tdikow mydidae morphological phylogeny hennig 2012
DESCRIPTION
Current (Jun 2012) phylogenetic hypothesis of relationships within Mydidae (Diptra: Asiloidea)TRANSCRIPT
1
Phylogeny of Apioceridae and Mydidae (Insecta: Diptera) based on morphological characters of adult flies
Torsten DikowBiodiversity Synthesis Center Field Museum of Natural History, Chicago, IL, USA
2
Asilidae
Apiocera (Pyrocera)
Apiocera (Ripidosyrma)
Apiocera (Anypenus)
Apiocera (Apiocera)
Rhaphiomidinae
Megascelinae
Rhopaliinae
Cacatuopyginae
Leptomydinae
Syllegomydinae
Anomalomydinae
Diochlistinae
Ectyphinae
Apiophorinae
Mydinae
Yeates & Irwin 1996 – Apioceridae: cladistic reappraisal and biogeography› 6 Apioceridae exemplar species › 13 Mydidae exemplar species › 77 morphological characters
. introduction
Asilidae
Apiocera (Pyrocera)
Apiocera (Ripidosyrma)
Apiocera (Anypenus)
Apiocera (Apiocera)
Rhaphiomidinae
Megascelinae
Rhopaliinae
Cacatuopyginae
Leptomydinae
Syllegomydinae
Anomalomydinae
Diochlistinae
Ectyphinae
Apiophorinae
Mydinae
Pre-1996 = non-monophyletic Apioceridae Post-1996 = monophyletic Apioceridae
Apioceridae
Mydidae
3
Apiocera (Pyrocera) western North America
Apiocera (Apiocera)
Australia
Apiocera (Ripidosyrma) South Africa
Apiocera (Anypenus)Argentina + Chile
Tongamya
Megascelus
Neorhaphiomidas
Mitrodetus
Diochlistus
Apiophora, Midacritus
Miltinus
Apioceridae
Mydidae: Megascelinae
Mydidae: Diochlistinae
Mydidae: Apiophorinae
. introduction
Yeates & Irwin 1996 – Exclusively southern connection taxa = Gondwanan
4
Apiocera (Apiocera) moerens © Peter Chew
Apioceridae – “flower-loving flies”Adult flies flower-feeders◊
larvae possibly predatory◊
138 species and 1 genus ◊ (4 subgenera)
length = 15 – 30 mm◊
occur only in ◊ › Argentina and Chile – 4 species › Australia – 67 species › South Africa – 3 species › western North America (Canada, Mexico, USA) – 64 species
Apiocera bilineata © Greg Ballmer
. introduction
5. introduction
Mydidae: Opomydas townsendi
Mydidae – “mydas flies”Adult flies flower-feeders, some do not feed at all◊
Larvae possibly predatory◊
472 species and 66 genera world-wide◊
length = 9 – 60 mm ◊ (wing length up to 52 mm)
World-wide primarily in arid regions◊
Rarely collected – Afrotropical Region most speciose◊
Oldest definitive fossil ◊ † Cretomydas santanensis › Crato Formation, Brazil › Cretaceous: Aptian – Albian › ≈ 112 myo
Mydidae: Eremomidas arabicus
6
Taxon Sampling27 ◊ Apiocera species (= 19.6 %)
134 Mydidae species◊ (= 28.4 %) › 46 genera (= 69.7 %)
› 7 tribal taxa (= 87.5 %) › 11 subfamily taxa (= 100.0 %)
Outgroup taxa ◊ › 37 Asiloidea species › 3 Nemestrinoidea › 1 Tabanomorpha
collecting localities of specimens included
.. methods
Apioceridae
Mydidae
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Morphological Characters211 discrete, parsimony informative characters◊
36 coded separately for female and male ◊ semaphoronts
tagma # characters remarksHead 44Thorax 103
46 chaetotaxy, sclerites, etc.24 wing33 leg
Abdomen 6213 sclerite shapes, chaetotaxy, etc.30 ? terminalia19 / terminalia
.. methods
8
Phylogenetic AnalysisData assembly and visualization ◊
› Mesquite (version 2.75) › FigTree (version 1.3.1)
› TreeMap3 (version 3.0b)
Tree Search ◊ › TNT (version 1.1 June 2012) › Ratchet, TreeDrifting
› all multi-state characters non-additive › all characters equally weighted
Bremer Support ◊ › TNT (version 1.1 June 2012)
Mydidae: Heteromydas bicolor © Greg Ballmer
Mydidae: Eremomidas arabicus © Drew Gardner
.. methods
9
Rhopalia annulata
Hoplistomerus nobilis
Dolichogaster brevicornis
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Apiocera Ap sp Stuart M
Mitrodetus microglossa
Apiocera An obscura
Miltinus maculipennis
Megascelus nigricornis
Hispanomydas hispanicus
Opomydas townsendi
Arenomydas callosus
Midacritus stuardoanus M
Nomoneuroides natalensis
Vespiodes phaios
Afroleptomydas C sp Stampriet
Syllegomydas S elachys
Apiocera Ap similis
Nemomydas pantherinus
Afroleptomydas A suffusipennis
Haplomydas crassipesLachnocorynus sp nov Kotwa M
Eremomidas sp nov Tiega M
Stratiomydas lividus
Syllegomydas sp nov Ruacana
Leptomydas sardous
Limata tenuicornis
Apiocera Ap pica
Cephalocera sp PapendorpCephalocera sp Ratelfontein
Nemestrinus nigrovillosus
Ectyphus pinguis
Nemomydas melanopogon
Neorhaphiomidas sp Yanchep
Hessemydas daugeroni
Apiophora quadricinctata
Nothomydas picketti
Bombylius major
Apiocera P mexicana M
Tongamya miranda
Prorates sp Escalante
Pseudonomoneura micheneri
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Mydaselpis sp nov Gamsberg M
Cleamydas gen nov sp nov Ugab F
Namibimydas stuckenbergi
Cretomydas santanensisNeorhaphiomidas sp Mandurah
Apiocera P barri
Anomalomydas australicus F
Euscelidia pulchra
Rhaphiomidas parkeri
Nomoneura caffra
Pseudonomoneura hirta
Apiocera P parahydra
Rhaphiomidas maehleri
Neorhaphiomidas sp Mingenew M
Nemomydas gruenbergi M
Namadytes sp Gobabeb M
Apiocera Ap brevicornis
Perissocerus dumonti F
Neolophonotus bimaculatus
Afroleptomydas C sp Gobabeb
Neorhaphiomidas hardyi
Apiocera An philippii
Perissocerus transcaspicus
Diochlistus melleipennis
Syllegomydas S algiricus
Apiocera P arena
Paramydas igniticornis
Apiocera Ap tonnoiri
Lasiopogon cinctus
Pseudorhopalia mirandai
Apiocera P painteri
Nomoneura paradoxa
Syllegomydas N heothinos
Apiocera Ap sp Wurarga
Afroleptomydas A omeri
Machimus occidentalis
Megascelus melanoproctus
Pegesimallus laticornis
Apiocera Ap imminuta
Eulonchus sp Six Rivers
Cacatuopyga ruficornis M
Nemomydas lamia
Leptomydas lusitanicus
Megascelus albovillosus
Apiocera P bilineata
Syllegomydas N arnoldi M
Midacritus wagenknechti
Afroleptomydas A paganus
Holcocephala calva
Apiocera P sp Colnett M
Ectyphus amboseli M
Halterorchis sp nov Cape Recife M
Mitrodetus nanoglossa M
Ospriocerus aeacus
Leptomydas turcicus
Pseudonomoneura bajaensis
Cephalocera longirostris
Eremomidas arabicusHalterorchis karooensis F
Apiocera Ap maritima
Syll gen nov sp nov minute proboscis F
Apiocera P aldrichi
Ectyphus armipes
Gonioscelis ventralis
Anomalomydas mackerrasi
Laphria aktis
Hessemydas parkeri M
Diochlistinae Argentina F
Miltinus viduatus
Neolaparopsis puncturatus
Heteromydas bicolor
Cacatuopyga basifascia
Pilica formidolosa
Leptomydas rapti M
Afroleptomydas A consanguinensis
Syllegomydas S proximus
Rhopalia olivieri
Apsilocephala longistyla
Apiocera P haruspex
Philonicus albiceps
Namibimydas psamminos
Syllegomydas S vittatus
Rhopalia cincta
Leptomydas corsicanus
Parectyphus namibiensis
Vespiodes leopolditertii
Mitrodetus dimidiatus
Rhaphiomidas hasbroucki
Namadytes vansoni
Phyllomydas bruesii
Mydaselpis sp nov Daan
Apiophora paulseni
Plyomydas sp nov Tandil
Neorhynchocephalus volaticus
Apiocera Ap asilica
Messiasia californica M
Emphysomera pallidapex
Apiocera Ap minor
Syll gen nov sp nov Gobabeb
Nemomydas brachyrhynchus
Eremomidas bek
Hessemydas seyrigi
Apiocera P parkeri
Diochlistus gracilis
Syllegomydas N astrictus
Messiasia painteri
Mydaselpis ngurumani
Apiocera R braunsi
Apiocera Ap sp Lloyd
Syllegomydas N dispar
Cephalocera sp Sors Sors
Diogmites grossus
Apiocera R alastor M
Eremohaplomydas desertorum
Hemigephyra atra
Plyomydas peruviensis M
Leptomydas notos
Arenomydas caerulescens
Syllegomydas S lineatus
Mydas clavatusMydas tricinctus
Damalis monochaetes
Ommatius tibialis
Syllegomydas S palestinensis
Stenomydas gen nov sp nov Panne M
Apiocera Ap hardyi
Dipalta serpentina
Arenomydas partitus
Leptogaster cylindrica
Apiocera P martinorum
Afroleptomydas A sp Clanwilliam
Arenomydas namaquensis M
Apiocera Ap newmani
Afroleptomydas A rufithorax
Miltinus minutus
Mydas xanthopterus
Cephalocera imitata M
Phyllomydas parvulus
Afroleptomydas C mauricei
Stichopogon punctum
Miltinus stenogaster
Toxophora maxima
Diochlistus auripennis M
Afroleptomydas A opacicinctus
Afroleptomydas A inhacae
Dioctria hyalipennis
Phycus frommeri
Apioceridae
Mydidae
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... new phylogenetic hypothesis
Results1040 most parsimonious trees◊
1845 steps◊
strict consensus ◊ › several polytomies › not well-resolved in derived ingroup
strict consensus of 1040 treesBremer 3 – 9Bremer ≥ 10
Rhopalia annulata
Hoplistomerus nobilis
Dolichogaster brevicornis
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Apiocera Ap sp Stuart M
Mitrodetus microglossa
Apiocera An obscura
Miltinus maculipennis
Megascelus nigricornis
