1




Phylogenetic relationships within Mydidae based on
morphology of imagines (Diptera: Asiloidea)




Torsten Dikow
Bio...
. introduction                                      2

Mydidae – “mydas flies”
  ◊ Imagines flower-feeders, some do not fe...
. introduction                                                     3

Taxa overview
  Taxon                       Known Di...
4

Overview of presentation
  ◊ Previous Phylogenetic Hypotheses


  ◊ New Phylogenetic Hypothesis


  ◊ Character Evoluti...
.. previous phylogenetic hypotheses                                              5

Wilcox & Papavero 1974                ...
.. previous phylogenetic hypotheses                 6

Yeates & Irwin 1996                                Irwin & Wiegmann...
.. previous phylogenetic hypotheses                                7

Dikow 2009                                          ...
... new phylogenetic hypothesis                                 8

Taxon Sampling
  ◊ 94 Mydidae species (= 19,9 %)

  ◊ H...
... new phylogenetic hypothesis                                  9

Morphological Characters
  ◊ 186 discrete, parsimony i...
... new phylogenetic hypothesis                 10

Phylogenetic Analysis
  ◊ Morphological data
    › MacClade (version 4...
... new phylogenetic hypothesis                                   11
                                                     ...
... new phylogenetic hypothesis                               12
                                                         ...
... new phylogenetic hypothesis                                                  13

Apiophorinae non-monophyly
          ...
... new phylogenetic hypothesis                            14

Leptomydinae non-monophyly
                                ...
... new phylogenetic hypothesis                                             15

Syllegomydinae non-monophyly + tribal taxa...
.... character evolution                                               16
                                                ...
.... character evolution                                                17
                                               ...
.... summary + future                           18

Summary
 ◊ Mydidae monophyletic

 ◊ (Mydidae + Apioceridae) + Asilidae...
19

Acknowledgements
 ◊ Museum curators and collectors who made specimens available

 ◊ U.S. National Science Foundation (...
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TDikow Mydidae morphological phylogeny ICD 2010

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TDikow Mydidae morphological phylogeny ICD 2010

