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Effect of taxon sampling in morphological phylogenetic analyses

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Torsten Dikow
Torsten Dikow
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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 Washington, DC Smithsonian National Museum of Natural History
. introduction 2 Taxon sampling in cladistic studies ◊ integral and important component –> data + taxa ◊ too often data have priority while taxa are “neglected” › morphology – based on “unlimited” number of taxa in natural history collections › traditional molecular – based on more recently collected, limited number of taxa (95% ethanol) › genome molecular – based on newly collected, very limited number of taxa (RNAlater, DNAgard) ◊ taxon sampling examined in two morphological studies –> exclude or include?
. introduction 3 Taxon sampling in cladistic studies ◊ a posteriori taxon exclusion › Was the addition of taxa “necessary” to obtain the same hypothesis of relationships? › Does one need to include more than one species per genus? › example: two small families with dense generic sampling ◊ a posteriori taxon inclusion › Would the addition of taxa corroborate the postulated hypothesis of relationships? › Does the inclusion of additional species of a genus alter its position? › example: a very diverse group with limited generic sampling
. introduction 4 Taxa ◊ Apioceridae ◊ Mydidae › 143 species › rarely collected › Australia, Chile, western North America, South Africa Apiocera (Apiocera) pica © J. Hort ◊ Asilidae › 473 species › rarely collected › world-wide primarily in arid regions Eremomidas arabicus © D. Gardner › > 7,500 species › collected by specialists and non-specialists alike › speciose world-wide Scleropogon duncani
. introduction 5 Minimum ages + relationships ◊ Crato Formation, Brazil › Cretaceous: Aptian – Albian › ≈ 112 myo › Asilidae: † Araripogon axelrodi › Mydidae: † Cretomydas santanensis Asilidae: † Araripogon axelrodi Asilidae Mydidae Apioceridae Mydidae: † Cretomydas santanensis
6 Overview ◊ Methods + data-sets ◊ a posteriori taxon exclusion › Apioceridae biogeography › Mydidae female ovipositor morphology ◊ a posteriori taxon inclusion › Asilidae: Bathypogoninae position Trichardis picta Prytanomyia albida
7 Methods + data-sets ◊ Mesquite 2.75 ◊ TNT 1.1 (January 2013) ◊ 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; taxon Apioceridae all Apioceridae 1 Apioceridae 2 Apioceridae 5 Mydidae all Mydidae 1 Mydidae 2 Mydidae 5 Asilidae all Bathypogon 2 Bathypogon 4 # species in analysis 30 21.7 % 4 2.9 % 8 5.8 % 14 10.1 % 135 28.4 % 49 10.4 % 80 16.9 % 117 24.7 % 158 2.1 % 159 2.1 % 161 2.1 % # genera in analysis 4 100 % 51 77.2 % 140 25.9 %
.. a posteriori taxon exclusion – Apioceridae Apioceridae all 8 Apioceridae 1 Apiocera An obscura Apiocera An philippii Apiocera An philippii Apiocera Ap pica Apiocera Ap helenae Apiocera P arena Apiocera Ap imminuta Apiocera R braunsi Apiocera Ap tonnoiri Apiocera R alastor M length 1787 steps Apiocera R braunsi Apiocera Ap goerlingi 306 trees Apiocera Ap asilica strict consensus shown Apiocera Ap brevicornis Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani 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 Apioceridae 2 Apiocera An obscura Apiocera An philippii Apiocera Ap minor Apiocera Ap pica Apiocera P arena Apiocera P bilineata Apiocera R alastor M Apiocera R braunsi Afrotropical length 1807 steps 376 trees strict consensus shown Nearctic + Neotropical (Mexico: Veracruz) length 1839 steps 346 trees strict consensus shown monophyletic subgenus Chilean Apiocera = Bremer ≥ 8 = Bremer 3 – 7 = Bremer 1 – 2 Apioceridae 5 Apiocera An obscura Apiocera An philippii Apiocera Ap tonnoiri Apiocera R alastor M Apiocera R braunsi Apiocera Ap hardyi Apiocera Ap maritima Apiocera Ap minor Apiocera Ap pica Apiocera P acuticauda Apiocera P arena Apiocera P parkeri Apiocera P bilineata Apiocera P mexicana M length 1819 steps 345 trees strict consensus shown
.. a posteriori taxon exclusion – Apioceridae 9 Apioceridae – Yeates & Irwin 1996 Apioceridae Apiocera (Pyrocera) Apiocera (Ripidosyrma) Apiocera (Anypenus) Apiocera (Apiocera) Yeates & Irwin 1996 western North America South Africa Argentina / Chile Australia area relationships
.. a posteriori taxon exclusion – Apioceridae 10 Apioceridae – current hypothesis Apioceridae Apiocera (Pyrocera) Apiocera (Pyrocera) Apiocera (Ripidosyrma) Apiocera (Ripidosyrma) Apiocera (Anypenus) western North America South Africa Argentina / Chile Apiocera (Anypenus) 1 Apiocera (Anypenus) 2 Apiocera (Apiocera) Apiocera (Apiocera) 1 Apiocera (Apiocera) 2 Apiocera (Apiocera) ... Yeates & Irwin 1996 current hypothesis Australia area relationships
