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TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
TDikow Mydidae morphological phylogeny Hennig 2012
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TDikow Mydidae morphological phylogeny Hennig 2012

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Current (Jun 2012) phylogenetic hypothesis of relationships within Mydidae (Diptra: Asiloidea)

Current (Jun 2012) phylogenetic hypothesis of relationships within Mydidae (Diptra: Asiloidea)

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  • 1. 1Phylogeny of Apioceridae and Mydidae (Insecta: Diptera)based on morphological characters of adult fliesTorsten DikowBiodiversity Synthesis CenterField Museum of Natural History, Chicago, IL, USA
  • 2. . introduction 2Yeates & 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 3Yeates & 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 4Apioceridae – “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 5Mydidae – “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 6Taxon 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 7Morphological 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 8Phylogenetic 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 nigrovillosusResults 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 parkeriPlacement 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 FNon-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 RhaphiomidinaeMonophyly 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
  • 12. ... new phylogenetic hypothesis 12Earliest branchings within Mydidae Rhaphiomidinae Rhaphiomidas hasbroucki 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 remaining Mydidae Mydidae: Rhaphiomidas auratus © Greg Ballmer 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 Mydidae: Anomalomydas mackerrasi © Jean Hort Mydidae: Neorhaphiomidas sp. © Fred Hort 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
  • 13. .... study of topology 13What 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?
  • 14. .... study of topology 14Sexual 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
  • 15. .... study of topology 15 Rhaphiomidas hasbroucki Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas maehleri Rhaphiomidas parkeriSeparate analyses – results 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 Neorhaphiomidas hardyi Megascelus melanoproctus Neorhaphiomidas sp Mandurah Megascelus nigricornis Cretomydas santanensis Cretomydas santanensis Diochlistus auripennis M 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 Parectyphus namibiensis Ectyphus armipes 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 Paramydas igniticornis Cacatuopyga ruficornis M Ectyphus amboseli M Cacatuopyga basifascia Ectyphus pinguis Mydas clavatus Males Females 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 Dolichogaster brevicornis Perissocerus transcaspicus Rhopalia olivieri Phyllomydas bruesii 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 Perissocerus transcaspicus Eremomidas bek 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 lamia Nemomydas melanopogon 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 Notosyllegomydas brincki Nomoneuroides natalensis 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 Arenomydas partitus Syllegomydas S palestinensis 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 Cephalocera sp Papendorp Pseudonomoneura hirta Cephalocera sp Sors Sors Pseudonomoneura micheneri 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 Eremohaplomydas desertorum Afroleptomydas A consanguinensis Lachnocorynus sp nov Kotwa M Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Afroleptomydas A opacicinctus Dipalta serpentina Toxophora maxima Prorates sp Escalante Arenomydas caerulescens Apsilocephala longistyla Hemigephyra atra Phycus frommeri Afroleptomydas A sp Clanwilliam Diogmites grossus Pegesimallus laticornis Arenomydas callosus Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Ospriocerus aeacus Gonioscelis ventralis Afroleptomydas A sp Obib Stichopogon punctum Arenomydas namaquensis M Lasiopogon cinctus Damalis monochaetes Holcocephala calva Hoplistomerus nobilis Pilica formidolosa Arenomydas partitus Afroleptomydas C sp Koeroegab Laphria aktis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps Mydaselpis sp nov Daan Afroleptomydas A omeri Apiocera R alastor M Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana M Apiocera P barri Apiocera P martinorum Mydaselpis sp nov Gamsberg M Afroleptomydas A rufithorax Apiocera P parahydra Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Vespiodes leopolditertii Apiocera P parkeri Pseudonomoneura bajaensis Apiocera P sp Colnett M Apiocera Ap imminuta Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Vespiodes phaios Apiocera Ap sp Wurarga Apiocera Ap asilica Eremomidas arabicus Apiocera Ap brevicornis Apiocera Ap sp Stuart M Apiocera Ap hardyi Nothomydas picketti Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Halterorchis karooensis F Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Namibimydas psamminos Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Namadytes vansoni Tongamya miranda Megascelus albovillosus Namibimydas stuckenbergi Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Syllegomydas N