  1. 1. 1 Phylogenetic relationships within Mydidae based on morphology of imagines (Diptera: Asiloidea) Torsten Dikow Biodiversity Synthesis Center Field Museum of Natural History
  2. 2. . introduction 2 Mydidae – “mydas flies” ◊ Imagines flower-feeders, some do not feed at all ◊ Larvae possibly predatory ◊ 472* species and 66 genera world-wide ◊ 11 subfamily taxa ◊ Imagines 9 – 60 mm (wing length up to 52 mm) Opomydas townsendi ◊ World-wide primarily in arid regions ◊ Oldest definitive fossil Cretomydas santanensis › Crato Formation, Brazil › ≈ 110 Million years * includes species currently being described Cretomydas santanensis
  3. 3. . introduction 3 Taxa overview Taxon Known Diversity Known Distribution Genera Species Afr Aus Nea Neo Ori Pal Anomalomydinae 1 2 × Apiophorinae 5 29 × × Cacatuopyginae 2 6 × Diochlistinae 2 20 × × Ectyphinae 4 10* × × Anomalomydas mackerrasi Leptomydinae 6 54* × × × × × Megascelinae 3 15* × × × Mydinae 12 84 × × Rhaphiomidinae 1 23* × Rhopaliinae 4 21 × × × Syllegomydinae 25 206* × × × unplaced 1 2 * includes species currently being described Cacatuopyga basifascia
  4. 4. 4 Overview of presentation ◊ Previous Phylogenetic Hypotheses ◊ New Phylogenetic Hypothesis ◊ Character Evolution Miltinus stenogaster © Shaun Winterton ◊ Summary + Future Stratiomydas lividus © Charles Melton
  5. 5. .. previous phylogenetic hypotheses 5 Wilcox & Papavero 1974 Artigas & Papavero 1990 › morphology › morphology › similarity ae › similarity ae ae in m e in in e yd go na ae g yd rina ae yd e py om lle di tin na iin Sy my Le ae uo o is al hi ph al in hl at om o yp op io pt oc Diochlistinae remaining Mydidae ac t Ap Rh An Ec M Di C Mitrodetus dentitarsis © Steve Marshall
  6. 6. .. previous phylogenetic hypotheses 6 Yeates & Irwin 1996 Irwin & Wiegmann 2001 › morphology › 28S rDNA › parsimony (successive weighting) › maximum likelihood Asilidae Scenopinidae Apiocera (Pyrocera) Therevidae Apiocera (Ripidosyrma) Apiocera (Anypenus) phillippi Apiocera (Anypenus) Apiocera (Apiocera) sp 1 Apiocera (Apiocera) Apiocera (Apiocera) sp 2 Rhaphiomidinae Apiocera (Pyrocera) haruspex Megascelinae Apiocera (Pyrocera) sp Rhopaliinae Rhaphiomidas terminalis Rhaphiomidinae Cacatuopyginae Megascelus albovillosus Leptomydinae Megascelinae Tongamya stuckenbergi Syllegomydinae Diochlistus hackeri Diochlistinae Anomalomydinae Rhopalia sp Rhopaliinae Diochlistinae Ectyphinae Mitrodetus australis Diochlistinae Apiophorinae Nemomydas pantherinus Leptomydinae Mydinae Arenomydas callosus Syllegomydinae
  7. 7. .. previous phylogenetic hypotheses 7 Dikow 2009 Dikow unpublished presented at ICD6 in Fukuoka › morphology › 18S + 28S rDNA, CAD › parsimony › parsimony (POY) Bombylius major Apsilocephala longistylus Prorates sp (Esacalante) Phycus frommeri ... Asilidae Hemigephyra atra Apiocera (Apiocera) sp (Barakula) Apiocera aldrichi Apiocera (Pyrocera) haruspex Apioceridae Apioceridae Apiocera (Pyrocera) painteri Apiocera painteri Stichopogon punctum Saropogon luteus Rhaphiomidas maehleri Rhaphiomidinae Laxenecera albicincta Hoplistomerus nobilis Asilidae Neorhaphiomidas sp (Mandurah) Megascelinae Gonioscelis ventralis Leptogaster cylindrica Opomydas townsendi Ectyphinae Asilus crabroniformis Promachus amastrus Mitrodetus dentitarsis Diochlistinae Emphysomera pallidapex Rhaphiomidas xanthos Rhaphiomidinae Mitrodetus dentitarsis Diochlistinae Mydas clavatus Mydinae Opomydas townsendi Ectyphinae Miltinus cardinalis Apiophorinae Pseudonomoneura hirta Mydas clavatus Mydinae Leptomydinae Mahafalymydas wiegmanni Nemomydas brachyrhynchus Syllegomydas sp (Nizzanem) Afroleptomydas sp (Clanwilliam) Syllegomydinae Afroleptomydas sp (Clanwilliam) Syllegomydinae Afroleptomydas sp (RSA) Cephalocera imitata Cephalocera sp (Montegu) Nemomydas brachyrhynchus Nemomydas sp (Tesopaco) Leptomydinae
  8. 8. ... new phylogenetic hypothesis 8 Taxon Sampling ◊ 94 Mydidae species (= 19,9 %) ◊ Higher-level taxa › 40 genera (= 61,5 %) › 7 tribal taxa (= 87,5 %) › 11 subfamily taxa (= 100 %) collecting localities of specimens included ◊ Outgroup taxa › 37 Asiloidea species Apioceridae (11), Apsilocephalidae (1), Asilidae (19), Bombyliidae (3), Scenopinidae (1), Therevidae (2) › 1 Acroceridae, 2 Nemestrinidae › 1 Tabanidae
  9. 9. ... new phylogenetic hypothesis 9 Morphological Characters ◊ 186 discrete, parsimony informative characters tagma # characters remarks Head 42 Thorax 83 39 chaetotaxy, sclerites, etc. 21 wing 23 leg Abdomen 61 16 sclerite shapes, chaetotaxy, etc. 25 ? terminalia 20 / genitalia
  10. 10. ... new phylogenetic hypothesis 10 Phylogenetic Analysis ◊ Morphological data › MacClade (version 4.08) › WinClada (version 1.0.08) › all characters non-additive ◊ Tree Search › TNT (version 1.1 June 2010) Eremomidas arabicus © Drew Gardner › Ratchet, TreeDrifting, TreeFusing, Sectorial Searches (Goloboff 1999, Nixon 1999) › regular parsimony › implied weighting (k = 3) (Goloboff 1993) Heteromydas bicolor © Greg Ballmer
  11. 11. ... new phylogenetic hypothesis 11 Limata tenuicornis Tabanidae Results & Sister-group Neorhynchocephalus volaticus Nemestrinus nigrovillosus Nemestrinidae Eulonchus sp. (SixRivers) Acroceridae Prorates sp. (Escalante) Scenopinidae Toxophora maxima Bombylius major Bombyliidae Dipalta serpentina Apsilocephala longistyla Apsilocephalidae Hemigephyra atra Phycus frommeri Therevidae ◊ Parsimony search Hoplistomerus nobilis Pilica formidolosa Laphria aktis Neolophonotus bimaculatus Emphysomera pallidapex Ommatius tibialis Machimus occidentalis Asilidae Philonicus albiceps › 421 trees, 1485 steps Stichopogon punctum Lasiopogon cinctus Diogmites grossus Ospriocerus aeacus Gonioscelis ventralis Damalis monochaetes Pegesimallus laticornis Leptogaster cylindrica › strict consensus with large polytomy Euscelidia pulchra Dioctria hyalipennis Holcocephala calva Apiocera (Pyrocera) bilineata Apiocera (Ripidosyrma) braunsi Apioceridae Apiocera (Pyrocera) aldrichi Apiocera (Pyrocera) painteri Apiocera (Pyrocera) arena Apiocera (Pyrocera) parkeri in derived ingroup Apiocera (Anypenus) obscura Apiocera (Anypenus) philippii Apiocera (Apiocera) sp. (Wurarga) Apiocera (Apiocera) sp. (Maranoa) Apiocera (Apiocera) sp. (Stuart) Rhaphiomidas hasbroucki Rhaphiomidas parkeri Rhaphiomidas maehleri Tongamya miranda Neorhaphiomidas sp. (Mandurah) Neorhaphiomidas setosus ◊ Implied weights Megascelus albovillosus Megascelus nigricornis Rhopalia olivieri Rhopalia annulata Perissocerus dumonti Anomalomydas mackerrasi Cacatuopyga basifascia Cacatuopyga ruficornis Messiasia californica › 6 trees, fit 95.58644 Phyllomydas parvulus Mydas clavatus Mydas xanthopterus Paramydas igniticornis Stratiomydas lividus Apiophora quadricinctata Apiophora paulseni Miltinus maculipennis › = 1556 steps Miltinus viduatus Miltinus stenogaster Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Ectyphus pinguis Ectyphus sp. nov. (Amboseli) Haplomydas crassipes › strict consensus well-resolved Eremohaplomydas desertorum Eremohaplomydas sp. nov. (Kotwa) Eremomidas arabicus Eremomidas sp. nov. (Tiega) Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Pseudonomoneura micheneri Pseudonomoneura hirta Hessemydas parkeri Hessemydas seyrigi Nemomydas venosus Mydidae Nemomydas brachyrhynchus Leptomydas gruenbergi Nemomydas melanopogon Leptomydas corsicanus Leptomydas sardous Leptomydas notos sp. nov. Leptomydas rapti sp. nov. Leptomydas turcicus Eremomidas bek Leptomydas lusitanicus Mydaselpis ngurumani sp. nov. Vespiodes lepolditertii Limata tenuicornis volaticus Neorhynchocephalus Vespiodes phaios sp. nov. Nemestrinus nigrovillosus Eulonchus sp. (SixRivers) Prorates sp. (Escalante) Toxophora maxima Mydaselpis sp. nov. (Daan) Bombylius major Dipalta serpentina Apsilocephalaatra Hemigephyra longistyla Mydaselpis sp. nov. (Gamsberg) Phycus frommeri Hoplistomerus nobilis Pilica formidolosa Laphria aktis bimaculatus Namadytes maculiventris Neolophonotus Emphysomera pallidapex Ommatius tibialis Machimus occidentalis Namadytes prozeskyi Philonicus albiceps Stichopogoncinctus Lasiopogon punctum Diogmites grossus Cephalocera imitata Ospriocerus aeacus Gonioscelis ventralis Damalis monochaetes Pegesimalluscylindrica Cephalocera longirostris Leptogaster laticornis Euscelidia pulchra Cephalocera sp. (Papendorp) Dioctria hyalipennis Holcocephala calva bilineata Apiocera (Pyrocera) Apiocera (Ripidosyrma) braunsi Apiocera (Pyrocera) aldrichi Neolaparopsis puncturatus Apiocera (Pyrocera) painteri Apiocera (Pyrocera) arena Apiocera (Pyrocera) obscura Apiocera (Anypenus)parkeri Arenomydas caerulescens Apiocera (Anypenus)sp. (Wurarga) philippii Apiocera (Apiocera) sp. (Maranoa) Apiocera (Apiocera) Apiocera (Apiocera) sp. (Stuart) Arenomydas callosus Rhaphiomidas hasbroucki Rhaphiomidas parkeri Rhaphiomidas maehleri Tongamya miranda (Mandurah) Arenomydas namaquensis Neorhaphiomidas sp. Neorhaphiomidas setosus Megascelus albovillosus Megascelus nigricornis Nomoneura caffra Rhopalia olivieri Rhopalia annulata Perissocerus dumonti Anomalomydas mackerrasi Nomoneura paradoxa Cacatuopyga basifascia Cacatuopyga ruficornis Messiasia californica Phyllomydas parvulus Nomoneuroides natalensis Mydas clavatus Mydas xanthopterus Paramydas igniticornis Stratiomydas lividus Halterorchis sp. nov. (Cape Recife) Apiophora quadricinctata Apiophora paulseni Miltinus maculipennis Miltinus viduatus gen. nov. + sp. nov. (Panne) Miltinus stenogaster Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis gen. nov. + sp. nov. (Ugab) Ectyphus pinguis (Amboseli) Ectyphus sp. crassipes Haplomydas nov. Eremohaplomydas desertorum gen. nov. + sp. nov. (Gobabeb) Eremohaplomydas sp. nov. (Kotwa) Eremomidas arabicus(Tiega) Eremomidas sp. nov. Namibimydas sp. nov. (Agate) Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Pseudonomoneura micheneri Nothomydas sp. nov. (Skerpioenkop) Pseudonomoneura Hessemydas seyrigihirta parkeri Hessemydas venosus Nemomydas brachyrhynchus Afroleptomydas A consanguinensis Nemomydas gruenbergi Leptomydas Nemomydas corsicanus Leptomydas melanopogon Afroleptomydas A suffusipennis Leptomydas sardous nov. Leptomydas notos sp. Leptomydas rapti sp. Leptomydas turcicus nov. Afroleptomydas C mauricei Eremomidas lusitanicus Leptomydas bek Mydaselpislepolditertii sp. nov. Vespiodes ngurumani Afroleptomydas A sp. (Clanwilliam) Vespiodes phaios sp. nov. Mydaselpis sp. nov. (Daan) Mydaselpis sp. nov. (Gamsberg) Namadytes maculiventris Afroleptomydas C sp. (Stampriet) Namadytes prozeskyi Cephalocera imitata Cephalocera longirostris Cephalocera sp. (Papendorp) Syllegomydas sp. nov. (Ruacana) Neolaparopsis puncturatus Arenomydas caerulescens Arenomydas callosus Arenomydascaffra Syllegomydas N arnoldi Nomoneura namaquensis Nomoneura paradoxa Syllegomydas N dispar Nomoneuroides nov. (Cape sp. natalensis Halterorchissp. nov. (Panne)Recife) gen. nov. + sp. nov. (Ugab) gen. nov. + gen. nov. + sp. nov. (Gobabeb) Syllegomydas S palestinensis strict consensus tree from implied weights analysis Namibimydassp. nov. (Skerpioenkop) Nothomydas sp. nov. (Agate) Afroleptomydas A consanguinensis Afroleptomydas A suffusipennis Syllegomydas N heothinos sp. nov. Afroleptomydas C sp. (Clanwilliam) Afroleptomydas A mauricei Afroleptomydas C sp. Syllegomydas sp. nov.(Stampriet) Syllegomydas S algiricus Syllegomydas N dispar(Ruacana) arnoldi Syllegomydas N palestinensis Syllegomydas S Syllegomydas N heothinos Syllegomydas S elachys sp. nov. Syllegomydas S algiricus sp. nov. Syllegomydas S astrictus sp. nov. Syllegomydas N elachys sp. nov. Syllegomydas N astrictus sp. nov.
