TDikow Mydidae morphological phylogeny Hennig 2012
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Current (Jun 2012) phylogenetic hypothesis of relationships within Mydidae (Diptra: Asiloidea)
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TDikow Mydidae morphological phylogeny Hennig 2012
- 1. 1 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
- 2. . 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 Leptomydinae Mydidae Cacatuopyginae Leptomydinae Syllegomydinae Syllegomydinae Anomalomydinae Anomalomydinae Diochlistinae Diochlistinae Ectyphinae Ectyphinae Apiophorinae Apiophorinae Mydinae Mydinae
- 3. . introduction 3 Yeates & Irwin 1996 – Exclusively southern connection taxa = Gondwanan Apiocera (Pyrocera) western North America Apiocera (Ripidosyrma) South Africa Tongamya Apioceridae Mydidae: Megascelinae Apiocera (Anypenus) Megascelus Argentina + Chile Mitrodetus Apiophora, Midacritus Mydidae: Diochlistinae Mydidae: Apiophorinae Apiocera (Apiocera) Neorhaphiomidas Diochlistus Australia Miltinus
- 4. . 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 Apiocera (Apiocera) moerens © Peter Chew › Argentina and Chile – 4 species › Australia – 67 species › South Africa – 3 species › western North America (Canada, Mexico, USA) – 64 species Apiocera bilineata © Greg Ballmer
- 5. . 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
- 6. .. 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
- 7. .. 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
- 8. .. 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) Mydidae: Eremomidas arabicus © Drew Gardner › Ratchet, TreeDrifting › all multi-state characters non-additive › all characters equally weighted ◊ Bremer Support › TNT (version 1.1 June 2012) Mydidae: Heteromydas bicolor © Greg Ballmer
- 9. ... new phylogenetic hypothesis 9 Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticus Nemestrinus nigrovillosus Results 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 ◊ 1040 most parsimonious trees 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 ◊ 1845 steps Apiocera P parahydra Apiocera P haruspex • Apioceridae 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 ◊ strict consensus 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 › several polytomies 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 › not well-resolved in derived ingroup • 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 • Mydidae 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 •• Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticus Namadytes vansoni Neolaparopsis puncturatus Nemestrinus nigrovillosus Bombylius major Dipalta serpentina •• Haplomydas crassipes Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Eremohaplomydas desertorum Phycus frommeri Diogmites grossus Pegesimallus laticornis • Dioctria hyalipennis Lachnocorynus sp nov Kotwa M Leptogaster cylindrica Euscelidia pulchra Ospriocerus aeacus Gonioscelis ventralis Mydaselpis sp nov Daan Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Mydaselpis sp nov Gamsberg M Holcocephala calva Hoplistomerus nobilis Pilica formidolosa Laphria aktis Vespiodes leopolditertii Emphysomera pallidapex Ommatius tibialis • Neolophonotus bimaculatus Machimus occidentalis Vespiodes phaios Philonicus albiceps Apiocera R alastor M Apiocera R braunsi Mydaselpis ngurumani Apiocera P bilineata Apiocera P mexicana M Apiocera P barri Apiocera P martinorum Arenomydas caerulescens Apiocera P parahydra Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Arenomydas callosus Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett M Arenomydas namaquensis M Apiocera Ap imminuta Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii • Arenomydas partitus Apiocera Ap sp Wurarga Apiocera Ap asilica Apiocera Ap brevicornis Apiocera Ap sp Stuart M Syll gen nov sp nov Gobabeb Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Notosyllegomydas brincki Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Eremomidas sp nov Tiega M Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Eremomidas arabicus Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Halterorchis karooensis F Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Afroleptomydas A inhacae Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Afroleptomydas A paganus Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Afroleptomydas A suffusipennis Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Apiophora quadricinctata Afroleptomydas C sp Gobabeb Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Syllegomydas sp nov Ruacana Opomydas townsendi Miltinus stenogaster Miltinus viduatus • Miltinus maculipennis Stenomydas gen nov sp nov Panne M Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Afroleptomydas A consanguinensis Stratiomydas lividus Mydas clavatus Mydas tricinctus Afroleptomydas A opacicinctus Mydas xanthopterus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Afroleptomydas C mauricei Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Afroleptomydas C sp Stampriet Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Afroleptomydas A sp Clanwilliam Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Afroleptomydas A sp Obib Hessemydas daugeroni Hessemydas seyrigi Nemomydas melanopogon Leptomydas notos Afroleptomydas C sp Koeroegab Nemomydas pantherinus Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Afroleptomydas A omeri Eremomidas