astrictus Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Mitrodetus dimidiatus Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Syllegomydas N dispar Anomalomydas mackerrasi Mitrodetus dentitarsis Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Mitrodetus microglossa Syllegomydas N heothinos Apiophora quadricinctata Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Mitrodetus nanoglossa M Miltinus stenogaster Syllegomydas S elachys Miltinus viduatus Miltinus maculipennis Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia Cephalocera imitata M Cacatuopyga ruficornis M Syllegomydas S lineatus Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Cephalocera sp Ratelfontein Dolichogaster brevicornis Phyllomydas bruesii Syllegomydas S palestinensis Phyllomydas parvulus Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Cephalocera longirostris Messiasia painteri Pseudorhopalia mirandai Syllegomydas S proximus Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Cephalocera sp Papendorp Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Syllegomydas S vittatus Hessemydas seyrigi Nemomydas melanopogon Cephalocera sp Sors Sors Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Nemomydas venosus Syllegomydas sp nov Ruacana Leptomydas lusitanicus Eremomidas bek Afroleptomydas A consanguinensis Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Syll gen nov sp nov minute proboscis F Leptomydas sardous Afroleptomydas A inhacae Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Hispanomydas hispanicus Nothomydas picketti Afroleptomydas A opacicinctus Namibimydas psamminos Namibimydas stuckenbergi Cephalocera imitata M Cephalocera sp Ratelfontein Cephalocera longirostris Afroleptomydas A paganus Cephalocera sp Papendorp Syllegomydas S algiricus Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M monophyletic Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Afroleptomydas A sp Clanwilliam Namadytes sp Gobabeb M Diochlistinae Argentina F Namadytes vansoni Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Afroleptomydas A sp Obib Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mitrodetus microglossa Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Afroleptomydas C mauricei Arenomydas caerulescens Arenomydas callosus Mitrodetus dentitarsis Arenomydas namaquensis M Arenomydas partitus Syll gen nov sp nov Gobabeb Afroleptomydas C sp Gobabeb Notosyllegomydas brincki Eremomidas sp nov Tiega M Eremomidas arabicus Mitrodetus dimidiatus Halterorchis karooensis F Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas C sp Koeroegab Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb Syllegomydas sp nov Ruacana Stenomydas gen nov sp nov Panne M Namibimydas psamminos Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas C sp Stampriet Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Syllegomydinae strict consensus of 2989 trees strict consensus of 2226 trees Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Namibimydas stuckenbergi Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Hispanomydas hispanicus Syllegomydas S algiricus Nothomydas picketti Syllegomydas S elachys Afroleptomydas A rufithorax Syllegomydas S palestinensis Syllegomydas S proximus Syllegomydas S vittatus Syllegomydas N astrictus Syllegomydas N heothinos Afroleptomydas A suffusipennis Cleamydas gen nov sp nov Ugab F 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
  • 16. .... study of topology 16Separate analyses – TreeMap3 Males Females ◊ fully resolved trees needed › 1 mp tree chosen ◊ same # of taxa needed › species linked within genus ◊ tanglegram not “untangled”
  • 17. .... study of topology 17 Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeriExcluding dimorphic characters 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 ◊ all 134 taxa Diochlistus gracilis Diochlistus melleipennis Midacritus stuardoanus M Midacritus wagenknechti Pseudorhopalia mirandai Anomalomydas mackerrasi Anomalomydas australicus F Apiophora paulseni Apiophora quadricinctata Heteromydas bicolor Opomydas townsendi Miltinus maculipennis ◊ 175 characters (- 36) Miltinus minutus Miltinus stenogaster Miltinus viduatus Parectyphus namibiensis Ectyphus armipes Ectyphus amboseli M Ectyphus pinguis Rhopalia annulata Rhopalia cincta › including 30 ? terminalia characters Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Paramydas igniticornis Plyomydas peruviensis M Plyomydas sp nov Tandil Dolichogaster brevicornis Messiasia californica M › including 19 / terminalia characters Messiasia painteri Mydas clavatus Mydas tricinctus Mydas xanthopterus Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruficornis M Phyllomydas bruesii Phyllomydas parvulus Eremomidas bek Leptomydas corsicanus Leptomydas lusitanicus ◊ 3000 mp trees Leptomydas notos Leptomydas rapti M Leptomydas sardous Leptomydas turcicus Nemomydas brachyrhynchus Nemomydas gruenbergi M Nemomydas lamia Nemomydas melanopogon Nemomydas pantherinus Nemomydas venosus Afroleptomydas A inhacae Afroleptomydas A omeri ◊ 1367 steps Afroleptomydas A paganus Afroleptomydas A rufithorax Afroleptomydas A suffusipennis Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib 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 Hispanomydas hispanicus Syllegomydas S algiricus Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Namadytes