorpha Perissommatidae Bibionomorpha Nemestrinidae Neodiptera Xylophagidae Tabanomorpha Acroceridae Orthorrhapha Hilarimorphidae Bombyliidae Stratiomyomorpha Asilidae Brachycera Asiloidea Mydidae > 8,100 species Empidoidea Apioceridae Apystomyiidae > 15,500 species Eremoneura Therevidae Platypezoidea Scenopinidae Syrphidae Cyclorrhapha Apsilocephalidae Pipunculidae Acalyptratae I Evocoidae Acalyptratae II Schizophora Ephydroidea Calyptratae phylogeny from Wiegmann et al. 2011 (simplified), images Apiocera alleni, Leptomydas lusitanicus, Cyrtopogon lutatius, Echthodopa formosa © G. Ballmer, V. Jacinto, M. Thomas
- 3. . introduction 3 Apioceridae – “flower-loving flies” ◊ adult flies flower-feeders ◊ size = 15 – 30 mm ◊ 138 species in 1 genus (4 subgenera) ◊ occur only in › Argentina and Chile –> 4 species › Australia –> 67 species Apiocera (Apiocera) pica © J. Hort › South Africa –> 3 species › western North America (Canada, Mexico, USA) –> 64 species Apiocera (Apiocera) sp. © J. Hort
- 4. . introduction 4 Mydidae – “mydas flies” ◊ adult flies 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
- 5. . introduction 5 Yeates & Irwin 1996 – study of Apioceridae relationships Apioceridae Apiocera (Pyrocera) western North America area relationships Apiocera (Ripidosyrma) South Africa Apiocera (Anypenus) Argentina / Chile Apiocera (Apiocera) Australia
- 6. . introduction 6 Yeates & Irwin 1996 – study of Apioceridae relationships Apioceridae Apiocera (Pyrocera) western North America area relationships Mydidae Rhaphiomidas Apiocera (Ripidosyrma) South Africa Tongamya Apiocera (Anypenus) Argentina / Chile Megascelus Neorhaphiomidas Apiocera (Apiocera) Australia remaining Mydidae
- 7. .. phylogenetic analysis 7 Taxon Sampling ◊ 134 Mydidae species (= 28.4 %) › 46 genera (= 69.7 %) › 7 tribal taxa (= 87.5 %) › 11 subfamily taxa (= 100.0 %) ◊ 27 Apiocera species (= 19.6 %) Mydidae ◊ Outgroup taxa › 19 Asilidae species › 7 Asiloidea species › 3 Nemestrinoidea › 1 Tabanomorpha Apioceridae collecting localities of specimens included
- 8. .. phylogenetic analysis 8 Characters ◊ 214 discrete, parsimony informative characters ◊ 36 non-genitalic features sexually dimorphic tagma # characters remarks Head 44 10 sexually dimorphic Thorax 103 22 sexually dimorphic 47 chaetotaxy, sclerites etc. 23 wing 33 leg Abdomen 67 4 sexually dimorphic 11 chaetotaxy, sclerites etc. 30 ? terminalia 26 / terminalia ◊ DNA data are being gathered
- 9. .. phylogenetic analysis 9 Characters – sexual dimorphism Syllegomydas vittatus ? 36 characters / › 30 char. ? terminalia › 19 char. / terminalia
- 10. .. phylogenetic analysis 10 Phylogenetic analysis ◊ TNT 1.1 (July 2012) Goloboff et al. 2008 ◊ exhaustive search of tree space (MacBook Pro 2.2 GHz Intel Core i7) command(s) time (min) # re-arrangements examined mult300 01:04 8,383,595,488 xmult=rss hits 20 fuse 2 drift 3 ratchet 10 multiply consense 10 12:31 79,691,144,513 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 ◊ DIVA 1.2 Ronquist 1994, Nylander Mac version
- 11. ... results 11 Results Limata tenuicornis F Eulonchus sp Six Rivers M Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Dipalta serpentina Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra ◊ 440 most parsimonious trees Phycus frommeri Asilidae Diogmites grossus Ospriocerus aeacus Gonioscelis ventralis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps Hoplistomerus nobilis Pilica formidolosa Laphria aktis Dioctria hyalipennis Pegesimallus laticornis Leptogaster cylindrica Euscelidia pulchra Stichopogon punctum ◊ 1837 steps Lasiopogon cinctus Damalis monochaetes Holcocephala calva Apiocera Ap imminuta Apioceridae Apiocera Ap tonnoiri Apiocera R alastor M Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana M 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 ◊ strict consensus Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga 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 › several polytomies Apiocera Ap newmani Rhaphiomidas hasbroucki Rhaphiomidas maehleri Mydidae Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah › not well-resolved are: Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus viduatus Miltinus maculipennis › earliest branchings within Apioceridae Miltinus minutus Ectyphus armipes Parectyphus namibiensis Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Paramydas igniticornis Plyomydas peruviensis M Plyomydas sp nov Tandil Dolichogaster brevicornis