... a posteriori taxon exclusion – Mydidae Mydidae 1 11 Mydidae 2 Rhaphiomidas maehleri Megascelus albovillosus Neorhaphiomidas hardyi Tongamya miranda Diochlistus gracilis Cretomydas santanensis Anomalomydas mackerrasi Midacritus wagenknechti Paramydas igniticornis Cacatuopyga basifascia Plyomydas peruviensis M Dolichogaster brevicornis Messiasia painteri Mydas clavatus Phyllomydas parvulus Stratiomydas lividus Pseudorhopalia mirandai Perissocerus transcaspicus Rhopalia olivieri Apiophora paulseni Miltinus viduatus Ectyphus pinguis Parectyphus namibiensis Heteromydas bicolor Opomydas townsendi Eremomidas bek Hessemydas daugeroni Leptomydas lusitanicus Nemomydas venosus Afroleptomydas A rufithorax Arenomydas caerulescens Cephalocera longirostris Eremohaplomydas desertorum Haplomydas crassipes Lachnocorynus sp nov Kotwa M Halterorchis karooensis F Namadytes vansoni Neolaparopsis puncturatus Nomoneura caffra Nomoneuroides natalensis Notosyllegomydas brincki Hispanomydas hispanicus Syllegomydas S algiricus Mydaselpis sp nov Daan Vespiodes phaios Pseudonomoneura hirta Mitrodetus dentitarsis Namibimydas stuckenbergi Nothomydas picketti length 1566 steps 589 trees strict consensus shown monophyletic higher taxon = Bremer ≥ 8 = Bremer 3 – 7 = Bremer 1 – 2 Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Apiophora paulseni Apiophora quadricinctata Miltinus minutus Miltinus viduatus Ectyphus armipes Ectyphus pinguis Parectyphus namibiensis Heteromydas bicolor Opomydas townsendi Paramydas igniticornis Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Mydas clavatus Mydas xanthopterus Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruficornis M Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia annulata Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Nemomydas brachyrhynchus Nemomydas venosus Eremomidas bek Leptomydas lusitanicus Hessemydas daugeroni Hessemydas seyrigi Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Namadytes cimbebasiensis Namadytes vansoni Afroleptomydas A rufithorax Afroleptomydas C mauricei Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S vittatus Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Cephalocera longirostris Cephalocera sp Papendorp Notosyllegomydas brincki Eremomidas arabicus Halterorchis karooensis F Leptomydas sardous Pseudonomoneura hirta Pseudonomoneura micheneri Mitrodetus dentitarsis Mitrodetus dimidiatus Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi length 1673 steps 335 trees strict consensus shown
... a posteriori taxon exclusion – Mydidae Mydidae 5 12 Mydidae all Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Apiophora paulseni Apiophora quadricinctata Midacritus stuardoanus M Midacritus wagenknechti Miltinus sp. (Northern Territory) Ectyphus armipes Parectyphus namibiensis Pseudorhopalia mirandai Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Anomalomydas mackerrasi Anomalomydas australicus F Anomalomydas sp nov Simpson F Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Paramydas igniticornis Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Mydas clavatus Mydas tricinctus Mydas xanthopterus Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruficornis M Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Eremomidas bek Hessemydas daugeroni Hessemydas parkeri M Hessemydas seyrigi Leptomydas lusitanicus Leptomydas notos Leptomydas rapti M Leptomydas sardous Leptomydas turcicus Nemomydas brachyrhynchus Nemomydas lamia Nemomydas melanopogon Nemomydas pantherinus Nemomydas venosus Pseudonomoneura bajaensis Hispanomydas hispanicus Mydaselpis ngurumani Neolaparopsis puncturatus Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Notosyllegomydas brincki Syllegomydas N dispar Syllegomydas S algiricus Syllegomydas S palestinensis Syllegomydas S proximus Syllegomydas S vittatus Syll gen nov sp nov minute proboscis F Pseudonomoneura hirta Pseudonomoneura micheneri Namadytes cimbebasiensis Namadytes vansoni Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus Afroleptomydas A inhacae Afroleptomydas A opacicinctus Afroleptomydas C mauricei Afroleptomydas A omeri Afroleptomydas A rufithorax Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata M Cephalocera sp Ratelfontein length 1801 steps 771 trees strict consensus shown monophyletic higher taxon = Bremer ≥ 8 = Bremer 3 – 7 = Bremer 1 – 2 Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Midacritus stuardoanus M Midacritus wagenknechti Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Ectyphus armipes Parectyphus namibiensis Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Messiasia californica M Messiasia painteri Plyomydas peruviensis M Plyomydas sp nov Tandil Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Anomalomydas australicus F Anomalomydas mackerrasi Anomalomydas sp nov Simpson F Pseudorhopalia mirandai Miltinus sp. (Northern Territory) Rhopalia cincta Rhopalia annulata Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Nemomydas venosus Nemomydas brachyrhynchus Nemomydas melanopogon Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Leptomydas notos Nemomydas pantherinus Pseudonomoneura hirta Pseudonomoneura micheneri Nemomydas lamia Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Notosyllegomydas brincki Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata M Cephalocera sp Ratelfontein Namadytes cimbebasiensis Namadytes vansoni Mydaselpis ngurumani Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Afroleptomydas A suffusipennis Afroleptomydas A paganus Afroleptomydas A inhacae Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Afroleptomydas C sp Gobabeb Syllegomydas S lineatus Hispanomydas hispanicus Syllegomydas N astrictus 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 Halterorchis karooensis F Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus length 1839 steps 346 trees strict consensus shown
... a posteriori taxon exclusion – Mydidae 13 Female oviposition strategy Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis Stratiomydas lividus Mydas clavatus female ovipositor dorsal, Cacatuopyga basifascia Mydas tricinctus Mydas xanthopterus Messiasia californica Messiasia painteri female ovipositor dorsal, Leptomydas lusitanicus Mydinae - New World Mydidae all length 1839 steps 346 trees strict consensus shown Mydas type Cacatuopyga type Phyllomydas type = Bremer ≥ 8 = Bremer 3 – 7 = Bremer 1 – 2 Plyomydas peruviensis Plyomydas sp nov Tandil Dolichogaster brevicornis Phyllomydas bruesii female ovipositor dorsal, Messiasia painteri Phyllomydas parvulus = tergite 10 = tergite 9 = tergite 8 = tergite 9+10 scale lines = 1 mm female ovipositor dorsal, Phyllomydas parvulus