bek Leptomydas rapti M Leptomydas turcicus Afroleptomydas A rufithorax Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Nemomydas lamia Syllegomydas S lineatus Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Hispanomydas hispanicus Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Syllegomydas S algiricus Cephalocera imitata M Cephalocera sp Ratelfontein Cephalocera longirostris Cephalocera sp Papendorp Syllegomydas S elachys Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M Nomoneura caffra Nomoneura paradoxa Syllegomydas S palestinensis Nomoneuroides natalensis Bremer 3 – 9 Namadytes sp Gobabeb M Namadytes vansoni Syllegomydas S proximus Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Syllegomydas S vittatus Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii • Vespiodes phaios Syllegomydas N astrictus Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Syllegomydas N heothinos Arenomydas partitus Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Syll gen nov sp nov minute proboscis F Eremomidas sp nov Tiega M •• Eremomidas arabicus Halterorchis karooensis F Afroleptomydas A inhacae Syllegomydas N arnoldi M 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 Diochlistinae Argentina F Afroleptomydas C mauricei • Afroleptomydas C sp Stampriet Bremer ≥ 10 strict consensus of 1040 trees Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Mitrodetus nanoglossa M Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Mitrodetus microglossa Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Mitrodetus dentitarsis Syllegomydas S palestinensis Syllegomydas S proximus Syllegomydas S vittatus Syllegomydas N astrictus Mitrodetus dimidiatus 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
- 10. ... new phylogenetic hypothesis 10 Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Placement of † Cretomydas santanensis Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M ≈ 112 myo Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah 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 † Cretomydas santanensis, Brazil Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Non-monophyly of Diochlistinae 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 Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Eremomidas sp nov Tiega M Dipalta serpentina Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Phycus frommeri Eremomidas arabicus Diogmites grossus Pegesimallus laticornis Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Halterorchis karooensis F Ospriocerus aeacus Gonioscelis ventralis Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Afroleptomydas A inhacae Holcocephala calva Hoplistomerus nobilis Pilica formidolosa Laphria aktis Emphysomera pallidapex Ommatius tibialis Afroleptomydas A paganus Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps Apiocera R alastor M Apiocera R braunsi Afroleptomydas A suffusipennis Apiocera P bilineata Apiocera P mexicana M Apiocera P barri Apiocera P martinorum Apiocera P parahydra Afroleptomydas C sp Gobabeb Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Syllegomydas sp nov Ruacana Apiocera P sp Colnett M Apiocera Ap imminuta Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga Stenomydas gen nov sp nov Panne M Apiocera Ap asilica Apiocera Ap brevicornis Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Afroleptomydas A consanguinensis Apiocera Ap sp Lloyd Apiocera Ap similis Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Afroleptomydas A opacicinctus Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Afroleptomydas C mauricei Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Afroleptomydas C sp Stampriet Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Afroleptomydas A sp Clanwilliam Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Afroleptomydas A sp Obib Apiophora paulseni Apiophora quadricinctata Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Afroleptomydas C sp Koeroegab Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Paramydas igniticornis Afroleptomydas A omeri Cacatuopyga basifascia Cacatuopyga ruficornis M Stratiomydas lividus Mydas clavatus Mydas tricinctus Afroleptomydas A rufithorax Mydas xanthopterus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Plyomydas sp nov Tandil Syllegomydas S lineatus Messiasia californica M Messiasia painteri Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Hispanomydas hispanicus Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Syllegomydas S algiricus Hessemydas seyrigi Nemomydas melanopogon Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Syllegomydas S elachys Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Syllegomydas S palestinensis Nemomydas gruenbergi M Leptomydas sardous Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Syllegomydas S proximus Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Syllegomydas S vittatus Cephalocera imitata M Cephalocera sp Ratelfontein Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M Syllegomydas N astrictus Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Syllegomydas N heothinos Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Syll gen nov sp nov minute proboscis F Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Syllegomydas N arnoldi M Arenomydas callosus Arenomydas namaquensis M Arenomydas partitus Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Eremomidas sp nov Tiega M Syllegomydas N dispar