Hispanomydas hispanicus
Opomydas townsendi
Arenomydas callosus
Midacritus stuardoanus M
Nomoneuroides natalensis
Vespiodes phaios
Afroleptomydas C sp Stampriet
Syllegomydas S elachys
Apiocera Ap similis
Nemomydas pantherinus
Afroleptomydas A suffusipennis
Haplomydas crassipesLachnocorynus sp nov Kotwa M
Eremomidas sp nov Tiega M
Stratiomydas lividus
Syllegomydas sp nov Ruacana
Leptomydas sardous
Limata tenuicornis
Apiocera Ap pica
Cephalocera sp PapendorpCephalocera sp Ratelfontein
Nemestrinus nigrovillosus
Ectyphus pinguis
Nemomydas melanopogon
Neorhaphiomidas sp Yanchep
Hessemydas daugeroni
Apiophora quadricinctata
Nothomydas picketti
Bombylius major
Apiocera P mexicana M
Tongamya miranda
Prorates sp Escalante
Pseudonomoneura micheneri
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Mydaselpis sp nov Gamsberg M
Cleamydas gen nov sp nov Ugab F
Namibimydas stuckenbergi
Cretomydas santanensisNeorhaphiomidas sp Mandurah
Apiocera P barri
Anomalomydas australicus F
Euscelidia pulchra
Rhaphiomidas parkeri
Nomoneura caffra
Pseudonomoneura hirta
Apiocera P parahydra
Rhaphiomidas maehleri
Neorhaphiomidas sp Mingenew M
Nemomydas gruenbergi M
Namadytes sp Gobabeb M
Apiocera Ap brevicornis
Perissocerus dumonti F
Neolophonotus bimaculatus
Afroleptomydas C sp Gobabeb
Neorhaphiomidas hardyi
Apiocera An philippii
Perissocerus transcaspicus
Diochlistus melleipennis
Syllegomydas S algiricus
Apiocera P arena
Paramydas igniticornis
Apiocera Ap tonnoiri
Lasiopogon cinctus
Pseudorhopalia mirandai
Apiocera P painteri
Nomoneura paradoxa
Syllegomydas N heothinos
Apiocera Ap sp Wurarga
Afroleptomydas A omeri
Machimus occidentalis
Megascelus melanoproctus
Pegesimallus laticornis
Apiocera Ap imminuta
Eulonchus sp Six Rivers
Cacatuopyga ruficornis M
Nemomydas lamia
Leptomydas lusitanicus
Megascelus albovillosus
Apiocera P bilineata
Syllegomydas N arnoldi M
Midacritus wagenknechti
Afroleptomydas A paganus
Holcocephala calva
Apiocera P sp Colnett M
Ectyphus amboseli M
Halterorchis sp nov Cape Recife M
Mitrodetus nanoglossa M
Ospriocerus aeacus
Leptomydas turcicus
Pseudonomoneura bajaensis
Cephalocera longirostris
Eremomidas arabicusHalterorchis karooensis F
Apiocera Ap maritima
Syll gen nov sp nov minute proboscis F
Apiocera P aldrichi
Ectyphus armipes
Gonioscelis ventralis
Anomalomydas mackerrasi
Laphria aktis
Hessemydas parkeri M
Diochlistinae Argentina F
Miltinus viduatus
Neolaparopsis puncturatus
Heteromydas bicolor
Cacatuopyga basifascia
Pilica formidolosa
Leptomydas rapti M
Afroleptomydas A consanguinensis
Syllegomydas S proximus
Rhopalia olivieri
Apsilocephala longistyla
Apiocera P haruspex
Philonicus albiceps
Namibimydas psamminos
Syllegomydas S vittatus
Rhopalia cincta
Leptomydas corsicanus
Parectyphus namibiensis
Vespiodes leopolditertii
Mitrodetus dimidiatus
Rhaphiomidas hasbroucki
Namadytes vansoni
Phyllomydas bruesii
Mydaselpis sp nov Daan
Apiophora paulseni
Plyomydas sp nov Tandil
Neorhynchocephalus volaticus
Apiocera Ap asilica
Messiasia californica M
Emphysomera pallidapex
Apiocera Ap minor
Syll gen nov sp nov Gobabeb
Nemomydas brachyrhynchus
Eremomidas bek
Hessemydas seyrigi
Apiocera P parkeri
Diochlistus gracilis
Syllegomydas N astrictus
Messiasia painteri
Mydaselpis ngurumani
Apiocera R braunsi
Apiocera Ap sp Lloyd
Syllegomydas N dispar
Cephalocera sp Sors Sors
Diogmites grossus
Apiocera R alastor M
Eremohaplomydas desertorum
Hemigephyra atra
Plyomydas peruviensis M
Leptomydas notos
Arenomydas caerulescens
Syllegomydas S lineatus
Mydas clavatusMydas tricinctus
Damalis monochaetes
Ommatius tibialis
Syllegomydas S palestinensis
Stenomydas gen nov sp nov Panne M
Apiocera Ap hardyi
Dipalta serpentina
Arenomydas partitus
Leptogaster cylindrica
Apiocera P martinorum
Afroleptomydas A sp Clanwilliam
Arenomydas namaquensis M
Apiocera Ap newmani
Afroleptomydas A rufithorax
Miltinus minutus
Mydas xanthopterus
Cephalocera imitata M
Phyllomydas parvulus
Afroleptomydas C mauricei
Stichopogon punctum
Miltinus stenogaster
Toxophora maxima
Diochlistus auripennis M
Afroleptomydas A opacicinctus
Afroleptomydas A inhacae
Dioctria hyalipennis
Phycus frommeri
10
Rhopalia annulata
Dolichogaster brevicornis
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Mitrodetus microglossa
Miltinus maculipennis
Megascelus nigricornis
Hispanomydas hispanicus
Opomydas townsendi
Arenomydas callosus
Midacritus stuardoanus M
Nomoneuroides natalensis
Vespiodes phaios
Afroleptomydas C sp Stampriet
Syllegomydas S elachys
Nemomydas pantherinus
Afroleptomydas A suffusipennis
Haplomydas crassipesLachnocorynus sp nov Kotwa M
Eremomidas sp nov Tiega M
Stratiomydas lividus
Syllegomydas sp nov Ruacana
Leptomydas sardous
Cephalocera sp PapendorpCephalocera sp Ratelfontein
Ectyphus pinguis
Nemomydas melanopogon
Neorhaphiomidas sp Yanchep
Hessemydas daugeroni
Apiophora quadricinctata
Nothomydas picketti
Tongamya miranda
Pseudonomoneura micheneri
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Mydaselpis sp nov Gamsberg M
Cleamydas gen nov sp nov Ugab F
Namibimydas stuckenbergi
Cretomydas santanensisNeorhaphiomidas sp Mandurah
Anomalomydas australicus F
Rhaphiomidas parkeri
Nomoneura caffra
Pseudonomoneura hirta
Rhaphiomidas maehleri
Neorhaphiomidas sp Mingenew M
Nemomydas gruenbergi M
Namadytes sp Gobabeb M
Perissocerus dumonti F
Afroleptomydas C sp Gobabeb
Neorhaphiomidas hardyi
Perissocerus transcaspicus
Diochlistus melleipennis
Syllegomydas S algiricus
Paramydas igniticornis
Pseudorhopalia mirandai
Nomoneura paradoxa
Syllegomydas N heothinos
Afroleptomydas A omeri
Megascelus melanoproctus
Cacatuopyga ruficornis M
Nemomydas lamia
Leptomydas lusitanicus
Megascelus albovillosus
Syllegomydas N arnoldi M
Midacritus wagenknechti
Afroleptomydas A paganus
Ectyphus amboseli M
Halterorchis sp nov Cape Recife M
Mitrodetus nanoglossa M
Leptomydas turcicus
Pseudonomoneura bajaensis
Cephalocera longirostris
Eremomidas arabicusHalterorchis karooensis F
Syll gen nov sp nov minute proboscis F
Ectyphus armipes
Anomalomydas mackerrasi
Hessemydas parkeri M
Diochlistinae Argentina F
Miltinus viduatus
Neolaparopsis puncturatus
Heteromydas bicolor
Cacatuopyga basifascia
Leptomydas rapti M
Afroleptomydas A consanguinensis
Syllegomydas S proximus
Rhopalia olivieri
Namibimydas psamminos
Syllegomydas S vittatus
Rhopalia cincta
Leptomydas corsicanus
Parectyphus namibiensis
Vespiodes leopolditertii
Mitrodetus dimidiatus
Rhaphiomidas hasbroucki
Namadytes vansoni
Phyllomydas bruesii
Mydaselpis sp nov Daan
Apiophora paulseni
Plyomydas sp nov TandilMessiasia californica M
Syll gen nov sp nov Gobabeb
Nemomydas brachyrhynchus
Eremomidas bek
Hessemydas seyrigi
Diochlistus gracilis
Syllegomydas N astrictus
Messiasia painteri
Mydaselpis ngurumani
Syllegomydas N dispar
Cephalocera sp Sors Sors
Eremohaplomydas desertorum
Plyomydas peruviensis M
Leptomydas notos
Arenomydas caerulescens
Syllegomydas S lineatus
Mydas clavatusMydas tricinctus
Syllegomydas S palestinensis
Stenomydas gen nov sp nov Panne M
Arenomydas partitus
Afroleptomydas A sp Clanwilliam
Arenomydas namaquensis M
Afroleptomydas A rufithorax
Miltinus minutus
Mydas xanthopterus
Cephalocera imitata M
Phyllomydas parvulus
Afroleptomydas C mauricei
Miltinus stenogaster
Diochlistus auripennis M
Afroleptomydas A opacicinctus
Afroleptomydas A inhacae
Australia
Argentina + Chile
† Cretomydas santanensis, Brazil
≈ 112 myoPlacement of † Cretomydas santanensis
Diochlistus auripennis, Australia © Peter Chew
... new phylogenetic hypothesis
Rhopalia annulata
Hoplistomerus nobilis
Dolichogaster brevicornis
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Apiocera Ap sp Stuart M
Mitrodetus microglossa
Apiocera An obscura
Miltinus maculipennis
Megascelus nigricornis
Hispanomydas hispanicus
Opomydas townsendi
Arenomydas callosus
Midacritus stuardoanus M
Nomoneuroides natalensis
Vespiodes phaios
Afroleptomydas C sp Stampriet
Syllegomydas S elachys
Apiocera Ap similis
Nemomydas pantherinus
Afroleptomydas A suffusipennis
Haplomydas crassipesLachnocorynus sp nov Kotwa M
Eremomidas sp nov Tiega M
Stratiomydas lividus
Syllegomydas sp nov Ruacana
Leptomydas sardous
Limata tenuicornis
Apiocera Ap pica
Cephalocera sp PapendorpCephalocera sp Ratelfontein
Nemestrinus nigrovillosus
Ectyphus pinguis
Nemomydas melanopogon
Neorhaphiomidas sp Yanchep
Hessemydas daugeroni
Apiophora quadricinctata
Nothomydas picketti
Bombylius major
Apiocera P mexicana M