... a posteriori taxon exclusion – Mydidae 14 Female oviposition strategy – Mydinae Mydidae all (14 species) Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Messiasia californica Messiasia painteri Plyomydas peruviensis Plyomydas sp nov Tandil Dolichogaster brevicornis Mydidae 2 (13 species) Paramydas igniticornis Plyomydas peruviensis Plyomydas sp nov Tandil Messiasia californica Messiasia painteri Mydas clavatus Mydas xanthopterus Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruficornis Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Mydas type Cacatuopyga type Phyllomydas type = Bremer ≥ 8 = Bremer 3 – 7 = Bremer 1 – 2 Mydidae 5 (14 species) Phyllomydas bruesii Phyllomydas parvulus Paramydas igniticornis Plyomydas peruviensis Plyomydas sp nov Tandil Messiasia californica Messiasia painteri Mydas clavatus Mydas tricinctus Mydas xanthopterus Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruficornis Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus
.... a posteriori taxon inclusion – Asilidae 15 Bathypogoninae Asilidae all ◊ newly postulated because of unique position › Australian Bathypogon › originally only a single species included Bathypogon Phellinae Perasis transvaalensis Zabrops tagax Protometer sp. (El Tuparro) Psilocurus modestus Laphystia aegyptiaca 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 Choerades bella Laphria flava Laphria aktis Lamyra gulo Stiphrolamyra angularis Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Afroestricus chiastoneurus Ommatius tibialis Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus Lycomya germainii Pogonioefferia pogonias Proctacanthus philadelphicus Dasophrys crenulatus Eichoichemus pyrrhomystax Lochmorhynchus albicans Myaptex brachyptera Neolophonotus bimaculatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Dysmachus trigonus Philonicus albiceps Congomochtherus inachus Heligmonevra laevis Tolmerus atricapillus Zosteria rosevillensis Asilus crabroniformis Asilus sericeus Neomochtherus pallipes Afromochtherus mendax Clephydroneura sundaica Clephydroneura sp. (Kepong) Colepia rufiventris Neoitamus cyanurus Threnia carbonaria Megaphorus pulchrus Promachus amastrus Apoclea algira Philodicus tenuipes Bathypogon nigrinus Phellus olgae Obelophorus terebratus Tillobroma punctipennis Hypenetes stigmatias Lycostommyia albifacies Coleomyia setigera Ancylorhynchus fulvicollis Scylaticus costalis Connomyia varipennis Prolepsis tristis Creolestes nigribarbis Microstylum sp. (Karkloof) Daspletis stenoura Plesiomma sp. (Guanacaste) Ospriocerus aeacus Scleropogon subulatus Stenopogon sabaudus Stenopogon rufibarbis Gonioscelis ventralis Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Acnephalum cylindricum Sisyrnodytes sp. (Gamka) Ablautus coquilletti Willistonina bilineata Trichoura sp. (Tierberg) Oligopogon sp. (Cape Recife) Broticosia paramonovi Dicolonus simplex Echthodopa pubera Myelaphus melas Nannodioctria sp. (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Acronyches maya Beameromyia bifida Euscelidia pulchra Tipulogaster glabrata Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Damalis sp. (Palatupana) Damalis monochaetes Damalis annulata Rhipidocephala sp. (Harold Johnson) Trigonomima sp. (anamaliensis) Holcocephala abdominalis Holcocephala calva Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Holopogon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis length 2760 steps 460 trees strict consensus shown Bathypogon sp. Bathypogon nigrinus
.... a posteriori taxon inclusion – Asilidae 16 Bathypogon 2 Perasis transvaalensis Zabrops tagax Protometer sp. (El Tuparro) Psilocurus modestus Laphystia aegyptiaca 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 Choerades bella Laphria flava Laphria aktis Lamyra gulo Stiphrolamyra angularis Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Afroestricus chiastoneurus Ommatius tibialis Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus Lycomya germainii Pogonioefferia pogonias Proctacanthus philadelphicus Dasophrys crenulatus Eichoichemus pyrrhomystax Lochmorhynchus albicans Myaptex brachyptera Neolophonotus bimaculatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Dysmachus trigonus Philonicus albiceps Congomochtherus inachus Heligmonevra laevis Tolmerus atricapillus Zosteria rosevillensis Asilus crabroniformis Asilus sericeus Neomochtherus pallipes Afromochtherus mendax Clephydroneura sundaica Clephydroneura sp. (Kepong) Colepia rufiventris Neoitamus cyanurus Threnia carbonaria Megaphorus pulchrus Promachus amastrus Apoclea algira Philodicus tenuipes Bathypogon nigrinus Bathypogon rubellus Phellus olgae Obelophorus terebratus Tillobroma punctipennis Hypenetes stigmatias Lycostommyia albifacies Coleomyia setigera Ancylorhynchus fulvicollis Scylaticus costalis Connomyia varipennis Prolepsis tristis Creolestes nigribarbis Microstylum sp. (Karkloof) Daspletis stenoura Plesiomma sp. (Guanacaste) Ospriocerus aeacus Scleropogon subulatus Stenopogon sabaudus Stenopogon rufibarbis Gonioscelis ventralis Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Acnephalum cylindricum Sisyrnodytes sp. (Gamka) Ablautus