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TDikow Apioceridae + Mydidae morphological phylogeny ESA 2011
 

TDikow Apioceridae + Mydidae morphological phylogeny ESA 2011

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Current (Nov 2011) phylogenetic hypothesis of relationships within Apioceridae and Mydidae (Diptra: Asiloidea)

Current (Nov 2011) phylogenetic hypothesis of relationships within Apioceridae and Mydidae (Diptra: Asiloidea)

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    TDikow Apioceridae + Mydidae morphological phylogeny ESA 2011 TDikow Apioceridae + Mydidae morphological phylogeny ESA 2011 Presentation Transcript

    • 1Phylogeny and historical biogeography of Apioceridaeand Mydidae inferred from morphological characters ofimagines (Diptera: Asiloidea)Torsten DikowBiodiversity Synthesis CenterField Museum of Natural History, Chicago, IL
    • . 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 monophyletic Apioceridae Asilidae Asilidae Apiocera (Pyrocera) Apiocera (Pyrocera) Apiocera (Ripidosyrma) Apiocera (Ripidosyrma) Apiocera (Anypenus) Apiocera (Anypenus) Apiocera (Apiocera) Apiocera (Apiocera) Rhaphiomidinae Rhaphiomidinae Megascelinae Megascelinae Rhopaliinae Rhopaliinae Cacatuopyginae Cacatuopyginae Leptomydinae Leptomydinae Syllegomydinae Syllegomydinae Anomalomydinae Anomalomydinae Diochlistinae Diochlistinae Ectyphinae Ectyphinae Apiophorinae Apiophorinae Mydinae Mydinae
    • . 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
    • . introduction 4Apioceridae – “flower-loving flies” ◊ Imagines flower-feeders & larvae possibly predatory ◊ 138 species and 1 genus (4 subgenera) ◊ Imagines 15 – 30 mm ◊ occur only in › Argentina and Chile – 4 species › Australia – 67 species Apiocera (Apiocera) moerens © Peter Chew › South Africa – 3 species › western North America (Canada, Mexico, USA) – 64 species ◊ Fossil † Protapiocera 3 species › † Protapioceridae – sister-group of Apioceridae? › Upper Yixian Formation, north-eastern China › Jurassic: Tithonian › ≈ 145 – 150 myo † Protapiocera convergens Zhang et al. 2004
    • . introduction 5Mydidae – “mydas flies” ◊ Imagines flower-feeders, some do not feed at all ◊ Larvae possibly predatory ◊ 469 species and 66 genera world-wide ◊ Imagines 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: † Cretomydas santanensis
    • 6Overview of presentation ◊ Set out to postulate cladistic relationships within Mydidae ◊ Methods ◊ New Phylogenetic Hypothesis Apiocera (Pyrocera) bilineata © Greg Ballmer ◊ Biogeographically interesting taxa ◊ Summary + Future Miltinus stenogaster © Shaun Winterton
    • .. methods 7Taxon Sampling ◊ 27 Apiocera species (= 19.6 %) › 2 Apiocera (Anypenus) (= 50.0 %) › 11 Apiocera (Apiocera) (= 16.4 %) › 11 Apiocera (Pyrocera) (= 17.2 %) › 2 Apiocera (Ripidosyrma) (= 66.7 %) ◊ 120 Mydidae species (= 25.6 %) Apioceridae › 46 genera (= 69.7 %) › 7 tribal taxa (= 87.5 %) › 11 subfamily taxa (= 100.0 %) ◊ Outgroup taxa › 37 Asiloidea species › 3 Nemestrinoidea Mydidae › 1 Tabanomorpha collecting localities of specimens included
    • .. methods 8Morphological Characters ◊ 206 discrete, parsimony informative characters tagma # characters remarks Head 42 Thorax 101 45 chaetotaxy, sclerites, etc. 23 wing 33 leg Abdomen 62 11 sclerite shapes, chaetotaxy, etc. 32 ? terminalia 20 / genitalia
    • .. methods 9Phylogenetic Analysis ◊ Morphological data › Mesquite (version 2.75) › MacClade (version 4.08) › WinClada / Asado (version 1.89) ◊ Tree Search › TNT (version 1.1 August 2011) Mydidae: Eremomidas arabicus © Drew Gardner › Ratchet, TreeDrifting (Nixon 1999, Goloboff 1999) › all multi-state characters non-additive › all characters equally weighted ◊ Bremer Support › TNT (version 1.1 August 2011) Mydidae: Heteromydas bicolor © Greg Ballmer
    • ... new phylogenetic hypothesis 10 Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticusResults Nemestrinus nigrovillosus Bombylius major Dipalta serpentina Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Phycus frommeri Diogmites grossus Ospriocerus aeacus Gonioscelis ventralis Hoplistomerus nobilis Pilica formidolosa Laphria aktis ◊ 428 most parsimonious trees Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps Pegesimallus laticornis Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Stichopogon punctum Lasiopogon cinctus • Damalis monochaetes Holcocephala calva Apiocera R alastor male • Apiocera R braunsi ◊ 1714 steps Apiocera P bilineata Apiocera P mexicana