Effect of Taxon Sampling in Morphological Phylogenetic Analyses: Insights from Three Fly Taxa with Different Diversification Rates
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1 Effect of taxon sampling in morphological phylogenetic analyses: insights from three fly taxa with different diversification rates Apioceridae: Apiocera sp. © J. Hort Asilidae: Prytanomyia albida Mydidae: Miltinus minutus © J. Hort Torsten Dikow National Museum of Natural History Smithsonian Washington, DC, U.S.A. National Museum of Natural History . introduction 2 Taxon sampling in cladistic analyses ◊ integral and important component -> data + taxa ◊ too often data have priority while taxa are “neglected” ◊ cladistic analyses of insects › morphology – based on museum specimens › traditional molecular – based on more recently collected material › genome molecular – based on newly collected material (RNAlater, DNAgard) ◊ a posteriori taxon exclusion › Was the addition of taxa “necessary” to obtain the same hypothesis of relationships? ◊ a posteriori taxon inclusion › Would the addition of taxa corroborate the postulated hypothesis of relationships? . introduction 3 Diptera = 159,294 valid species (Pape et al. 2011) Deuterophlebiidae Nymphomyiidae Tipulomorpha Ptychopteridae Psychodomorpha Culicomorpha Perissommatidae Bibionomorpha Nemestrinidae Neodiptera Xylophagidae Tabanomorpha Acroceridae Orthorrhapha Hilarimorphidae Bombyliidae Stratiomyomorpha Asilidae Brachycera Asiloidea Mydidae > 8,100 species Empidoidea Apioceridae Apystomyiidae > 15,500 species Eremoneura Therevidae Platypezoidea Scenopinidae Syrphidae Cyclorrhapha Pipunculidae Apsilocephalidae Acalyptratae I Evocoidae Acalyptratae II Schizophora Ephydroidea Calyptratae phylogeny from Wiegmann et al. 2011 (simplified), images Apiocera alleni, Leptomydas lusitanicus, Cyrtopogon lutatius, Echthodopa formosa © G. Ballmer, V. Jacinto, M. Thomas . introduction 4 Apioceridae – “flower-loving flies” ◊ adult flies flower-feeders ◊ size = 15 – 30 mm ◊ 138 species in 1 genus (4 subgenera) ◊ occur only in › Argentina and Chile –> 4 species › Australia –> 67 species Apiocera (Apiocera) pica © J. Hort › South Africa –> 3 species › western North America (Canada, Mexico, USA) –> 64 species ◊ rarely collected Apiocera (Apiocera) sp. © J. Hort . introduction 5 Mydidae – “mydas flies” ◊ adult flies flower-feeders › some adults do not feed at all ◊ size = 9 – 60 mm ◊ 473 species in 66 genera ◊ world-wide primarily in arid regions › southern Africa most speciose Opomydas townsendi ◊ rarely collected Eremomidas arabicus © D. Gardner . introduction 6 Asilidae – “robber flies” or “assassin flies” ◊ adult flies predatory ◊ size = 5 – 60 mm ◊ > 7,500 species in 540 genera ◊ speciose world-wide › arid & semi-arid environments › tropical environments Scleropogon duncani ◊ regularly collected by specialists and non-specialists alike Ceraturgus fasciatus © M. Thomas . introduction 7 Fossils ◊ Crato Formation, Brazil › Cretaceous: Aptian – Albian › ≈ 112 myo › Asilidae – † Araripogon axelrodi › Mydidae – † Cretomydas santanensis Asilidae: † Araripogon axelrodi Mydidae: † Cretomydas santanensis 8 Overview ◊ Methods ◊ a