Phylogeny of Apioceridae and Mydidae (Insecta: Diptera) Based on Morphological Characters of Adult Flies

Phylogeny of Apioceridae and Mydidae (Insecta: Diptera) based on morphological characters of adult flies

Torsten Dikow Biodiversity Synthesis Center Field Museum of Natural History, Chicago, IL, USA . introduction 2 Yeates & Irwin 1996 – Apioceridae: cladistic reappraisal and biogeography › 6 Apioceridae exemplar species › 13 Mydidae exemplar species › 77 morphological characters Pre-1996 = non-monophyletic Apioceridae Post-1996 = monophyletic Apioceridae Asilidae Asilidae Apiocera (Pyrocera) Apiocera (Pyrocera) Apiocera (Ripidosyrma) Apioceridae Apiocera (Ripidosyrma) Apiocera (Anypenus) Apiocera (Anypenus) Apiocera (Apiocera) Apiocera (Apiocera) Rhaphiomidinae Rhaphiomidinae Megascelinae Megascelinae Rhopaliinae Rhopaliinae Cacatuopyginae Cacatuopyginae Leptomydinae Mydidae Leptomydinae Syllegomydinae Syllegomydinae Anomalomydinae Anomalomydinae Diochlistinae Diochlistinae Ectyphinae Ectyphinae Apiophorinae Apiophorinae Mydinae Mydinae . introduction 3 Yeates & Irwin 1996 – Exclusively southern connection taxa = Gondwanan Apiocera (Pyrocera) western North America

Apiocera (Ripidosyrma) Tongamya South Africa

Apioceridae

Mydidae: Megascelinae Apiocera (Anypenus) Megascelus Mitrodetus Argentina + Chile Apiophora, Midacritus

Mydidae: Diochlistinae

Mydidae: Apiophorinae Apiocera (Apiocera) Neorhaphiomidas Diochlistus Australia Miltinus . introduction 4 Apioceridae – “flower-loving flies” ◊ Adult flies flower-feeders ◊ larvae possibly predatory

◊ 138 species and 1 genus (4 subgenera) ◊ length = 15 – 30 mm ◊ occur only in › Argentina and Chile – 4 species Apiocera (Apiocera) moerens © Peter Chew › Australia – 67 species › South Africa – 3 species › western North America (Canada, Mexico, USA) – 64 species

Apiocera bilineata © Greg Ballmer . introduction 5 Mydidae – “mydas flies” ◊ Adult flies flower-feeders, some do not feed at all ◊ Larvae possibly predatory ◊ 472 species and 66 genera world-wide

◊ length = 9 – 60 mm (wing length up to 52 mm) ◊ World-wide primarily in arid regions Mydidae: Opomydas townsendi ◊ Rarely collected – Afrotropical Region most speciose ◊ Oldest definitive fossil† Cretomydas santanensis › Crato Formation, Brazil › Cretaceous: Aptian – Albian › ≈ 112 myo

Mydidae: Eremomidas arabicus .. methods 6 Taxon Sampling ◊ 27 Apiocera species (= 19.6 %)

◊ 134 Mydidae species (= 28.4 %) › 46 genera (= 69.7 %) › 7 tribal taxa (= 87.5 %) › 11 subfamily taxa (= 100.0 %) Apioceridae ◊ Outgroup taxa › 37 Asiloidea species › 3 Nemestrinoidea › 1 Tabanomorpha

Mydidae

collecting localities of specimens included .. methods 7 Morphological Characters ◊ 211 discrete, parsimony informative characters ◊ 36 coded separately for female and male semaphoronts

tagma # characters remarks Head 44 Thorax 103 46 chaetotaxy, sclerites, etc. 24 wing 33 leg Abdomen 62 13 sclerite shapes, chaetotaxy, etc. 30 ? terminalia 19 / terminalia .. methods 8 Phylogenetic Analysis ◊ Data assembly and visualization › Mesquite (version 2.75) › FigTree (version 1.3.1) › TreeMap3 (version 3.0b) ◊ Tree Search › TNT (version 1.1 June 2012) › Ratchet, TreeDrifting Mydidae: Eremomidas arabicus © Drew Gardner › all multi-state characters non-additive › all characters equally weighted ◊ Bremer Support › TNT (version 1.1 June 2012)