male Apiocera P parahydra • Apiocera P barri Apiocera P martinorum Apiocera P haruspex •• Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett male Apioceridae • Apiocera Ap imminuta Apiocera Ap tonnoiri Apiocera An obscura • Apiocera An philippii •• Apiocera Ap sp Wurarga ◊ Apioceridae, Rhaphiomidinae, and Apiocera Ap asilica Apiocera Ap brevicornis Apiocera Ap sp Stuart male •• Apiocera Ap hardyi Apiocera Ap newmani Apiocera Ap sp Lloyd • Apiocera Ap maritima Apiocera Ap similis Rhaphiomidas hasbroucki Rhaphiomidas maehleri • • Rhaphiomidas parkeri Megascelinae as in Yeates & Irwin 1996 Tongamya miranda • Megascelus albovillosus Megascelus nigricornis Neorhaphiomidas setosa male • •• Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis male • Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi • Anomalomydas australicus female Paramydas igniticornis Plyomydas peruviensis male Plyomydas sp nov Tandil Messiasia californica male Messiasia painteri ◊ several polytomies • Midacritus stuardoanus male • Midacritus wagenknechti • Cacatuopyga basifascia Cacatuopyga ruficornis male Heteromydas bicolor Opomydas townsendi • Parectyphus namibiensis Ectyphus armipes • Ectyphus amboseli male Ectyphus pinguis Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus maculipennis Miltinus viduatus Mydas clavatus ◊ strict consensus not well-resolved in Mydas xanthopterus • Stratiomydas lividus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus • Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti female Perissocerus transcaspicus derived ingroup Hessemydas parkeri male Hessemydas seyrigi Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Mydaselpis ngurumani Arenomydas partitus Mydidae Arenomydas caerulescens Arenomydas callosus •• Arenomydas namaquensis male Neolaparopsis puncturatus Haplomydas crassipes •• Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa male • Mydaselpis sp nov Daan • Mydaselpis sp nov Gamsberg male Vespiodes leopolditertii Vespiodes phaios Nemomydas brachyrhynchus Nemomydas venosus Leptomydas corsicanus Nemomydas gruenbergi male Leptomydas lusitanicus Eremomidas bek Leptomydas rapti male • Leptomydas turcicus •• Leptomydas sardous Leptomydas notos Nemomydas pantherinus Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticus Pseudonomoneura hirta Pseudonomoneura micheneri Nemestrinus nigrovillosus Bombylius major Dipalta serpentina • Toxophora maxima Nemomydas lamia Prorates sp Escalante Apsilocephala longistyla Hemigephyra atra Notosyllegomydas brincki Phycus frommeri Diogmites grossus Ospriocerus aeacus Gonioscelis ventralis Namibimydas stuckenbergi sp nov Hoplistomerus nobilis Pilica formidolosa Laphria aktis •• Emphysomera pallidapex Nothomydas picketti sp nov Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps Cephalocera longirostris Pegesimallus laticornis Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Cephalocera sp Papendorp Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Cephalocera sp Sors Sors Holcocephala calva Apiocera R alastor male Apiocera R braunsi Apiocera P bilineata Cephalocera imitata male Apiocera P mexicana male Apiocera P parahydra • Apiocera P barri Apiocera P martinorum Cephalocera sp Ratelfontein Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Nemomydas melanopogon Apiocera P parkeri Apiocera P sp Colnett male Apiocera Ap imminuta Apiocera Ap tonnoiri Namadytes sp Gobabeb male Apiocera An obscura Apiocera An philippii Apiocera Ap sp Wurarga Namadytes vansoni Apiocera Ap asilica Apiocera Ap brevicornis Apiocera Ap sp Stuart male Apiocera Ap hardyi Afroleptomydas C mauricei Apiocera Ap newmani Apiocera Ap sp Lloyd Apiocera Ap maritima Apiocera Ap similis Afroleptomydas C sp Gobabeb Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Afroleptomydas C sp Stampriet Megascelus albovillosus Megascelus nigricornis Neorhaphiomidas setosa male Neorhaphiomidas sp Mandurah Afroleptomydas A inhacae Cretomydas santanensis Diochlistus auripennis male Diochlistus gracilis Afroleptomydas A consanguinensis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus female Paramydas igniticornis Afroleptomydas A opacicinctus Plyomydas peruviensis male Plyomydas sp nov Tandil Messiasia californica male Messiasia painteri Afroleptomydas A paganus Midacritus stuardoanus male Midacritus wagenknechti Cacatuopyga basifascia Cacatuopyga ruficornis male Afroleptomydas A suffusipennis Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Ectyphus armipes Afroleptomydas A sp Clanwilliam