1

Phylogeny and historical biogeography of Apioceridae and Mydidae inferred from morphological characters of imagines (Diptera: Asiloidea)

Torsten DikowBiodiversity Synthesis Center Field Museum of Natural History, Chicago, IL

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 monophyletic Apioceridae

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”Imagines flower-feeders & larvae possibly predatory◊

138 species and 1 genus ◊ (4 subgenera)

Imagines 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

Fossil ◊ † Protapiocera 3 species › † Protapioceridae – sister-group of Apioceridae? › Upper Yixian Formation, north-eastern China › Jurassic: Tithonian › ≈ 145 – 150 myo † Protapiocera convergens Zhang et al. 2004

. introduction

5. introduction

Mydidae: Opomydas townsendi

Mydidae – “mydas flies”Imagines flower-feeders, some do not feed at all◊

Larvae possibly predatory◊

469 species and 66 genera world-wide◊

Imagines 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: † Cretomydas santanensis

6

Overview of presentationSet out to postulate cladistic relationships within Mydidae ◊

Methods ◊

New Phylogenetic Hypothesis ◊

Biogeographically interesting taxa ◊

Summary + Future◊

Miltinus stenogaster © Shaun Winterton

Apiocera (Pyrocera) bilineata © Greg Ballmer

7

Taxon Sampling27 ◊ Apiocera species (= 19.6 %)

› 2 Apiocera (Anypenus) (= 50.0 %)

› 11 Apiocera (Apiocera) (= 16.4 %)

› 11 Apiocera (Pyrocera) (= 17.2 %)

› 2 Apiocera (Ripidosyrma) (= 66.7 %)

120 Mydidae species◊ (= 25.6 %) › 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

8

Morphological Characters206 discrete, parsimony informative characters ◊

tagma # characters remarksHead 42Thorax 101

45 chaetotaxy, sclerites, etc.23 wing33 leg

Abdomen 6211 sclerite shapes, chaetotaxy, etc.32 ? terminalia20 / genitalia

.. methods

9

Phylogenetic AnalysisMorphological data ◊

› Mesquite (version 2.75) › MacClade (version 4.08) › WinClada / Asado (version 1.89)

Tree Search ◊ › TNT (version 1.1 August 2011) › Ratchet, TreeDrifting (Nixon 1999, Goloboff 1999)

› all multi-state characters non-additive › all characters equally weighted

Bremer Support ◊ › TNT (version 1.1 August 2011)