sp Gobabeb M Dipalta serpentina Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Phycus frommeri Namadytes vansoni Diogmites grossus Pegesimallus laticornis Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Syllegomydas N arnoldi M Ospriocerus aeacus Gonioscelis ventralis Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Syllegomydas N dispar Holcocephala calva Hoplistomerus nobilis Pilica formidolosa Laphria aktis Emphysomera pallidapex Hessemydas daugeroni Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps Apiocera R alastor M Apiocera R braunsi Hessemydas parkeri M Apiocera P bilineata Apiocera P mexicana M Apiocera P barri Apiocera P martinorum Apiocera P parahydra Hessemydas seyrigi Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Pseudonomoneura bajaensis Apiocera P sp Colnett M Apiocera Ap imminuta Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga Pseudonomoneura hirta Apiocera Ap asilica Apiocera Ap brevicornis Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Pseudonomoneura micheneri Apiocera Ap sp Lloyd Apiocera Ap similis Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Haplomydas crassipes Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Eremohaplomydas desertorum Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Lachnocorynus sp nov Kotwa M Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Arenomydas caerulescens Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Ectyphus armipes Parectyphus namibiensis Arenomydas callosus Apiophora paulseni Apiophora quadricinctata Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Arenomydas namaquensis M Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Paramydas igniticornis Arenomydas partitus Cacatuopyga basifascia Cacatuopyga ruficornis M Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydaselpis sp nov Daan Mydas xanthopterus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Plyomydas peruviensis M Mydaselpis sp nov Gamsberg M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Vespiodes leopolditertii Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Vespiodes phaios Hessemydas seyrigi Nemomydas melanopogon Leptomydas notos Nemomydas pantherinus Nemomydas brachyrhynchus Diochlistinae Argentina F Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Mitrodetus nanoglossa M Nemomydas gruenbergi M Leptomydas sardous Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Mitrodetus microglossa Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Mitrodetus dentitarsis Cephalocera imitata M Cephalocera sp Ratelfontein Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M Mitrodetus dimidiatus monophyletic Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Namadytes sp Gobabeb M Namadytes vansoni Cleamydas gen nov sp nov Ugab F Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Nothomydas picketti Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Namibimydas psamminos Arenomydas callosus Arenomydas namaquensis M Arenomydas partitus Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Eremomidas sp nov Tiega M Namibimydas stuckenbergi Eremomidas arabicus Halterorchis karooensis F Afroleptomydas A inhacae Afroleptomydas A paganus Afroleptomydas A suffusipennis Cephalocera imitata M Afroleptomydas C sp Gobabeb Syllegomydas sp nov Ruacana Stenomydas gen nov sp nov Panne M Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Cephalocera sp Ratelfontein Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Syllegomydinae strict consensus of 3000 trees Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Cephalocera longirostris Afroleptomydas A rufithorax Syllegomydas S lineatus Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Cephalocera sp Papendorp Syllegomydas S palestinensis Syllegomydas S proximus Syllegomydas S vittatus Syllegomydas N astrictus Syllegomydas N heothinos Cephalocera sp Sors Sors 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
  • 18. .... study of topology 18 Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeriTaxon pruning with IterPCR in TNT Tongamya miranda Megascelus albovillosus Megascelus mel Megascelus nigricornis Neorhaphiomidas Min Neorhaphiomidas Yan Neorhaphiomidas hardyi Neorhaphiomidas Man Cretomydas santanensis Diochlistus aur ◊ 132 taxa (- 2) Diochlistus gracilis Diochlistus melleipennis Anomalomydas mac Anomalomydas aus Midacritus stu Midacritus wag Ectyphus armipes Parectyphus namibiensis Apiophora paulseni › Syllegomydas palestinensis Apiophora qua Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Miltinus viduatus Miltinus mac › Arenomydas namaquensis (male only) Miltinus minutus Paramydas igniticornis Mydas clavatus Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruf Mydas tricinctus Mydas xanthopterus Messiasia californica M Messiasia painteri Plyomydas peruviensis M Plyomydas sp nov Tandil ◊ all 211 characters Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Perissocerus tra Hessemydas parkeri M Hessemydas daugeroni ◊ 1431 mp trees Hessemydas seyrigi Halterorchis Cape Recife Haplomydas crassipes Eremohaplomydas des Lachnocorynus Kotwa Nemomydas melanopogon Leptomydas notos Nemomydas pantherinus Nemomydas bra Nemomydas venosus Leptomydas lusitanicus Eremomidas bek ◊ 1819 steps Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gru Leptomydas sardous 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 