Ectyphus amboseli male Ectyphus pinguis Apiophora paulseni Afroleptomydas A sp Obib Bremer 3 – 8 Apiophora quadricinctata Miltinus stenogaster Miltinus maculipennis Miltinus viduatus Afroleptomydas C sp Koeroegab Mydas clavatus Mydas xanthopterus Stratiomydas lividus Dolichogaster brevicornis Afroleptomydas A omeri Phyllomydas bruesii Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia annulata Afroleptomydas A rufithorax Rhopalia cincta Rhopalia olivieri Perissocerus dumonti female Perissocerus transcaspicus Syllegomydas S lineatus Hessemydas parkeri male Hessemydas seyrigi Nomoneura caffra Syllegomydas S vittatus Nomoneura paradoxa Nomoneuroides natalensis Mydaselpis ngurumani Arenomydas partitus Syllegomydas S palestinensis Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis male Neolaparopsis puncturatus Syllegomydas N astrictus Haplomydas crassipes Eremohaplomydas desertorum Bremer ≥ 10 Lachnocorynus sp nov Kotwa male Mydaselpis sp nov Daan Syllegomydas N heothinos Mydaselpis sp nov Gamsberg male Vespiodes leopolditertii Vespiodes phaios Nemomydas brachyrhynchus Syllegomydas sp nov Ruacana Nemomydas venosus Leptomydas corsicanus Nemomydas gruenbergi male Syllegomydas N arnoldi male Leptomydas lusitanicus Eremomidas bek Leptomydas rapti male Leptomydas turcicus Syllegomydas N dispar Leptomydas sardous Leptomydas notos Nemomydas pantherinus Pseudonomoneura hirta Hispanomydas hispanicus Pseudonomoneura micheneri Nemomydas lamia Notosyllegomydas brincki Namibimydas stuckenbergi sp nov Syllegomydas S algiricus Nothomydas picketti sp nov Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors ••• • Syllegomydas S elachys Cephalocera imitata male Cephalocera sp Ratelfontein Nemomydas melanopogon Syllegomydas S proximus Namadytes sp Gobabeb male Namadytes vansoni Afroleptomydas C mauricei Afroleptomydas C sp Gobabeb Halterorchis karooensis female Afroleptomydas C sp Stampriet Afroleptomydas A inhacae Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Eremomidas arabicus Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Eremomidas sp nov Tiega male •• Afroleptomydas C sp Koeroegab strict consensus of 428 trees Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Mitrodetus sp nov Argentina female Syllegomydas S vittatus Syllegomydas S palestinensis Syllegomydas N astrictus Mitrodetus nanoglossa male Syllegomydas N heothinos Syllegomydas sp nov Ruacana Syllegomydas N arnoldi male Syllegomydas N dispar Mitrodetus dentitarsis Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S proximus Mitrodetus dimidiatus male Halterorchis karooensis female Eremomidas arabicus Eremomidas sp nov Tiega male Mitrodetus sp nov Argentina female Mitrodetus nanoglossa male Mitrodetus dentitarsis Mitrodetus dimidiatus male
    • ... new phylogenetic hypothesis 11 Apiocera subgenera Apiocera R alastor male Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana male Apiocera P parahydra Apiocera P barri Apiocera P martinorum Apiocera P haruspex Apiocera P aldrichi South Africa Apiocera P painteri Apiocera P arena Apiocera P parkeriwestern North America Apiocera P sp Colnett male Apiocera Ap imminuta Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Apiocera (Pyrocera) alleni © Greg Ballmer Apiocera Ap sp Wurarga Apiocera Ap asilica Apiocera Ap brevicornis Apiocera Ap sp Stuart male Apiocera Ap hardyi Apiocera Ap newmani Apiocera Ap sp Lloyd Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticus Nemestrinus nigrovillosus Bombylius major Dipalta serpentina Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Chile Apiocera Ap maritima Apiocera Ap similis Hemigephyra atra Phycus frommeri Diogmites grossus Ospriocerus aeacus Gonioscelis ventralis Hoplistomerus nobilis Pilica formidolosa Laphria aktis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps Pegesimallus laticornis Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Holcocephala calva Apiocera R alastor male Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana male Apiocera P parahydra Apiocera P barri Apiocera P martinorum Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett male 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 male Apiocera Ap hardyi Apiocera Ap newmani Apiocera Ap sp Lloyd Apiocera Ap maritima Apiocera Ap similis Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus nigricornis Neorhaphiomidas setosa male Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis male Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus female Paramydas igniticornis