Messiasia californica M › derived Mydidae ingroup Messiasia painteri Mydas clavatus Mydas tricinctus Mydas xanthopterus Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruficornis M Phyllomydas bruesii Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Halterorchis sp nov Cape Recife M Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Leptomydas notos Nemomydas pantherinus Nemomydas melanopogon Namadytes cimbebasiensis Namadytes vansoni Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Nemomydas lamia Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb Hispanomydas hispanicus Mydaselpis ngurumani Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Syllegomydas S algiricus Limata tenuicornis F Eulonchus sp Six Rivers M Syllegomydas S lineatus Syllegomydas S palestinensis Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Dipalta serpentina Syllegomydas S proximus Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Syllegomydas S vittatus Phycus frommeri Diogmites grossus Ospriocerus aeacus Syllegomydas sp nov Ruacana Gonioscelis ventralis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Stenomydas gen nov sp nov Panne M Machimus occidentalis Philonicus albiceps Hoplistomerus nobilis Pilica formidolosa Pseudonomoneura hirta Laphria aktis Dioctria hyalipennis Pegesimallus laticornis Leptogaster cylindrica Pseudonomoneura micheneri Euscelidia pulchra Stichopogon punctum Lasiopogon cinctus Afroleptomydas C mauricei Damalis monochaetes Holcocephala calva Apiocera Ap imminuta Apiocera Ap tonnoiri Afroleptomydas C sp Stampriet Apiocera R alastor M Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana M Pseudonomoneura bajaensis Apiocera P barri Apiocera P martinorum Apiocera P parahydra Eremomidas sp nov Tiega M Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Eremomidas arabicus Apiocera P parkeri Apiocera P sp Colnett M Apiocera An obscura Apiocera An philippii Halterorchis karooensis F Apiocera Ap sp Wurarga Apiocera Ap asilica Apiocera Ap brevicornis Apiocera Ap sp Stuart M Neolaparopsis puncturatus Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Mydaselpis sp nov Daan Apiocera Ap similis Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Mydaselpis sp nov Gamsberg M Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Vespiodes leopolditertii Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Vespiodes phaios Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Afroleptomydas A consanguinensis Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Afroleptomydas A opacicinctus Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Afroleptomydas A sp Clanwilliam Midacritus wagenknechti Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Afroleptomydas A sp Obib Miltinus viduatus Miltinus maculipennis Miltinus minutus Ectyphus armipes Afroleptomydas C sp Koeroegab Parectyphus namibiensis Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Afroleptomydas A omeri Opomydas townsendi Paramydas igniticornis Plyomydas peruviensis M Afroleptomydas A rufithorax Plyomydas sp nov Tandil Dolichogaster brevicornis Messiasia californica M Messiasia painteri Arenomydas partitus Mydas clavatus Mydas tricinctus Mydas xanthopterus Stratiomydas lividus Arenomydas caerulescens Cacatuopyga basifascia Cacatuopyga ruficornis M Phyllomydas bruesii Arenomydas callosus Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Arenomydas namaquensis M Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Notosyllegomydas brincki strict consensus topology (2121 steps) Hessemydas daugeroni Hessemydas seyrigi Halterorchis sp nov Cape Recife M Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Leptomydas notos Syll gen nov sp nov Gobabeb Cephalocera longirostris Nemomydas pantherinus Nemomydas melanopogon Namadytes cimbebasiensis Namadytes vansoni Cephalocera sp Papendorp Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Cephalocera sp Sors Sors Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Cephalocera imitata M Nemomydas gruenbergi M Leptomydas sardous Nemomydas lamia Afroleptomydas A inhacae Cephalocera sp Ratelfontein Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb Hispanomydas hispanicus Syllegomydas S elachys Mydaselpis ngurumani Nomoneura caffra Nomoneura paradoxa Syllegomydas N astrictus Nomoneuroides natalensis Bremer ≥ 3 Syllegomydas S algiricus Syllegomydas S lineatus Syllegomydas S palestinensis Namibimydas psamminos Syllegomydas S proximus Syllegomydas S vittatus Syllegomydas sp nov Ruacana