Eremomidas arabicus Halterorchis karooensis F Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Diochlistinae Argentina F Afroleptomydas C sp Gobabeb Syllegomydas sp nov Ruacana Stenomydas gen nov sp nov Panne M Afroleptomydas A consanguinensis Argentina + Chile Afroleptomydas A opacicinctus Mitrodetus nanoglossa M Afroleptomydas C mauricei Afroleptomydas C sp Stampriet strict consensus of 1040 trees Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Mitrodetus microglossa Afroleptomydas A rufithorax Syllegomydas S lineatus Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Mitrodetus dentitarsis Syllegomydas S palestinensis Syllegomydas S proximus Syllegomydas S vittatus Syllegomydas N astrictus Syllegomydas N heothinos Diochlistus auripennis, Australia © Peter Chew Mitrodetus dimidiatus 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
- 11. ... new phylogenetic hypothesis 11 Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Rhaphiomidinae Monophyly of subfamily taxa Tongamya miranda Megascelus albovillosus Megascelinae Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Anomalomydinae 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 Cacatuopyginae Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M 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 Syllegomydinae: Cephalocera imitata 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 Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Eremomidas sp nov Tiega M Dipalta serpentina Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Phycus frommeri Eremomidas arabicus Diogmites grossus Pegesimallus laticornis Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Halterorchis karooensis F Ospriocerus aeacus Gonioscelis ventralis Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Afroleptomydas A inhacae Holcocephala calva Hoplistomerus nobilis Pilica formidolosa Laphria aktis Emphysomera pallidapex Ommatius tibialis Afroleptomydas A paganus Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps Apiocera R alastor M Apiocera R braunsi Afroleptomydas A suffusipennis Apiocera P bilineata Apiocera P mexicana M Apiocera P barri Apiocera P martinorum Apiocera P parahydra Afroleptomydas C sp Gobabeb Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Syllegomydas sp nov Ruacana Apiocera P sp Colnett M Apiocera Ap imminuta Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga Stenomydas gen nov sp nov Panne M Apiocera Ap asilica Apiocera Ap brevicornis Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Afroleptomydas A consanguinensis Apiocera Ap sp Lloyd Apiocera Ap similis Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Afroleptomydas A opacicinctus Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Afroleptomydas C mauricei Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Afroleptomydas C sp Stampriet Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Afroleptomydas A sp Clanwilliam Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Afroleptomydas A sp Obib Apiophora paulseni Apiophora quadricinctata Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Afroleptomydas C sp Koeroegab Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Paramydas igniticornis Afroleptomydas A omeri Cacatuopyga basifascia Cacatuopyga ruficornis M Stratiomydas lividus Mydas clavatus Mydas tricinctus Afroleptomydas A rufithorax Mydas xanthopterus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Plyomydas sp nov Tandil Syllegomydas S lineatus Messiasia californica M Messiasia painteri Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Hispanomydas hispanicus Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Syllegomydas S algiricus Hessemydas seyrigi Nemomydas melanopogon Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Syllegomydas S elachys Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Syllegomydas S palestinensis Nemomydas gruenbergi M Leptomydas sardous Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Syllegomydas S proximus Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Syllegomydas S vittatus Cephalocera imitata M Cephalocera sp Ratelfontein Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M Syllegomydas N astrictus monophyletic Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Syllegomydas N heothinos Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Syll gen nov sp nov minute proboscis F Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Syllegomydas N arnoldi M Arenomydas callosus Arenomydas namaquensis M Arenomydas partitus Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Eremomidas sp nov Tiega M Syllegomydas N dispar Eremomidas arabicus Halterorchis karooensis F Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Diochlistinae Argentina F Afroleptomydas C sp Gobabeb Syllegomydas sp nov Ruacana Stenomydas gen nov sp nov Panne M Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Mitrodetus nanoglossa M Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Syllegomydinae (43 % species diversity) strict consensus of 1040 trees Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Mitrodetus microglossa Afroleptomydas A rufithorax Syllegomydas S lineatus Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Mitrodetus dentitarsis Syllegomydas S palestinensis Syllegomydas S proximus Syllegomydas S vittatus Syllegomydas N astrictus Syllegomydas N heothinos Mitrodetus dimidiatus 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