Plyomydas peruviensis male Plyomydas sp nov Tandil Messiasia californica male Messiasia painteri Midacritus stuardoanus male Midacritus wagenknechti Cacatuopyga basifascia Cacatuopyga ruficornis male Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Ectyphus armipes Ectyphus amboseli male Ectyphus pinguis Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus maculipennis Miltinus viduatus Mydas clavatus Mydas xanthopterus Stratiomydas lividus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti female Perissocerus transcaspicus Hessemydas parkeri male Australian Apiocera (Apiocera) Hessemydas seyrigi Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Mydaselpis ngurumani Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis male Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa male Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg male Vespiodes leopolditertii Vespiodes phaios Nemomydas brachyrhynchus Nemomydas venosus Leptomydas corsicanus Nemomydas gruenbergi male Leptomydas lusitanicus Eremomidas bek Leptomydas rapti male Leptomydas turcicus Leptomydas sardous Leptomydas notos Nemomydas pantherinus Pseudonomoneura hirta Pseudonomoneura micheneri Nemomydas lamia Notosyllegomydas brincki Namibimydas stuckenbergi sp nov Nothomydas picketti sp nov Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata male Cephalocera sp Ratelfontein Nemomydas melanopogon Namadytes sp Gobabeb male Namadytes vansoni Afroleptomydas C mauricei Afroleptomydas C sp Gobabeb Afroleptomydas C sp Stampriet Afroleptomydas A inhacae Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas A sp Clanwilliam Apiocera (Apiocera) sp. © Farhan Bokhari Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Syllegomydas S vittatus Syllegomydas S palestinensis Syllegomydas N astrictus Syllegomydas N heothinos Syllegomydas sp nov Ruacana Syllegomydas N arnoldi male Syllegomydas N dispar Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S proximus Halterorchis karooensis female Eremomidas arabicus Eremomidas sp nov Tiega male Mitrodetus sp nov Argentina female Mitrodetus nanoglossa male Mitrodetus dentitarsis Mitrodetus dimidiatus male
    • ... new phylogenetic hypothesis 12Placement of † Cretomydas santanensis Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus ≈ 112 myo Megascelus nigricornis Neorhaphiomidas setosa male Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis male Diochlistus gracilis Diochlistus melleipennis Australia Anomalomydas mackerrasi Anomalomydas australicus female Paramydas igniticornis Plyomydas peruviensis male Plyomydas sp nov Tandil Messiasia californica male Messiasia painteri Midacritus stuardoanus male Midacritus wagenknechti Cacatuopyga basifascia Cacatuopyga ruficornis male Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Ectyphus armipes Ectyphus amboseli male Ectyphus pinguis Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus maculipennis Miltinus viduatus Mydas clavatus Mydas xanthopterus Stratiomydas lividus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti female Perissocerus transcaspicus Hessemydas parkeri male Hessemydas seyrigi Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Mydaselpis ngurumani Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis male Neolaparopsis puncturatus Haplomydas crassipes † Cretomydas santanensis, Brazil Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa male Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg male Vespiodes leopolditertii Vespiodes phaios Nemomydas brachyrhynchus Nemomydas venosus Leptomydas corsicanus Nemomydas gruenbergi male Leptomydas lusitanicus Eremomidas bek Leptomydas rapti male Leptomydas turcicus Leptomydas sardous Leptomydas notos Nemomydas pantherinus Pseudonomoneura hirta Pseudonomoneura micheneri Nemomydas lamia Notosyllegomydas brincki Namibimydas stuckenbergi sp nov Nothomydas picketti sp nov Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata male Cephalocera sp Ratelfontein Nemomydas melanopogon Namadytes sp Gobabeb male Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticus Nemestrinus nigrovillosus Namadytes vansoni Afroleptomydas C mauricei Bombylius major Dipalta serpentina Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Afroleptomydas C sp Gobabeb Hemigephyra atra Phycus frommeri Diogmites grossus Ospriocerus aeacus Gonioscelis ventralis Hoplistomerus nobilis Afroleptomydas C sp Stampriet Pilica formidolosa Laphria aktis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Afroleptomydas A inhacae Machimus