Apiocera

Transkript

Apiocera
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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