  12. 12. ... new phylogenetic hypothesis 12 Rhaphiomidas hasbroucki Rhaphiomidinae Monophyly & Rhaphiomidinae Rhaphiomidas parkeri Rhaphiomidas maehleri Tongamya miranda Neorhaphiomidas sp. (Mandurah) Neorhaphiomidas setosus Megascelus albovillosus Megascelinae Megascelus nigricornis Rhopalia olivieri Rhopalia annulata Rhopaliinae Perissocerus dumonti Anomalomydas mackerrasi Anomalomydinae Cacatuopyga basifascia Cacatuopyga ruficornis Cacatuopyginae Messiasia californica Phyllomydas parvulus Mydas clavatus Mydas xanthopterus Mydinae Paramydas igniticornis Stratiomydas lividus Apiophora quadricinctata Apiophora paulseni Miltinus maculipennis Miltinus viduatus Apiophorinae Miltinus stenogaster Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Ectyphus pinguis Ectyphinae Ectyphus sp. nov. (Amboseli) Haplomydas crassipes Eremohaplomydas desertorum Eremohaplomydas sp. nov. (Kotwa) Syllegomydinae Eremomidas arabicus Eremomidas sp. nov. (Tiega) Leptomydinae Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Diochlistinae Pseudonomoneura micheneri Pseudonomoneura hirta Hessemydas parkeri Hessemydas seyrigi Nemomydas venosus Nemomydas brachyrhynchus Rhaphiomidas auratus © Greg Ballmer Leptomydas gruenbergi Nemomydas melanopogon Leptomydas corsicanus Leptomydinae Leptomydas sardous Leptomydas notos sp. nov. Leptomydas rapti sp. nov. Leptomydas turcicus Eremomidas bek Leptomydas lusitanicus Mydaselpis ngurumani sp. nov. Vespiodes lepolditertii Vespiodes phaios sp. nov. Mydaselpis sp. nov. (Daan) Mydaselpis sp. nov. (Gamsberg) Namadytes maculiventris Namadytes prozeskyi Cephalocera imitata Cephalocera longirostris Cephalocera sp. (Papendorp) Neolaparopsis puncturatus Arenomydas caerulescens Arenomydas callosus Limata tenuicornis volaticus Neorhynchocephalus Nemestrinus nigrovillosus Eulonchus sp. (SixRivers) Prorates sp. (Escalante) Arenomydas namaquensis Nomoneura caffra Toxophora maxima Bombylius major Dipalta serpentina Apsilocephalaatra Hemigephyra longistyla Nomoneura paradoxa Phycus frommeri Hoplistomerus nobilis Pilica formidolosa Laphria aktis bimaculatus Neolophonotus Emphysomera pallidapex Nomoneuroides natalensis Ommatius tibialis Syllegomydinae Machimus occidentalis Philonicus albiceps Stichopogoncinctus Lasiopogon punctum Diogmites grossus Halterorchis sp. nov. (Cape Recife) Ospriocerus aeacus Gonioscelis ventralis Damalis monochaetes Pegesimalluscylindrica Leptogaster laticornis Euscelidia pulchra Dioctria hyalipennis Holcocephala calva bilineata Apiocera (Pyrocera) Apiocera (Ripidosyrma) braunsi Apiocera (Pyrocera) aldrichi gen. nov. + sp. nov. (Panne) gen. nov. + sp. nov. (Ugab) Apiocera (Pyrocera) painteri Apiocera (Pyrocera) arena Apiocera (Pyrocera) obscura Apiocera (Anypenus)parkeri Apiocera (Anypenus)sp. (Wurarga) philippii Apiocera (Apiocera) sp. (Maranoa) gen. nov. + sp. nov. (Gobabeb) Apiocera (Apiocera) Apiocera (Apiocera) sp. (Stuart) Rhaphiomidas hasbroucki Rhaphiomidas parkeri monophyletic Rhaphiomidas maehleri Tongamya miranda (Mandurah) Neorhaphiomidas sp. Neorhaphiomidas setosus Megascelus albovillosus Megascelus nigricornis Rhopalia olivieri Namibimydas sp. nov. (Agate) Nothomydas sp. nov. (Skerpioenkop) Rhopalia annulata Perissocerus dumonti Anomalomydas mackerrasi Cacatuopyga basifascia Cacatuopyga ruficornis Messiasia californica Afroleptomydas A consanguinensis Phyllomydas parvulus Mydas clavatus Mydas xanthopterus Paramydas igniticornis Stratiomydas lividus Apiophora quadricinctata Afroleptomydas A suffusipennis Apiophora paulseni Miltinus maculipennis Miltinus viduatus Miltinus stenogaster Heteromydas bicolor Afroleptomydas C mauricei Opomydas townsendi Parectyphus namibiensis Ectyphus pinguis (Amboseli) Ectyphus sp. crassipes Haplomydas nov. Eremohaplomydas desertorum Afroleptomydas A sp. (Clanwilliam) Eremohaplomydas sp. nov. (Kotwa) Eremomidas arabicus(Tiega) non-monophyletic Eremomidas sp. nov. Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Afroleptomydas C sp. (Stampriet) Pseudonomoneura micheneri Pseudonomoneura Hessemydas seyrigihirta parkeri Hessemydas venosus Nemomydas brachyrhynchus Syllegomydas sp. nov. (Ruacana) Nemomydas gruenbergi Leptomydas Nemomydas corsicanus Leptomydas melanopogon Leptomydas sardous nov. Leptomydas notos sp. Syllegomydas N arnoldi Leptomydas rapti sp. Leptomydas turcicus nov. Eremomidas lusitanicus Leptomydas bek Mydaselpislepolditertii sp. nov. Vespiodes ngurumani Syllegomydas N dispar Vespiodes phaios sp. nov. Mydaselpis sp. nov. (Daan) Mydaselpis sp. nov. (Gamsberg) Namadytes maculiventris Namadytes prozeskyi Syllegomydas S palestinensis Cephalocera imitata Cephalocera longirostris Cephalocera sp. (Papendorp) Neolaparopsis puncturatus Arenomydas caerulescens Arenomydas callosus Syllegomydas N heothinos sp. nov. Arenomydascaffra Nomoneura namaquensis Nomoneura paradoxa Nomoneuroides nov. (Cape sp. natalensis Halterorchissp. nov. (Panne)Recife) gen. nov. + sp. nov. (Ugab) Syllegomydas S algiricus gen. nov. + gen. nov. + sp. nov. (Gobabeb) Namibimydassp. nov. (Skerpioenkop) Nothomydas sp. nov. (Agate) Afroleptomydas A consanguinensis Afroleptomydas A suffusipennis Afroleptomydas C sp. (Clanwilliam) Afroleptomydas A mauricei Afroleptomydas C sp. Syllegomydas sp. nov.(Stampriet) Syllegomydas N dispar(Ruacana) arnoldi Syllegomydas S elachys sp. nov. Syllegomydas N astrictus sp. nov. Syllegomydas N palestinensis Syllegomydas S Syllegomydas N heothinos Syllegomydas S algiricus sp. nov. Syllegomydas S astrictus sp. nov. Syllegomydas N elachys sp. nov.