Stenomydas gen nov sp nov Panne M Namibimydas stuckenbergi Pseudonomoneura hirta Pseudonomoneura micheneri Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Nothomydas picketti Pseudonomoneura bajaensis Eremomidas sp nov Tiega M Eremomidas arabicus Cleamydas gen nov sp nov Ugab F Halterorchis karooensis F Neolaparopsis puncturatus Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Syll gen nov sp nov minute proboscis F Vespiodes leopolditertii Vespiodes phaios Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Syllegomydas N arnoldi M Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Syllegomydas N dispar Afroleptomydas A omeri Afroleptomydas A rufithorax Arenomydas partitus Arenomydas caerulescens Syllegomydas N heothinos Arenomydas callosus Arenomydas namaquensis M Notosyllegomydas brincki Syll gen nov sp nov Gobabeb Diochlistinae Argentina F Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata M Cephalocera sp Ratelfontein Syllegomydas S elachys Syllegomydas N astrictus Mitrodetus nanoglossa M Mitrodetus microglossa Namibimydas psamminos Namibimydas stuckenbergi Nothomydas picketti Cleamydas gen nov sp nov Ugab F Mitrodetus dentitarsis Syll gen nov sp nov minute proboscis F Syllegomydas N arnoldi M Syllegomydas N dispar Syllegomydas N heothinos Mitrodetus dimidiatus Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus
- 12. ... results 12 Placement of † Cretomydas santanensis Mydidae Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis ◊ explicitly included in matrix Neorhaphiomidas hardyi ≈ 112 myo Neorhaphiomidas sp Mandurah Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti ◊ 80% missing data Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Ectyphus armipes Ectyphus amboseli M Ectyphus pinguis Parectyphus namibiensis Heteromydas bicolor Opomydas townsendi ◊ wing venation very similar to Diochlistus Paramydas igniticornis Messiasia californica M Messiasia painteri Plyomydas sp nov Tandil Plyomydas peruviensis M Cacatuopyga basifascia Cacatuopyga ruficornis M Mydas clavatus Mydas tricinctus (Australia) and Rhaphiomidas (wNA) Mydas xanthopterus Stratiomydas lividus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia cincta Rhopalia annulata Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Halterorchis sp nov Cape Recife M Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Leptomydas notos Nemomydas pantherinus Nemomydas melanopogon Namadytes cimbebasiensis Namadytes vansoni Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Pseudonomoneura hirta Pseudonomoneura micheneri Pseudonomoneura bajaensis Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F Nemomydas lamia Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Notosyllegomydas brincki Syll gen nov sp nov Gobabeb Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata M Cephalocera sp Ratelfontein Mydaselpis ngurumani Neolaparopsis puncturatus Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Nomoneura caffra Nomoneura paradoxa Limata tenuicornis F Eulonchus sp Six Rivers M Nomoneuroides natalensis Afroleptomydas A suffusipennis Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Dipalta serpentina Toxophora maxima Afroleptomydas A paganus Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Phycus frommeri Diogmites grossus Ospriocerus aeacus Afroleptomydas A inhacae Gonioscelis ventralis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Afroleptomydas A consanguinensis Philonicus albiceps Hoplistomerus nobilis Pilica formidolosa Laphria aktis Dioctria hyalipennis Afroleptomydas A opacicinctus Pegesimallus laticornis Leptogaster cylindrica Euscelidia pulchra Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Afroleptomydas A sp Clanwilliam Holcocephala calva Apiocera Ap imminuta Apiocera Ap tonnoiri Apiocera R alastor M Apiocera R braunsi Afroleptomydas C sp Koeroegab Apiocera P bilineata Apiocera P mexicana M Apiocera P barri Apiocera P martinorum Apiocera P parahydra Afroleptomydas A sp Obib Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett M Afroleptomydas A omeri Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga † Cretomydas santanensis, Crato Formation, Brazil Apiocera Ap asilica Apiocera Ap brevicornis Afroleptomydas A rufithorax Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Afroleptomydas C mauricei Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Afroleptomydas C sp Stampriet Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Afroleptomydas C sp