coquilletti Willistonina bilineata Trichoura sp. (Tierberg) Oligopogon sp. (Cape Recife) Broticosia paramonovi Dicolonus simplex Echthodopa pubera Myelaphus melas Nannodioctria sp. (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Acronyches maya Beameromyia bifida Euscelidia pulchra Tipulogaster glabrata Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Damalis sp. (Palatupana) Damalis monochaetes Damalis annulata Rhipidocephala sp. (Harold Johnson) Trigonomima sp. (anamaliensis) Holcocephala abdominalis Holcocephala calva Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Holopogon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis length 2763 steps 391 trees strict consensus shown Bathypogon 4 Bathypogon nigrinus Bathypogon rubellus Phellus olgae Obelophorus terebratus Tillobroma punctipennis Hypenetes stigmatias Lycostommyia albifacies Coleomyia setigera Ancylorhynchus fulvicollis Scylaticus costalis Connomyia varipennis Prolepsis tristis Creolestes nigribarbis Microstylum sp. (Karkloof) Daspletis stenoura Plesiomma sp. (Guanacaste) Ospriocerus aeacus Scleropogon subulatus Stenopogon sabaudus Stenopogon rufibarbis Gonioscelis ventralis Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens remaining Asilidae Bathypogon Phellinae = Bremer ≥ 8 = Bremer 3 – 7 = Bremer 1 – 2 Perasis transvaalensis Zabrops tagax Protometer sp. (El Tuparro) Psilocurus modestus Laphystia aegyptiaca 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 Choerades bella Laphria flava Laphria aktis Lamyra gulo Stiphrolamyra angularis Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Afroestricus chiastoneurus Ommatius tibialis Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus Lycomya germainii Pogonioefferia pogonias Proctacanthus philadelphicus Dasophrys crenulatus Eichoichemus pyrrhomystax Lochmorhynchus albicans Myaptex brachyptera Neolophonotus bimaculatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Dysmachus trigonus Philonicus albiceps Congomochtherus inachus Heligmonevra laevis Tolmerus atricapillus Zosteria rosevillensis Asilus crabroniformis Asilus sericeus Neomochtherus pallipes Afromochtherus mendax Clephydroneura sundaica Clephydroneura sp. (Kepong) Colepia rufiventris Neoitamus cyanurus Threnia carbonaria Megaphorus pulchrus Promachus amastrus Apoclea algira Philodicus tenuipes Bathypogon nigrinus Bathypogon rubellus Bathypogon robustus Bathypogon sp. (Lake King) Phellus olgae Obelophorus terebratus Tillobroma punctipennis Hypenetes stigmatias Lycostommyia albifacies Coleomyia setigera Ancylorhynchus fulvicollis Scylaticus costalis Connomyia varipennis Prolepsis tristis Creolestes nigribarbis Microstylum sp. (Karkloof) Daspletis stenoura Plesiomma sp. (Guanacaste) Ospriocerus aeacus Scleropogon subulatus Stenopogon sabaudus Stenopogon rufibarbis Gonioscelis ventralis Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Acnephalum cylindricum Sisyrnodytes sp. (Gamka) Ablautus coquilletti Willistonina bilineata Trichoura sp. (Tierberg) Oligopogon sp. (Cape Recife) Broticosia paramonovi Dicolonus simplex Echthodopa pubera Myelaphus melas Nannodioctria sp. (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Acronyches maya Beameromyia bifida Euscelidia pulchra Tipulogaster glabrata Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Damalis sp. (Palatupana) Damalis monochaetes Damalis annulata Rhipidocephala sp. (Harold Johnson) Trigonomima sp. (anamaliensis) Holcocephala abdominalis Holcocephala calva Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Holopogon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis length 2771 steps 323 trees strict consensus shown Bathypogon nigrinus Bathypogon rubellus Bathypogon robustus Bathypogon sp. (Lake King) Phellus olgae Obelophorus terebratus Tillobroma punctipennis Hypenetes stigmatias Lycostommyia albifacies Coleomyia setigera Ancylorhynchus fulvicollis Scylaticus costalis Connomyia varipennis Prolepsis tristis Creolestes nigribarbis Microstylum sp. (Karkloof) Daspletis stenoura Plesiomma sp. (Guanacaste) Ospriocerus aeacus Scleropogon subulatus Stenopogon sabaudus Stenopogon rufibarbis Gonioscelis ventralis Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens remaining Asilidae
..... conclusion 17 Conclusion ◊ strive to include as many taxa as feasible ◊ Apioceridae › biogeographic hypothesis failed test ◊ Mydidae › discussion of evolution of ovipositor only feasible with hypothesis derived from full data-set › secondary signal (from elsewhere in the tree) emerges ◊ Asilidae: Bathypogoninae › inclusion of one and three additional taxa corroborates unique position
...... acknowledgements 18 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 ◊ Willi Hennig Society – TNT Smithsonian National Museum of Natural History

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Effect of taxon sampling in morphological phylogenetic analyses

  • 1. 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 Washington, DC Smithsonian National Museum of Natural History
  • 2. . introduction 2 Taxon sampling in cladistic studies ◊ integral and important component –> data + taxa ◊ too often data have priority while taxa are “neglected” › morphology – based on “unlimited” number of taxa in natural history collections › traditional molecular – based on more recently collected, limited number of taxa (95% ethanol) › genome molecular – based on newly collected, very limited number of taxa (RNAlater, DNAgard) ◊ taxon sampling examined in two morphological studies –> exclude or include?