  13. 13. ... new phylogenetic hypothesis 13 Apiophorinae non-monophyly Rhaphiomidas hasbroucki Rhaphiomidas parkeri Rhaphiomidas maehleri Tongamya miranda Neorhaphiomidas sp. (Mandurah) Neorhaphiomidas setosus Megascelus albovillosus Megascelus nigricornis Rhopalia olivieri Rhopalia annulata Perissocerus dumonti Anomalomydas mackerrasi Cacatuopyga basifascia Cacatuopyga ruficornis Messiasia californica Phyllomydas parvulus Mydas clavatus Mydas xanthopterus Paramydas igniticornis Stratiomydas lividus Apiophora quadricinctata Apiophora paulseni Miltinus maculipennis Miltinus viduatus Miltinus stenogaster Paramydas igniticornis Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Ectyphus pinguis Ectyphus sp. nov. (Amboseli) ... Limata tenuicornis volaticus Neorhynchocephalus Nemestrinus nigrovillosus Eulonchus sp. (SixRivers) Prorates sp. (Escalante) Toxophora maxima Bombylius major Dipalta serpentina Apsilocephalaatra Hemigephyra longistyla Phycus frommeri Hoplistomerus nobilis Pilica formidolosa Laphria aktis bimaculatus Neolophonotus Emphysomera pallidapex Ommatius tibialis Machimus occidentalis Philonicus albiceps Stichopogoncinctus Lasiopogon punctum Diogmites grossus Ospriocerus aeacus Gonioscelis ventralis Damalis monochaetes Pegesimalluscylindrica Leptogaster laticornis Euscelidia pulchra Dioctria hyalipennis Holcocephala calva bilineata Apiocera (Pyrocera) Apiocera (Ripidosyrma) braunsi Apiocera (Pyrocera) aldrichi Apiocera (Pyrocera) painteri Apiocera (Pyrocera) arena Apiocera (Pyrocera) obscura Apiocera (Anypenus)parkeri Apiocera (Anypenus)sp. (Wurarga) philippii Apiocera (Apiocera) sp. (Maranoa) Apiocera (Apiocera) Apiocera (Apiocera) sp. (Stuart) Rhaphiomidas hasbroucki Rhaphiomidas parkeri Apiophorinae Rhaphiomidas maehleri Tongamya miranda (Mandurah) Neorhaphiomidas sp. Neorhaphiomidas setosus Megascelus albovillosus Megascelus nigricornis Rhopalia olivieri Rhopalia annulata Perissocerus dumonti Anomalomydas mackerrasi Cacatuopyga basifascia Cacatuopyga ruficornis Messiasia californica Phyllomydas parvulus Mydas clavatus Mydas xanthopterus Paramydas igniticornis Stratiomydas lividus Apiophora quadricinctata Apiophora paulseni Miltinus maculipennis Miltinus viduatus Miltinus stenogaster Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Ectyphus pinguis (Amboseli) Ectyphus sp. crassipes Haplomydas nov. Eremohaplomydas desertorum Eremohaplomydas sp. nov. (Kotwa) Eremomidas arabicus(Tiega) Mydinae Eremomidas sp. nov. Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Pseudonomoneura micheneri Pseudonomoneura Hessemydas seyrigihirta parkeri Hessemydas venosus Nemomydas brachyrhynchus Nemomydas gruenbergi Leptomydas Nemomydas corsicanus Leptomydas melanopogon Leptomydas sardous nov. Leptomydas notos sp. Leptomydas rapti sp. Leptomydas turcicus nov. Eremomidas lusitanicus Leptomydas bek Mydaselpislepolditertii sp. nov. Vespiodes ngurumani Vespiodes phaios sp. nov. Mydaselpis sp. nov. (Daan) Mydaselpis sp. nov. (Gamsberg) Namadytes maculiventris Namadytes prozeskyi Cephalocera imitata Cephalocera longirostris Cephalocera sp. (Papendorp) Neolaparopsis puncturatus Arenomydas caerulescens Arenomydas callosus Arenomydascaffra Nomoneura namaquensis Nomoneura paradoxa Nomoneuroides nov. (Cape sp. natalensis Halterorchissp. nov. (Panne)Recife) gen. nov. + sp. nov. (Ugab) Miltinus viduatus © Gerhard Weber gen. nov. + gen. nov. + sp. nov. (Gobabeb) Namibimydassp. nov. (Skerpioenkop) Nothomydas sp. nov. (Agate) Afroleptomydas A consanguinensis Afroleptomydas A suffusipennis Afroleptomydas C sp. (Clanwilliam) Afroleptomydas A mauricei Afroleptomydas C sp. Syllegomydas sp. nov.(Stampriet) Syllegomydas N dispar(Ruacana) arnoldi Syllegomydas N palestinensis Syllegomydas S Syllegomydas N heothinos Syllegomydas S algiricus sp. nov. Syllegomydas S astrictus sp. nov. Syllegomydas N elachys sp. nov.
  14. 14. ... new phylogenetic hypothesis 14 Leptomydinae non-monophyly Haplomydas crassipes Eremohaplomydas desertorum Eremohaplomydas sp. nov. (Kotwa) Eremomidas arabicus Eremomidas sp. nov. (Tiega) Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Pseudonomoneura micheneri Pseudonomoneura hirta Hessemydas parkeri Hessemydas seyrigi Nemomydas venosus Nemomydas brachyrhynchus Leptomydas gruenbergi Nemomydas melanopogon Leptomydas corsicanus Leptomydas sardous Leptomydas notos sp. nov. Leptomydas rapti sp. nov. Leptomydas turcicus Eremomidas bek Leptomydas lusitanicus Mydaselpis ngurumani sp. nov. Vespiodes lepolditertii Vespiodes phaios sp. nov. Mydaselpis sp. nov. (Daan) Mydaselpis sp. nov. (Gamsberg) Eremomidas arabicus © Drew Gardner Namadytes maculiventris Namadytes prozeskyi Cephalocera imitata Cephalocera longirostris Cephalocera sp. (Papendorp) Neolaparopsis puncturatus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Halterorchis sp. nov. (Cape Recife) Limata tenuicornis volaticus gen. nov. + sp. nov. (Panne) gen. nov. + sp. nov. (Ugab) Neorhynchocephalus Nemestrinus nigrovillosus Eulonchus sp. (SixRivers) Prorates sp. (Escalante) Toxophora maxima Bombylius major Dipalta serpentina gen. nov. + sp. nov. (Gobabeb) Apsilocephalaatra Hemigephyra longistyla Phycus frommeri Hoplistomerus nobilis Pilica formidolosa Laphria aktis bimaculatus Neolophonotus Emphysomera pallidapex Namibimydas sp. nov. (Agate) Ommatius tibialis Machimus occidentalis Philonicus albiceps Stichopogoncinctus Lasiopogon punctum Diogmites grossus Ospriocerus aeacus Nothomydas sp. nov. (Skerpioenkop) Gonioscelis ventralis Damalis monochaetes Pegesimalluscylindrica Leptogaster laticornis Euscelidia pulchra Dioctria hyalipennis Holcocephala calva bilineata Apiocera (Pyrocera) Afroleptomydas A consanguinensis Apiocera (Ripidosyrma) braunsi Apiocera (Pyrocera) aldrichi Apiocera (Pyrocera) painteri Apiocera (Pyrocera) arena Apiocera (Pyrocera) obscura Apiocera (Anypenus)parkeri Apiocera (Anypenus)sp. (Wurarga) philippii Afroleptomydas A suffusipennis Apiocera (Apiocera) sp. (Maranoa) Apiocera (Apiocera) Apiocera (Apiocera) sp. (Stuart) Rhaphiomidas hasbroucki Rhaphiomidas parkeri Rhaphiomidas maehleri Tongamya miranda (Mandurah) Neorhaphiomidas sp. Afroleptomydas C mauricei Neorhaphiomidas setosus Megascelus albovillosus Megascelus nigricornis Rhopalia olivieri Rhopalia annulata Perissocerus dumonti Anomalomydas mackerrasi Afroleptomydas A sp. (Clanwilliam) Cacatuopyga basifascia Cacatuopyga ruficornis Messiasia californica Phyllomydas parvulus Mydas clavatus Mydas xanthopterus Paramydas igniticornis Stratiomydas lividus Afroleptomydas C sp. (Stampriet) Apiophora quadricinctata Apiophora paulseni Miltinus maculipennis Miltinus viduatus Miltinus stenogaster Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Syllegomydas sp. nov. (Ruacana) Ectyphus pinguis (Amboseli) Ectyphus sp. crassipes Haplomydas