Gobabeb Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Stenomydas gen nov sp nov Panne M Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Syllegomydas S lineatus Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Syllegomydas sp nov Ruacana Miltinus minutus Ectyphus armipes Parectyphus namibiensis Ectyphus amboseli M Ectyphus pinguis Hispanomydas hispanicus Heteromydas bicolor Opomydas townsendi Paramydas igniticornis Plyomydas peruviensis M Plyomydas sp nov Tandil Syllegomydas S vittatus Dolichogaster brevicornis Messiasia californica M Messiasia painteri Mydas clavatus Mydas tricinctus Mydas xanthopterus Syllegomydas S palestinensis Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruficornis M Phyllomydas bruesii Phyllomydas parvulus Syllegomydas S algiricus Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Syllegomydas S proximus Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Halterorchis sp nov Cape Recife M Haplomydas crassipes Syllegomydas S elachys Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Leptomydas notos Nemomydas pantherinus Nemomydas melanopogon Syllegomydas N astrictus Namadytes cimbebasiensis Namadytes vansoni Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Namibimydas psamminos Eremomidas bek Leptomydas rapti M Leptomydas turcicus 1 of the mp trees Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Namibimydas stuckenbergi Nemomydas lamia Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb Nothomydas picketti Hispanomydas hispanicus Mydaselpis ngurumani Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Cleamydas gen nov sp nov Ugab F Syllegomydas S algiricus Syllegomydas S lineatus Syllegomydas S palestinensis Syllegomydas S proximus Syllegomydas S vittatus Syllegomydas sp nov Ruacana Syll gen nov sp nov minute proboscis F Stenomydas gen nov sp nov Panne M Pseudonomoneura hirta Pseudonomoneura micheneri Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Syllegomydas N arnoldi M Pseudonomoneura bajaensis Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F Neolaparopsis puncturatus Syllegomydas N dispar Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Syllegomydas N heothinos Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Diochlistinae Argentina F Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Notosyllegomydas brincki Mitrodetus nanoglossa M Syll gen nov sp nov Gobabeb Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata M Cephalocera sp Ratelfontein Mitrodetus microglossa Syllegomydas S elachys Syllegomydas N astrictus Namibimydas psamminos Namibimydas stuckenbergi Nothomydas picketti Mitrodetus dentitarsis Cleamydas gen nov sp nov Ugab F Syll gen nov sp nov minute proboscis F Syllegomydas N arnoldi M Syllegomydas N dispar Syllegomydas N heothinos Mitrodetus dimidiatus Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus
- 13. ... results 13 Placement of † Cretomydas santanensis Mydidae Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas hardyi ≈ 112 myo Neorhaphiomidas sp Mandurah Diochlistinae Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Australia Midacritus stuardoanus M Midacritus wagenknechti Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Ectyphus armipes Ectyphus amboseli M Ectyphus pinguis Parectyphus namibiensis Heteromydas bicolor Opomydas townsendi Paramydas igniticornis Messiasia californica M Messiasia painteri Plyomydas sp nov Tandil Plyomydas peruviensis M Cacatuopyga basifascia Cacatuopyga ruficornis M Mydas clavatus † Cretomydas santanensis, Crato Formation, Brazil Mydas tricinctus Mydas xanthopterus Stratiomydas lividus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia cincta Rhopalia annulata Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Non-monophyly of Diochlistinae Hessemydas daugeroni Hessemydas seyrigi Halterorchis sp nov Cape Recife M Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Leptomydas notos Nemomydas pantherinus Nemomydas melanopogon Namadytes cimbebasiensis Namadytes vansoni Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Pseudonomoneura hirta Pseudonomoneura micheneri Pseudonomoneura bajaensis Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F Nemomydas lamia Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Notosyllegomydas brincki Syll gen nov sp nov Gobabeb Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata M Cephalocera sp Ratelfontein Mydaselpis ngurumani Neolaparopsis puncturatus Diochlistus auripennis, Australia © P. Chew Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Nomoneura