Mydidae: Heteromydas bicolor © Greg Ballmer ... new phylogenetic hypothesis 9

Limata tenuicornis Namadytes sp Gobabeb M Eulonchus sp Six Rivers Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Namadytes vansoni Dipalta serpentina Toxophora maxima Prorates sp Escalante • Neolaparopsis puncturatus Apsilocephala longistyla Hemigephyra atra Phycus frommeri Diogmites grossus Haplomydas crassipes Pegesimallus laticornis Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Eremohaplomydas desertorum Ospriocerus aeacus Gonioscelis ventralis Stichopogon punctum Lachnocorynus sp nov Kotwa M Lasiopogon cinctus • Damalis monochaetes Holcocephala calva Hoplistomerus nobilis • Mydaselpis sp nov Daan Pilica formidolosa Laphria aktis Emphysomera pallidapex Ommatius tibialis Mydaselpis sp nov Gamsberg M Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps • Vespiodes leopolditertii Apiocera R alastor M Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana M • Vespiodes phaios Apiocera P barri Apiocera P martinorum Apiocera P parahydra Apiocera P haruspex • Mydaselpis ngurumani Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Arenomydas caerulescens Apiocera P sp Colnett M Apiocera Ap imminuta Apiocera Ap tonnoiri Arenomydas callosus Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga Apiocera Ap asilica Arenomydas namaquensis M Apiocera Ap brevicornis Apiocera Ap sp Stuart M • Apiocera Ap hardyi Apiocera Ap pica Arenomydas partitus Apiocera Ap sp Lloyd Apiocera Ap similis Apiocera Ap maritima Apiocera Ap minor Syll gen nov sp nov Gobabeb Apiocera Ap newmani Rhaphiomidas hasbroucki Rhaphiomidas maehleri Notosyllegomydas brincki Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Eremomidas sp nov Tiega M Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Eremomidas arabicus Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Halterorchis karooensis F Diochlistus gracilis • Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Afroleptomydas A inhacae Midacritus stuardoanus M Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Afroleptomydas A paganus Apiophora paulseni Apiophora quadricinctata Ectyphus amboseli M Afroleptomydas A suffusipennis Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Afroleptomydas C sp Gobabeb Miltinus viduatus Miltinus maculipennis Miltinus minutus Paramydas igniticornis Syllegomydas sp nov Ruacana Cacatuopyga basifascia Cacatuopyga ruficornis M Stratiomydas lividus Mydas clavatus Stenomydas gen nov sp nov Panne M Mydas tricinctus Mydas xanthopterus Dolichogaster brevicornis Afroleptomydas A consanguinensis Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Plyomydas sp nov Tandil Afroleptomydas A opacicinctus Messiasia californica M Messiasia painteri Pseudorhopalia mirandai Rhopalia annulata Afroleptomydas C mauricei Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Afroleptomydas C sp Stampriet Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi • Afroleptomydas A sp Clanwilliam Nemomydas melanopogon Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Afroleptomydas A sp Obib Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Afroleptomydas C sp Koeroegab Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Afroleptomydas A omeri Leptomydas sardous Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Afroleptomydas A rufithorax Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Nothomydas picketti Namibimydas psamminos Syllegomydas S lineatus Namibimydas stuckenbergi Cephalocera imitata M Cephalocera sp Ratelfontein Hispanomydas hispanicus Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M Syllegomydas S algiricus Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Namadytes sp Gobabeb M Syllegomydas S elachys Namadytes vansoni Neolaparopsis puncturatus Haplomydas crassipes Syllegomydas S palestinensis Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Syllegomydas S proximus Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Syllegomydas S vittatus Arenomydas callosus Arenomydas namaquensis M Arenomydas partitus Syll gen nov sp nov Gobabeb Syllegomydas N astrictus Notosyllegomydas brincki Eremomidas sp nov Tiega M Eremomidas arabicus Syllegomydas N heothinos Halterorchis karooensis F Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Syll gen nov sp nov minute proboscis F Afroleptomydas C sp Gobabeb Syllegomydas sp nov Ruacana Stenomydas gen nov sp nov Panne M Afroleptomydas A consanguinensis Syllegomydas N arnoldi M Afroleptomydas A opacicinctus Bremer 3 – 9 Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Syllegomydas N dispar Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri • Diochlistinae Argentina F Afroleptomydas A rufithorax Syllegomydas S lineatus Hispanomydas hispanicus Syllegomydas S algiricus Mitrodetus nanoglossa M Syllegomydas S elachys Syllegomydas S palestinensis Syllegomydas S proximus Mitrodetus microglossa Syllegomydas S vittatus Syllegomydas N astrictus • Syllegomydas N heothinos Syll gen nov sp nov minute proboscis F Mitrodetus dentitarsis Syllegomydas N arnoldi M Syllegomydas N dispar • Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus dimidiatus Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus Bremer ≥ 10 strict consensus of 1040 trees • ... new phylogenetic hypothesis 10

Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Placement of † Cretomydas santanensis Neorhaphiomidas sp Mandurah ≈ 112 myo Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Australia Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Apiophora quadricinctata Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Nemomydas melanopogon Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta † Cretomydas santanensis, Brazil Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Cephalocera imitata M Cephalocera sp Ratelfontein Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Non-monophyly of Diochlistinae Halterorchis sp nov Cape Recife M Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Arenomydas partitus