nov. Eremohaplomydas desertorum Eremohaplomydas sp. nov. (Kotwa) Eremomidas arabicus(Tiega) Eremomidas sp. nov. Syllegomydas N arnoldi Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Pseudonomoneura micheneri Pseudonomoneura Hessemydas seyrigihirta parkeri Hessemydas venosus Nemomydas brachyrhynchus Syllegomydas N dispar Nemomydas gruenbergi Leptomydas Nemomydas corsicanus Leptomydas melanopogon Leptomydas sardous nov. Leptomydas notos sp. Leptomydas rapti sp. Leptomydas turcicus nov. Syllegomydas S palestinensis Eremomidas lusitanicus Leptomydas bek Mydaselpislepolditertii sp. nov. Vespiodes ngurumani Vespiodes phaios sp. nov. Mydaselpis sp. nov. (Daan) Mydaselpis sp. nov. (Gamsberg) Syllegomydas N heothinos sp. nov. Namadytes maculiventris Namadytes prozeskyi Cephalocera imitata Cephalocera longirostris Cephalocera sp. (Papendorp) Neolaparopsis puncturatus Arenomydas caerulescens Syllegomydas S algiricus Arenomydas callosus Arenomydascaffra Nomoneura namaquensis Nomoneura paradoxa Nomoneuroides nov. (Cape sp. natalensis Halterorchissp. nov. (Panne)Recife) gen. nov. + sp. nov. (Ugab) Leptomydas lusitanicus © Valter Jacinto gen. nov. + Syllegomydas S elachys sp. nov. gen. nov. + sp. nov. (Gobabeb) Namibimydassp. nov. (Skerpioenkop) Nothomydas sp. nov. (Agate) Afroleptomydas A consanguinensis Afroleptomydas A suffusipennis Afroleptomydas C sp. (Clanwilliam) Afroleptomydas A mauricei Afroleptomydas C sp. Syllegomydas sp. nov.(Stampriet) Syllegomydas N astrictus sp. nov. Syllegomydas N dispar(Ruacana) arnoldi Syllegomydas N palestinensis Syllegomydas S Syllegomydas N heothinos Syllegomydas S algiricus sp. nov. Syllegomydas S astrictus sp. nov. Syllegomydas N elachys sp. nov.
  15. 15. ... new phylogenetic hypothesis 15 Syllegomydinae non-monophyly + tribal taxa Haplomydas crassipes Eremohaplomydas desertorum Eremohaplomydas sp. nov. (Kotwa) Eremomidas arabicus Eremomidas sp. nov. (Tiega) Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Pseudonomoneura micheneri Pseudonomoneura hirta Hessemydas parkeri Hessemydas seyrigi Nemomydas venosus Nemomydas brachyrhynchus Leptomydas gruenbergi Nemomydas melanopogon Leptomydas corsicanus Leptomydas sardous Leptomydas notos sp. nov. Leptomydas rapti sp. nov. Leptomydas turcicus Eremomidas bek Leptomydas lusitanicus Mydaselpis ngurumani sp. nov. Vespiodes lepolditertii Vespiodes phaios sp. nov. Mydaselpis sp. nov. (Daan) Mydaselpis sp. nov. (Gamsberg) Haplomydas crassipes Mydaselpidini Namadytes maculiventris Namadytes prozeskyi Cephalocera imitata Cephalocera longirostris Cephalocera sp. (Papendorp) Neolaparopsis puncturatus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Halterorchis sp. nov. (Cape Recife) Limata tenuicornis volaticus Cephalocerini gen. nov. + sp. nov. (Panne) gen. nov. + sp. nov. (Ugab) Neorhynchocephalus Nemestrinus nigrovillosus Eulonchus sp. (SixRivers) Prorates sp. (Escalante) Toxophora maxima Bombylius major Dipalta serpentina gen. nov. + sp. nov. (Gobabeb) Apsilocephalaatra Hemigephyra longistyla Phycus frommeri Hoplistomerus nobilis Pilica formidolosa Laphria aktis bimaculatus Neolophonotus Emphysomera pallidapex Namibimydas sp. nov. (Agate) Ommatius tibialis Machimus occidentalis Philonicus albiceps Stichopogoncinctus Lasiopogon punctum Diogmites grossus Ospriocerus aeacus Nothomydas sp. nov. (Skerpioenkop) Gonioscelis ventralis Damalis monochaetes Pegesimalluscylindrica Leptogaster laticornis Euscelidia pulchra Dioctria hyalipennis Holcocephala calva bilineata Apiocera (Pyrocera) Afroleptomydas A consanguinensis Apiocera (Ripidosyrma) braunsi Apiocera (Pyrocera) aldrichi Apiocera (Pyrocera) painteri Apiocera (Pyrocera) arena Apiocera (Pyrocera) obscura Apiocera (Anypenus)parkeri Apiocera (Anypenus)sp. (Wurarga) philippii Afroleptomydas A suffusipennis Apiocera (Apiocera) sp. (Maranoa) Apiocera (Apiocera) Apiocera (Apiocera) sp. (Stuart) Rhaphiomidas hasbroucki Rhaphiomidas parkeri Rhaphiomidas maehleri Tongamya miranda (Mandurah) Neorhaphiomidas sp. Afroleptomydas C mauricei Neorhaphiomidas setosus Megascelus albovillosus Megascelus nigricornis Rhopalia olivieri Rhopalia annulata Perissocerus dumonti Anomalomydas mackerrasi Afroleptomydas A sp. (Clanwilliam) Cacatuopyga basifascia Cacatuopyga ruficornis Messiasia californica Phyllomydas parvulus Mydas clavatus Mydas xanthopterus Paramydas igniticornis Stratiomydas lividus Afroleptomydas C sp. (Stampriet) Apiophora quadricinctata Apiophora paulseni Miltinus maculipennis Miltinus viduatus Miltinus stenogaster Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Halterorchini Syllegomydas sp. nov. (Ruacana) Ectyphus pinguis (Amboseli) Ectyphus sp. crassipes Haplomydas nov. Eremohaplomydas desertorum Eremohaplomydas sp. nov. (Kotwa) Eremomidas arabicus(Tiega) Eremomidas sp. nov. Syllegomydas N arnoldi Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Pseudonomoneura micheneri Pseudonomoneura Hessemydas seyrigihirta parkeri Hessemydas venosus Nemomydas brachyrhynchus Syllegomydas N dispar Nemomydas gruenbergi Leptomydas Nemomydas corsicanus Leptomydas melanopogon Leptomydas sardous nov. Leptomydas notos sp. Leptomydas rapti sp. Leptomydas turcicus nov. Syllegomydas S palestinensis Eremomidas lusitanicus Leptomydas bek Mydaselpislepolditertii sp. nov. Vespiodes ngurumani Vespiodes phaios sp. nov. Mydaselpis sp. nov. (Daan) Mydaselpis sp. nov. (Gamsberg) Syllegomydas N heothinos sp. nov. Namadytes maculiventris Namadytes prozeskyi Syllegomydini Cephalocera imitata Cephalocera longirostris Cephalocera sp. (Papendorp) Neolaparopsis puncturatus Arenomydas caerulescens Syllegomydas S algiricus Arenomydas callosus Arenomydascaffra Nomoneura namaquensis Nomoneura paradoxa Nomoneuroides nov. (Cape sp. natalensis Halterorchissp. nov. (Panne)Recife) gen. nov. + sp. nov. (Ugab) Cephalocera imitata gen. nov. + Syllegomydas S elachys sp. nov. gen. nov. + sp. nov. (Gobabeb) Namibimydassp. nov. (Skerpioenkop) Nothomydas sp. nov. (Agate) Afroleptomydas A consanguinensis Afroleptomydas A suffusipennis Afroleptomydas C sp. (Clanwilliam) Afroleptomydas A mauricei Afroleptomydas C sp. Syllegomydas sp. nov.(Stampriet) Syllegomydas N astrictus sp. nov. Syllegomydas N dispar(Ruacana) arnoldi Syllegomydas N palestinensis Syllegomydas S Syllegomydas N heothinos Syllegomydas S algiricus sp. nov. Syllegomydas S astrictus sp. nov. Syllegomydas N elachys sp. nov.