posteriori taxon exclusion › Apioceridae biogeography › Mydidae relationships + female oviposition ◊ a posteriori taxon inclusion/addition › Asilidae: Bathypogoninae status Trichardis picta Prytanomyia albida 9 Methods taxon # known # species in analysis # known # genera analysis species genera Apioceridae all 138 30 21.7 % 4 4 100 % Apioceridae 1 4 2.9 % Apioceridae 2 8 5.8 % Apioceridae 5 14 10.1 % Mydidae all 473 135 28.4 % 66 51 77.2 % Mydidae 1 49 10.4 % Mydidae 2 80 16.9 % Mydidae 5 117 24.7 % Asilidae > 7,500 158 2.1 % 540 140 25.9 % Asilidae 2 159 2.1 % Asilidae 4 161 2.1 % ◊ TNT 1.1 (January 2013) Goloboff et al. 2008 ◊ exhaustive search (Mac mini 2.3 GHz quad-core Intel Core i7, 16 GB RAM) › mult1000; › xmult=rss hits 20 fuse 2 drift 3 ratchet 10 multiply consense 10; .. a posteriori taxon exclusion 10 Apioceridae – Yeates & Irwin 1996 Apioceridae Apiocera (Pyrocera) western North America area relationships Apiocera (Ripidosyrma) South Africa Apiocera (Anypenus) Argentina / Chile Apiocera (Apiocera) Australia .. a posteriori taxon exclusion 11 Apioceridae Apioceridae 1 Apioceridae 2 Apioceridae 5 Apioceridae all Apiocera An philippii Apiocera An obscura Apiocera An obscura Apiocera An obscura Apiocera Ap pica Apiocera An philippii Apiocera An philippii Apiocera An philippii Apiocera P arena Apiocera Ap minor Apiocera Ap tonnoiri Apiocera Ap helenae Apiocera R braunsi Apiocera Ap pica Apiocera R alastor M Apiocera Ap imminuta Apiocera P arena Apiocera R braunsi Apiocera P bilineata Apiocera Ap hardyi Apiocera Ap tonnoiri Apiocera R alastor M Apiocera Ap maritima Apiocera R alastor M Apiocera R braunsi Apiocera Ap minor Apiocera R braunsi Apiocera Ap pica Apiocera Ap goerlingi Apiocera P acuticauda Apiocera Ap asilica Apiocera P arena Apiocera Ap brevicornis Apiocera P parkeri Apiocera P bilineata Apiocera Ap sp Stuart M Apiocera P mexicana M Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani Apiocera P bilineata Apiocera P mexicana M Apiocera P acuticauda 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 length 1787 steps length 1807 steps length 1819 steps length 1839 steps 306 trees 376 trees 345 trees 346 trees monophyletic subgenus .. a posteriori taxon exclusion 12 Rhaphiomidas_maehleri Mydidae Rhaphiomidas_parkeri Mydidae 2 Tongamya_miranda Megascelus_albovillosus Mydidae 1 Megascelus_melanoproctus Neorhaphiomidas_hardyi Rhaphiomidas maehleri Neorhaphiomidas_sp_Mandurah Megascelus albovillosus Cretomydas_santanensis Diochlistus_gracilis Neorhaphiomidas hardyi Diochlistus_melleipennis Tongamya miranda Anomalomydas_mackerrasi Diochlistus gracilis Anomalomydas_australicus_F Cretomydas santanensis Midacritus_stuardoanus_M Midacritus_wagenknechti Anomalomydas mackerrasi Apiophora_paulseni Midacritus wagenknechti Apiophora_quadricinctata Paramydas igniticornis Miltinus_minutus Cacatuopyga basifascia Miltinus_viduatus Ectyphus_armipes Plyomydas peruviensis M Ectyphus_pinguis Dolichogaster brevicornis Parectyphus_namibiensis Messiasia painteri Heteromydas_bicolor Opomydas_townsendi Mydas clavatus Paramydas_igniticornis Phyllomydas parvulus Plyomydas_peruviensis_M Stratiomydas lividus Plyomydas_sp_nov_Tandil Pseudorhopalia