  • 3. . introduction 3 Taxon sampling in cladistic studies ◊ a posteriori taxon exclusion › Was the addition of taxa “necessary” to obtain the same hypothesis of relationships? › Does one need to include more than one species per genus? › example: two small families with dense generic sampling ◊ a posteriori taxon inclusion › Would the addition of taxa corroborate the postulated hypothesis of relationships? › Does the inclusion of additional species of a genus alter its position? › example: a very diverse group with limited generic sampling
  • 4. . introduction 4 Taxa ◊ Apioceridae ◊ Mydidae › 143 species › rarely collected › Australia, Chile, western North America, South Africa Apiocera (Apiocera) pica © J. Hort ◊ Asilidae › 473 species › rarely collected › world-wide primarily in arid regions Eremomidas arabicus © D. Gardner › > 7,500 species › collected by specialists and non-specialists alike › speciose world-wide Scleropogon duncani
  • 5. . introduction 5 Minimum ages + relationships ◊ Crato Formation, Brazil › Cretaceous: Aptian – Albian › ≈ 112 myo › Asilidae: † Araripogon axelrodi › Mydidae: † Cretomydas santanensis Asilidae: † Araripogon axelrodi Asilidae Mydidae Apioceridae Mydidae: † Cretomydas santanensis
  • 6. 6 Overview ◊ Methods + data-sets ◊ a posteriori taxon exclusion › Apioceridae biogeography › Mydidae female ovipositor morphology ◊ a posteriori taxon inclusion › Asilidae: Bathypogoninae position Trichardis picta Prytanomyia albida
  • 7. 7 Methods + data-sets ◊ Mesquite 2.75 ◊ TNT 1.1 (January 2013) ◊ 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; taxon Apioceridae all Apioceridae 1 Apioceridae 2 Apioceridae 5 Mydidae all Mydidae 1 Mydidae 2 Mydidae 5 Asilidae all Bathypogon 2 Bathypogon 4 # species in analysis 30 21.7 % 4 2.9 % 8 5.8 % 14 10.1 % 135 28.4 % 49 10.4 % 80 16.9 % 117 24.7 % 158 2.1 % 159 2.1 % 161 2.1 % # genera in analysis 4 100 % 51 77.2 % 140 25.9 %
  • 8. .. a posteriori taxon exclusion – Apioceridae Apioceridae all 8 Apioceridae 1 Apiocera An obscura Apiocera An philippii Apiocera An philippii Apiocera Ap pica Apiocera Ap helenae Apiocera P arena Apiocera Ap imminuta Apiocera R braunsi Apiocera Ap tonnoiri Apiocera R alastor M length 1787 steps Apiocera R braunsi Apiocera Ap goerlingi 306 trees Apiocera Ap asilica strict consensus shown Apiocera Ap brevicornis Apiocera Ap sp Stuart M Apiocera Ap hardyi Apiocera Ap pica Apiocera Ap sp Lloyd Apiocera Ap similis Apiocera Ap maritima Apiocera Ap minor Apiocera Ap newmani 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 Apioceridae 2 Apiocera An obscura Apiocera An philippii Apiocera Ap minor Apiocera Ap pica Apiocera P arena Apiocera P bilineata Apiocera R alastor M Apiocera R braunsi Afrotropical length 1807 steps 376 trees strict consensus shown Nearctic + Neotropical (Mexico: Veracruz) length 1839 steps 346 trees strict consensus shown monophyletic subgenus Chilean Apiocera = Bremer ≥ 8 = Bremer 3 – 7 = Bremer 1 – 2 Apioceridae 5 Apiocera An obscura Apiocera An philippii Apiocera Ap tonnoiri Apiocera R alastor M Apiocera R braunsi Apiocera Ap hardyi Apiocera Ap maritima Apiocera Ap minor Apiocera Ap pica Apiocera P acuticauda Apiocera P arena Apiocera P parkeri Apiocera P bilineata Apiocera P mexicana M length 1819 steps 345 trees strict consensus shown
  • 9. .. a posteriori taxon exclusion – Apioceridae 9 Apioceridae – Yeates & Irwin 1996 Apioceridae Apiocera (Pyrocera) Apiocera (Ripidosyrma) Apiocera (Anypenus) Apiocera (Apiocera) Yeates & Irwin 1996 western North America South Africa Argentina / Chile Australia area relationships
  • 10. .. a posteriori taxon exclusion – Apioceridae 10 Apioceridae – current hypothesis Apioceridae Apiocera (Pyrocera) Apiocera (Pyrocera) Apiocera (Ripidosyrma) Apiocera (Ripidosyrma) Apiocera (Anypenus) western North America South Africa Argentina / Chile Apiocera (Anypenus) 1 Apiocera (Anypenus) 2 Apiocera (Apiocera) Apiocera (Apiocera) 1 Apiocera (Apiocera) 2 Apiocera (Apiocera) ... Yeates & Irwin 1996 current hypothesis Australia area relationships
  • 11. ... a posteriori taxon exclusion – Mydidae Mydidae 1 11 Mydidae 2 Rhaphiomidas maehleri Megascelus albovillosus Neorhaphiomidas hardyi Tongamya miranda Diochlistus gracilis Cretomydas santanensis Anomalomydas mackerrasi Midacritus wagenknechti Paramydas igniticornis Cacatuopyga basifascia Plyomydas peruviensis M Dolichogaster brevicornis Messiasia painteri Mydas clavatus Phyllomydas parvulus Stratiomydas lividus Pseudorhopalia mirandai Perissocerus transcaspicus Rhopalia olivieri Apiophora paulseni Miltinus viduatus Ectyphus pinguis Parectyphus namibiensis Heteromydas bicolor Opomydas townsendi Eremomidas bek Hessemydas daugeroni Leptomydas lusitanicus Nemomydas venosus Afroleptomydas A rufithorax Arenomydas caerulescens Cephalocera longirostris Eremohaplomydas desertorum Haplomydas crassipes Lachnocorynus sp nov Kotwa M Halterorchis karooensis F Namadytes vansoni Neolaparopsis puncturatus Nomoneura caffra Nomoneuroides natalensis Notosyllegomydas brincki Hispanomydas hispanicus Syllegomydas S algiricus Mydaselpis sp nov Daan Vespiodes phaios Pseudonomoneura hirta Mitrodetus dentitarsis Namibimydas stuckenbergi Nothomydas picketti length 1566 steps 589 trees strict consensus shown monophyletic higher taxon = Bremer ≥ 8 = Bremer 3 – 7 = Bremer 1 – 2 Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus gracilis Diochlistus melleipennis Anomalomydas mackerrasi Anomalomydas australicus F Midacritus stuardoanus M Midacritus wagenknechti Apiophora paulseni Apiophora quadricinctata Miltinus minutus Miltinus viduatus Ectyphus armipes Ectyphus pinguis Parectyphus