  16. 16. .... character evolution 16 Rhaphiomidas hasbroucki Character evolution: head + female Rhaphiomidas parkeri Rhaphiomidas maehleri Tongamya miranda Neorhaphiomidas sp. (Mandurah) Neorhaphiomidas setosus Megascelus albovillosus Megascelus nigricornis Rhopalia olivieri Rhopalia annulata Perissocerus dumonti Anomalomydas mackerrasi Cacatuopyga basifascia Cacatuopyga ruficornis Messiasia californica Phyllomydas parvulus Mydas clavatus Mydas xanthopterus Paramydas igniticornis Stratiomydas lividus Apiophora quadricinctata Apiophora paulseni Miltinus maculipennis Miltinus viduatus Miltinus stenogaster Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Ectyphus pinguis Ectyphus sp. nov. (Amboseli) Haplomydas crassipes Eremohaplomydas desertorum Eremohaplomydas sp. nov. (Kotwa) Eremomidas arabicus Eremomidas sp. nov. (Tiega) Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Pseudonomoneura micheneri Pseudonomoneura hirta Hessemydas parkeri Hessemydas seyrigi Nemomydas venosus Nemomydas brachyrhynchus gen. nov. + sp. nov. Gobabeb Leptomydas gruenbergi Nemomydas melanopogon Leptomydas corsicanus Leptomydas sardous Leptomydas notos sp. nov. Leptomydas rapti sp. nov. Leptomydas turcicus Eremomidas bek Leptomydas lusitanicus Mydaselpis ngurumani sp. nov. Vespiodes lepolditertii Vespiodes phaios sp. nov. Mydaselpis sp. nov. (Daan) Mydaselpis sp. nov. (Gamsberg) Namadytes maculiventris Namadytes prozeskyi Cephalocera imitata Cephalocera longirostris Cephalocera sp. (Papendorp) Neolaparopsis puncturatus Arenomydas caerulescens Arenomydas callosus Limata tenuicornis volaticus Neorhynchocephalus Nemestrinus nigrovillosus Eulonchus sp. (SixRivers) Prorates sp. (Escalante) Arenomydas namaquensis Nomoneura caffra Toxophora maxima Bombylius major Dipalta serpentina Apsilocephalaatra Hemigephyra longistyla Nomoneura paradoxa Phycus frommeri acanthophorite spurs absent Hoplistomerus nobilis Pilica formidolosa Laphria aktis bimaculatus Neolophonotus Emphysomera pallidapex Nomoneuroides natalensis Ommatius tibialis Machimus occidentalis Philonicus albiceps Stichopogoncinctus Lasiopogon punctum Diogmites grossus Halterorchis sp. nov. (Cape Recife) Ospriocerus aeacus Gonioscelis ventralis Damalis monochaetes Pegesimalluscylindrica Leptogaster laticornis Euscelidia pulchra Dioctria hyalipennis Holcocephala calva bilineata Apiocera (Pyrocera) Apiocera (Ripidosyrma) braunsi Apiocera (Pyrocera) aldrichi gen. nov. + sp. nov. (Panne) gen. nov. + sp. nov. (Ugab) Apiocera (Pyrocera) painteri Apiocera (Pyrocera) arena Apiocera (Pyrocera) obscura Apiocera (Anypenus)parkeri Apiocera (Anypenus)sp. (Wurarga) philippii Apiocera (Apiocera) sp. (Maranoa) gen. nov. + sp. nov. (Gobabeb) Apiocera (Apiocera) Apiocera (Apiocera) sp. (Stuart) Rhaphiomidas hasbroucki Rhaphiomidas parkeri proboscis vestigial Rhaphiomidas maehleri Tongamya miranda (Mandurah) Neorhaphiomidas sp. Neorhaphiomidas setosus Megascelus albovillosus Megascelus nigricornis Rhopalia olivieri Namibimydas sp. nov. (Agate) Nothomydas sp. nov. (Skerpioenkop) Rhopalia annulata Perissocerus dumonti Anomalomydas mackerrasi Cacatuopyga basifascia Cacatuopyga ruficornis Messiasia californica Afroleptomydas A consanguinensis Phyllomydas parvulus Mydas clavatus Mydas xanthopterus Paramydas igniticornis Stratiomydas lividus Apiophora quadricinctata Afroleptomydas A suffusipennis Apiophora paulseni Miltinus maculipennis Miltinus viduatus Miltinus stenogaster Heteromydas bicolor Afroleptomydas C mauricei Opomydas townsendi Parectyphus namibiensis Ectyphus pinguis (Amboseli) Ectyphus sp. crassipes Haplomydas nov. Eremohaplomydas desertorum Afroleptomydas A sp. (Clanwilliam) Eremohaplomydas sp. nov. (Kotwa) Eremomidas arabicus(Tiega) proboscis and acanthophorite spurs Eremomidas sp. nov. Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Afroleptomydas C sp. (Stampriet) Pseudonomoneura micheneri Pseudonomoneura Hessemydas seyrigihirta parkeri Hessemydas venosus Nemomydas brachyrhynchus Syllegomydas sp. nov. (Ruacana) Nemomydas gruenbergi Leptomydas Nemomydas corsicanus Leptomydas melanopogon Leptomydas sardous nov. Leptomydas notos sp. Syllegomydas N arnoldi Leptomydas rapti sp. Leptomydas turcicus nov. Eremomidas lusitanicus Leptomydas bek Mydaselpislepolditertii sp. nov. Vespiodes ngurumani Syllegomydas N dispar Vespiodes phaios sp. nov. Mydaselpis sp. nov. (Daan) Mydaselpis sp. nov. (Gamsberg) Namadytes maculiventris Namadytes prozeskyi Syllegomydas S palestinensis Cephalocera imitata Cephalocera longirostris Cephalocera sp. (Papendorp) Neolaparopsis puncturatus Arenomydas caerulescens Arenomydas callosus well-developed Syllegomydas N heothinos sp. nov. Arenomydascaffra Nomoneura namaquensis Nomoneura paradoxa Nomoneuroides nov. (Cape sp. natalensis Halterorchissp. nov. (Panne)Recife) gen. nov. + sp. nov. (Ugab) Syllegomydas S algiricus gen. nov. + gen. nov. + sp. nov. (Gobabeb) Namibimydassp. nov. (Skerpioenkop) Nothomydas sp. nov. (Agate) Afroleptomydas A consanguinensis Afroleptomydas A suffusipennis Afroleptomydas C sp. (Clanwilliam) Afroleptomydas A mauricei Afroleptomydas C sp. Syllegomydas sp. nov.(Stampriet) Syllegomydas N dispar(Ruacana) arnoldi Syllegomydas S elachys sp. nov. Syllegomydas N astrictus sp. nov. Syllegomydas N palestinensis Syllegomydas S Syllegomydas N heothinos Syllegomydas S algiricus sp. nov. Syllegomydas S astrictus sp. nov. Syllegomydas N elachys sp. nov.