occidentalis Philonicus albiceps Pegesimallus laticornis Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Afroleptomydas A consanguinensis Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Holcocephala calva Apiocera R alastor male Afroleptomydas A opacicinctus Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana male Apiocera P parahydra Apiocera P barri Apiocera P martinorum Afroleptomydas A paganus Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Afroleptomydas A suffusipennis Apiocera P sp Colnett male Apiocera Ap imminuta Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Afroleptomydas A sp Clanwilliam Apiocera Ap sp Wurarga Apiocera Ap asilica Apiocera Ap brevicornis Apiocera Ap sp Stuart male Apiocera Ap hardyi Apiocera Ap newmani Afroleptomydas A sp Obib Apiocera Ap sp Lloyd Apiocera Ap maritima Apiocera Ap similis Rhaphiomidas hasbroucki Rhaphiomidas maehleri Afroleptomydas C sp Koeroegab Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus nigricornis Neorhaphiomidas setosa male Neorhaphiomidas sp Mandurah Afroleptomydas A omeri Cretomydas santanensis Diochlistus auripennis male Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Afroleptomydas A rufithorax Anomalomydas australicus female Paramydas igniticornis Plyomydas peruviensis male Plyomydas sp nov Tandil Messiasia californica male Syllegomydas S lineatus Messiasia painteri Midacritus stuardoanus male Midacritus wagenknechti Cacatuopyga basifascia Cacatuopyga ruficornis male Heteromydas bicolor Syllegomydas S vittatus Opomydas townsendi Parectyphus namibiensis Ectyphus armipes Ectyphus amboseli male Ectyphus pinguis Syllegomydas S palestinensis Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus maculipennis Miltinus viduatus Mydas clavatus Syllegomydas N astrictus Mydas xanthopterus Stratiomydas lividus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Syllegomydas N heothinos Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti female Perissocerus transcaspicus Syllegomydas sp nov Ruacana Hessemydas parkeri male Hessemydas seyrigi Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Syllegomydas N arnoldi male Mydaselpis ngurumani Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis male Syllegomydas N dispar Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa male Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg male Hispanomydas hispanicus Vespiodes leopolditertii Vespiodes phaios Nemomydas brachyrhynchus Nemomydas venosus Leptomydas corsicanus Syllegomydas S algiricus Nemomydas gruenbergi male Leptomydas lusitanicus Eremomidas bek Leptomydas rapti male Leptomydas turcicus Leptomydas sardous Syllegomydas S elachys Leptomydas notos Nemomydas pantherinus Pseudonomoneura hirta Pseudonomoneura micheneri Nemomydas lamia Syllegomydas S proximus Notosyllegomydas brincki Namibimydas stuckenbergi sp nov Nothomydas picketti sp nov Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Halterorchis karooensis female Cephalocera imitata male Cephalocera sp Ratelfontein Nemomydas melanopogon Namadytes sp Gobabeb male Namadytes vansoni Eremomidas arabicus Afroleptomydas C mauricei Afroleptomydas C sp Gobabeb Afroleptomydas C sp Stampriet Afroleptomydas A inhacae Afroleptomydas A consanguinensis Eremomidas sp nov Tiega male Afroleptomydas A opacicinctus Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas A sp Clanwilliam Diochlistus auripennis, Australia © Peter Chew Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Mitrodetus sp nov Argentina female Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Argentina + Chile Syllegomydas S vittatus Syllegomydas S palestinensis Mitrodetus nanoglossa male Syllegomydas N astrictus Syllegomydas N heothinos Syllegomydas sp nov Ruacana Syllegomydas N arnoldi male Syllegomydas N dispar Hispanomydas hispanicus Mitrodetus dentitarsis Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S proximus Halterorchis karooensis female Eremomidas arabicus Mitrodetus dimidiatus male Eremomidas sp nov Tiega male Mitrodetus sp nov Argentina female Mitrodetus nanoglossa male Mitrodetus dentitarsis Mitrodetus dimidiatus male