namibiensis Heteromydas bicolor Opomydas townsendi Paramydas igniticornis Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Mydas clavatus Mydas xanthopterus Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruficornis M Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Pseudorhopalia mirandai Rhopalia annulata Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Nemomydas brachyrhynchus Nemomydas venosus Eremomidas bek Leptomydas lusitanicus Hessemydas daugeroni Hessemydas seyrigi Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Namadytes cimbebasiensis Namadytes vansoni Afroleptomydas A rufithorax Afroleptomydas C mauricei Hispanomydas hispanicus Syllegomydas S algiricus Syllegomydas S vittatus Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Vespiodes leopolditertii Vespiodes phaios Mydaselpis ngurumani Arenomydas caerulescens Arenomydas callosus Cephalocera longirostris Cephalocera sp Papendorp Notosyllegomydas brincki Eremomidas arabicus Halterorchis karooensis F Leptomydas sardous Pseudonomoneura hirta Pseudonomoneura micheneri Mitrodetus dentitarsis Mitrodetus dimidiatus Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi length 1673 steps 335 trees strict consensus shown
  • 12. ... a posteriori taxon exclusion – Mydidae Mydidae 5 12 Mydidae all Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Apiophora paulseni Apiophora quadricinctata Midacritus stuardoanus M Midacritus wagenknechti Miltinus sp. (Northern Territory) Ectyphus armipes Parectyphus namibiensis Pseudorhopalia mirandai Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Anomalomydas mackerrasi Anomalomydas australicus F Anomalomydas sp nov Simpson F Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Rhopalia annulata Rhopalia cincta Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Paramydas igniticornis Plyomydas peruviensis M Plyomydas sp nov Tandil Messiasia californica M Messiasia painteri Mydas clavatus Mydas tricinctus Mydas xanthopterus Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruficornis M Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Eremomidas bek Hessemydas daugeroni Hessemydas parkeri M Hessemydas seyrigi Leptomydas lusitanicus Leptomydas notos Leptomydas rapti M Leptomydas sardous Leptomydas turcicus Nemomydas brachyrhynchus Nemomydas lamia Nemomydas melanopogon Nemomydas pantherinus Nemomydas venosus Pseudonomoneura bajaensis Hispanomydas hispanicus Mydaselpis ngurumani Neolaparopsis puncturatus Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Notosyllegomydas brincki Syllegomydas N dispar Syllegomydas S algiricus Syllegomydas S palestinensis Syllegomydas S proximus Syllegomydas S vittatus Syll gen nov sp nov minute proboscis F Pseudonomoneura hirta Pseudonomoneura micheneri Namadytes cimbebasiensis Namadytes vansoni Eremomidas sp nov Tiega M Eremomidas arabicus Halterorchis karooensis F Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Nothomydas picketti Namibimydas psamminos Namibimydas stuckenbergi Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus Afroleptomydas A inhacae Afroleptomydas A opacicinctus Afroleptomydas C mauricei Afroleptomydas A omeri Afroleptomydas A rufithorax Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata M Cephalocera sp Ratelfontein length 1801 steps 771 trees strict consensus shown monophyletic higher taxon = Bremer ≥ 8 = Bremer 3 – 7 = Bremer 1 – 2 Rhaphiomidas hasbroucki Rhaphiomidas maehleri Rhaphiomidas parkeri Tongamya miranda Megascelus albovillosus Megascelus melanoproctus Megascelus nigricornis Neorhaphiomidas sp Mingenew M Neorhaphiomidas sp Yanchep Neorhaphiomidas hardyi Neorhaphiomidas sp Mandurah Cretomydas santanensis Diochlistus auripennis M Diochlistus gracilis Diochlistus melleipennis Midacritus stuardoanus M Midacritus wagenknechti Apiophora paulseni Apiophora quadricinctata Miltinus stenogaster Miltinus viduatus Miltinus maculipennis Miltinus minutus Ectyphus armipes Parectyphus namibiensis Ectyphus amboseli M Ectyphus pinguis Heteromydas bicolor Opomydas townsendi Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis M Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Messiasia californica M Messiasia painteri Plyomydas peruviensis M Plyomydas sp nov Tandil Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Anomalomydas australicus F Anomalomydas mackerrasi Anomalomydas sp nov Simpson F Pseudorhopalia mirandai Miltinus sp. (Northern Territory) Rhopalia cincta Rhopalia annulata Rhopalia olivieri Perissocerus dumonti F Perissocerus transcaspicus Hessemydas parkeri M Hessemydas daugeroni Hessemydas seyrigi Nemomydas venosus Nemomydas brachyrhynchus Nemomydas melanopogon Leptomydas lusitanicus Eremomidas bek Leptomydas rapti M Leptomydas turcicus Leptomydas corsicanus Nemomydas gruenbergi M Leptomydas sardous Leptomydas notos Nemomydas pantherinus Pseudonomoneura hirta Pseudonomoneura micheneri Nemomydas lamia Arenomydas partitus Arenomydas caerulescens Arenomydas callosus Arenomydas namaquensis M Notosyllegomydas brincki Cephalocera longirostris Cephalocera sp Papendorp Cephalocera sp Sors Sors Cephalocera imitata M Cephalocera sp Ratelfontein Namadytes cimbebasiensis Namadytes vansoni Mydaselpis ngurumani Neolaparopsis puncturatus Haplomydas crassipes Eremohaplomydas desertorum Lachnocorynus sp nov Kotwa M Mydaselpis sp nov Daan Mydaselpis sp nov Gamsberg M Vespiodes leopolditertii Vespiodes phaios Nomoneura caffra Nomoneura paradoxa Nomoneuroides natalensis Afroleptomydas A suffusipennis Afroleptomydas A paganus Afroleptomydas A inhacae Afroleptomydas A consanguinensis Afroleptomydas A opacicinctus Afroleptomydas A sp Clanwilliam Afroleptomydas A sp Obib Afroleptomydas C sp Koeroegab Afroleptomydas A omeri Afroleptomydas A rufithorax Afroleptomydas C mauricei Afroleptomydas C sp Stampriet Afroleptomydas C sp Gobabeb Syllegomydas S lineatus Hispanomydas hispanicus Syllegomydas N astrictus 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 Halterorchis karooensis F Diochlistinae Argentina F Mitrodetus nanoglossa M Mitrodetus microglossa Mitrodetus dentitarsis Mitrodetus dimidiatus length 1839 steps 346 trees strict consensus shown