mirandai Messiasia_californica_M Messiasia_painteri Perissocerus transcaspicus Mydas_clavatus Rhopalia olivieri Mydas_xanthopterus Apiophora paulseni Stratiomydas_lividus Miltinus viduatus Cacatuopyga_basifascia Cacatuopyga_ruficornis_M Ectyphus pinguis Dolichogaster_brevicornis Parectyphus namibiensis Phyllomydas_bruesii Heteromydas bicolor Phyllomydas_parvulus Opomydas townsendi Pseudorhopalia_mirandai Rhopalia_annulata Eremomidas bek Rhopalia_olivieri Hessemydas daugeroni Perissocerus_dumonti_F Leptomydas lusitanicus Perissocerus_transcaspicus Nemomydas venosus Nemomydas_brachyrhynchus Nemomydas_venosus Afroleptomydas A rufithorax Eremomidas_bek Arenomydas caerulescens Leptomydas_lusitanicus Cephalocera longirostris Hessemydas_daugeroni Eremohaplomydas desertorum Hessemydas_seyrigi Nomoneura_caffra Haplomydas crassipes Nomoneura_paradoxa Lachnocorynus sp nov Kotwa M Nomoneuroides_natalensis Halterorchis karooensis F Namadytes_cimbebasiensis Namadytes vansoni Namadytes_vansoni Afroleptomydas_A_rufithorax Neolaparopsis puncturatus Afroleptomydas_C_mauricei Nomoneura caffra Hispanomydas_hispanicus Nomoneuroides natalensis Syllegomydas_S_algiricus Notosyllegomydas brincki Syllegomydas_S_vittatus Neolaparopsis_puncturatus Hispanomydas hispanicus Haplomydas_crassipes Syllegomydas S algiricus Eremohaplomydas_desertorum Mydaselpis sp nov Daan Lachnocorynus_sp_nov_Kotwa_M Vespiodes phaios Mydaselpis_sp_nov_Daan Vespiodes_leopolditertii Pseudonomoneura hirta Vespiodes_phaios length 1566 steps Mitrodetus dentitarsis Mydaselpis_ngurumani Namibimydas stuckenbergi Arenomydas_caerulescens Nothomydas picketti Arenomydas_callosus 589 trees Cephalocera_longirostris Cephalocera_sp_Papendorp Notosyllegomydas_brincki Eremomidas_arabicus Halterorchis_karooensis_F Leptomydas_sardous Pseudonomoneura_hirta Pseudonomoneura_micheneri Mitrodetus_dentitarsis monophyletic higher taxon Mitrodetus_dimidiatus length 1673 steps Nothomydas_picketti Namibimydas_psamminos Australian Diochlistinae 335 trees Namibimydas_stuckenbergi .. a posteriori taxon exclusion 13 Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Mydidae Tongamya miranda Megascelus albovillosus Mydidae all Megascelus melanoproctus Megascelus nigricornis Rhaphiomidas hasbroucki Neorhaphiomidas sp Mingenew M Rhaphiomidas maehleri Rhaphiomidas parkeri Neorhaphiomidas sp Yanchep Tongamya miranda Neorhaphiomidas hardyi Mydidae 5 Neorhaphiomidas sp Mandurah Megascelus albovillosus Megascelus melanoproctus Cretomydas santanensis Megascelus nigricornis Diochlistus auripennis M Neorhaphiomidas sp Mingenew M Diochlistus gracilis Neorhaphiomidas sp Yanchep Diochlistus melleipennis Neorhaphiomidas hardyi Midacritus stuardoanus M Neorhaphiomidas sp Mandurah Midacritus wagenknechti Cretomydas santanensis Diochlistus auripennis M Apiophora paulseni Diochlistus gracilis Apiophora quadricinctata Diochlistus melleipennis Miltinus stenogaster Apiophora paulseni Miltinus viduatus Apiophora quadricinctata Miltinus maculipennis Midacritus stuardoanus M Miltinus minutus Midacritus wagenknechti Ectyphus armipes Miltinus sp. (Northern Territory) Parectyphus namibiensis Ectyphus armipes Parectyphus namibiensis Ectyphus