    • ... new phylogenetic hypothesis 13Monophyly of subfamily taxa Rhaphiomidinae Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelinae Megascelus albovillosus Megascelus nigricornis Neorhaphiomidas setosa male Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis male Diochlistus gracilis Diochlistus melleipennis * Diochlistinae Anomalomydas mackerrasi Anomalomydas australicus female Paramydas igniticornis Anomalomydinae Plyomydas peruviensis male Plyomydas sp nov Tandil Messiasia californica male * Mydinae * Apiophorinae Messiasia painteri Midacritus stuardoanus male Midacritus wagenknechti Cacatuopyga basifascia Cacatuopyga ruficornis male Heteromydas bicolor Cacatuopyginae Opomydas townsendi Parectyphus namibiensis Ectyphus armipes Ectyphus amboseli male * Ectyphinae Ectyphus pinguis Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus maculipennis Miltinus viduatus * Apiophorinae Mydas clavatus Mydas xanthopterus Stratiomydas lividus Dolichogaster brevicornis Phyllomydas bruesii * Mydinae Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti female * Rhopaliinae Perissocerus transcaspicus * Leptomydinae Hessemydas parkeri male Hessemydas seyrigi Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Mydaselpis ngurumani Arenomydas partitus * Syllegomydinae Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis male Neolaparopsis puncturatus Haplomydas crassipes Syllegomydinae: Cephalocera imitata Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa male Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg male Vespiodes leopolditertii Vespiodes phaios Nemomydas brachyrhynchus Nemomydas venosus Leptomydas corsicanus Nemomydas gruenbergi male Leptomydas lusitanicus Eremomidas bek Leptomydas rapti male Leptomydas turcicus Leptomydas sardous Leptomydas notos * Leptomydinae Nemomydas pantherinus Pseudonomoneura hirta Pseudonomoneura micheneri Nemomydas lamia Notosyllegomydas brincki Namibimydas stuckenbergi sp nov Nothomydas picketti sp nov Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata male Cephalocera sp Ratelfontein Nemomydas melanopogon Namadytes sp Gobabeb male Limata tenuicornis Eulonchus sp Six Rivers Neorhynchocephalus volaticus Nemestrinus nigrovillosus Namadytes vansoni Afroleptomydas C mauricei Bombylius major Dipalta serpentina Toxophora maxima Prorates sp Escalante Apsilocephala longistyla Afroleptomydas C sp Gobabeb Hemigephyra atra Phycus frommeri Diogmites grossus Ospriocerus aeacus Gonioscelis ventralis Hoplistomerus nobilis Afroleptomydas C sp Stampriet Pilica formidolosa Laphria aktis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Afroleptomydas A inhacae Machimus occidentalis Philonicus albiceps Pegesimallus laticornis Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Afroleptomydas A consanguinensis Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Holcocephala calva Apiocera R alastor male Afroleptomydas A opacicinctus Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana male Apiocera P parahydra Apiocera P barri Apiocera P martinorum Afroleptomydas A paganus * Syllegomydinae Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri monophyletic Apiocera P arena Apiocera P parkeri Afroleptomydas A suffusipennis Apiocera P sp Colnett male Apiocera Ap imminuta Apiocera Ap tonnoiri Apiocera An obscura Apiocera An philippii Afroleptomydas A sp Clanwilliam Apiocera Ap sp Wurarga Apiocera Ap asilica Apiocera Ap brevicornis Apiocera Ap sp Stuart male Apiocera Ap hardyi Apiocera Ap newmani Afroleptomydas A sp Obib Apiocera Ap sp Lloyd Apiocera Ap maritima Apiocera Ap similis Rhaphiomidas hasbroucki Rhaphiomidas maehleri Afroleptomydas C sp Koeroegab Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus nigricornis Neorhaphiomidas setosa male Neorhaphiomidas sp Mandurah Afroleptomydas A omeri Cretomydas santanensis Diochlistus auripennis male Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Afroleptomydas A rufithorax Anomalomydas australicus female Paramydas igniticornis Plyomydas peruviensis male Plyomydas sp nov Tandil Messiasia californica male Syllegomydas S lineatus Messiasia painteri Midacritus stuardoanus male Midacritus wagenknechti Cacatuopyga basifascia Cacatuopyga ruficornis male Heteromydas bicolor Syllegomydas S vittatus Opomydas townsendi Parectyphus namibiensis * non-monophyletic Ectyphus armipes Ectyphus amboseli male Ectyphus pinguis Syllegomydas S palestinensis Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus maculipennis Miltinus viduatus Mydas clavatus Syllegomydas N astrictus Mydas xanthopterus Stratiomydas lividus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Syllegomydas N heothinos Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti female Perissocerus transcaspicus Syllegomydas sp nov Ruacana Hessemydas parkeri male Hessemydas seyrigi Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Syllegomydas N arnoldi male Mydaselpis