Mydidae: Heteromydas bicolor © Greg Ballmer

Mydidae: Eremomidas arabicus © Drew Gardner

.. methods

10Limata tenuicornisEulonchus sp Six RiversNeorhynchocephalus volaticusNemestrinus nigrovillosusBombylius majorDipalta serpentinaToxophora maximaProrates sp EscalanteApsilocephala longistylaHemigephyra atraPhycus frommeriDiogmites grossusOspriocerus aeacusGonioscelis ventralisHoplistomerus nobilisPilica formidolosaLaphria aktisEmphysomera pallidapexOmmatius tibialisNeolophonotus bimaculatusMachimus occidentalisPhilonicus albicepsPegesimallus laticornisDioctria hyalipennisLeptogaster cylindricaEuscelidia pulchraStichopogon punctumLasiopogon cinctusDamalis monochaetesHolcocephala calvaApiocera R alastor maleApiocera R braunsiApiocera P bilineataApiocera P mexicana maleApiocera P parahydraApiocera P barriApiocera P martinorumApiocera P haruspexApiocera P aldrichiApiocera P painteriApiocera P arenaApiocera P parkeriApiocera P sp Colnett maleApiocera Ap imminutaApiocera Ap tonnoiriApiocera An obscuraApiocera An philippiiApiocera Ap sp WurargaApiocera Ap asilicaApiocera Ap brevicornisApiocera Ap sp Stuart maleApiocera Ap hardyiApiocera Ap newmaniApiocera Ap sp LloydApiocera Ap maritimaApiocera Ap similisRhaphiomidas hasbrouckiRhaphiomidas maehleriRhaphiomidas parkeriTongamya mirandaMegascelus albovillosusMegascelus nigricornisNeorhaphiomidas setosa maleNeorhaphiomidas sp MandurahCretomydas santanensisDiochlistus auripennis maleDiochlistus gracilisDiochlistus melleipennisAnomalomydas mackerrasiAnomalomydas australicus femaleParamydas igniticornisPlyomydas peruviensis malePlyomydas sp nov TandilMessiasia californica maleMessiasia painteriMidacritus stuardoanus maleMidacritus wagenknechtiCacatuopyga basifasciaCacatuopyga ruficornis maleHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus armipesEctyphus amboseli maleEctyphus pinguisApiophora paulseniApiophora quadricinctataMiltinus stenogasterMiltinus maculipennisMiltinus viduatusMydas clavatusMydas xanthopterusStratiomydas lividusDolichogaster brevicornisPhyllomydas bruesiiPhyllomydas parvulusPseudorhopalia mirandaiRhopalia annulataRhopalia cinctaRhopalia olivieriPerissocerus dumonti femalePerissocerus transcaspicusHessemydas parkeri maleHessemydas seyrigiNomoneura caffraNomoneura paradoxaNomoneuroides natalensisMydaselpis ngurumaniArenomydas partitusArenomydas caerulescensArenomydas callosusArenomydas namaquensis maleNeolaparopsis puncturatusHaplomydas crassipesEremohaplomydas desertorumLachnocorynus sp nov Kotwa maleMydaselpis sp nov DaanMydaselpis sp nov Gamsberg maleVespiodes leopolditertiiVespiodes phaiosNemomydas brachyrhynchusNemomydas venosusLeptomydas corsicanusNemomydas gruenbergi maleLeptomydas lusitanicusEremomidas bekLeptomydas rapti maleLeptomydas turcicusLeptomydas sardousLeptomydas notosNemomydas pantherinusPseudonomoneura hirtaPseudonomoneura micheneriNemomydas lamiaNotosyllegomydas brinckiNamibimydas stuckenbergi sp novNothomydas picketti sp novCephalocera longirostrisCephalocera sp PapendorpCephalocera sp Sors SorsCephalocera imitata maleCephalocera sp RatelfonteinNemomydas melanopogonNamadytes sp Gobabeb maleNamadytes vansoniAfroleptomydas C mauriceiAfroleptomydas C sp GobabebAfroleptomydas C sp StamprietAfroleptomydas A inhacaeAfroleptomydas A consanguinensisAfroleptomydas A opacicinctusAfroleptomydas A paganusAfroleptomydas A suffusipennisAfroleptomydas A sp ClanwilliamAfroleptomydas A sp ObibAfroleptomydas C sp KoeroegabAfroleptomydas A omeriAfroleptomydas A rufithoraxSyllegomydas S lineatusSyllegomydas S vittatusSyllegomydas S palestinensisSyllegomydas N astrictusSyllegomydas N heothinosSyllegomydas sp nov RuacanaSyllegomydas N arnoldi maleSyllegomydas N disparHispanomydas hispanicusSyllegomydas S algiricusSyllegomydas S elachysSyllegomydas S proximusHalterorchis karooensis femaleEremomidas arabicusEremomidas sp nov Tiega maleMitrodetus sp nov Argentina femaleMitrodetus nanoglossa maleMitrodetus dentitarsisMitrodetus dimidiatus male