Limata tenuicornis Eulonchus sp Six Rivers Syll gen nov sp nov Gobabeb Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Dipalta serpentina Toxophora maxima Notosyllegomydas brincki Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Phycus frommeri Diogmites grossus Pegesimallus laticornis Eremomidas sp nov Tiega M Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Ospriocerus aeacus Gonioscelis ventralis Eremomidas arabicus Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Holcocephala calva Hoplistomerus nobilis Halterorchis karooensis F Pilica formidolosa Laphria aktis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Afroleptomydas A inhacae Philonicus albiceps Apiocera R alastor M Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana M Afroleptomydas A paganus Apiocera P barri Apiocera P martinorum Apiocera P parahydra Apiocera P haruspex Apiocera P aldrichi Afroleptomydas A suffusipennis Apiocera P painteri Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett M Apiocera Ap imminuta Afroleptomydas C sp Gobabeb Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga Apiocera Ap asilica Apiocera Ap brevicornis Syllegomydas sp nov Ruacana Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Stenomydas gen nov sp nov Panne M Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Rhaphiomidas hasbroucki Rhaphiomidas maehleri Afroleptomydas A consanguinensis Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Afroleptomydas A opacicinctus Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Afroleptomydas C mauricei Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Afroleptomydas C sp Stampriet Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Apiophora quadricinctata Afroleptomydas A sp Clanwilliam Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Miltinus viduatus Afroleptomydas A sp Obib Miltinus maculipennis Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Afroleptomydas C sp Koeroegab Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Dolichogaster brevicornis Afroleptomydas A omeri Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Afroleptomydas A rufithorax Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Syllegomydas S lineatus Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Nemomydas melanopogon Hispanomydas hispanicus Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Syllegomydas S algiricus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Syllegomydas S elachys Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Syllegomydas S palestinensis Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Cephalocera imitata M Cephalocera sp Ratelfontein Syllegomydas S proximus Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M Nomoneura caffra Nomoneura paradoxa Syllegomydas S vittatus Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Neolaparopsis puncturatus Haplomydas crassipes Syllegomydas N astrictus Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Syllegomydas N heothinos Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Arenomydas partitus Syll gen nov sp nov minute proboscis F Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F Syllegomydas N arnoldi M Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb Syllegomydas sp nov Ruacana Syllegomydas N dispar Stenomydas gen nov sp nov Panne M Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Diochlistinae Argentina F Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Mitrodetus nanoglossa M Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S palestinensis Syllegomydas S proximus Mitrodetus microglossa Syllegomydas S vittatus Syllegomydas N astrictus Syllegomydas N heothinos Syll gen nov sp nov minute proboscis F Syllegomydas N arnoldi M Mitrodetus dentitarsis Syllegomydas N dispar Diochlistinae Argentina F Mitrodetus nanoglossa M Argentina + Chile Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus strict consensus of 1040 trees Mitrodetus dimidiatus Diochlistus auripennis, Australia © Peter Chew ... new phylogenetic hypothesis 11

Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Rhaphiomidinae Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Megascelinae Neorhaphiomidas hardyi Monophyly of subfamily taxa Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Anomalomydinae Midacritus stuardoanus M Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Apiophora quadricinctata Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Cacatuopyginae Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Mydinae Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Nemomydas melanopogon Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Syllegomydinae: Cephalocera imitata Cleamydas gen nov sp nov Ugab F Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Cephalocera imitata M Cephalocera sp Ratelfontein Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Arenomydas partitus

Limata tenuicornis Eulonchus sp Six Rivers Syll gen nov sp nov Gobabeb Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Dipalta serpentina Toxophora maxima Notosyllegomydas brincki Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Phycus frommeri Diogmites grossus Pegesimallus laticornis Eremomidas sp nov Tiega M Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Ospriocerus aeacus Gonioscelis ventralis Eremomidas arabicus Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Holcocephala calva Hoplistomerus nobilis Halterorchis karooensis F Pilica formidolosa Laphria aktis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Afroleptomydas A inhacae Philonicus albiceps Apiocera R alastor M Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana M Afroleptomydas A paganus Apiocera P barri Apiocera P martinorum Apiocera P parahydra Apiocera P haruspex Apiocera P aldrichi Afroleptomydas A suffusipennis Apiocera P painteri Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett M Apiocera Ap imminuta Afroleptomydas C sp Gobabeb Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga Apiocera Ap asilica Apiocera Ap brevicornis Syllegomydas sp nov Ruacana Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Stenomydas gen nov sp nov Panne M Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Rhaphiomidas hasbroucki Rhaphiomidas maehleri Afroleptomydas A consanguinensis Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Afroleptomydas A opacicinctus Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Afroleptomydas C mauricei Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Afroleptomydas C sp Stampriet Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Apiophora quadricinctata Afroleptomydas A sp Clanwilliam Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Miltinus viduatus Afroleptomydas A sp Obib Miltinus maculipennis Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Afroleptomydas C sp Koeroegab Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Dolichogaster brevicornis Afroleptomydas A omeri Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Afroleptomydas A rufithorax Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Syllegomydas S lineatus Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Nemomydas melanopogon Hispanomydas hispanicus Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Syllegomydas S algiricus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Syllegomydas S elachys Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Syllegomydas S palestinensis Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Cephalocera imitata M Cephalocera sp Ratelfontein Syllegomydas S proximus Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M Nomoneura caffra Nomoneura paradoxa Syllegomydas S vittatus Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Neolaparopsis puncturatus Haplomydas crassipes Syllegomydas N astrictus Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Syllegomydas N heothinos Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Syll gen nov sp nov minute proboscis F Arenomydas partitus Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F Syllegomydas N arnoldi M Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb monophyletic Syllegomydas sp nov Ruacana Syllegomydas N dispar Stenomydas gen nov sp nov Panne M Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Diochlistinae Argentina F Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Mitrodetus nanoglossa M Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S palestinensis Syllegomydas S proximus Mitrodetus microglossa Syllegomydas S vittatus Syllegomydas N astrictus Syllegomydas N heothinos Syll gen nov sp nov minute proboscis F Syllegomydas N arnoldi M Mitrodetus dentitarsis Syllegomydas N dispar Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus Mitrodetus dimidiatus Syllegomydinae (43 % species diversity) strict consensus of 1040 trees ... new phylogenetic hypothesis 12 Earliest branchings within Mydidae Rhaphiomidas hasbroucki Rhaphiomidinae Rhaphiomidas maehleri western North America Rhaphiomidas parkeri Tongamya miranda Namibia + South Africa Megascelus albovillosus Megascelinae Megascelus melanoproctus Chile Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Australia Neorhaphiomidas sp Mandurah part Diochlistinae † Cretomydas santanensis Brazil Diochlistus auripennis M Diochlistus gracilis Australia Diochlistus melleipennis Anomalomydinae Anomalomydas mackerrasi Anomalomydas australicus F Australia

Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Dipalta serpentina Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Phycus frommeri Diogmites grossus Pegesimallus laticornis Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Ospriocerus aeacus Gonioscelis ventralis Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Holcocephala calva Hoplistomerus nobilis Pilica formidolosa Laphria aktis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps 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 Apiocera Ap imminuta Apiocera Ap tonnoiri 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 Apiocera Ap newmani Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda 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 Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Apiophora quadricinctata Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Nemomydas melanopogon Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Cephalocera imitata M Cephalocera sp Ratelfontein Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Arenomydas partitus Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb Syllegomydas sp nov Ruacana Stenomydas gen nov sp nov Panne M Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S palestinensis Syllegomydas S proximus Syllegomydas S vittatus Syllegomydas N astrictus Syllegomydas N heothinos Syll gen nov sp nov minute proboscis F Syllegomydas N arnoldi M Syllegomydas N dispar Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus Mydidae: Anomalomydas mackerrasi © Jean Hort Mydidae: Neorhaphiomidas sp. © Fred Hort .... study of topology 13 What could be the reason for the poorly resolved topology? ◊ Is the taxon sampling inappropriate? ◊ Does a weighted analysis (PIWE) improve resolution? ◊ Does the presence of sexually dimorphic characters add “noise”? ◊ Does taxon pruning with IterPCR improve resolution? ◊ Did the apomorphic Mydidae experience a sudden radiation intractable for the morphological data used? .... study of topology 14 Sexual dimorphism – separate analyses Syllegomydas vittatus ? 36 characters / Males only Females only

› 128 taxa (- 6) › 114 taxa (- 20)

› 172 characters (- 39) › 162 characters (- 49) › 30 char. ? terminalia › 19 char. / terminalia › 2989 mp trees › 2226 mp trees › 1373 steps › 1362 steps .... study of topology 15