Tongamya miranda
Prorates sp Escalante
Pseudonomoneura micheneri
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Mydaselpis sp nov Gamsberg M
Cleamydas gen nov sp nov Ugab F
Namibimydas stuckenbergi
Cretomydas santanensisNeorhaphiomidas sp Mandurah
Apiocera P barri
Anomalomydas australicus F
Euscelidia pulchra
Rhaphiomidas parkeri
Nomoneura caffra
Pseudonomoneura hirta
Apiocera P parahydra
Rhaphiomidas maehleri
Neorhaphiomidas sp Mingenew M
Nemomydas gruenbergi M
Namadytes sp Gobabeb M
Apiocera Ap brevicornis
Perissocerus dumonti F
Neolophonotus bimaculatus
Afroleptomydas C sp Gobabeb
Neorhaphiomidas hardyi
Apiocera An philippii
Perissocerus transcaspicus
Diochlistus melleipennis
Syllegomydas S algiricus
Apiocera P arena
Paramydas igniticornis
Apiocera Ap tonnoiri
Lasiopogon cinctus
Pseudorhopalia mirandai
Apiocera P painteri
Nomoneura paradoxa
Syllegomydas N heothinos
Apiocera Ap sp Wurarga
Afroleptomydas A omeri
Machimus occidentalis
Megascelus melanoproctus
Pegesimallus laticornis
Apiocera Ap imminuta
Eulonchus sp Six Rivers
Cacatuopyga ruficornis M
Nemomydas lamia
Leptomydas lusitanicus
Megascelus albovillosus
Apiocera P bilineata
Syllegomydas N arnoldi M
Midacritus wagenknechti
Afroleptomydas A paganus
Holcocephala calva
Apiocera P sp Colnett M
Ectyphus amboseli M
Halterorchis sp nov Cape Recife M
Mitrodetus nanoglossa M
Ospriocerus aeacus
Leptomydas turcicus
Pseudonomoneura bajaensis
Cephalocera longirostris
Eremomidas arabicusHalterorchis karooensis F
Apiocera Ap maritima
Syll gen nov sp nov minute proboscis F
Apiocera P aldrichi
Ectyphus armipes
Gonioscelis ventralis
Anomalomydas mackerrasi
Laphria aktis
Hessemydas parkeri M
Diochlistinae Argentina F
Miltinus viduatus
Neolaparopsis puncturatus
Heteromydas bicolor
Cacatuopyga basifascia
Pilica formidolosa
Leptomydas rapti M
Afroleptomydas A consanguinensis
Syllegomydas S proximus
Rhopalia olivieri
Apsilocephala longistyla
Apiocera P haruspex
Philonicus albiceps
Namibimydas psamminos
Syllegomydas S vittatus
Rhopalia cincta
Leptomydas corsicanus
Parectyphus namibiensis
Vespiodes leopolditertii
Mitrodetus dimidiatus
Rhaphiomidas hasbroucki
Namadytes vansoni
Phyllomydas bruesii
Mydaselpis sp nov Daan
Apiophora paulseni
Plyomydas sp nov Tandil
Neorhynchocephalus volaticus
Apiocera Ap asilica
Messiasia californica M
Emphysomera pallidapex
Apiocera Ap minor
Syll gen nov sp nov Gobabeb
Nemomydas brachyrhynchus
Eremomidas bek
Hessemydas seyrigi
Apiocera P parkeri
Diochlistus gracilis
Syllegomydas N astrictus
Messiasia painteri
Mydaselpis ngurumani
Apiocera R braunsi
Apiocera Ap sp Lloyd
Syllegomydas N dispar
Cephalocera sp Sors Sors
Diogmites grossus
Apiocera R alastor M
Eremohaplomydas desertorum
Hemigephyra atra
Plyomydas peruviensis M
Leptomydas notos
Arenomydas caerulescens
Syllegomydas S lineatus
Mydas clavatusMydas tricinctus
Damalis monochaetes
Ommatius tibialis
Syllegomydas S palestinensis
Stenomydas gen nov sp nov Panne M
Apiocera Ap hardyi
Dipalta serpentina
Arenomydas partitus
Leptogaster cylindrica
Apiocera P martinorum
Afroleptomydas A sp Clanwilliam
Arenomydas namaquensis M
Apiocera Ap newmani
Afroleptomydas A rufithorax
Miltinus minutus
Mydas xanthopterus
Cephalocera imitata M
Phyllomydas parvulus
Afroleptomydas C mauricei
Stichopogon punctum
Miltinus stenogaster
Toxophora maxima
Diochlistus auripennis M
Afroleptomydas A opacicinctus
Afroleptomydas A inhacae
Dioctria hyalipennis
Phycus frommeri
Non-monophyly of Diochlistinae
strict consensus of 1040 trees
11... new phylogenetic hypothesis
monophyletic
Syllegomydinae: Cephalocera imitata
Monophyly of subfamily taxa
Syllegomydinae (43 % species diversity)
Rhopalia annulata
Dolichogaster brevicornis
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Mitrodetus microglossa
Miltinus maculipennis
Megascelus nigricornis
Hispanomydas hispanicus
Opomydas townsendi
Arenomydas callosus
Midacritus stuardoanus M
Nomoneuroides natalensis
Vespiodes phaios
Afroleptomydas C sp Stampriet
Syllegomydas S elachys
Nemomydas pantherinus
Afroleptomydas A suffusipennis
Haplomydas crassipesLachnocorynus sp nov Kotwa M
Eremomidas sp nov Tiega M
Stratiomydas lividus
Syllegomydas sp nov Ruacana
Leptomydas sardous
Cephalocera sp PapendorpCephalocera sp Ratelfontein
Ectyphus pinguis
Nemomydas melanopogon
Neorhaphiomidas sp Yanchep
Hessemydas daugeroni
Apiophora quadricinctata
Nothomydas picketti
Tongamya miranda
Pseudonomoneura micheneri
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Mydaselpis sp nov Gamsberg M
Cleamydas gen nov sp nov Ugab F
Namibimydas stuckenbergi
Cretomydas santanensisNeorhaphiomidas sp Mandurah
Anomalomydas australicus F
Rhaphiomidas parkeri
Nomoneura caffra
Pseudonomoneura hirta
Rhaphiomidas maehleri
Neorhaphiomidas sp Mingenew M
Nemomydas gruenbergi M
Namadytes sp Gobabeb M
Perissocerus dumonti F
Afroleptomydas C sp Gobabeb
Neorhaphiomidas hardyi
Perissocerus transcaspicus
Diochlistus melleipennis
Syllegomydas S algiricus
Paramydas igniticornis
Pseudorhopalia mirandai
Nomoneura paradoxa
Syllegomydas N heothinos
Afroleptomydas A omeri
Megascelus melanoproctus
Cacatuopyga ruficornis M
Nemomydas lamia
Leptomydas lusitanicus
Megascelus albovillosus
Syllegomydas N arnoldi M
Midacritus wagenknechti
Afroleptomydas A paganus
Ectyphus amboseli M
Halterorchis sp nov Cape Recife M
Mitrodetus nanoglossa M
Leptomydas turcicus
Pseudonomoneura bajaensis
Cephalocera longirostris
Eremomidas arabicusHalterorchis karooensis F
Syll gen nov sp nov minute proboscis F
Ectyphus armipes
Anomalomydas mackerrasi
Hessemydas parkeri M
Diochlistinae Argentina F
Miltinus viduatus
Neolaparopsis puncturatus
Heteromydas bicolor
Cacatuopyga basifascia
Leptomydas rapti M
Afroleptomydas A consanguinensis
Syllegomydas S proximus
Rhopalia olivieri
Namibimydas psamminos
Syllegomydas S vittatus
Rhopalia cincta
Leptomydas corsicanus
Parectyphus namibiensis
Vespiodes leopolditertii
Mitrodetus dimidiatus
Rhaphiomidas hasbroucki
Namadytes vansoni
Phyllomydas bruesii
Mydaselpis sp nov Daan
Apiophora paulseni
Plyomydas sp nov TandilMessiasia californica M
Syll gen nov sp nov Gobabeb
Nemomydas brachyrhynchus
Eremomidas bek
Hessemydas seyrigi
Diochlistus gracilis
Syllegomydas N astrictus
Messiasia painteri
Mydaselpis ngurumani
Syllegomydas N dispar
Cephalocera sp Sors Sors
Eremohaplomydas desertorum
Plyomydas peruviensis M
Leptomydas notos
Arenomydas caerulescens
Syllegomydas S lineatus
Mydas clavatusMydas tricinctus
Syllegomydas S palestinensis
Stenomydas gen nov sp nov Panne M
Arenomydas partitus
Afroleptomydas A sp Clanwilliam
Arenomydas namaquensis M
Afroleptomydas A rufithorax
Miltinus minutus
Mydas xanthopterus
Cephalocera imitata M
Phyllomydas parvulus
Afroleptomydas C mauricei
Miltinus stenogaster
Diochlistus auripennis M
Afroleptomydas A opacicinctus
Afroleptomydas A inhacae
Rhaphiomidinae
Megascelinae
Anomalomydinae
Cacatuopyginae
Mydinae
Rhopalia annulata
Hoplistomerus nobilis
Dolichogaster brevicornis
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Apiocera Ap sp Stuart M
Mitrodetus microglossa
Apiocera An obscura
Miltinus maculipennis
Megascelus nigricornis
Hispanomydas hispanicus
Opomydas townsendi
Arenomydas callosus
Midacritus stuardoanus M
Nomoneuroides natalensis
Vespiodes phaios
Afroleptomydas C sp Stampriet
Syllegomydas S elachys
Apiocera Ap similis
Nemomydas pantherinus
Afroleptomydas A suffusipennis
Haplomydas crassipesLachnocorynus sp nov Kotwa M
Eremomidas sp nov Tiega M
Stratiomydas lividus
Syllegomydas sp nov Ruacana
Leptomydas sardous
Limata tenuicornis
Apiocera Ap pica
Cephalocera sp PapendorpCephalocera sp Ratelfontein
Nemestrinus nigrovillosus
Ectyphus pinguis
Nemomydas melanopogon
Neorhaphiomidas sp Yanchep
Hessemydas daugeroni
Apiophora quadricinctata
Nothomydas picketti
Bombylius major
Apiocera P mexicana M
Tongamya miranda
Prorates sp Escalante
Pseudonomoneura micheneri
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Mydaselpis sp nov Gamsberg M
Cleamydas gen nov sp nov Ugab F
Namibimydas stuckenbergi
Cretomydas santanensisNeorhaphiomidas sp Mandurah
Apiocera P barri
Anomalomydas australicus F
Euscelidia pulchra
Rhaphiomidas parkeri
Nomoneura caffra
Pseudonomoneura hirta
Apiocera P parahydra
Rhaphiomidas maehleri
Neorhaphiomidas sp Mingenew M
Nemomydas gruenbergi M
Namadytes sp Gobabeb M
Apiocera Ap brevicornis
Perissocerus dumonti F
Neolophonotus bimaculatus
Afroleptomydas C sp Gobabeb