caffra Nomoneura paradoxa Limata tenuicornis F Eulonchus sp Six Rivers M Nomoneuroides natalensis Afroleptomydas A suffusipennis Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Dipalta serpentina Toxophora maxima Afroleptomydas A paganus Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Phycus frommeri Diogmites grossus Ospriocerus aeacus Afroleptomydas A inhacae Gonioscelis ventralis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Afroleptomydas A consanguinensis Philonicus albiceps Hoplistomerus nobilis Pilica formidolosa Laphria aktis Dioctria hyalipennis Afroleptomydas A opacicinctus Pegesimallus laticornis Leptogaster cylindrica Euscelidia pulchra Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Afroleptomydas A sp Clanwilliam Holcocephala calva Apiocera Ap imminuta Apiocera Ap tonnoiri Apiocera R alastor M Apiocera R braunsi Afroleptomydas C sp Koeroegab Apiocera P bilineata Apiocera P mexicana M Apiocera P barri Apiocera P martinorum Apiocera P parahydra Afroleptomydas A sp Obib Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett M Afroleptomydas A omeri Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga Apiocera Ap asilica Apiocera Ap brevicornis Afroleptomydas A rufithorax Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Afroleptomydas C mauricei Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Afroleptomydas C sp Stampriet Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Afroleptomydas C sp Gobabeb Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Stenomydas gen nov sp nov Panne M Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Syllegomydas S lineatus Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Syllegomydas sp nov Ruacana Miltinus minutus Ectyphus armipes Parectyphus namibiensis Ectyphus amboseli M Ectyphus pinguis Hispanomydas hispanicus Heteromydas bicolor Opomydas townsendi Paramydas igniticornis Plyomydas peruviensis M Plyomydas sp nov Tandil Syllegomydas S vittatus Dolichogaster brevicornis Messiasia californica M Messiasia painteri Mydas clavatus Mydas tricinctus Mydas xanthopterus Syllegomydas S palestinensis Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruficornis M Phyllomydas bruesii Phyllomydas parvulus Syllegomydas S algiricus Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Syllegomydas S proximus Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Halterorchis sp nov Cape Recife M Haplomydas crassipes Syllegomydas S elachys Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Leptomydas notos Nemomydas pantherinus Nemomydas melanopogon Syllegomydas N astrictus Namadytes cimbebasiensis Namadytes vansoni Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Namibimydas psamminos Eremomidas bek Leptomydas rapti M Leptomydas turcicus 1 of the mp trees Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Namibimydas stuckenbergi Nemomydas lamia Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb Nothomydas picketti Hispanomydas hispanicus Mydaselpis ngurumani Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Cleamydas gen nov sp nov Ugab F Syllegomydas S algiricus Syllegomydas S lineatus Syllegomydas S palestinensis Syllegomydas S proximus Syllegomydas S vittatus Syllegomydas sp nov Ruacana Syll gen nov sp nov minute proboscis F Stenomydas gen nov sp nov Panne M Pseudonomoneura hirta Pseudonomoneura micheneri Diochlistinae Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Syllegomydas N arnoldi M Pseudonomoneura bajaensis Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F Neolaparopsis puncturatus Syllegomydas N dispar Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Afroleptomydas A consanguinensis Mitdrodetus dentitarsis, Chile © S. Marshall Afroleptomydas A opacicinctus Syllegomydas N heothinos Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Diochlistinae Argentina F Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Argentina/Chile Notosyllegomydas brincki Mitrodetus nanoglossa M Syll gen nov sp nov Gobabeb Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata M Cephalocera sp Ratelfontein Mitrodetus microglossa Syllegomydas S elachys Syllegomydas N astrictus Namibimydas psamminos Namibimydas stuckenbergi Nothomydas picketti Mitrodetus dentitarsis Cleamydas gen nov sp nov Ugab F Syll gen nov sp nov minute proboscis F Syllegomydas N arnoldi M Syllegomydas N dispar Syllegomydas N heothinos Mitrodetus dimidiatus Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus