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Effect of taxon sampling in morphological phylogenetic analyses: insights from three taxa with different diversification rates

Apioceridae: sp. © J. Hort : 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 › 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 Neodiptera Orthorrhapha Stratiomyomorpha Asilidae Mydidae > 8,100 species Apioceridae Apystomyiidae > 15,500 species Eremoneura Syrphidae Cyclorrhapha I Evocoidae Acalyptratae II

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 ” ◊ 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 amboseli M Pseudorhopalia mirandai Ectyphus pinguis Ectyphus amboseli M Heteromydas bicolor Ectyphus pinguis Opomydas townsendi Heteromydas bicolor Paramydas igniticornis Opomydas townsendi Cacatuopyga basifascia Anomalomydas mackerrasi Cacatuopyga ruficornis M Anomalomydas australicus F Stratiomydas lividus Anomalomydas sp nov Simpson F Mydas clavatus Miltinus stenogaster Miltinus viduatus Mydas tricinctus Miltinus maculipennis Mydas xanthopterus Miltinus minutus Messiasia californica M Rhopalia annulata Messiasia painteri Rhopalia cincta Plyomydas peruviensis M Rhopalia olivieri Plyomydas sp nov Tandil Perissocerus dumonti F Dolichogaster brevicornis Perissocerus transcaspicus Phyllomydas bruesii Paramydas igniticornis Plyomydas peruviensis M Phyllomydas parvulus Plyomydas sp nov Tandil Anomalomydas australicus F Messiasia californica M Anomalomydas mackerrasi Messiasia painteri Anomalomydas sp nov Simpson F Mydas clavatus Pseudorhopalia mirandai Mydas tricinctus Miltinus sp. (Northern Territory) Mydas xanthopterus Rhopalia cincta Stratiomydas lividus Rhopalia annulata Cacatuopyga basifascia Rhopalia olivieri Cacatuopyga ruficornis M Dolichogaster brevicornis Perissocerus dumonti F Phyllomydas bruesii Perissocerus transcaspicus Phyllomydas parvulus Hessemydas parkeri M Eremomidas bek Hessemydas daugeroni Hessemydas daugeroni Hessemydas seyrigi Hessemydas parkeri M Nemomydas venosus Hessemydas seyrigi Nemomydas brachyrhynchus Leptomydas lusitanicus Nemomydas melanopogon Leptomydas notos Leptomydas lusitanicus Leptomydas rapti M Leptomydas sardous Eremomidas bek Leptomydas turcicus Leptomydas rapti M Nemomydas brachyrhynchus Leptomydas turcicus Nemomydas lamia Leptomydas corsicanus Nemomydas melanopogon Nemomydas gruenbergi M Nemomydas pantherinus Leptomydas sardous Nemomydas venosus Leptomydas notos Pseudonomoneura bajaensis Nemomydas pantherinus Hispanomydas hispanicus Mydaselpis ngurumani Pseudonomoneura hirta Neolaparopsis puncturatus Pseudonomoneura micheneri Nomoneura caffra Nemomydas lamia Nomoneura paradoxa Arenomydas partitus Nomoneuroides natalensis Arenomydas caerulescens Notosyllegomydas brincki Arenomydas callosus Syllegomydas N dispar Arenomydas namaquensis M Syllegomydas S algiricus Notosyllegomydas brincki Syllegomydas S palestinensis Cephalocera longirostris Syllegomydas S proximus Syllegomydas S vittatus Cephalocera sp Papendorp Syll gen nov sp nov minute proboscis F Cephalocera sp Sors Sors Pseudonomoneura hirta Cephalocera imitata M Pseudonomoneura micheneri Cephalocera sp Ratelfontein Namadytes cimbebasiensis Namadytes cimbebasiensis Namadytes vansoni Namadytes vansoni Eremomidas sp nov Tiega M Mydaselpis ngurumani Eremomidas arabicus Neolaparopsis puncturatus Halterorchis karooensis F Haplomydas crassipes Haplomydas crassipes Eremohaplomydas desertorum Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Lachnocorynus sp nov Kotwa M Nothomydas picketti Mydaselpis sp nov Daan Namibimydas psamminos Mydaselpis sp nov Gamsberg M Namibimydas stuckenbergi Vespiodes leopolditertii Arenomydas partitus Vespiodes phaios Arenomydas caerulescens Nomoneura caffra Arenomydas callosus Nomoneura paradoxa Arenomydas namaquensis M Mydaselpis sp nov Daan Nomoneuroides natalensis Mydaselpis sp nov Gamsberg M Afroleptomydas A suffusipennis Vespiodes leopolditertii Afroleptomydas A paganus Vespiodes phaios Afroleptomydas A inhacae Diochlistinae Argentina F Afroleptomydas A consanguinensis Mitrodetus nanoglossa M Afroleptomydas A opacicinctus Mitrodetus microglossa Afroleptomydas A sp Clanwilliam Mitrodetus dentitarsis Afroleptomydas A sp Obib Mitrodetus dimidiatus Afroleptomydas C sp Koeroegab Afroleptomydas A inhacae Afroleptomydas A opacicinctus Afroleptomydas A omeri Afroleptomydas C mauricei Afroleptomydas A rufithorax Afroleptomydas A omeri Afroleptomydas C mauricei Afroleptomydas A rufithorax Afroleptomydas C sp Stampriet length 1801 steps Cephalocera longirostris Afroleptomydas C sp Gobabeb Cephalocera sp Papendorp Syllegomydas S lineatus Cephalocera sp Sors Sors Hispanomydas hispanicus Cephalocera imitata M Syllegomydas N astrictus 771 trees Cephalocera sp Ratelfontein Syllegomydas N heothinos Syllegomydas S algiricus Syllegomydas S elachys Syllegomydas S palestinensis Syllegomydas S proximus Syllegomydas S vittatus Syllegomydas sp nov Ruacana Syll gen nov sp nov minute proboscis F Syllegomydas N arnoldi M Syllegomydas N dispar Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Pseudonomoneura bajaensis Eremomidas sp nov Tiega M Eremomidas arabicus monophyletic higher taxon Halterorchis karooensis F length 1839 steps Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Australian Diochlistinae 346 trees Mitrodetus dimidiatus .. a posteriori taxon exclusion 14 Mydidae – Female oviposition strategy Mydidae all Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis Stratiomydas lividus Mydas clavatus / ovipositor dorsal, Cacatuopyga basifascia Mydas tricinctus Mydas xanthopterus Messiasia californica / ovipositor dorsal, Leptomydas lusitanicus Messiasia painteri Plyomydas peruviensis Plyomydas sp nov Tandil Dolichogaster brevicornis Phyllomydas bruesii / ovipositor dorsal, Messiasia painteri Phyllomydas parvulus

= tergite 10 = tergite 9 = tergite 8 = tergite 9+10 scale line = 1 mm / ovipositor dorsal, Phyllomydas parvulus .. a posteriori taxon exclusion 15 Mydidae – Female oviposition strategy

Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis Stratiomydas lividus Mydas clavatus / ovipositor dorsal, Cacatuopyga basifascia Mydas tricinctus Mydas xanthopterus Messiasia californica / ovipositor dorsal, Leptomydas lusitanicus Messiasia painteri Plyomydas peruviensis Plyomydas sp nov Tandil Dolichogaster brevicornis Phyllomydas bruesii / ovipositor dorsal, Messiasia painteri Phyllomydas parvulus Mydidae 2 Mydidae 5 Paramydas igniticornis Paramydas igniticornis Plyomydas peruviensis Plyomydas peruviensis Plyomydas sp nov Tandil Plyomydas sp nov Tandil Messiasia californica Messiasia californica Messiasia painteri Messiasia painteri / ovipositor dorsal, Phyllomydas parvulus Mydas clavatus Mydas clavatus Mydas xanthopterus Mydas tricinctus Stratiomydas lividus Mydas xanthopterus Cacatuopyga basifascia Stratiomydas lividus Cacatuopyga ruficornis Cacatuopyga basifascia Dolichogaster brevicornis Cacatuopyga ruficornis Phyllomydas bruesii Dolichogaster brevicornis Phyllomydas parvulus Phyllomydas bruesii Phyllomydas parvulus ... a posteriori taxon inclusion 16 Asilidae - Bathypogoninae

Perasis transvaalensis Zabrops tagax Protometer sp. (El Tuparro) Psilocurus modestus Laphystia tolandi Hoplistomerus nobilis Trichardis testacea Laxenecera albicincta Nusa infumata Smeryngolaphria numitor Orthogonis scapularis Goneccalypsis argenteoviridis Loewinella nigripes Aphestia annulipes Atomosia puella Cerotainia albipilosa Maira aenea Lampria clavipes Laphria flava Choerades bella Lamyra gulo Stiphrolamyra angularis Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris Proagonistes praeceps Blepharotes splendidissimus Lycomya germainii Cophinopoda chinensis Michotamia aurata Emphysomera pallidapex Afroestricus chiastoneurus Ommatius tibialis Megaphorus pulchrus Promachus amastrus Apoclea algira Philodicus tenuipes Pogonioefferia pogonias Dasophrys crenulatus Eichoichemus pyrrhomystax Myaptex brachyptera Neolophonotus bimaculatus Nomomyia murina Proctacanthus philadelphicus Lochmorhynchus albicans Machimus occidentalis Dysmachus trigonus Tolmerus atricapillus Neomochtherus pallipes Afromochtherus mendax Philonicus albiceps Clephydroneura sundaica Congomochtherus inachus Heligmonevra laevis Zosteria rosevillensis Asilus crabroniformis Satanas gigas Colepia rufiventris Neoitamus cyanurus Threnia carbonaria Bathypogon nigrinus Phellus olgae Obelophorus terebratus Gonioscelis ventralis Stenopogon sabaudus Bathypogon sp. Ospriocerus aeacus Scleropogon subulatus Prolepsis tristis Microstylum sp. (Karkloof) Daspletis stenoura Connomyia varipennis Plesiomma sp. (Guanacaste) Ancylorhynchus fulvicollis Scylaticus costalis Ablautus coquilletti Acnephalum cylindricum Sisyrnodytes sp. (Gamka) Willistonina bilineata Trichoura sp. (Tierberg) Coleomyia setigera Lestomyia fraudiger Dasypogon diadema Saropogon dispar Thereutria amaraca Archilestris magnificus Blepharepium cajennensis Neodiogmites melanogaster Diogmites grossus Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Tillobroma punctipennis Creolestes nigribarbis Hypenetes stigmatias Lycostommyia albifacies Cyrtopogon rattus Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Chrysopogon pilosifacies Codula limbipennis Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis Leptarthrus brevirostris Holopogon priscus Afroholopogon peregrinus Dasycyrton gibbosus Oligopogon sp. (Cape Recife) Lasiopogon cinctus Townsendia albomacula Stichopogon punctum Lissoteles aquilonius Broticosia paramonovi Dicolonus simplex Echthodopa pubera Myelaphus melas Dioctria atricapillus Nannodioctria sp. (formicaphaga) Eudioctria albius Damalis monochaetes Rhipidocephala sp. (Harold Johnson) Trigonomima sp. (anamaliensis) Holcocephala abdominalis Acronyches maya length 2760 steps Beameromyia bifida Euscelidia pulchra Tipulogaster glabrata Leptogaster cylindrica Bathypogon 460 trees Lasiocnemus lugens Bathypogon nigrinus Phellinae ... a posteriori taxon inclusion 17 Asilidae - Bathypogoninae

Perasis transvaalensis Zabrops tagax Perasis transvaalensis Protometer sp. (El Tuparro) Zabrops tagax Psilocurus modestus Protometer sp. (El Tuparro) Laphystia aegyptiaca Psilocurus modestus Laphystia tolandi Laphystia aegyptiaca Hoplistomerus nobilis Laphystia tolandi Trichardis testacea Hoplistomerus nobilis Laxenecera albicincta Trichardis testacea Nusa infumata Laxenecera albicincta Smeryngolaphria numitor Nusa infumata Orthogonis scapularis Smeryngolaphria numitor Goneccalypsis argenteoviridis Orthogonis scapularis Loewinella nigripes Goneccalypsis argenteoviridis Aphestia annulipes Loewinella nigripes Atomosia puella Aphestia annulipes Cerotainia albipilosa Atomosia puella Maira aenea Cerotainia albipilosa Lampria clavipes Maira aenea Choerades bella Lampria clavipes Laphria flava Choerades bella Laphria aktis Laphria flava Lamyra gulo Laphria aktis Stiphrolamyra angularis Lamyra gulo Andrenosoma cyrtoxys Stiphrolamyra angularis Hyperechia nigripennis Andrenosoma cyrtoxys Pilica formidolosa Hyperechia nigripennis Dasyllis funebris Pilica formidolosa Proagonistes praeceps Dasyllis funebris Cophinopoda chinensis Proagonistes praeceps Michotamia aurata Cophinopoda chinensis Ommatius incurvatus Michotamia aurata Afroestricus chiastoneurus Ommatius incurvatus Ommatius tibialis Afroestricus chiastoneurus Emphysomera conopsoides Ommatius tibialis Emphysomera pallidapex Emphysomera conopsoides Blepharotes splendidissimus Emphysomera pallidapex Lycomya germainii Blepharotes splendidissimus Pogonioefferia pogonias Lycomya germainii Proctacanthus philadelphicus Pogonioefferia pogonias Dasophrys crenulatus Proctacanthus philadelphicus Eichoichemus pyrrhomystax Dasophrys crenulatus Lochmorhynchus albicans Eichoichemus pyrrhomystax Myaptex brachyptera Lochmorhynchus albicans Neolophonotus bimaculatus Myaptex brachyptera Nomomyia murina Neolophonotus bimaculatus Satanas gigas Nomomyia murina Machimus chrysitis Satanas gigas Machimus occidentalis Machimus chrysitis Dysmachus trigonus Machimus occidentalis Philonicus albiceps Dysmachus trigonus Congomochtherus inachus Philonicus albiceps Heligmonevra laevis Congomochtherus inachus Tolmerus atricapillus Heligmonevra laevis Zosteria rosevillensis Tolmerus atricapillus Asilus crabroniformis Zosteria rosevillensis Asilus sericeus Asilus crabroniformis Neomochtherus pallipes Asilus sericeus Afromochtherus mendax Neomochtherus pallipes Clephydroneura sundaica Afromochtherus mendax Clephydroneura sp. (Kepong) Clephydroneura sundaica Colepia rufiventris Clephydroneura sp. (Kepong) Neoitamus cyanurus Colepia rufiventris Threnia carbonaria Neoitamus cyanurus Megaphorus pulchrus Threnia carbonaria Promachus amastrus Megaphorus pulchrus Apoclea algira Promachus amastrus Philodicus tenuipes Apoclea algira Bathypogon nigrinus Philodicus tenuipes Bathypogon rubellus Bathypogon nigrinus Phellus olgae Bathypogon rubellus Obelophorus terebratus Microstylum sp. (Karkloof) Tillobroma punctipennis Prolepsis tristis Hypenetes stigmatias Scylaticus costalis Lycostommyia albifacies Tillobroma punctipennis Coleomyia setigera Willistonina bilineata Ancylorhynchus fulvicollis Sisyrnodytes sp. (Gamka) Scylaticus costalis Stenopogon rufibarbis Connomyia varipennis Ospriocerus aeacus Prolepsis tristis Ablautus coquilletti Creolestes nigribarbis Creolestes nigribarbis Microstylum sp. (Karkloof) Rhabdogaster pedion Daspletis stenoura Archilestris magnificus Plesiomma sp. (Guanacaste) Hypenetes stigmatias Ospriocerus aeacus Phellus olgae Scleropogon subulatus Acnephalum cylindricum Stenopogon sabaudus Ancylorhynchus fulvicollis Stenopogon rufibarbis Obelophorus terebratus Gonioscelis ventralis Plesiomma sp. (Guanacaste) Lestomyia fraudiger Daspletis stenoura Dasypogon diadema Lestomyia fraudiger Saropogon dispar Dasypogon diadema Saropogon luteus Saropogon dispar Thereutria amaraca Saropogon luteus Archilestris magnificus Thereutria amaraca Neodiogmites melanogaster Nannocyrtopogon lestomyiformis Diogmites grossus Neodiogmites melanogaster Blepharepium cajennensis Diogmites grossus Molobratia teutonus Blepharepium cajennensis Pegesimallus laticornis Molobratia teutonus Senobasis corsair Pegesimallus laticornis Cyrtophrys attenuatus Senobasis corsair Deromyia fuscipennis Cyrtophrys attenuatus Megapoda labiata Deromyia fuscipennis Pseudorus distendens Megapoda labiata Acnephalum cylindricum Pseudorus distendens Sisyrnodytes sp. (Gamka) Stenopogon sabaudus Ablautus coquilletti Scleropogon subulatus Willistonina bilineata Gonioscelis ventralis Trichoura sp. (Tierberg) Coleomyia setigera Oligopogon sp. (Cape Recife) Lycostommyia albifacies Broticosia paramonovi Trichoura sp. (Tierberg) Dicolonus simplex Broticosia paramonovi Echthodopa pubera Dicolonus simplex Myelaphus melas Echthodopa pubera Nannodioctria sp. (formicaphaga) Bathypogon sp. (Lake King) Eudioctria albius Nannodioctria sp. (formicaphaga) Dioctria atricapillus Eudioctria albius Dioctria hyalipennis Dioctria atricapillus Dioctria rufipes Dioctria hyalipennis Acronyches maya Dioctria rufipes Beameromyia bifida Acronyches maya Euscelidia pulchra Beameromyia bifida Tipulogaster glabrata Euscelidia pulchra Leptogaster arida Tipulogaster glabrata Leptogaster cylindrica Leptogaster arida Lasiocnemus lugens Leptogaster cylindrica Damalis sp. (Palatupana) Lasiocnemus lugens Damalis monochaetes Damalis monochaetes Damalis annulata Lissoteles aquilonius Rhipidocephala sp. (Harold Johnson) Townsendia albomacula Trigonomima sp. (anamaliensis) Holcocephala abdominalis Holcocephala abdominalis Damalis annulata Holcocephala calva Damalis sp. (Palatupana) Lasiopogon aldrichii Trigonomima sp. (anamaliensis) Lasiopogon cinctus Stichopogon punctum Townsendia albomacula Stichopogon trifasciatus Stichopogon elegantulus Lasiopogon cinctus Stichopogon trifasciatus Oligopogon sp. (Cape Recife) Stichopogon punctum Stichopogon elegantulus Lissoteles aquilonius Connomyia varipennis Cyrtopogon lateralis Lasiopogon aldrichii Cyrtopogon rattus Ceraturgus fasciatus Afroholopogon peregrinus Myelaphus melas Dasycyrton gibbosus Cyrtopogon lateralis Leptarthrus brevirostris Cyrtopogon rattus Holopogon priscus Leptarthrus brevirostris Chrysopogon pilosifacies Dasycyrton gibbosus Codula limbipennis Chrysopogon pilosifacies Austrosaropogon nigrinus Codula limbipennis Brachyrhopala ruficornis Austrosaropogon nigrinus Cabasa pulchella Brachyrhopala ruficornis Ceraturgus fasciatus Cabasa pulchella length 2763 steps Nannocyrtopogon lestomyiformis Bathypogon robustus Nicocles politus length 2771 steps Holopogon priscus Cophura brevicornis Nicocles politus Rhabdogaster pedion Cophura brevicornis Heteropogon manicatus Metapogon punctipennis 391 trees Metapogon punctipennis Afroholopogon peregrinus Bathypogon 443 trees Heteropogon manicatus Phellinae .... conclusion 18 Conclusion ◊ strive to include as many taxa as feasible ◊ Apioceridae biogeography failed test ◊ Mydidae relationships › Chilean Diochlistinae group “already” with only 1 generic representative › ovipositor evolution › secondary signal emerges ◊ Asilidae: Bathypogoninae › inclusion of 1 additional taxon corroborates status › inclusion of 3 additional taxa causes “problems” ..... acknowledgements 19 Acknowledgements ◊ museum curators – collectors – photographers ◊ funding sources: › U.S. National Science Foundation REVSYS DEB-0919333 › John D. and Catherine T. MacArthur Foundation (Biodiversity Synthesis Group of the Encyclopedia of Life) › Field Museum of Natural History

Smithsonian National Museum of Natural History