Neorhaphiomidas hardyi
Apiocera An philippii
Perissocerus transcaspicus
Diochlistus melleipennis
Syllegomydas S algiricus
Apiocera P arena
Paramydas igniticornis
Apiocera Ap tonnoiri
Lasiopogon cinctus
Pseudorhopalia mirandai
Apiocera P painteri
Nomoneura paradoxa
Syllegomydas N heothinos
Apiocera Ap sp Wurarga
Afroleptomydas A omeri
Machimus occidentalis
Megascelus melanoproctus
Pegesimallus laticornis
Apiocera Ap imminuta
Eulonchus sp Six Rivers
Cacatuopyga ruficornis M
Nemomydas lamia
Leptomydas lusitanicus
Megascelus albovillosus
Apiocera P bilineata
Syllegomydas N arnoldi M
Midacritus wagenknechti
Afroleptomydas A paganus
Holcocephala calva
Apiocera P sp Colnett M
Ectyphus amboseli M
Halterorchis sp nov Cape Recife M
Mitrodetus nanoglossa M
Ospriocerus aeacus
Leptomydas turcicus
Pseudonomoneura bajaensis
Cephalocera longirostris
Eremomidas arabicusHalterorchis karooensis F
Apiocera Ap maritima
Syll gen nov sp nov minute proboscis F
Apiocera P aldrichi
Ectyphus armipes
Gonioscelis ventralis
Anomalomydas mackerrasi
Laphria aktis
Hessemydas parkeri M
Diochlistinae Argentina F
Miltinus viduatus
Neolaparopsis puncturatus
Heteromydas bicolor
Cacatuopyga basifascia
Pilica formidolosa
Leptomydas rapti M
Afroleptomydas A consanguinensis
Syllegomydas S proximus
Rhopalia olivieri
Apsilocephala longistyla
Apiocera P haruspex
Philonicus albiceps
Namibimydas psamminos
Syllegomydas S vittatus
Rhopalia cincta
Leptomydas corsicanus
Parectyphus namibiensis
Vespiodes leopolditertii
Mitrodetus dimidiatus
Rhaphiomidas hasbroucki
Namadytes vansoni
Phyllomydas bruesii
Mydaselpis sp nov Daan
Apiophora paulseni
Plyomydas sp nov Tandil
Neorhynchocephalus volaticus
Apiocera Ap asilica
Messiasia californica M
Emphysomera pallidapex
Apiocera Ap minor
Syll gen nov sp nov Gobabeb
Nemomydas brachyrhynchus
Eremomidas bek
Hessemydas seyrigi
Apiocera P parkeri
Diochlistus gracilis
Syllegomydas N astrictus
Messiasia painteri
Mydaselpis ngurumani
Apiocera R braunsi
Apiocera Ap sp Lloyd
Syllegomydas N dispar
Cephalocera sp Sors Sors
Diogmites grossus
Apiocera R alastor M
Eremohaplomydas desertorum
Hemigephyra atra
Plyomydas peruviensis M
Leptomydas notos
Arenomydas caerulescens
Syllegomydas S lineatus
Mydas clavatusMydas tricinctus
Damalis monochaetes
Ommatius tibialis
Syllegomydas S palestinensis
Stenomydas gen nov sp nov Panne M
Apiocera Ap hardyi
Dipalta serpentina
Arenomydas partitus
Leptogaster cylindrica
Apiocera P martinorum
Afroleptomydas A sp Clanwilliam
Arenomydas namaquensis M
Apiocera Ap newmani
Afroleptomydas A rufithorax
Miltinus minutus
Mydas xanthopterus
Cephalocera imitata M
Phyllomydas parvulus
Afroleptomydas C mauricei
Stichopogon punctum
Miltinus stenogaster
Toxophora maxima
Diochlistus auripennis M
Afroleptomydas A opacicinctus
Afroleptomydas A inhacae
Dioctria hyalipennis
Phycus frommeri
strict consensus of 1040 trees
12
Rhopalia annulata
Hoplistomerus nobilis
Dolichogaster brevicornis
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Apiocera Ap sp Stuart M
Mitrodetus microglossa
Apiocera An obscura
Miltinus maculipennis
Megascelus nigricornis
Hispanomydas hispanicus
Opomydas townsendi
Arenomydas callosus
Midacritus stuardoanus M
Nomoneuroides natalensis
Vespiodes phaios
Afroleptomydas C sp Stampriet
Syllegomydas S elachys
Apiocera Ap similis
Nemomydas pantherinus
Afroleptomydas A suffusipennis
Haplomydas crassipesLachnocorynus sp nov Kotwa M
Eremomidas sp nov Tiega M
Stratiomydas lividus
Syllegomydas sp nov Ruacana
Leptomydas sardous
Limata tenuicornis
Apiocera Ap pica
Cephalocera sp PapendorpCephalocera sp Ratelfontein
Nemestrinus nigrovillosus
Ectyphus pinguis
Nemomydas melanopogon
Neorhaphiomidas sp Yanchep
Hessemydas daugeroni
Apiophora quadricinctata
Nothomydas picketti
Bombylius major
Apiocera P mexicana M
Tongamya miranda
Prorates sp Escalante
Pseudonomoneura micheneri
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Mydaselpis sp nov Gamsberg M
Cleamydas gen nov sp nov Ugab F
Namibimydas stuckenbergi
Cretomydas santanensisNeorhaphiomidas sp Mandurah
Apiocera P barri
Anomalomydas australicus F
Euscelidia pulchra
Rhaphiomidas parkeri
Nomoneura caffra
Pseudonomoneura hirta
Apiocera P parahydra
Rhaphiomidas maehleri
Neorhaphiomidas sp Mingenew M
Nemomydas gruenbergi M
Namadytes sp Gobabeb M
Apiocera Ap brevicornis
Perissocerus dumonti F
Neolophonotus bimaculatus
Afroleptomydas C sp Gobabeb
Neorhaphiomidas hardyi
Apiocera An philippii
Perissocerus transcaspicus
Diochlistus melleipennis
Syllegomydas S algiricus
Apiocera P arena
Paramydas igniticornis
Apiocera Ap tonnoiri
Lasiopogon cinctus
Pseudorhopalia mirandai
Apiocera P painteri
Nomoneura paradoxa
Syllegomydas N heothinos
Apiocera Ap sp Wurarga
Afroleptomydas A omeri
Machimus occidentalis
Megascelus melanoproctus
Pegesimallus laticornis
Apiocera Ap imminuta
Eulonchus sp Six Rivers
Cacatuopyga ruficornis M
Nemomydas lamia
Leptomydas lusitanicus
Megascelus albovillosus
Apiocera P bilineata
Syllegomydas N arnoldi M
Midacritus wagenknechti
Afroleptomydas A paganus
Holcocephala calva
Apiocera P sp Colnett M
Ectyphus amboseli M
Halterorchis sp nov Cape Recife M
Mitrodetus nanoglossa M
Ospriocerus aeacus
Leptomydas turcicus
Pseudonomoneura bajaensis
Cephalocera longirostris
Eremomidas arabicusHalterorchis karooensis F
Apiocera Ap maritima
Syll gen nov sp nov minute proboscis F
Apiocera P aldrichi
Ectyphus armipes
Gonioscelis ventralis
Anomalomydas mackerrasi
Laphria aktis
Hessemydas parkeri M
Diochlistinae Argentina F
Miltinus viduatus
Neolaparopsis puncturatus
Heteromydas bicolor
Cacatuopyga basifascia
Pilica formidolosa
Leptomydas rapti M
Afroleptomydas A consanguinensis
Syllegomydas S proximus
Rhopalia olivieri
Apsilocephala longistyla
Apiocera P haruspex
Philonicus albiceps
Namibimydas psamminos
Syllegomydas S vittatus
Rhopalia cincta
Leptomydas corsicanus
Parectyphus namibiensis
Vespiodes leopolditertii
Mitrodetus dimidiatus
Rhaphiomidas hasbroucki
Namadytes vansoni
Phyllomydas bruesii
Mydaselpis sp nov Daan
Apiophora paulseni
Plyomydas sp nov Tandil
Neorhynchocephalus volaticus
Apiocera Ap asilica
Messiasia californica M
Emphysomera pallidapex
Apiocera Ap minor
Syll gen nov sp nov Gobabeb
Nemomydas brachyrhynchus
Eremomidas bek
Hessemydas seyrigi
Apiocera P parkeri
Diochlistus gracilis
Syllegomydas N astrictus
Messiasia painteri
Mydaselpis ngurumani
Apiocera R braunsi
Apiocera Ap sp Lloyd
Syllegomydas N dispar
Cephalocera sp Sors Sors
Diogmites grossus
Apiocera R alastor M
Eremohaplomydas desertorum
Hemigephyra atra
Plyomydas peruviensis M
Leptomydas notos
Arenomydas caerulescens
Syllegomydas S lineatus
Mydas clavatusMydas tricinctus
Damalis monochaetes
Ommatius tibialis
Syllegomydas S palestinensis
Stenomydas gen nov sp nov Panne M
Apiocera Ap hardyi
Dipalta serpentina
Arenomydas partitus
Leptogaster cylindrica
Apiocera P martinorum
Afroleptomydas A sp Clanwilliam
Arenomydas namaquensis M
Apiocera Ap newmani
Afroleptomydas A rufithorax
Miltinus minutus
Mydas xanthopterus
Cephalocera imitata M
Phyllomydas parvulus
Afroleptomydas C mauricei
Stichopogon punctum
Miltinus stenogaster
Toxophora maxima
Diochlistus auripennis M
Afroleptomydas A opacicinctus
Afroleptomydas A inhacae
Dioctria hyalipennis
Phycus frommeri
Mydidae: Rhaphiomidas auratus © Greg Ballmer
Mydidae: Anomalomydas mackerrasi © Jean Hort Mydidae: Neorhaphiomidas sp. © Fred Hort
Earliest branchings within Mydidae
... new phylogenetic hypothesis
Megascelus nigricornisNeorhaphiomidas sp Yanchep
Tongamya miranda
† Cretomydas santanensisNeorhaphiomidas sp Mandurah
Anomalomydas australicus F
Rhaphiomidas parkeriRhaphiomidas maehleri
Neorhaphiomidas sp Mingenew MNeorhaphiomidas hardyi
Diochlistus melleipennis
Megascelus melanoproctusMegascelus albovillosus
Anomalomydas mackerrasi
Rhaphiomidas hasbroucki
Diochlistus gracilisDiochlistus auripennis M
western North AmericaNamibia + South Africa
BrazilAustralia
Australia
remaining Mydidae
Rhaphiomidinae
Megascelinae
Anomalomydinae
part Diochlistinae
Australia
Chile
13
What could be the reason for the poorly resolved topology?Is the taxon sampling inappropriate?◊
Does a weighted analysis (PIWE) improve resolution?◊
Does the presence of sexually dimorphic characters add “noise”?◊
Does taxon pruning with IterPCR improve resolution?◊
Did the apomorphic Mydidae experience a sudden radiation intractable for the morphological ◊ data used?
.... study of topology
14
Sexual dimorphism – separate analysesSyllegomydas vittatus
? /Males only
› 128 taxa (- 6)
› 172 characters (- 39)
› 30 char. ? terminalia
› 2989 mp trees
› 1373 steps
Females only
› 114 taxa (- 20)
› 162 characters (- 49)
› 19 char. / terminalia
› 2226 mp trees
› 1362 steps
36 characters
.... study of topology
15
Rhopalia olivieri
Nomoneura caffraNomoneura paradoxa
Phyllomydas bruesii
Afroleptomydas A omeri
Mydas tricinctus
Rhopalia annulata
Afroleptomydas A inhacae
Ectyphus pinguis
Midacritus wagenknechti
Syllegomydas S palestinensis
Pseudonomoneura bajaensis
Nothomydas picketti
Namibimydas psamminos
Leptomydas corsicanus
Afroleptomydas A sp Clanwilliam
Syllegomydas N heothinos
Syll gen nov sp nov minute proboscis F
Apiophora paulseni
Leptomydas turcicus
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Cleamydas gen nov sp nov Ugab F
Mydas xanthopterus
Perissocerus dumonti F
Afroleptomydas A sp Obib
Vespiodes phaios
Syllegomydas S proximus
Syllegomydas N astrictus
Neorhaphiomidas hardyi
Anomalomydas mackerrasi
Cephalocera longirostris
Leptomydas sardous
Notosyllegomydas brincki
Miltinus minutus
Stratiomydas lividus
Mitrodetus dentitarsis
Namadytes vansoni
Nemomydas melanopogon
Cephalocera sp Ratelfontein
Diochlistus melleipennis
Rhaphiomidas parkeri
Paramydas igniticornis
Afroleptomydas A rufithorax
Miltinus stenogaster
Megascelus melanoproctus
Syllegomydas sp nov Ruacana
Neorhaphiomidas sp Mandurah
Afroleptomydas C mauricei
Afroleptomydas A suffusipennis
Perissocerus transcaspicus
Afroleptomydas A consanguinensis
Neolaparopsis puncturatus
Phyllomydas parvulus
Cretomydas santanensis
Afroleptomydas C sp Gobabeb
Hispanomydas hispanicus
Haplomydas crassipes
Neorhaphiomidas sp Yanchep
Diochlistinae Argentina F
Pseudorhopalia mirandai
Syll gen nov sp nov Gobabeb
Syllegomydas N dispar
Arenomydas partitus
Arenomydas callosus
Rhaphiomidas maehleri
Cephalocera sp Papendorp
Anomalomydas australicus F
Vespiodes leopolditertii
Mitrodetus microglossa
Diochlistus gracilis
Plyomydas sp nov Tandil
Hessemydas daugeroni
Nemomydas lamia
Syllegomydas S elachys
Heteromydas bicolor
Cacatuopyga basifascia
Eremomidas bek
Namibimydas stuckenbergi
Mydas clavatus
Rhaphiomidas hasbroucki
Nomoneuroides natalensis
Cephalocera sp Sors Sors
Opomydas townsendi
Rhopalia cincta
Mitrodetus dimidiatus
Halterorchis karooensis F
Leptomydas notos
Megascelus nigricornis
Syllegomydas S lineatus
Afroleptomydas C sp Stampriet
Messiasia painteri
Mydaselpis sp nov Daan
Mydaselpis ngurumani
Syllegomydas S algiricus
Nemomydas brachyrhynchus
Arenomydas caerulescens
Afroleptomydas A opacicinctus
Eremomidas arabicus
Tongamya mirandaMegascelus albovillosus
Hessemydas seyrigi
Syllegomydas S vittatus
Apiophora quadricinctata
Pseudonomoneura micheneriPseudonomoneura hirta
Miltinus maculipennis
Parectyphus namibiensis
Ectyphus armipes
Dolichogaster brevicornis
Afroleptomydas A paganus
Eremohaplomydas desertorum
Miltinus viduatus
Leptomydas lusitanicus
Nemomydas pantherinus
Namadytes vansoni
Diochlistus auripennis M
Midacritus wagenknechti
Arenomydas callosus
Rhopalia cincta
Afroleptomydas A rufithorax
Nemomydas lamia
Megascelus melanoproctus
Neorhaphiomidas sp Yanchep
Perissocerus transcaspicus
Neorhaphiomidas sp Mingenew M
Diochlistus gracilis
Lachnocorynus sp nov Kotwa M
Plyomydas sp nov Tandil
Namibimydas psamminos
Syllegomydas N astrictus
Cephalocera longirostris
Rhaphiomidas hasbroucki
Rhopalia olivieri
Cacatuopyga ruficornis M
Stenomydas gen nov sp nov Panne M
Mydaselpis ngurumani
Syllegomydas S algiricus
Afroleptomydas A consanguinensis
Nemomydas pantherinus
Mydas clavatus
Hessemydas daugeroni
Afroleptomydas A opacicinctus
Phyllomydas bruesii
Afroleptomydas C mauricei
Midacritus stuardoanus M
Mitrodetus nanoglossa M
Messiasia californica M
Nemomydas gruenbergi M
Cephalocera sp Papendorp
Mydas tricinctus
Opomydas townsendi
Afroleptomydas C sp Stampriet
Plyomydas peruviensis M
Anomalomydas mackerrasi
Pseudorhopalia mirandai
Ectyphus armipes
Neorhaphiomidas sp Mandurah
Eremohaplomydas desertorum
Paramydas igniticornis
Namibimydas stuckenbergi
Leptomydas rapti M
Messiasia painteri
Pseudonomoneura micheneri
Dolichogaster brevicornis
Eremomidas bek
Cephalocera sp Sors Sors
Arenomydas caerulescens
Tongamya miranda
Mydaselpis sp nov Gamsberg M
Heteromydas bicolor
Leptomydas sardous
Phyllomydas parvulus
Neolaparopsis puncturatus
Syllegomydas N dispar
Afroleptomydas A inhacae
Vespiodes leopolditertii
Afroleptomydas C sp Gobabeb
Miltinus viduatus
Cephalocera imitata M
Hessemydas parkeri M
Nemomydas venosus
Leptomydas lusitanicus
Afroleptomydas A sp Clanwilliam
Miltinus minutus
Nemomydas brachyrhynchus
Afroleptomydas C sp Koeroegab
Apiophora paulseni
Namadytes sp Gobabeb M
Hessemydas seyrigi
Mitrodetus dimidiatus
Parectyphus namibiensis
Afroleptomydas A suffusipennis
Apiophora quadricinctata
Rhopalia annulata
Stratiomydas lividus
Neorhaphiomidas hardyi
Nothomydas picketti
Mydas xanthopterus
Miltinus stenogaster
Vespiodes phaios
Syllegomydas S vittatus
Mydaselpis sp nov Daan
Syllegomydas S elachys
Syllegomydas S proximus
Cretomydas santanensis
Haplomydas crassipes
Syllegomydas N arnoldi M
Diochlistus melleipennis
Nomoneura caffra
Cephalocera sp Ratelfontein
Nemomydas melanopogon
Megascelus nigricornis
Miltinus maculipennis
Hispanomydas hispanicus
Syllegomydas S lineatus
Arenomydas namaquensis M
Mitrodetus microglossa
Syllegomydas sp nov Ruacana
Eremomidas arabicus
Nomoneuroides natalensis
Syll gen nov sp nov Gobabeb
Pseudonomoneura hirta
Nomoneura paradoxa
Ectyphus amboseli M
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Cacatuopyga basifascia
Afroleptomydas A omeri
Afroleptomydas A paganus
Rhaphiomidas maehleri
Syllegomydas N heothinos
Eremomidas sp nov Tiega M
Arenomydas partitus
Leptomydas corsicanusLeptomydas notos
Leptomydas turcicus
Syllegomydas S palestinensis
Pseudonomoneura bajaensis
Rhaphiomidas parkeriMegascelus albovillosus
Halterorchis sp nov Cape Recife M
Ectyphus pinguis
Males Females
strict consensus of 2989 trees strict consensus of 2226 treesmonophyleticSyllegomydinae
Rhopalia annulata
Hoplistomerus nobilis
Dolichogaster brevicornis
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Apiocera Ap sp Stuart M
Mitrodetus microglossa
Apiocera An obscura
Miltinus maculipennis
Megascelus nigricornis
Hispanomydas hispanicus
Opomydas townsendi
Arenomydas callosus
Midacritus stuardoanus M
Nomoneuroides natalensis
Vespiodes phaios
Afroleptomydas C sp Stampriet
Syllegomydas S elachys
Apiocera Ap similis
Nemomydas pantherinus
Afroleptomydas A suffusipennis
Haplomydas crassipesLachnocorynus sp nov Kotwa M
Eremomidas sp nov Tiega M
Stratiomydas lividus
Syllegomydas sp nov Ruacana
Leptomydas sardous
Limata tenuicornis
Apiocera Ap pica
Cephalocera sp PapendorpCephalocera sp Ratelfontein
Nemestrinus nigrovillosus
Ectyphus pinguis
Nemomydas melanopogon
Neorhaphiomidas sp Yanchep
Hessemydas daugeroni
Apiophora quadricinctata
Nothomydas picketti
Bombylius major
Apiocera P mexicana M
Tongamya miranda
Prorates sp Escalante
Pseudonomoneura micheneri
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Mydaselpis sp nov Gamsberg M
Cleamydas gen nov sp nov Ugab F
Namibimydas stuckenbergi
Cretomydas santanensisNeorhaphiomidas sp Mandurah
Apiocera P barri
Anomalomydas australicus F
Euscelidia pulchra
Rhaphiomidas parkeri
Nomoneura caffra
Pseudonomoneura hirta
Apiocera P parahydra
Rhaphiomidas maehleri
Neorhaphiomidas sp Mingenew M
Nemomydas gruenbergi M
Namadytes sp Gobabeb M
Apiocera Ap brevicornis
Perissocerus dumonti F
Neolophonotus bimaculatus
Afroleptomydas C sp Gobabeb
Neorhaphiomidas hardyi
Apiocera An philippii
Perissocerus transcaspicus
Diochlistus melleipennis
Syllegomydas S algiricus
Apiocera P arena
Paramydas igniticornis
Apiocera Ap tonnoiri
Lasiopogon cinctus
Pseudorhopalia mirandai
Apiocera P painteri
Nomoneura paradoxa
Syllegomydas N heothinos
Apiocera Ap sp Wurarga
Afroleptomydas A omeri
Machimus occidentalis
Megascelus melanoproctus
Pegesimallus laticornis
Apiocera Ap imminuta
Eulonchus sp Six Rivers
Cacatuopyga ruficornis M
Nemomydas lamia
Leptomydas lusitanicus
Megascelus albovillosus
Apiocera P bilineata
Syllegomydas N arnoldi M
Midacritus wagenknechti
Afroleptomydas A paganus
Holcocephala calva
Apiocera P sp Colnett M
Ectyphus amboseli M
Halterorchis sp nov Cape Recife M
Mitrodetus nanoglossa M
Ospriocerus aeacus
Leptomydas turcicus
Pseudonomoneura bajaensis
Cephalocera longirostris
Eremomidas arabicusHalterorchis karooensis F
Apiocera Ap maritima
Syll gen nov sp nov minute proboscis F
Apiocera P aldrichi
Ectyphus armipes
Gonioscelis ventralis
Anomalomydas mackerrasi
Laphria aktis
Hessemydas parkeri M
Diochlistinae Argentina F
Miltinus viduatus
Neolaparopsis puncturatus
Heteromydas bicolor
Cacatuopyga basifascia
Pilica formidolosa
Leptomydas rapti M
Afroleptomydas A consanguinensis
Syllegomydas S proximus
Rhopalia olivieri
Apsilocephala longistyla
Apiocera P haruspex
Philonicus albiceps
Namibimydas psamminos
Syllegomydas S vittatus
Rhopalia cincta
Leptomydas corsicanus
Parectyphus namibiensis
Vespiodes leopolditertii
Mitrodetus dimidiatus
Rhaphiomidas hasbroucki
Namadytes vansoni
Phyllomydas bruesii
Mydaselpis sp nov Daan
Apiophora paulseni
Plyomydas sp nov Tandil
Neorhynchocephalus volaticus
Apiocera Ap asilica
Messiasia californica M
Emphysomera pallidapex
Apiocera Ap minor
Syll gen nov sp nov Gobabeb
Nemomydas brachyrhynchus
Eremomidas bek
Hessemydas seyrigi
Apiocera P parkeri
Diochlistus gracilis
Syllegomydas N astrictus
Messiasia painteri
Mydaselpis ngurumani
Apiocera R braunsi
Apiocera Ap sp Lloyd
Syllegomydas N dispar
Cephalocera sp Sors Sors
Diogmites grossus
Apiocera R alastor M
Eremohaplomydas desertorum
Hemigephyra atra
Plyomydas peruviensis M
Leptomydas notos
Arenomydas caerulescens
Syllegomydas S lineatus
Mydas clavatusMydas tricinctus
Damalis monochaetes
Ommatius tibialis
Syllegomydas S palestinensis
Stenomydas gen nov sp nov Panne M
Apiocera Ap hardyi
Dipalta serpentina
Arenomydas partitus
Leptogaster cylindrica
Apiocera P martinorum
Afroleptomydas A sp Clanwilliam
Arenomydas namaquensis M
Apiocera Ap newmani
Afroleptomydas A rufithorax
Miltinus minutus
Mydas xanthopterus
Cephalocera imitata M
Phyllomydas parvulus
Afroleptomydas C mauricei
Stichopogon punctum
Miltinus stenogaster
Toxophora maxima
Diochlistus auripennis M
Afroleptomydas A opacicinctus
Afroleptomydas A inhacae
Dioctria hyalipennis
Phycus frommeri
Separate analyses – results
.... study of topology
16
Males FemalesSeparate analyses – TreeMap3fully resolved trees needed ◊
› 1 mp tree chosen
same # of taxa needed ◊ › species linked within genus
tanglegram not “untangled”◊
.... study of topology
17
Rhopalia annulata
Miltinus stenogaster
Nothomydas picketti
Afroleptomydas A omeri
Diochlistus gracilis
Notosyllegomydas brinckiSyllegomydas N astrictus
Nemomydas venosus
Ectyphus amboseli M
Mydas clavatus
Cretomydas santanensis
Nemomydas gruenbergi M
Afroleptomydas A sp ClanwilliamAfroleptomydas A sp Obib
Syllegomydas N arnoldi M
Syll gen nov sp nov Gobabeb
Syllegomydas N disparHessemydas parkeri M
Afroleptomydas C sp Koeroegab
Neorhaphiomidas hardyi
Leptomydas turcicus
Pseudonomoneura hirta
Miltinus maculipennis
Neolaparopsis puncturatus
Nomoneuroides natalensis
Pseudonomoneura micheneri
Nemomydas melanopogon
Dolichogaster brevicornis
Nemomydas brachyrhynchus
Lachnocorynus sp nov Kotwa M
Namadytes sp Gobabeb M
Afroleptomydas A rufithorax
Eremomidas sp nov Tiega M
Nomoneura caffra
Anomalomydas australicus F
Rhaphiomidas parkeri
Syllegomydas S elachys
Rhaphiomidas hasbroucki
Plyomydas peruviensis M
Leptomydas sardous
Tongamya miranda
Apiophora quadricinctata
Afroleptomydas A opacicinctus
Vespiodes phaios
Mydas tricinctus
Parectyphus namibiensis
Plyomydas sp nov Tandil
Rhopalia cincta
Syllegomydas S vittatus
Megascelus melanoproctus
Cephalocera longirostris
Nemomydas pantherinus
Phyllomydas parvulusLeptomydas corsicanus
Pseudorhopalia mirandai
Mitrodetus dentitarsis
Megascelus nigricornis
Rhopalia olivieri
Diochlistus melleipennis
Cephalocera sp Papendorp
Eremomidas arabicus
Mydaselpis ngurumani
Mydaselpis sp nov Daan
Neorhaphiomidas sp Mandurah
Syllegomydas S palestinensis
Hessemydas seyrigi
Phyllomydas bruesii
Afroleptomydas A suffusipennis
Syll gen nov sp nov minute proboscis F
Megascelus albovillosus
Hessemydas daugeroni
Messiasia painteri
Syllegomydas S algiricus
Stratiomydas lividus
Afroleptomydas A paganus
Cleamydas gen nov sp nov Ugab F
Eremomidas bek
Afroleptomydas A consanguinensis
Diochlistinae Argentina F
Apiophora paulseni
Opomydas townsendi
Mitrodetus nanoglossa M
Miltinus minutus
Leptomydas rapti M
Namadytes vansoni
Namibimydas stuckenbergiNamibimydas psamminos
Paramydas igniticornis
Cacatuopyga basifascia
Midacritus stuardoanus M
Nemomydas lamia
Arenomydas caerulescens
Syllegomydas sp nov Ruacana
Cephalocera sp Sors Sors
Halterorchis karooensis F
Neorhaphiomidas sp Mingenew M
Ectyphus pinguis
Hispanomydas hispanicus
Diochlistus auripennis M
Afroleptomydas C mauricei
Miltinus viduatus
Rhaphiomidas maehleri
Pseudonomoneura bajaensis
Cephalocera imitata M
Heteromydas bicolor
Syllegomydas S lineatus
Mydas xanthopterus
Ectyphus armipes
Leptomydas notos
Arenomydas partitus
Anomalomydas mackerrasi
Cacatuopyga ruficornis M
Arenomydas callosus
Messiasia californica M
Mitrodetus dimidiatus
Cephalocera sp Ratelfontein
Neorhaphiomidas sp Yanchep
Halterorchis sp nov Cape Recife M
Nomoneura paradoxa
Haplomydas crassipes
Vespiodes leopolditertii
Afroleptomydas C sp GobabebAfroleptomydas C sp Stampriet
Perissocerus dumonti F
Leptomydas lusitanicus
Midacritus wagenknechti
Perissocerus transcaspicus
Mitrodetus microglossa
Syllegomydas N heothinos
Eremohaplomydas desertorum
Afroleptomydas A inhacae
Syllegomydas S proximus
Mydaselpis sp nov Gamsberg M
Stenomydas gen nov sp nov Panne M
Arenomydas namaquensis M
monophyleticSyllegomydinae
Rhopalia annulata
Hoplistomerus nobilis
Dolichogaster brevicornis
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Apiocera Ap sp Stuart M
Mitrodetus microglossa
Apiocera An obscura
Miltinus maculipennis
Megascelus nigricornis
Hispanomydas hispanicus
Opomydas townsendi
Arenomydas callosus
Midacritus stuardoanus M
Nomoneuroides natalensis
Vespiodes phaios
Afroleptomydas C sp Stampriet
Syllegomydas S elachys
Apiocera Ap similis
Nemomydas pantherinus
Afroleptomydas A suffusipennis
Haplomydas crassipesLachnocorynus sp nov Kotwa M
Eremomidas sp nov Tiega M
Stratiomydas lividus
Syllegomydas sp nov Ruacana
Leptomydas sardous
Limata tenuicornis
Apiocera Ap pica
Cephalocera sp PapendorpCephalocera sp Ratelfontein
Nemestrinus nigrovillosus
Ectyphus pinguis
Nemomydas melanopogon
Neorhaphiomidas sp Yanchep
Hessemydas daugeroni
Apiophora quadricinctata
Nothomydas picketti
Bombylius major
Apiocera P mexicana M
Tongamya miranda
Prorates sp Escalante
Pseudonomoneura micheneri
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Mydaselpis sp nov Gamsberg M
Cleamydas gen nov sp nov Ugab F
Namibimydas stuckenbergi
Cretomydas santanensisNeorhaphiomidas sp Mandurah
Apiocera P barri
Anomalomydas australicus F
Euscelidia pulchra
Rhaphiomidas parkeri
Nomoneura caffra
Pseudonomoneura hirta
Apiocera P parahydra
Rhaphiomidas maehleri
Neorhaphiomidas sp Mingenew M
Nemomydas gruenbergi M
Namadytes sp Gobabeb M
Apiocera Ap brevicornis
Perissocerus dumonti F
Neolophonotus bimaculatus
Afroleptomydas C sp Gobabeb
Neorhaphiomidas hardyi
Apiocera An philippii
Perissocerus transcaspicus
Diochlistus melleipennis
Syllegomydas S algiricus
Apiocera P arena
Paramydas igniticornis
Apiocera Ap tonnoiri
Lasiopogon cinctus
Pseudorhopalia mirandai
Apiocera P painteri
Nomoneura paradoxa
Syllegomydas N heothinos
Apiocera Ap sp Wurarga
Afroleptomydas A omeri
Machimus occidentalis
Megascelus melanoproctus
Pegesimallus laticornis
Apiocera Ap imminuta
Eulonchus sp Six Rivers
Cacatuopyga ruficornis M
Nemomydas lamia
Leptomydas lusitanicus
Megascelus albovillosus
Apiocera P bilineata
Syllegomydas N arnoldi M
Midacritus wagenknechti
Afroleptomydas A paganus
Holcocephala calva
Apiocera P sp Colnett M
Ectyphus amboseli M
Halterorchis sp nov Cape Recife M
Mitrodetus nanoglossa M
Ospriocerus aeacus
Leptomydas turcicus
Pseudonomoneura bajaensis
Cephalocera longirostris
Eremomidas arabicusHalterorchis karooensis F
Apiocera Ap maritima
Syll gen nov sp nov minute proboscis F
Apiocera P aldrichi
Ectyphus armipes
Gonioscelis ventralis
Anomalomydas mackerrasi
Laphria aktis
Hessemydas parkeri M
Diochlistinae Argentina F
Miltinus viduatus
Neolaparopsis puncturatus
Heteromydas bicolor
Cacatuopyga basifascia
Pilica formidolosa
Leptomydas rapti M
Afroleptomydas A consanguinensis
Syllegomydas S proximus
Rhopalia olivieri
Apsilocephala longistyla
Apiocera P haruspex
Philonicus albiceps
Namibimydas psamminos
Syllegomydas S vittatus
Rhopalia cincta
Leptomydas corsicanus
Parectyphus namibiensis
Vespiodes leopolditertii
Mitrodetus dimidiatus
Rhaphiomidas hasbroucki
Namadytes vansoni
Phyllomydas bruesii
Mydaselpis sp nov Daan
Apiophora paulseni
Plyomydas sp nov Tandil
Neorhynchocephalus volaticus
Apiocera Ap asilica
Messiasia californica M
Emphysomera pallidapex
Apiocera Ap minor
Syll gen nov sp nov Gobabeb
Nemomydas brachyrhynchus
Eremomidas bek
Hessemydas seyrigi
Apiocera P parkeri
Diochlistus gracilis
Syllegomydas N astrictus
Messiasia painteri
Mydaselpis ngurumani
Apiocera R braunsi
Apiocera Ap sp Lloyd
Syllegomydas N dispar
Cephalocera sp Sors Sors
Diogmites grossus
Apiocera R alastor M
Eremohaplomydas desertorum
Hemigephyra atra
Plyomydas peruviensis M
Leptomydas notos
Arenomydas caerulescens
Syllegomydas S lineatus
Mydas clavatusMydas tricinctus
Damalis monochaetes
Ommatius tibialis
Syllegomydas S palestinensis
Stenomydas gen nov sp nov Panne M
Apiocera Ap hardyi
Dipalta serpentina
Arenomydas partitus
Leptogaster cylindrica
Apiocera P martinorum
Afroleptomydas A sp Clanwilliam
Arenomydas namaquensis M
Apiocera Ap newmani
Afroleptomydas A rufithorax
Miltinus minutus
Mydas xanthopterus
Cephalocera imitata M
Phyllomydas parvulus
Afroleptomydas C mauricei
Stichopogon punctum
Miltinus stenogaster
Toxophora maxima
Diochlistus auripennis M
Afroleptomydas A opacicinctus
Afroleptomydas A inhacae
Dioctria hyalipennis
Phycus frommeri
Excluding dimorphic charactersall 134 taxa◊
175 characters ◊ (- 36)
› including 30 ? terminalia characters › including 19 / terminalia characters
3000 mp trees◊
1367 steps◊
strict consensus of 3000 trees
.... study of topology
18
Cephalocera imitata M
Phyllomydas bruesii
Syllegomydas S elachys
Ectyphus pinguis
Arenomydas callosus
Diochlistus melleipennis
Leptomydas turcicus
Afr C sp Gobabeb
Cacatuopyga ruf
Haplomydas crassipes
Afr A rufithorax
Miltinus minutus
Afr C sp Stampriet
Paramydas igniticornis
Diochlistus aur
Midacritus stu
Neorhaphiomidas Yan
Dolichogaster brevicornis
Syllegomydas N heothinosSyll proboscis F
Mydaselpis sp nov Daan
Midacritus wag
Hessemydas seyrigi
Mitrodetus nanoglossa M
Rhopalia olivieri
Anomalomydas mac
Ectyphus armipes
Hessemydas parkeri M
Mitrodetus microglossa
Arenomydas cae
Afr A opacicinctus
Plyomydas sp nov Tandil
Rhaphiomidas hasbroucki
Pseudorhopalia mirandai
Namadytes vansoni
Pseudonomoneura baj
Stenomydas Panne M
Hessemydas daugeroni
Nomoneuroides natalensis
Mitrodetus dentitarsis
Pseudonomoneura hirta
Namibimydas stuckenbergi
Megascelus albovillosus
Perissocerus tra
Halterorchis Cape Recife
Syll gen nov sp nov Gobabeb
Syllegomydas sp nov Ruacana
Notosyllegomydas brincki
Apiophora qua
Megascelus nigricornis
Lachnocorynus Kotwa
Neorhaphiomidas hardyi
Opomydas townsendi
Miltinus mac
Nemomydas venosus
Syllegomydas N dispar
Messiasia californica M
Nomoneura paradoxa
Cephalocera sp Rat
Nemomydas melanopogon
Neorhaphiomidas Man
Mydaselpis Gamsberg
Neorhaphiomidas Min
Heteromydas bicolor
Syllegomydas N astrictus
Nemomydas bra
Mitrodetus dimidiatus
Rhopalia cincta
Hispanomydas hispanicus
Parectyphus namibiensis
Perissocerus dumonti F
Anomalomydas aus
Eremohaplomydas des
Afr C sp Koeroegab
Cephalocera lon
Phyllomydas parvulus
Syllegomydas S lineatus
Miltinus stenogaster
Eremomidas sp nov
Afr A inhacae
Cleamydas Ugab F
Stratiomydas lividus
Eremomidas arabicus
Cacatuopyga basifascia
Vespiodes leopolditertii
Neolaparopsis puncturatus
Megascelus mel
Diochlistinae Argentina F
Namadytes sp Gobabeb M
Rhopalia annulata
Tongamya miranda
Leptomydas sardous
Syllegomydas S proximus
Leptomydas rapti M
Mydaselpis ngurumani
Leptomydas lusitanicus
Arenomydas partitus
Cephalocera sp Pap
Nomoneura caffra
Afr A paganus
Syllegomydas S algiricus
Syllegomydas N arnoldi M
Messiasia painteri
Miltinus viduatus
Leptomydas notos
Nemomydas gru
Mydas xanthopterus
Vespiodes phaios
Apiophora paulseni
Afr A omeri
Cephalocera sp Sors Sors
Cretomydas santanensis
Ectyphus amboseli M
Halterorchis karooensis F
Afr A sp Clanwilliam
Syllegomydas S vittatus
Namibimydas psamminos
Rhaphiomidas parkeri
Afr A consanguinensis
Rhaphiomidas maehleri
Afr C mauricei
Nemomydas lamiaPseudonomoneura mic
Mydas clavatus
Nemomydas pantherinus
Nothomydas picketti
Diochlistus gracilis
Mydas tricinctus
Afr A suffusipennis
Afr A sp Obib
Leptomydas corsicanus
Plyomydas peruviensis M
Eremomidas bek
monophyleticSyllegomydinae
Rhopalia annulata
Hoplistomerus nobilis
Dolichogaster brevicornis
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Apiocera Ap sp Stuart M
Mitrodetus microglossa
Apiocera An obscura
Miltinus maculipennis
Megascelus nigricornis
Hispanomydas hispanicus
Opomydas townsendi
Arenomydas callosus
Midacritus stuardoanus M
Nomoneuroides natalensis
Vespiodes phaios
Afroleptomydas C sp Stampriet
Syllegomydas S elachys
Apiocera Ap similis
Nemomydas pantherinus
Afroleptomydas A suffusipennis
Haplomydas crassipesLachnocorynus sp nov Kotwa M
Eremomidas sp nov Tiega M
Stratiomydas lividus
Syllegomydas sp nov Ruacana
Leptomydas sardous
Limata tenuicornis
Apiocera Ap pica
Cephalocera sp PapendorpCephalocera sp Ratelfontein
Nemestrinus nigrovillosus
Ectyphus pinguis
Nemomydas melanopogon
Neorhaphiomidas sp Yanchep
Hessemydas daugeroni
Apiophora quadricinctata
Nothomydas picketti
Bombylius major
Apiocera P mexicana M
Tongamya miranda
Prorates sp Escalante
Pseudonomoneura micheneri
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Mydaselpis sp nov Gamsberg M
Cleamydas gen nov sp nov Ugab F
Namibimydas stuckenbergi
Cretomydas santanensisNeorhaphiomidas sp Mandurah
Apiocera P barri
Anomalomydas australicus F
Euscelidia pulchra
Rhaphiomidas parkeri
Nomoneura caffra
Pseudonomoneura hirta
Apiocera P parahydra
Rhaphiomidas maehleri
Neorhaphiomidas sp Mingenew M
Nemomydas gruenbergi M
Namadytes sp Gobabeb M
Apiocera Ap brevicornis
Perissocerus dumonti F
Neolophonotus bimaculatus
Afroleptomydas C sp Gobabeb
Neorhaphiomidas hardyi
Apiocera An philippii
Perissocerus transcaspicus
Diochlistus melleipennis
Syllegomydas S algiricus
Apiocera P arena
Paramydas igniticornis
Apiocera Ap tonnoiri
Lasiopogon cinctus
Pseudorhopalia mirandai
Apiocera P painteri
Nomoneura paradoxa
Syllegomydas N heothinos
Apiocera Ap sp Wurarga
Afroleptomydas A omeri
Machimus occidentalis
Megascelus melanoproctus
Pegesimallus laticornis
Apiocera Ap imminuta
Eulonchus sp Six Rivers
Cacatuopyga ruficornis M
Nemomydas lamia
Leptomydas lusitanicus
Megascelus albovillosus
Apiocera P bilineata
Syllegomydas N arnoldi M
Midacritus wagenknechti
Afroleptomydas A paganus
Holcocephala calva
Apiocera P sp Colnett M
Ectyphus amboseli M
Halterorchis sp nov Cape Recife M
Mitrodetus nanoglossa M
Ospriocerus aeacus
Leptomydas turcicus
Pseudonomoneura bajaensis
Cephalocera longirostris
Eremomidas arabicusHalterorchis karooensis F
Apiocera Ap maritima
Syll gen nov sp nov minute proboscis F
Apiocera P aldrichi
Ectyphus armipes
Gonioscelis ventralis
Anomalomydas mackerrasi
Laphria aktis
Hessemydas parkeri M
Diochlistinae Argentina F
Miltinus viduatus
Neolaparopsis puncturatus
Heteromydas bicolor
Cacatuopyga basifascia
Pilica formidolosa
Leptomydas rapti M
Afroleptomydas A consanguinensis
Syllegomydas S proximus
Rhopalia olivieri
Apsilocephala longistyla
Apiocera P haruspex
Philonicus albiceps
Namibimydas psamminos
Syllegomydas S vittatus
Rhopalia cincta
Leptomydas corsicanus
Parectyphus namibiensis
Vespiodes leopolditertii
Mitrodetus dimidiatus
Rhaphiomidas hasbroucki
Namadytes vansoni
Phyllomydas bruesii
Mydaselpis sp nov Daan
Apiophora paulseni
Plyomydas sp nov Tandil
Neorhynchocephalus volaticus
Apiocera Ap asilica
Messiasia californica M
Emphysomera pallidapex
Apiocera Ap minor
Syll gen nov sp nov Gobabeb
Nemomydas brachyrhynchus
Eremomidas bek
Hessemydas seyrigi
Apiocera P parkeri
Diochlistus gracilis
Syllegomydas N astrictus
Messiasia painteri
Mydaselpis ngurumani
Apiocera R braunsi
Apiocera Ap sp Lloyd
Syllegomydas N dispar
Cephalocera sp Sors Sors
Diogmites grossus
Apiocera R alastor M
Eremohaplomydas desertorum
Hemigephyra atra
Plyomydas peruviensis M
Leptomydas notos
Arenomydas caerulescens
Syllegomydas S lineatus
Mydas clavatusMydas tricinctus
Damalis monochaetes
Ommatius tibialis
Syllegomydas S palestinensis
Stenomydas gen nov sp nov Panne M
Apiocera Ap hardyi
Dipalta serpentina
Arenomydas partitus
Leptogaster cylindrica
Apiocera P martinorum
Afroleptomydas A sp Clanwilliam
Arenomydas namaquensis M
Apiocera Ap newmani
Afroleptomydas A rufithorax
Miltinus minutus
Mydas xanthopterus
Cephalocera imitata M
Phyllomydas parvulus
Afroleptomydas C mauricei
Stichopogon punctum
Miltinus stenogaster
Toxophora maxima
Diochlistus auripennis M
Afroleptomydas A opacicinctus
Afroleptomydas A inhacae
Dioctria hyalipennis
Phycus frommeri
Taxon pruning with IterPCR in TNT132 taxa ◊ (- 2)
› Syllegomydas palestinensis › Arenomydas namaquensis (male only)
all 211 characters◊
1431 mp trees◊
1819 steps◊
strict consensus of 1431 trees
.... study of topology
19
Rhopalia annulata
Dolichogaster brevicornis
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Mitrodetus microglossa
Miltinus maculipennis
Megascelus nigricornis
Hispanomydas hispanicus
Opomydas townsendi
Arenomydas callosus
Midacritus stuardoanus M
Nomoneuroides natalensis
Vespiodes phaios
Afroleptomydas C sp Stampriet
Syllegomydas S elachys
Nemomydas pantherinus
Afroleptomydas A suffusipennis
Haplomydas crassipesLachnocorynus sp nov Kotwa M
Eremomidas sp nov Tiega M
Stratiomydas lividus
Syllegomydas sp nov Ruacana
Leptomydas sardous
Cephalocera sp PapendorpCephalocera sp Ratelfontein
Ectyphus pinguis
Nemomydas melanopogon
Neorhaphiomidas sp Yanchep
Hessemydas daugeroni
Apiophora quadricinctata
Nothomydas picketti
Tongamya miranda
Pseudonomoneura micheneri
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Mydaselpis sp nov Gamsberg M
Cleamydas gen nov sp nov Ugab F
Namibimydas stuckenbergi
Cretomydas santanensisNeorhaphiomidas sp Mandurah
Anomalomydas australicus F
Rhaphiomidas parkeri
Nomoneura caffra
Pseudonomoneura hirta
Rhaphiomidas maehleri
Neorhaphiomidas sp Mingenew M
Nemomydas gruenbergi M
Namadytes sp Gobabeb M
Perissocerus dumonti F
Afroleptomydas C sp Gobabeb
Neorhaphiomidas hardyi
Perissocerus transcaspicus
Diochlistus melleipennis
Syllegomydas S algiricus
Paramydas igniticornis
Pseudorhopalia mirandai
Nomoneura paradoxa
Syllegomydas N heothinos
Afroleptomydas A omeri
Megascelus melanoproctus
Cacatuopyga ruficornis M
Nemomydas lamia
Leptomydas lusitanicus
Megascelus albovillosus
Syllegomydas N arnoldi M
Midacritus wagenknechti
Afroleptomydas A paganus
Ectyphus amboseli M
Halterorchis sp nov Cape Recife M
Mitrodetus nanoglossa M
Leptomydas turcicus
Pseudonomoneura bajaensis
Cephalocera longirostris
Eremomidas arabicusHalterorchis karooensis F
Syll gen nov sp nov minute proboscis F
Ectyphus armipes
Anomalomydas mackerrasi
Hessemydas parkeri M
Diochlistinae Argentina F
Miltinus viduatus
Neolaparopsis puncturatus
Heteromydas bicolor
Cacatuopyga basifascia
Leptomydas rapti M
Afroleptomydas A consanguinensis
Syllegomydas S proximus
Rhopalia olivieri
Namibimydas psamminos
Syllegomydas S vittatus
Rhopalia cincta
Leptomydas corsicanus
Parectyphus namibiensis
Vespiodes leopolditertii
Mitrodetus dimidiatus
Rhaphiomidas hasbroucki
Namadytes vansoni
Phyllomydas bruesii
Mydaselpis sp nov Daan
Apiophora paulseni
Plyomydas sp nov TandilMessiasia californica M
Syll gen nov sp nov Gobabeb
Nemomydas brachyrhynchus
Eremomidas bek
Hessemydas seyrigi
Diochlistus gracilis
Syllegomydas N astrictus
Messiasia painteri
Mydaselpis ngurumani
Syllegomydas N dispar
Cephalocera sp Sors Sors
Eremohaplomydas desertorum
Plyomydas peruviensis M
Leptomydas notos
Arenomydas caerulescens
Syllegomydas S lineatus
Mydas clavatusMydas tricinctus
Syllegomydas S palestinensis
Stenomydas gen nov sp nov Panne M
Arenomydas partitus
Afroleptomydas A sp Clanwilliam
Arenomydas namaquensis M
Afroleptomydas A rufithorax
Miltinus minutus
Mydas xanthopterus
Cephalocera imitata M
Phyllomydas parvulus
Afroleptomydas C mauricei
Miltinus stenogaster
Diochlistus auripennis M
Afroleptomydas A opacicinctus
Afroleptomydas A inhacae
monophyleticSyllegomydinae
Rhopalia annulata
Hoplistomerus nobilis
Dolichogaster brevicornis
Mitrodetus dentitarsis
Notosyllegomydas brincki
Afroleptomydas A sp Obib
Apiocera Ap sp Stuart M
Mitrodetus microglossa
Apiocera An obscura
Miltinus maculipennis
Megascelus nigricornis
Hispanomydas hispanicus
Opomydas townsendi
Arenomydas callosus
Midacritus stuardoanus M
Nomoneuroides natalensis
Vespiodes phaios
Afroleptomydas C sp Stampriet
Syllegomydas S elachys
Apiocera Ap similis
Nemomydas pantherinus
Afroleptomydas A suffusipennis
Haplomydas crassipesLachnocorynus sp nov Kotwa M
Eremomidas sp nov Tiega M
Stratiomydas lividus
Syllegomydas sp nov Ruacana
Leptomydas sardous
Limata tenuicornis
Apiocera Ap pica
Cephalocera sp PapendorpCephalocera sp Ratelfontein
Nemestrinus nigrovillosus
Ectyphus pinguis
Nemomydas melanopogon
Neorhaphiomidas sp Yanchep
Hessemydas daugeroni
Apiophora quadricinctata
Nothomydas picketti
Bombylius major
Apiocera P mexicana M
Tongamya miranda
Prorates sp Escalante
Pseudonomoneura micheneri
Afroleptomydas C sp Koeroegab
Nemomydas venosus
Mydaselpis sp nov Gamsberg M
Cleamydas gen nov sp nov Ugab F
Namibimydas stuckenbergi
Cretomydas santanensisNeorhaphiomidas sp Mandurah
Apiocera P barri
Anomalomydas australicus F
Euscelidia pulchra
Rhaphiomidas parkeri
Nomoneura caffra
Pseudonomoneura hirta
Apiocera P parahydra
Rhaphiomidas maehleri
Neorhaphiomidas sp Mingenew M
Nemomydas gruenbergi M
Namadytes sp Gobabeb M
Apiocera Ap brevicornis
Perissocerus dumonti F
Neolophonotus bimaculatus
Afroleptomydas C sp Gobabeb
Neorhaphiomidas hardyi
Apiocera An philippii
Perissocerus transcaspicus
Diochlistus melleipennis
Syllegomydas S algiricus
Apiocera P arena
Paramydas igniticornis
Apiocera Ap tonnoiri
Lasiopogon cinctus
Pseudorhopalia mirandai
Apiocera P painteri
Nomoneura paradoxa
Syllegomydas N heothinos
Apiocera Ap sp Wurarga
Afroleptomydas A omeri
Machimus occidentalis
Megascelus melanoproctus
Pegesimallus laticornis
Apiocera Ap imminuta
Eulonchus sp Six Rivers
Cacatuopyga ruficornis M
Nemomydas lamia
Leptomydas lusitanicus
Megascelus albovillosus
Apiocera P bilineata
Syllegomydas N arnoldi M
Midacritus wagenknechti
Afroleptomydas A paganus
Holcocephala calva
Apiocera P sp Colnett M
Ectyphus amboseli M
Halterorchis sp nov Cape Recife M
Mitrodetus nanoglossa M
Ospriocerus aeacus
Leptomydas turcicus
Pseudonomoneura bajaensis
Cephalocera longirostris
Eremomidas arabicusHalterorchis karooensis F
Apiocera Ap maritima
Syll gen nov sp nov minute proboscis F
Apiocera P aldrichi
Ectyphus armipes
Gonioscelis ventralis
Anomalomydas mackerrasi
Laphria aktis
Hessemydas parkeri M
Diochlistinae Argentina F
Miltinus viduatus
Neolaparopsis puncturatus
Heteromydas bicolor
Cacatuopyga basifascia
Pilica formidolosa
Leptomydas rapti M
Afroleptomydas A consanguinensis
Syllegomydas S proximus
Rhopalia olivieri
Apsilocephala longistyla
Apiocera P haruspex
Philonicus albiceps
Namibimydas psamminos
Syllegomydas S vittatus
Rhopalia cincta
Leptomydas corsicanus
Parectyphus namibiensis
Vespiodes leopolditertii
Mitrodetus dimidiatus
Rhaphiomidas hasbroucki
Namadytes vansoni
Phyllomydas bruesii
Mydaselpis sp nov Daan
Apiophora paulseni
Plyomydas sp nov Tandil
Neorhynchocephalus volaticus
Apiocera Ap asilica
Messiasia californica M
Emphysomera pallidapex
Apiocera Ap minor
Syll gen nov sp nov Gobabeb
Nemomydas brachyrhynchus
Eremomidas bek
Hessemydas seyrigi
Apiocera P parkeri
Diochlistus gracilis
Syllegomydas N astrictus
Messiasia painteri
Mydaselpis ngurumani
Apiocera R braunsi
Apiocera Ap sp Lloyd
Syllegomydas N dispar
Cephalocera sp Sors Sors
Diogmites grossus
Apiocera R alastor M
Eremohaplomydas desertorum
Hemigephyra atra
Plyomydas peruviensis M
Leptomydas notos
Arenomydas caerulescens
Syllegomydas S lineatus
Mydas clavatusMydas tricinctus
Damalis monochaetes
Ommatius tibialis
Syllegomydas S palestinensis
Stenomydas gen nov sp nov Panne M
Apiocera Ap hardyi
Dipalta serpentina
Arenomydas partitus
Leptogaster cylindrica
Apiocera P martinorum
Afroleptomydas A sp Clanwilliam
Arenomydas namaquensis M
Apiocera Ap newmani
Afroleptomydas A rufithorax
Miltinus minutus
Mydas xanthopterus
Cephalocera imitata M
Phyllomydas parvulus
Afroleptomydas C mauricei
Stichopogon punctum
Miltinus stenogaster
Toxophora maxima
Diochlistus auripennis M
Afroleptomydas A opacicinctus
Afroleptomydas A inhacae
Dioctria hyalipennis
Phycus frommeri
strict consensus of 1040 trees
Preferred hypothesis – all data and semaphorontsMydidae is an old taxon◊
more recently evolved taxa appear to have speciated with little ◊ morphological diversification
?◊ terminalia within speciose Syllegomydinae taxa are virtually identical
Steps for further enhancing topology and resolution◊ › addition of internal head and thoracic characters › addition of molecular data
.... study of topology
20..... summary
Summary
Mydidae: Phyllomydas parvulus © Lyn Atherton
Apioceridae + Mydidae monophyletic◊
(Apioceridae + Mydidae) + Asilidae◊
earliest branchings within Mydidae well-supported◊
speciose taxa Leptomydinae and Syllegomydinae not ◊ monophyletic
subfamily taxa relationships still not resolved◊
additional work needed and in particular molecular data◊
21
AcknowledgementsMuseum curators and collectors who made specimens available◊
Photographers◊
U.S. National Science Foundation (REVSYS DEB-0919333)◊