ngurumani Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis male Syllegomydas N dispar Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa male Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg male Hispanomydas hispanicus Vespiodes leopolditertii Vespiodes phaios Nemomydas brachyrhynchus Nemomydas venosus Leptomydas corsicanus Syllegomydas S algiricus Nemomydas gruenbergi male Syllegomydinae (43 % species diversity) Leptomydas lusitanicus Eremomidas bek Leptomydas rapti male Leptomydas turcicus Leptomydas sardous Syllegomydas S elachys Leptomydas notos Nemomydas pantherinus Pseudonomoneura hirta Pseudonomoneura micheneri Nemomydas lamia Syllegomydas S proximus Notosyllegomydas brincki Namibimydas stuckenbergi sp nov Nothomydas picketti sp nov Cephalocera longirostris Cephalocera sp Papendorp * Leptomydinae Cephalocera sp Sors Sors Halterorchis karooensis female Cephalocera imitata male Cephalocera sp Ratelfontein Nemomydas melanopogon Namadytes sp Gobabeb male Namadytes vansoni Eremomidas arabicus Afroleptomydas C mauricei Afroleptomydas C sp Gobabeb Afroleptomydas C sp Stampriet Afroleptomydas A inhacae Afroleptomydas A consanguinensis Eremomidas sp nov Tiega male Afroleptomydas A opacicinctus Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Mitrodetus sp nov Argentina female Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Syllegomydas S vittatus * Diochlistinae Syllegomydas S palestinensis Mitrodetus nanoglossa male Syllegomydas N astrictus Syllegomydas N heothinos Syllegomydas sp nov Ruacana Syllegomydas N arnoldi male Syllegomydas N dispar Hispanomydas hispanicus Mitrodetus dentitarsis Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S proximus Halterorchis karooensis female Eremomidas arabicus Mitrodetus dimidiatus male Eremomidas sp nov Tiega male Mitrodetus sp nov Argentina female Mitrodetus nanoglossa male Mitrodetus dentitarsis Mitrodetus dimidiatus male
    • ... new phylogenetic hypothesis 14Earliest branchings within Mydidae Rhaphiomidinae Rhaphiomidas hasbroucki Rhaphiomidas maehleri western North America Rhaphiomidas parkeri Megascelinae Tongamya miranda Namibia + South Africa Megascelus albovillosus Megascelus nigricornis Chile Neorhaphiomidas setosa male Australia Neorhaphiomidas sp Mandurah * Diochlistinae Cretomydas santanensis Diochlistus auripennis male Brazil Diochlistus gracilis Australia Diochlistus melleipennis Anomalomydinae Anomalomydas mackerrasi Anomalomydas australicus female 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 Ospriocerus aeacus Gonioscelis ventralis Hoplistomerus nobilis Pilica formidolosa Laphria aktis Emphysomera pallidapex Ommatius tibialis Neolophonotus bimaculatus Machimus occidentalis Philonicus albiceps Pegesimallus laticornis Dioctria hyalipennis Leptogaster cylindrica Euscelidia pulchra Stichopogon punctum Lasiopogon cinctus Damalis monochaetes Holcocephala calva Apiocera R alastor male Apiocera R braunsi Apiocera P bilineata Apiocera P mexicana male Apiocera P parahydra Apiocera P barri Apiocera P martinorum Apiocera P haruspex Apiocera P aldrichi Apiocera P painteri Apiocera P arena Apiocera P parkeri Apiocera P sp Colnett male 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 male Apiocera Ap hardyi Apiocera Ap newmani Apiocera Ap sp Lloyd Apiocera Ap maritima Apiocera Ap similis Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus nigricornis Neorhaphiomidas setosa male Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis male Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus female Paramydas igniticornis Plyomydas peruviensis male Plyomydas sp nov Tandil Messiasia californica male Messiasia painteri Midacritus stuardoanus male Midacritus wagenknechti Cacatuopyga basifascia Cacatuopyga ruficornis male Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Ectyphus armipes Ectyphus amboseli male Ectyphus pinguis Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus maculipennis Miltinus viduatus Mydas clavatus Mydas xanthopterus Stratiomydas lividus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti female Perissocerus transcaspicus Hessemydas parkeri male Hessemydas seyrigi Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Mydaselpis ngurumani Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis male Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa male Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg male Vespiodes leopolditertii Vespiodes phaios Nemomydas brachyrhynchus Nemomydas venosus Leptomydas corsicanus