  • 13. ... a posteriori taxon exclusion – Mydidae 13 Female oviposition strategy Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis Stratiomydas lividus Mydas clavatus female ovipositor dorsal, Cacatuopyga basifascia Mydas tricinctus Mydas xanthopterus Messiasia californica Messiasia painteri female ovipositor dorsal, Leptomydas lusitanicus Mydinae - New World Mydidae all length 1839 steps 346 trees strict consensus shown Mydas type Cacatuopyga type Phyllomydas type = Bremer ≥ 8 = Bremer 3 – 7 = Bremer 1 – 2 Plyomydas peruviensis Plyomydas sp nov Tandil Dolichogaster brevicornis Phyllomydas bruesii female ovipositor dorsal, Messiasia painteri Phyllomydas parvulus = tergite 10 = tergite 9 = tergite 8 = tergite 9+10 scale lines = 1 mm female ovipositor dorsal, Phyllomydas parvulus
  • 14. ... a posteriori taxon exclusion – Mydidae 14 Female oviposition strategy – Mydinae Mydidae all (14 species) Paramydas igniticornis Cacatuopyga basifascia Cacatuopyga ruficornis Stratiomydas lividus Mydas clavatus Mydas tricinctus Mydas xanthopterus Messiasia californica Messiasia painteri Plyomydas peruviensis Plyomydas sp nov Tandil Dolichogaster brevicornis Mydidae 2 (13 species) Paramydas igniticornis Plyomydas peruviensis Plyomydas sp nov Tandil Messiasia californica Messiasia painteri Mydas clavatus Mydas xanthopterus Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruficornis Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus Mydas type Cacatuopyga type Phyllomydas type = Bremer ≥ 8 = Bremer 3 – 7 = Bremer 1 – 2 Mydidae 5 (14 species) Phyllomydas bruesii Phyllomydas parvulus Paramydas igniticornis Plyomydas peruviensis Plyomydas sp nov Tandil Messiasia californica Messiasia painteri Mydas clavatus Mydas tricinctus Mydas xanthopterus Stratiomydas lividus Cacatuopyga basifascia Cacatuopyga ruficornis Dolichogaster brevicornis Phyllomydas bruesii Phyllomydas parvulus
  • 15. .... a posteriori taxon inclusion – Asilidae 15 Bathypogoninae Asilidae all ◊ newly postulated because of unique position › Australian Bathypogon › originally only a single species included Bathypogon Phellinae Perasis transvaalensis Zabrops tagax Protometer sp. (El Tuparro) Psilocurus modestus Laphystia aegyptiaca 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 Choerades bella Laphria flava Laphria aktis Lamyra gulo Stiphrolamyra angularis Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Afroestricus chiastoneurus Ommatius tibialis Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus Lycomya germainii Pogonioefferia pogonias Proctacanthus philadelphicus Dasophrys crenulatus Eichoichemus pyrrhomystax Lochmorhynchus albicans Myaptex brachyptera Neolophonotus bimaculatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Dysmachus trigonus Philonicus albiceps Congomochtherus inachus Heligmonevra laevis Tolmerus atricapillus Zosteria rosevillensis Asilus crabroniformis Asilus sericeus Neomochtherus pallipes Afromochtherus mendax Clephydroneura sundaica Clephydroneura sp. (Kepong) Colepia rufiventris Neoitamus cyanurus Threnia carbonaria Megaphorus pulchrus Promachus amastrus Apoclea algira Philodicus tenuipes Bathypogon nigrinus Phellus olgae Obelophorus terebratus Tillobroma punctipennis Hypenetes stigmatias Lycostommyia albifacies Coleomyia setigera Ancylorhynchus fulvicollis Scylaticus costalis Connomyia varipennis Prolepsis tristis Creolestes nigribarbis Microstylum sp. (Karkloof) Daspletis stenoura Plesiomma sp. (Guanacaste) Ospriocerus aeacus Scleropogon subulatus Stenopogon sabaudus Stenopogon rufibarbis Gonioscelis ventralis Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Acnephalum cylindricum Sisyrnodytes sp. (Gamka) Ablautus coquilletti Willistonina bilineata Trichoura sp. (Tierberg) Oligopogon sp. (Cape Recife) Broticosia paramonovi Dicolonus simplex Echthodopa pubera Myelaphus melas Nannodioctria sp. (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Acronyches maya Beameromyia bifida Euscelidia pulchra Tipulogaster glabrata Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Damalis sp. (Palatupana) Damalis monochaetes Damalis annulata Rhipidocephala sp. (Harold Johnson) Trigonomima sp. (anamaliensis) Holcocephala abdominalis Holcocephala calva Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Holopogon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis length 2760 steps 460 trees strict consensus shown Bathypogon sp. Bathypogon nigrinus