Rhaphiomidas hasbroucki Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Rhaphiomidas parkeri Megascelus albovillosus Neorhaphiomidas sp Yanchep Megascelus melanoproctus Neorhaphiomidas hardyi Megascelus nigricornis Neorhaphiomidas sp Mandurah Neorhaphiomidas sp Mingenew M Tongamya miranda Neorhaphiomidas sp Yanchep Megascelus albovillosus Separate analyses – results Neorhaphiomidas hardyi Megascelus melanoproctus Neorhaphiomidas sp Mandurah Megascelus nigricornis Cretomydas santanensis Diochlistus auripennis M Cretomydas santanensis Diochlistus gracilis Diochlistus gracilis Diochlistus melleipennis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas mackerrasi Midacritus stuardoanus M Anomalomydas australicus F Midacritus wagenknechti Midacritus wagenknechti Miltinus stenogaster Ectyphus armipes Miltinus viduatus Ectyphus pinguis Paramydas igniticornis Ectyphus armipes Parectyphus namibiensis Parectyphus namibiensis Heteromydas bicolor Messiasia californica M Opomydas townsendi Messiasia painteri Apiophora paulseni Pseudorhopalia mirandai Apiophora quadricinctata Apiophora paulseni Miltinus stenogaster Apiophora quadricinctata Miltinus viduatus Miltinus maculipennis Miltinus maculipennis Miltinus minutus Miltinus minutus Cacatuopyga basifascia Cacatuopyga ruficornis M Paramydas igniticornis Ectyphus amboseli M Cacatuopyga basifascia Ectyphus pinguis Mydas clavatus Heteromydas bicolor Mydas tricinctus Opomydas townsendi Mydas xanthopterus Plyomydas peruviensis M Stratiomydas lividus Plyomydas sp nov Tandil Plyomydas sp nov Tandil Rhopalia annulata Messiasia painteri Rhopalia cincta Perissocerus transcaspicus Dolichogaster brevicornis Males Females Phyllomydas bruesii Rhopalia olivieri Dolichogaster brevicornis Phyllomydas parvulus Mydas clavatus Pseudorhopalia mirandai Mydas tricinctus Rhopalia annulata Mydas xanthopterus Rhopalia cincta Stratiomydas lividus Rhopalia olivieri Phyllomydas bruesii Perissocerus dumonti F Phyllomydas parvulus Eremomidas bek Perissocerus transcaspicus Leptomydas corsicanus Eremomidas bek Leptomydas lusitanicus Hessemydas daugeroni Leptomydas notos Hessemydas seyrigi Leptomydas rapti M Leptomydas corsicanus Leptomydas sardous Leptomydas lusitanicus Leptomydas turcicus Leptomydas sardous Nemomydas brachyrhynchus Leptomydas turcicus Nemomydas gruenbergi M Nemomydas brachyrhynchus Nemomydas lamia Nemomydas melanopogon Nemomydas lamia Nemomydas pantherinus Nemomydas melanopogon Nemomydas venosus Nemomydas venosus Pseudonomoneura bajaensis Eremohaplomydas desertorum Hispanomydas hispanicus Haplomydas crassipes Mydaselpis ngurumani Mydaselpis ngurumani Neolaparopsis puncturatus Neolaparopsis puncturatus Nomoneura caffra Nomoneuroides natalensis Nomoneura paradoxa Nomoneuroides natalensis Notosyllegomydas brincki Notosyllegomydas brincki Syll gen nov sp nov Gobabeb Syllegomydas N arnoldi M Leptomydas notos Syllegomydas N astrictus Nemomydas pantherinus Syllegomydas N dispar Pseudonomoneura hirta Syllegomydas N heothinos Pseudonomoneura micheneri Syllegomydas S algiricus Nomoneura caffra Syllegomydas S elachys Nomoneura paradoxa Syllegomydas S lineatus Syllegomydas S palestinensis Arenomydas partitus Syllegomydas S proximus Arenomydas caerulescens Syllegomydas S vittatus Arenomydas callosus Syllegomydas sp nov Ruacana Mydaselpis sp nov Daan Halterorchis sp nov Cape Recife M Vespiodes leopolditertii Stenomydas gen nov sp nov Panne M Vespiodes phaios Syll gen nov sp nov Gobabeb Cephalocera sp Ratelfontein Eremomidas arabicus Cephalocera longirostris Eremomidas sp nov Tiega M Pseudonomoneura hirta Cephalocera sp Papendorp Pseudonomoneura micheneri Cephalocera sp Sors Sors Namadytes sp Gobabeb M Afroleptomydas A suffusipennis Namadytes vansoni Afroleptomydas A inhacae Hessemydas parkeri M Afroleptomydas C mauricei Hessemydas daugeroni Afroleptomydas C sp Stampriet Hessemydas seyrigi Afroleptomydas A paganus Haplomydas crassipes Afroleptomydas C sp Gobabeb Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticus Nemestrinus nigrovillosus Eremohaplomydas desertorum Bombylius major Dipalta serpentina Afroleptomydas A consanguinensis Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Lachnocorynus sp nov Kotwa M Phycus frommeri Diogmites grossus Pegesimallus laticornis Afroleptomydas A opacicinctus Dioctria hyalipennis Leptogaster cylindrica Arenomydas caerulescens Euscelidia pulchra Ospriocerus aeacus Gonioscelis ventralis Stichopogon punctum Afroleptomydas A sp Clanwilliam Lasiopogon cinctus Damalis monochaetes Arenomydas callosus Holcocephala calva Hoplistomerus nobilis Pilica formidolosa Laphria aktis Afroleptomydas A sp Obib Emphysomera pallidapex Arenomydas namaquensis M Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps Apiocera R alastor M Afroleptomydas C sp Koeroegab Apiocera R braunsi Arenomydas partitus Apiocera P bilineata Apiocera P mexicana M Apiocera P barri Apiocera P martinorum Apiocera P parahydra Mydaselpis sp nov Daan Afroleptomydas A omeri Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett M Mydaselpis sp nov Gamsberg M Apiocera Ap imminuta Afroleptomydas A rufithorax Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga Vespiodes leopolditertii Apiocera Ap asilica Apiocera Ap brevicornis Pseudonomoneura bajaensis Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Vespiodes phaios Apiocera Ap similis Apiocera Ap maritima Eremomidas arabicus Apiocera Ap minor Apiocera Ap newmani Rhaphiomidas hasbroucki Rhaphiomidas maehleri Nothomydas picketti Rhaphiomidas parkeri Tongamya miranda Halterorchis karooensis F Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Namibimydas psamminos Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Namadytes vansoni Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Namibimydas stuckenbergi Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Syllegomydas N astrictus Midacritus stuardoanus M Mitrodetus dimidiatus Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Apiophora quadricinctata Syllegomydas N dispar Ectyphus amboseli M Mitrodetus dentitarsis Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Miltinus viduatus Mitrodetus microglossa Syllegomydas N heothinos Miltinus maculipennis Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Stratiomydas lividus Mitrodetus nanoglossa M Syllegomydas S elachys Mydas clavatus Mydas tricinctus Mydas xanthopterus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Cephalocera imitata M Syllegomydas S lineatus Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Pseudorhopalia mirandai Cephalocera sp Ratelfontein Rhopalia annulata Rhopalia cincta Syllegomydas S palestinensis Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Cephalocera longirostris Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Syllegomydas S proximus Nemomydas melanopogon Leptomydas notos Nemomydas pantherinus Cephalocera sp Papendorp Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Syllegomydas S vittatus Eremomidas bek Leptomydas rapti M Cephalocera sp Sors Sors Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Syllegomydas sp nov Ruacana Nemomydas lamia Pseudonomoneura bajaensis Afroleptomydas A consanguinensis Pseudonomoneura hirta Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Nothomydas picketti Syll gen nov sp nov minute proboscis F Namibimydas psamminos Namibimydas stuckenbergi Afroleptomydas A inhacae Cephalocera imitata M Cephalocera sp Ratelfontein Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Afroleptomydas A opacicinctus Hispanomydas hispanicus Halterorchis sp nov Cape Recife M Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Afroleptomydas A paganus Syllegomydas S algiricus Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Afroleptomydas A sp Clanwilliam Diochlistinae Argentina F Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Afroleptomydas A sp Obib Arenomydas namaquensis M Mitrodetus microglossa Arenomydas partitus Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Eremomidas sp nov Tiega M Afroleptomydas C mauricei Eremomidas arabicus Halterorchis karooensis F Mitrodetus dentitarsis Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb monophyletic Afroleptomydas C sp Gobabeb Syllegomydas sp nov Ruacana Stenomydas gen nov sp nov Panne M Afroleptomydas A consanguinensis Mitrodetus dimidiatus Afroleptomydas A opacicinctus Afroleptomydas C mauricei Afroleptomydas C sp Koeroegab Afroleptomydas C sp Stampriet Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Namibimydas psamminos Afroleptomydas A omeri Afroleptomydas A rufithorax Afroleptomydas C sp Stampriet Syllegomydas S lineatus Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Namibimydas stuckenbergi Syllegomydas S palestinensis Afroleptomydas A omeri Syllegomydas S proximus Syllegomydas S vittatus Syllegomydas N astrictus Syllegomydas N heothinos Syll gen nov sp nov minute proboscis F Syllegomydas N arnoldi M Afroleptomydas A rufithorax Nothomydas picketti Syllegomydas N dispar Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus Syllegomydinae strict consensus of 2989 trees Afroleptomydas A suffusipennis strict consensus of 2226 trees Cleamydas gen nov sp nov Ugab F .... study of topology 16 Separate analyses – TreeMap3 Males Females ◊ fully resolved trees needed › 1 mp tree chosen ◊ same # of taxa needed › species linked within genus ◊ tanglegram not “untangled” .... study of topology 17

Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Excluding dimorphic characters Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Midacritus stuardoanus M Midacritus wagenknechti Pseudorhopalia mirandai ◊ Anomalomydas mackerrasi all 134 taxa Anomalomydas australicus F Apiophora paulseni Apiophora quadricinctata Heteromydas bicolor Opomydas townsendi Miltinus maculipennis Miltinus minutus Miltinus stenogaster Miltinus viduatus Parectyphus namibiensis Ectyphus armipes ◊ (- 36) Ectyphus amboseli M 175 characters Ectyphus pinguis Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Paramydas igniticornis Plyomydas peruviensis M › including 30 ? terminalia characters Plyomydas sp nov Tandil Dolichogaster brevicornis Messiasia californica M Messiasia painteri Mydas clavatus Mydas tricinctus Mydas xanthopterus Stratiomydas lividus / Cacatuopyga basifascia › including 19 terminalia characters Cacatuopyga ruficornis M Phyllomydas bruesii Phyllomydas parvulus Eremomidas bek Leptomydas corsicanus Leptomydas lusitanicus Leptomydas notos Leptomydas rapti M Leptomydas sardous Leptomydas turcicus Nemomydas brachyrhynchus ◊ Nemomydas gruenbergi M 3000 mp trees Nemomydas lamia Nemomydas melanopogon Nemomydas pantherinus Nemomydas venosus Afroleptomydas A inhacae Afroleptomydas A omeri Afroleptomydas A paganus Afroleptomydas A rufithorax Afroleptomydas A suffusipennis Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib ◊ 1367 steps Afroleptomydas C mauricei Afroleptomydas C sp Gobabeb Afroleptomydas C sp Koeroegab Afroleptomydas C sp Stampriet Mydaselpis ngurumani Halterorchis karooensis F Neolaparopsis puncturatus Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Notosyllegomydas brincki Syllegomydas N astrictus Syllegomydas N heothinos Syllegomydas S elachys Syllegomydas S lineatus Syllegomydas S palestinensis Syllegomydas S proximus Syllegomydas S vittatus Syllegomydas sp nov Ruacana Halterorchis sp nov Cape Recife M Stenomydas gen nov sp nov Panne M Syll gen nov sp nov Gobabeb Syll gen nov sp nov minute proboscis F Eremomidas arabicus Eremomidas sp nov Tiega M Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus

Limata tenuicornis Eulonchus sp Six Rivers Hispanomydas hispanicus Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Dipalta serpentina Toxophora maxima Syllegomydas S algiricus Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Phycus frommeri Diogmites grossus Pegesimallus laticornis Namadytes sp Gobabeb M Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Ospriocerus aeacus Gonioscelis ventralis Namadytes vansoni Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Holcocephala calva Hoplistomerus nobilis Syllegomydas N arnoldi M Pilica formidolosa Laphria aktis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Syllegomydas N dispar Machimus occidentalis Philonicus albiceps Apiocera R alastor M Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana M Hessemydas daugeroni Apiocera P barri Apiocera P martinorum Apiocera P parahydra Apiocera P haruspex Apiocera P aldrichi Hessemydas parkeri M Apiocera P painteri Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett M Apiocera Ap imminuta Hessemydas seyrigi Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga Apiocera Ap asilica Apiocera Ap brevicornis Pseudonomoneura bajaensis Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Pseudonomoneura hirta Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Rhaphiomidas hasbroucki Rhaphiomidas maehleri Pseudonomoneura micheneri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Haplomydas crassipes Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Eremohaplomydas desertorum Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Lachnocorynus sp nov Kotwa M Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Apiophora quadricinctata Arenomydas caerulescens Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Miltinus viduatus Arenomydas callosus Miltinus maculipennis Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Arenomydas namaquensis M Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Dolichogaster brevicornis Arenomydas partitus Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Mydaselpis sp nov Daan Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Mydaselpis sp nov Gamsberg M Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Nemomydas melanopogon Vespiodes leopolditertii Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Vespiodes phaios Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Diochlistinae Argentina F Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Mitrodetus nanoglossa M Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Cephalocera imitata M Cephalocera sp Ratelfontein Mitrodetus microglossa Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M Nomoneura caffra Nomoneura paradoxa Mitrodetus dentitarsis Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Neolaparopsis puncturatus Haplomydas crassipes Mitrodetus dimidiatus Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Cleamydas gen nov sp nov Ugab F Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Nothomydas picketti Arenomydas partitus Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F Namibimydas psamminos Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb monophyletic Syllegomydas sp nov Ruacana Namibimydas stuckenbergi Stenomydas gen nov sp nov Panne M Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Cephalocera imitata M Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Cephalocera sp Ratelfontein Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S palestinensis Syllegomydas S proximus Cephalocera longirostris Syllegomydas S vittatus Syllegomydas N astrictus Syllegomydas N heothinos Syll gen nov sp nov minute proboscis F Syllegomydas N arnoldi M Cephalocera sp Papendorp Syllegomydas N dispar Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Cephalocera sp Sors Sors Mitrodetus dimidiatus Syllegomydinae strict consensus of 3000 trees .... study of topology 18

Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus mel Megascelus nigricornis Neorhaphiomidas Min Neorhaphiomidas Yan Neorhaphiomidas hardyi Taxon pruning with IterPCR in TNT Neorhaphiomidas Man Cretomydas santanensis Diochlistus aur Diochlistus gracilis Diochlistus melleipennis Anomalomydas mac Anomalomydas aus Midacritus stu ◊ (- 2) Midacritus wag 132 taxa Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Apiophora qua Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi › Syllegomydas palestinensis Miltinus stenogaster Miltinus viduatus Miltinus mac Miltinus minutus Paramydas igniticornis Mydas clavatus Stratiomydas lividus Cacatuopyga basifascia › Arenomydas namaquensis (male only) Cacatuopyga ruf Mydas tricinctus Mydas xanthopterus Messiasia californica M Messiasia painteri Plyomydas peruviensis M Plyomydas sp nov Tandil Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia annulata ◊ all 211 characters Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Perissocerus tra Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Halterorchis Cape Recife Haplomydas crassipes Eremohaplomydas des Lachnocorynus Kotwa ◊ Nemomydas melanopogon 1431 mp trees Leptomydas notos Nemomydas pantherinus Nemomydas bra Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gru Leptomydas sardous ◊ 1819 steps Pseudonomoneura baj Pseudonomoneura hirta Pseudonomoneura mic Nemomydas lamia Arenomydas partitus Arenomydas cae Arenomydas callosus Notosyllegomydas brincki Syll gen nov sp nov Gobabeb Cephalocera lon Cephalocera sp Pap Cephalocera sp Sors Sors Cephalocera imitata M Cephalocera sp Rat Mydaselpis ngurumani Neolaparopsis puncturatus Mydaselpis sp nov Daan Mydaselpis Gamsberg Vespiodes leopolditertii Vespiodes phaios Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Afr A suffusipennis Afr A consanguinensis Afr A inhacae Afr A opacicinctus Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticus Nemestrinus nigrovillosus Afr C sp Gobabeb Bombylius major Dipalta serpentina Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Afr C mauricei Phycus frommeri Diogmites grossus Pegesimallus laticornis Dioctria hyalipennis Leptogaster cylindrica Afr C sp Stampriet Euscelidia pulchra Ospriocerus aeacus Gonioscelis ventralis Stichopogon punctum Lasiopogon cinctus Afr A sp Clanwilliam Damalis monochaetes Holcocephala calva Hoplistomerus nobilis Pilica formidolosa Laphria aktis Emphysomera pallidapex Afr A sp Obib Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps Apiocera R alastor M Afr C sp Koeroegab Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana M Apiocera P barri Apiocera P martinorum Afr A omeri Apiocera P parahydra Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Afr A rufithorax Apiocera P sp Colnett M Apiocera Ap imminuta Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Afr A paganus Apiocera Ap sp Wurarga Apiocera Ap asilica Apiocera Ap brevicornis Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Syllegomydas sp nov Ruacana Apiocera Ap sp Lloyd Apiocera Ap similis Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Namadytes sp Gobabeb M Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Namadytes vansoni Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Stenomydas Panne M Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Syllegomydas S lineatus Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Syllegomydas S vittatus Apiophora quadricinctata Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Hispanomydas hispanicus Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Paramydas igniticornis Syllegomydas S algiricus Cacatuopyga basifascia Cacatuopyga ruficornis M Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Syllegomydas S proximus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Plyomydas sp nov Tandil Syllegomydas N astrictus Messiasia californica M Messiasia painteri Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Syllegomydas N heothinos Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Syllegomydas S elachys Nemomydas melanopogon Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Nemomydas venosus Syll proboscis F Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Syllegomydas N arnoldi M Leptomydas sardous Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Syllegomydas N dispar Cleamydas gen nov sp nov Ugab F Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Cephalocera imitata M Halterorchis karooensis F Cephalocera sp Ratelfontein Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M Nomoneura caffra Eremomidas arabicus Nomoneura paradoxa Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Neolaparopsis puncturatus Eremomidas sp nov Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Nothomydas picketti Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Cleamydas Ugab F Arenomydas partitus Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Eremomidas sp nov Tiega M Eremomidas arabicus Namibimydas psamminos Halterorchis karooensis F Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb monophyletic Syllegomydas sp nov Ruacana Namibimydas stuckenbergi Stenomydas gen nov sp nov Panne M Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Diochlistinae Argentina F Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Mitrodetus microglossa Syllegomydas S lineatus Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S palestinensis Syllegomydas S proximus Mitrodetus nanoglossa M Syllegomydas S vittatus Syllegomydas N astrictus Syllegomydas N heothinos Syll gen nov sp nov minute proboscis F Syllegomydas N arnoldi M Mitrodetus dentitarsis Syllegomydas N dispar Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus Mitrodetus dimidiatus Syllegomydinae strict consensus of 1431 trees .... study of topology 19

Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Preferred hypothesis – all data and semaphoronts Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M ◊ Midacritus wagenknechti Mydidae is an old taxon Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Apiophora quadricinctata Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Miltinus viduatus Miltinus maculipennis ◊ more recently evolved taxa appear to have speciated with little Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Dolichogaster brevicornis morphological diversification Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus ◊ ? terminalia within speciose Syllegomydinae taxa are Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Nemomydas melanopogon Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus virtually identical Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F ◊ Steps for further enhancing topology and resolution Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Cephalocera imitata M Cephalocera sp Ratelfontein Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M › addition of internal head and thoracic characters Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M › addition of molecular data Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Arenomydas partitus

Limata tenuicornis Eulonchus sp Six Rivers Syll gen nov sp nov Gobabeb Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Dipalta serpentina Toxophora maxima Notosyllegomydas brincki Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Phycus frommeri Diogmites grossus Pegesimallus laticornis Eremomidas sp nov Tiega M Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Ospriocerus aeacus Gonioscelis ventralis Eremomidas arabicus Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Holcocephala calva Hoplistomerus nobilis Halterorchis karooensis F Pilica formidolosa Laphria aktis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Afroleptomydas A inhacae Philonicus albiceps Apiocera R alastor M Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana M Afroleptomydas A paganus Apiocera P barri Apiocera P martinorum Apiocera P parahydra Apiocera P haruspex Apiocera P aldrichi Afroleptomydas A suffusipennis Apiocera P painteri Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett M Apiocera Ap imminuta Afroleptomydas C sp Gobabeb Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga Apiocera Ap asilica Apiocera Ap brevicornis Syllegomydas sp nov Ruacana Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Stenomydas gen nov sp nov Panne M Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Rhaphiomidas hasbroucki Rhaphiomidas maehleri Afroleptomydas A consanguinensis Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Afroleptomydas A opacicinctus Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Afroleptomydas C mauricei Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Afroleptomydas C sp Stampriet Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Apiophora quadricinctata Afroleptomydas A sp Clanwilliam Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Miltinus viduatus Afroleptomydas A sp Obib Miltinus maculipennis Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Afroleptomydas C sp Koeroegab Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Dolichogaster brevicornis Afroleptomydas A omeri Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Afroleptomydas A rufithorax Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Syllegomydas S lineatus Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Nemomydas melanopogon Hispanomydas hispanicus Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Syllegomydas S algiricus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Syllegomydas S elachys Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Syllegomydas S palestinensis Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Cephalocera imitata M Cephalocera sp Ratelfontein Syllegomydas S proximus Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M Nomoneura caffra Nomoneura paradoxa Syllegomydas S vittatus Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Neolaparopsis puncturatus Haplomydas crassipes Syllegomydas N astrictus Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Syllegomydas N heothinos Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Arenomydas partitus Syll gen nov sp nov minute proboscis F Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F Syllegomydas N arnoldi M Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb monophyletic Syllegomydas sp nov Ruacana Syllegomydas N dispar Stenomydas gen nov sp nov Panne M Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Diochlistinae Argentina F Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Mitrodetus nanoglossa M Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S palestinensis Syllegomydas S proximus Mitrodetus microglossa Syllegomydas S vittatus Syllegomydas N astrictus Syllegomydas N heothinos Syll gen nov sp nov minute proboscis F Syllegomydas N arnoldi M Mitrodetus dentitarsis Syllegomydas N dispar Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus Syllegomydinae strict consensus of 1040 trees Mitrodetus dimidiatus ..... summary 20 Summary ◊ Apioceridae + Mydidae monophyletic ◊ (Apioceridae + Mydidae) + Asilidae ◊ earliest branchings within Mydidae well-supported ◊ speciose taxa Leptomydinae and Syllegomydinae not monophyletic ◊ subfamily taxa relationships still not resolved ◊ additional work needed and in particular molecular data

Mydidae: Phyllomydas parvulus © Lyn Atherton 21 Acknowledgements ◊ Museum curators and collectors who made specimens available ◊ Photographers ◊ U.S. National Science Foundation (REVSYS DEB-0919333)