Nemomydas gruenbergi male Leptomydas lusitanicus Eremomidas bek Leptomydas rapti male Leptomydas turcicus Leptomydas sardous Leptomydas notos Nemomydas pantherinus Pseudonomoneura hirta Pseudonomoneura micheneri Nemomydas lamia Notosyllegomydas brincki Namibimydas stuckenbergi sp nov Nothomydas picketti sp nov Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata male Cephalocera sp Ratelfontein Nemomydas melanopogon Namadytes sp Gobabeb male Namadytes vansoni Afroleptomydas C mauricei Afroleptomydas C sp Gobabeb Afroleptomydas C sp Stampriet Afroleptomydas A inhacae Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas A sp Clanwilliam Mydidae: Anomalomydas mackerrasi © Jean Hort Mydidae: Neorhaphiomidas sp. © Fred Hort Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Syllegomydas S vittatus Syllegomydas S palestinensis Syllegomydas N astrictus Syllegomydas N heothinos Syllegomydas sp nov Ruacana Syllegomydas N arnoldi male Syllegomydas N dispar Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S proximus Halterorchis karooensis female Eremomidas arabicus Eremomidas sp nov Tiega male Mitrodetus sp nov Argentina female Mitrodetus nanoglossa male Mitrodetus dentitarsis Mitrodetus dimidiatus male
    • .... biogeographically interesting taxa 15Biogeographically interesting taxa Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus nigricornis Neorhaphiomidas setosa male Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis male Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus female Paramydas igniticornis Plyomydas peruviensis male Plyomydas sp nov Tandil Messiasia californica male * Apiophorinae Messiasia painteri Midacritus stuardoanus male Midacritus wagenknechti Cacatuopyga basifascia Cacatuopyga ruficornis male Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Ectyphus armipes Ectyphus amboseli male * Ectyphinae Ectyphus pinguis Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus maculipennis Miltinus viduatus * Apiophorinae Mydas clavatus Mydas xanthopterus Stratiomydas lividus Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Pseudorhopalia mirandai Mydidae: Apiophorinae Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti female * Rhopaliinae Perissocerus transcaspicus Hessemydas parkeri male Hessemydas seyrigi Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Mydaselpis ngurumani Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis male Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa male Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg male Vespiodes leopolditertii Vespiodes phaios Nemomydas brachyrhynchus Nemomydas venosus Leptomydas corsicanus Nemomydas gruenbergi male Leptomydas lusitanicus Eremomidas bek Leptomydas rapti male Leptomydas turcicus Leptomydas sardous Leptomydas notos Nemomydas pantherinus Pseudonomoneura hirta Pseudonomoneura micheneri Nemomydas lamia Notosyllegomydas brincki Mydidae: Ectyphinae Namibimydas stuckenbergi sp nov Nothomydas picketti sp nov Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata male Cephalocera sp Ratelfontein Nemomydas melanopogon Namadytes sp Gobabeb male Namadytes vansoni Afroleptomydas C mauricei Afroleptomydas C sp Gobabeb Afroleptomydas C sp Stampriet Afroleptomydas A inhacae Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas A paganus Afroleptomydas A suffusipennis Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Syllegomydas S lineatus Syllegomydas S vittatus Syllegomydas S palestinensis Syllegomydas N astrictus Syllegomydas N heothinos Syllegomydas sp nov Ruacana Syllegomydas N arnoldi male Syllegomydas N dispar Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S proximus Halterorchis karooensis female Eremomidas arabicus Eremomidas sp nov Tiega male Mydidae: Rhopaliinae Mitrodetus sp nov Argentina female Mitrodetus nanoglossa male Mitrodetus dentitarsis Mitrodetus dimidiatus male
    • .... summary + future 16Summary ◊ Apioceridae + Mydidae monophyletic ◊ (Apioceridae + Mydidae) + Asilidae ◊ Apiocera (Apiocera) not monophyletic ◊ earliest branchings within Mydidae well-supported ◊ subfamily taxa relationships still not resolved ◊ Future Research: › field work in Australia (Dec 2011) and Namibia (Feb 2012) › internal head characters › addition of molecular data › adding yet additional taxa? Mydidae: Phyllomydas parvulus © Lyn Atherton
    • 17Acknowledgements ◊ Museum curators and collectors who made specimens available ◊ Photographers ◊ U.S. National Science Foundation (REVSYS DEB-0919333) www.mydidae.tdvia.de