  • 16. .... a posteriori taxon inclusion – Asilidae 16 Bathypogon 2 Perasis transvaalensis Zabrops tagax Protometer sp. (El Tuparro) Psilocurus modestus Laphystia aegyptiaca 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 Choerades bella Laphria flava Laphria aktis Lamyra gulo Stiphrolamyra angularis Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Afroestricus chiastoneurus Ommatius tibialis Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus Lycomya germainii Pogonioefferia pogonias Proctacanthus philadelphicus Dasophrys crenulatus Eichoichemus pyrrhomystax Lochmorhynchus albicans Myaptex brachyptera Neolophonotus bimaculatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Dysmachus trigonus Philonicus albiceps Congomochtherus inachus Heligmonevra laevis Tolmerus atricapillus Zosteria rosevillensis Asilus crabroniformis Asilus sericeus Neomochtherus pallipes Afromochtherus mendax Clephydroneura sundaica Clephydroneura sp. (Kepong) Colepia rufiventris Neoitamus cyanurus Threnia carbonaria Megaphorus pulchrus Promachus amastrus Apoclea algira Philodicus tenuipes Bathypogon nigrinus Bathypogon rubellus Phellus olgae Obelophorus terebratus Tillobroma punctipennis Hypenetes stigmatias Lycostommyia albifacies Coleomyia setigera Ancylorhynchus fulvicollis Scylaticus costalis Connomyia varipennis Prolepsis tristis Creolestes nigribarbis Microstylum sp. (Karkloof) Daspletis stenoura Plesiomma sp. (Guanacaste) Ospriocerus aeacus Scleropogon subulatus Stenopogon sabaudus Stenopogon rufibarbis Gonioscelis ventralis Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Acnephalum cylindricum Sisyrnodytes sp. (Gamka) Ablautus coquilletti Willistonina bilineata Trichoura sp. (Tierberg) Oligopogon sp. (Cape Recife) Broticosia paramonovi Dicolonus simplex Echthodopa pubera Myelaphus melas Nannodioctria sp. (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Acronyches maya Beameromyia bifida Euscelidia pulchra Tipulogaster glabrata Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Damalis sp. (Palatupana) Damalis monochaetes Damalis annulata Rhipidocephala sp. (Harold Johnson) Trigonomima sp. (anamaliensis) Holcocephala abdominalis Holcocephala calva Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Holopogon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis length 2763 steps 391 trees strict consensus shown Bathypogon 4 Bathypogon nigrinus Bathypogon rubellus Phellus olgae Obelophorus terebratus Tillobroma punctipennis Hypenetes stigmatias Lycostommyia albifacies Coleomyia setigera Ancylorhynchus fulvicollis Scylaticus costalis Connomyia varipennis Prolepsis tristis Creolestes nigribarbis Microstylum sp. (Karkloof) Daspletis stenoura Plesiomma sp. (Guanacaste) Ospriocerus aeacus Scleropogon subulatus Stenopogon sabaudus Stenopogon rufibarbis Gonioscelis ventralis Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens remaining Asilidae Bathypogon Phellinae = Bremer ≥ 8 = Bremer 3 – 7 = Bremer 1 – 2 Perasis transvaalensis Zabrops tagax Protometer sp. (El Tuparro) Psilocurus modestus Laphystia aegyptiaca 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 Choerades bella Laphria flava Laphria aktis Lamyra gulo Stiphrolamyra angularis Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Afroestricus chiastoneurus Ommatius tibialis Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus Lycomya germainii Pogonioefferia pogonias Proctacanthus philadelphicus Dasophrys crenulatus Eichoichemus pyrrhomystax Lochmorhynchus albicans Myaptex brachyptera Neolophonotus bimaculatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Dysmachus trigonus Philonicus albiceps Congomochtherus inachus Heligmonevra laevis Tolmerus atricapillus Zosteria rosevillensis Asilus crabroniformis Asilus sericeus Neomochtherus pallipes Afromochtherus mendax Clephydroneura sundaica Clephydroneura sp. (Kepong) Colepia rufiventris Neoitamus cyanurus Threnia carbonaria Megaphorus pulchrus Promachus amastrus Apoclea algira Philodicus tenuipes Bathypogon nigrinus Bathypogon rubellus Bathypogon robustus Bathypogon sp. (Lake King) Phellus olgae Obelophorus terebratus Tillobroma punctipennis Hypenetes stigmatias Lycostommyia albifacies Coleomyia setigera Ancylorhynchus fulvicollis Scylaticus costalis Connomyia varipennis Prolepsis tristis Creolestes nigribarbis Microstylum sp. (Karkloof) Daspletis stenoura Plesiomma sp. (Guanacaste) Ospriocerus aeacus Scleropogon subulatus Stenopogon sabaudus Stenopogon rufibarbis Gonioscelis ventralis Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Acnephalum cylindricum Sisyrnodytes sp. (Gamka) Ablautus coquilletti Willistonina bilineata Trichoura sp. (Tierberg) Oligopogon sp. (Cape Recife) Broticosia paramonovi Dicolonus simplex Echthodopa pubera Myelaphus melas Nannodioctria sp. (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Acronyches maya Beameromyia bifida Euscelidia pulchra Tipulogaster glabrata Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Damalis sp. (Palatupana) Damalis monochaetes Damalis annulata Rhipidocephala sp. (Harold Johnson) Trigonomima sp. (anamaliensis) Holcocephala abdominalis Holcocephala calva Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Holopogon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis length 2771 steps 323 trees strict consensus shown Bathypogon nigrinus Bathypogon rubellus Bathypogon robustus Bathypogon sp. (Lake King) Phellus olgae Obelophorus terebratus Tillobroma punctipennis Hypenetes stigmatias Lycostommyia albifacies Coleomyia setigera Ancylorhynchus fulvicollis Scylaticus costalis Connomyia varipennis Prolepsis tristis Creolestes nigribarbis Microstylum sp. (Karkloof) Daspletis stenoura Plesiomma sp. (Guanacaste) Ospriocerus aeacus Scleropogon subulatus Stenopogon sabaudus Stenopogon rufibarbis Gonioscelis ventralis Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens remaining Asilidae
  • 17. ..... conclusion 17 Conclusion ◊ strive to include as many taxa as feasible ◊ Apioceridae › biogeographic hypothesis failed test ◊ Mydidae › discussion of evolution of ovipositor only feasible with hypothesis derived from full data-set › secondary signal (from elsewhere in the tree) emerges ◊ Asilidae: Bathypogoninae › inclusion of one and three additional taxa corroborates unique position
  • 18. ...... acknowledgements 18 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 ◊ Willi Hennig Society – TNT Smithsonian National Museum of Natural History