  17. 17. .... character evolution 17 Rhaphiomidas hasbroucki Character evolution: met tibia Rhaphiomidas parkeri Rhaphiomidas maehleri Tongamya miranda Neorhaphiomidas sp. (Mandurah) Neorhaphiomidas setosus Megascelus albovillosus Megascelus nigricornis Rhopalia olivieri Rhopalia annulata Perissocerus dumonti Anomalomydas mackerrasi Cacatuopyga basifascia Cacatuopyga ruficornis Messiasia californica Phyllomydas parvulus Mydas clavatus Mydas xanthopterus Paramydas igniticornis Stratiomydas lividus Apiophora quadricinctata Apiophora paulseni Miltinus maculipennis Miltinus viduatus Miltinus stenogaster Heteromydas bicolor Opomydas townsendi Parectyphus namibiensis Ectyphus pinguis Ectyphus sp. nov. (Amboseli) Haplomydas crassipes Eremohaplomydas desertorum Eremohaplomydas sp. nov. (Kotwa) Eremomidas arabicus Eremomidas sp. nov. (Tiega) Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Pseudonomoneura micheneri Pseudonomoneura hirta Hessemydas parkeri Hessemydas seyrigi Nemomydas venosus Nemomydas brachyrhynchus Diochlistus cardinalis Leptomydas gruenbergi Nemomydas melanopogon Leptomydas corsicanus Leptomydas sardous Leptomydas notos sp. nov. Leptomydas rapti sp. nov. Leptomydas turcicus Eremomidas bek Leptomydas lusitanicus Mydaselpis ngurumani sp. nov. Vespiodes lepolditertii Vespiodes phaios sp. nov. Mydaselpis sp. nov. (Daan) Mydaselpis sp. nov. (Gamsberg) Namadytes maculiventris Namadytes prozeskyi Cephalocera imitata Cephalocera longirostris Cephalocera sp. (Papendorp) Neolaparopsis puncturatus Arenomydas caerulescens Arenomydas callosus Limata tenuicornis volaticus Neorhynchocephalus Nemestrinus nigrovillosus Eulonchus sp. (SixRivers) Prorates sp. (Escalante) Arenomydas namaquensis Nomoneura caffra Toxophora maxima Bombylius major Dipalta serpentina Apsilocephalaatra Hemigephyra longistyla Nomoneura paradoxa Phycus frommeri ventral keel absent Hoplistomerus nobilis Pilica formidolosa Laphria aktis bimaculatus Neolophonotus Emphysomera pallidapex Nomoneuroides natalensis Ommatius tibialis Machimus occidentalis Philonicus albiceps Stichopogoncinctus Lasiopogon punctum Diogmites grossus Halterorchis sp. nov. (Cape Recife) Ospriocerus aeacus Gonioscelis ventralis Damalis monochaetes Pegesimalluscylindrica Leptogaster laticornis Euscelidia pulchra Dioctria hyalipennis Holcocephala calva bilineata Apiocera (Pyrocera) Apiocera (Ripidosyrma) braunsi Apiocera (Pyrocera) aldrichi gen. nov. + sp. nov. (Panne) gen. nov. + sp. nov. (Ugab) Apiocera (Pyrocera) painteri Apiocera (Pyrocera) arena Apiocera (Pyrocera) obscura Apiocera (Anypenus)parkeri Apiocera (Anypenus)sp. (Wurarga) philippii Apiocera (Apiocera) sp. (Maranoa) gen. nov. + sp. nov. (Gobabeb) Apiocera (Apiocera) Apiocera (Apiocera) sp. (Stuart) Rhaphiomidas hasbroucki Rhaphiomidas parkeri keel at least proximally Rhaphiomidas maehleri Tongamya miranda (Mandurah) Neorhaphiomidas sp. Neorhaphiomidas setosus Megascelus albovillosus Megascelus nigricornis Rhopalia olivieri Namibimydas sp. nov. (Agate) Nothomydas sp. nov. (Skerpioenkop) Rhopalia annulata Perissocerus dumonti Anomalomydas mackerrasi Cacatuopyga basifascia Cacatuopyga ruficornis Messiasia californica Afroleptomydas A consanguinensis Phyllomydas parvulus Mydas clavatus Mydas xanthopterus Paramydas igniticornis Stratiomydas lividus Apiophora quadricinctata Afroleptomydas A suffusipennis Apiophora paulseni Miltinus maculipennis Miltinus viduatus Miltinus stenogaster Heteromydas bicolor Afroleptomydas C mauricei Opomydas townsendi Parectyphus namibiensis Ectyphus pinguis (Amboseli) Ectyphus sp. crassipes Haplomydas nov. Eremohaplomydas desertorum Afroleptomydas A sp. (Clanwilliam) Eremohaplomydas sp. nov. (Kotwa) Eremomidas arabicus(Tiega) keel well-developed with spur Eremomidas sp. nov. Diochlistus gracilis Mitrodetus dentitarsis Mitrodetus dimidiatus Afroleptomydas C sp. (Stampriet) Pseudonomoneura micheneri Pseudonomoneura Hessemydas seyrigihirta parkeri Hessemydas venosus Nemomydas brachyrhynchus Syllegomydas sp. nov. (Ruacana) Nemomydas gruenbergi Leptomydas Nemomydas corsicanus Leptomydas melanopogon Leptomydas sardous nov. Leptomydas notos sp. Syllegomydas N arnoldi Leptomydas rapti sp. Leptomydas turcicus nov. Eremomidas lusitanicus Leptomydas bek Mydaselpislepolditertii sp. nov. Vespiodes ngurumani Syllegomydas N dispar Vespiodes phaios sp. nov. Mydaselpis sp. nov. (Daan) Mydaselpis sp. nov. (Gamsberg) Namadytes maculiventris Namadytes prozeskyi Syllegomydas S palestinensis Cephalocera imitata Cephalocera longirostris Cephalocera sp. (Papendorp) Neolaparopsis puncturatus Arenomydas caerulescens Arenomydas callosus Syllegomydas N heothinos sp. nov. Arenomydascaffra Nomoneura namaquensis Nomoneura paradoxa Nomoneuroides nov. (Cape sp. natalensis Halterorchissp. nov. (Panne)Recife) gen. nov. + sp. nov. (Ugab) Syllegomydas S algiricus gen. nov. + gen. nov. + sp. nov. (Gobabeb) Namibimydassp. nov. (Skerpioenkop) Nothomydas sp. nov. (Agate) Afroleptomydas A consanguinensis Afroleptomydas A suffusipennis Afroleptomydas C sp. (Clanwilliam) Afroleptomydas A mauricei Afroleptomydas C sp. Syllegomydas sp. nov.(Stampriet) Syllegomydas N dispar(Ruacana) arnoldi Syllegomydas S elachys sp. nov. Syllegomydas N astrictus sp. nov. Syllegomydas N palestinensis Syllegomydas S Syllegomydas N heothinos Syllegomydas S algiricus sp. nov. Syllegomydas S astrictus sp. nov. Syllegomydas N elachys sp. nov.
  18. 18. .... summary + future 18 Summary ◊ Mydidae monophyletic ◊ (Mydidae + Apioceridae) + Asilidae ◊ Afrotropical Eremomidas in Leptomydinae? ◊ Eremohaplomydas + Haplomydas in Syllegomydinae? ◊ Chilean Paramydas to Mydinae – Apiophorinae monophyly ◊ Future Research: › additional exemplar species › internal head characters › inclusion of †Cretomydas, †Mydas miocenicus, †Protapiocera Phyllomydas parvulus © Lyn Atherton
  19. 19. 19 Acknowledgements ◊ Museum curators and collectors who made specimens available ◊ U.S. National Science Foundation (REVSYS DEB-0919333) ◊ I would be happy to study additional specimens from your collection! www.mydidae.tdvia.de

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