Afr C sp Gobabeb Afr C mauricei Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Dipalta serpentina Toxophora maxima Afr C sp Stampriet Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Phycus frommeri Diogmites grossus Pegesimallus laticornis Afr A sp Clanwilliam Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Ospriocerus aeacus Gonioscelis ventralis Afr A sp Obib Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Holcocephala calva Hoplistomerus nobilis Pilica formidolosa Afr C sp Koeroegab Laphria aktis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Afr A omeri Philonicus albiceps Apiocera R alastor M Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana M Afr A rufithorax Apiocera P barri Apiocera P martinorum Apiocera P parahydra Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Afr A paganus Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett M Apiocera Ap imminuta Apiocera Ap tonnoiri Syllegomydas sp nov Ruacana Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga Apiocera Ap asilica Apiocera Ap brevicornis Namadytes sp Gobabeb M Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Apiocera Ap maritima Namadytes vansoni Apiocera Ap minor Apiocera Ap newmani Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Stenomydas Panne M Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Syllegomydas S lineatus Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Syllegomydas S vittatus Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Hispanomydas hispanicus Ectyphus armipes Parectyphus namibiensis Apiophora paulseni Apiophora quadricinctata Ectyphus amboseli M Ectyphus pinguis Syllegomydas S algiricus Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Syllegomydas S proximus Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Stratiomydas lividus Syllegomydas N astrictus Mydas clavatus Mydas tricinctus Mydas xanthopterus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Syllegomydas N heothinos Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Pseudorhopalia mirandai Syllegomydas S elachys Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Syll proboscis F Hessemydas daugeroni Hessemydas seyrigi Nemomydas melanopogon Leptomydas notos Nemomydas pantherinus Syllegomydas N arnoldi M Nemomydas brachyrhynchus Nemomydas venosus Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Syllegomydas N dispar Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Nemomydas lamia Pseudonomoneura bajaensis Halterorchis karooensis F Pseudonomoneura hirta Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Nothomydas picketti Namibimydas psamminos Eremomidas arabicus Namibimydas stuckenbergi Cephalocera imitata M Cephalocera sp Ratelfontein Cephalocera longirostris Cephalocera sp Papendorp Eremomidas sp nov Cephalocera sp Sors Sors Halterorchis sp nov Cape Recife M monophyletic Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Namadytes sp Gobabeb M Nothomydas picketti Namadytes vansoni Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Cleamydas Ugab F Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Namibimydas psamminos Arenomydas callosus Arenomydas namaquensis M Arenomydas partitus Syll gen nov sp nov Gobabeb Notosyllegomydas brincki Namibimydas stuckenbergi Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F Afroleptomydas A inhacae Afroleptomydas A paganus Diochlistinae Argentina F Afroleptomydas A suffusipennis Afroleptomydas C sp Gobabeb Syllegomydas sp nov Ruacana Stenomydas gen nov sp nov Panne M Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Mitrodetus microglossa Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Syllegomydinae strict consensus of 1431 trees Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Mitrodetus nanoglossa M Afroleptomydas A omeri 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
  • 19. .... study of topology 19 Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeriPreferred hypothesis – all data and semaphoronts 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 ◊ Mydidae is an old taxon 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 ◊ more recently evolved taxa appear to have speciated with little Heteromydas bicolor Opomydas townsendi Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Paramydas igniticornis Cacatuopyga basifascia morphological diversification 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 ◊ ? terminalia within speciose Syllegomydinae taxa are Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi virtually identical 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 ◊ Steps for further enhancing topology and resolution Nemomydas lamia Pseudonomoneura bajaensis Pseudonomoneura hirta Pseudonomoneura micheneri Cleamydas gen nov sp nov Ugab F Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi › addition of internal head and thoracic characters 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 › addition of molecular data 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 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
  • 20. ..... summary 20Summary ◊ 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. 21Acknowledgements ◊ Museum curators and collectors who made specimens available ◊ Photographers ◊ U.S. National Science Foundation (REVSYS DEB-0919333)

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