Apioceridae

Mydidae

••

•••

• •• •

••••

••

•

••

••

•••

•

• ••

•••

••

••

•

• •• ••

••

•

• • •

... new phylogenetic hypothesis

Limata tenuicornisEulonchus sp Six RiversNeorhynchocephalus volaticusNemestrinus nigrovillosusBombylius majorDipalta serpentinaToxophora maximaProrates sp EscalanteApsilocephala longistylaHemigephyra atraPhycus frommeriDiogmites grossusOspriocerus aeacusGonioscelis ventralisHoplistomerus nobilisPilica formidolosaLaphria aktisEmphysomera pallidapexOmmatius tibialisNeolophonotus bimaculatusMachimus occidentalisPhilonicus albicepsPegesimallus laticornisDioctria hyalipennisLeptogaster cylindricaEuscelidia pulchraStichopogon punctumLasiopogon cinctusDamalis monochaetesHolcocephala calvaApiocera R alastor maleApiocera R braunsiApiocera P bilineataApiocera P mexicana maleApiocera P parahydraApiocera P barriApiocera P martinorumApiocera P haruspexApiocera P aldrichiApiocera P painteriApiocera P arenaApiocera P parkeriApiocera P sp Colnett maleApiocera Ap imminutaApiocera Ap tonnoiriApiocera An obscuraApiocera An philippiiApiocera Ap sp WurargaApiocera Ap asilicaApiocera Ap brevicornisApiocera Ap sp Stuart maleApiocera Ap hardyiApiocera Ap newmaniApiocera Ap sp LloydApiocera Ap maritimaApiocera Ap similisRhaphiomidas hasbrouckiRhaphiomidas maehleriRhaphiomidas parkeriTongamya mirandaMegascelus albovillosusMegascelus nigricornisNeorhaphiomidas setosa maleNeorhaphiomidas sp MandurahCretomydas santanensisDiochlistus auripennis maleDiochlistus gracilisDiochlistus melleipennisAnomalomydas mackerrasiAnomalomydas australicus femaleParamydas igniticornisPlyomydas peruviensis malePlyomydas sp nov TandilMessiasia californica maleMessiasia painteriMidacritus stuardoanus maleMidacritus wagenknechtiCacatuopyga basifasciaCacatuopyga ruficornis maleHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus armipesEctyphus amboseli maleEctyphus pinguisApiophora paulseniApiophora quadricinctataMiltinus stenogasterMiltinus maculipennisMiltinus viduatusMydas clavatusMydas xanthopterusStratiomydas lividusDolichogaster brevicornisPhyllomydas bruesiiPhyllomydas parvulusPseudorhopalia mirandaiRhopalia annulataRhopalia cinctaRhopalia olivieriPerissocerus dumonti femalePerissocerus transcaspicusHessemydas parkeri maleHessemydas seyrigiNomoneura caffraNomoneura paradoxaNomoneuroides natalensisMydaselpis ngurumaniArenomydas partitusArenomydas caerulescensArenomydas callosusArenomydas namaquensis maleNeolaparopsis puncturatusHaplomydas crassipesEremohaplomydas desertorumLachnocorynus sp nov Kotwa maleMydaselpis sp nov DaanMydaselpis sp nov Gamsberg maleVespiodes leopolditertiiVespiodes phaiosNemomydas brachyrhynchusNemomydas venosusLeptomydas corsicanusNemomydas gruenbergi maleLeptomydas lusitanicusEremomidas bekLeptomydas rapti maleLeptomydas turcicusLeptomydas sardousLeptomydas notosNemomydas pantherinusPseudonomoneura hirtaPseudonomoneura micheneriNemomydas lamiaNotosyllegomydas brinckiNamibimydas stuckenbergi sp novNothomydas picketti sp novCephalocera longirostrisCephalocera sp PapendorpCephalocera sp Sors SorsCephalocera imitata maleCephalocera sp RatelfonteinNemomydas melanopogonNamadytes sp Gobabeb maleNamadytes vansoniAfroleptomydas C mauriceiAfroleptomydas C sp GobabebAfroleptomydas C sp StamprietAfroleptomydas A inhacaeAfroleptomydas A consanguinensisAfroleptomydas A opacicinctusAfroleptomydas A paganusAfroleptomydas A suffusipennisAfroleptomydas A sp ClanwilliamAfroleptomydas A sp ObibAfroleptomydas C sp KoeroegabAfroleptomydas A omeriAfroleptomydas A rufithoraxSyllegomydas S lineatusSyllegomydas S vittatusSyllegomydas S palestinensisSyllegomydas N astrictusSyllegomydas N heothinosSyllegomydas sp nov RuacanaSyllegomydas N arnoldi maleSyllegomydas N disparHispanomydas hispanicusSyllegomydas S algiricusSyllegomydas S elachysSyllegomydas S proximusHalterorchis karooensis femaleEremomidas arabicusEremomidas sp nov Tiega maleMitrodetus sp nov Argentina femaleMitrodetus nanoglossa maleMitrodetus dentitarsisMitrodetus dimidiatus male

Results428 most parsimonious trees◊

1714 steps◊

Apioceridae, Rhaphiomidinae, and ◊ Megascelinae as in Yeates & Irwin 1996

several polytomies◊

strict consensus not well-resolved in ◊ derived ingroup

strict consensus of 428 trees

Bremer 3 – 8Bremer ≥ 10

11

Apiocera R alastor maleApiocera R braunsiApiocera P bilineataApiocera P mexicana maleApiocera P parahydraApiocera P barriApiocera P martinorumApiocera P haruspexApiocera P aldrichiApiocera P painteriApiocera P arenaApiocera P parkeriApiocera P sp Colnett maleApiocera Ap imminutaApiocera Ap tonnoiriApiocera An obscuraApiocera An philippiiApiocera Ap sp WurargaApiocera Ap asilicaApiocera Ap brevicornisApiocera Ap sp Stuart maleApiocera Ap hardyiApiocera Ap newmaniApiocera Ap sp LloydApiocera Ap maritimaApiocera Ap similis

Limata tenuicornisEulonchus sp Six RiversNeorhynchocephalus volaticusNemestrinus nigrovillosusBombylius majorDipalta serpentinaToxophora maximaProrates sp EscalanteApsilocephala longistylaHemigephyra atraPhycus frommeriDiogmites grossusOspriocerus aeacusGonioscelis ventralisHoplistomerus nobilisPilica formidolosaLaphria aktisEmphysomera pallidapexOmmatius tibialisNeolophonotus bimaculatusMachimus occidentalisPhilonicus albicepsPegesimallus laticornisDioctria hyalipennisLeptogaster cylindricaEuscelidia pulchraStichopogon punctumLasiopogon cinctusDamalis monochaetesHolcocephala calvaApiocera R alastor maleApiocera R braunsiApiocera P bilineataApiocera P mexicana maleApiocera P parahydraApiocera P barriApiocera P martinorumApiocera P haruspexApiocera P aldrichiApiocera P painteriApiocera P arenaApiocera P parkeriApiocera P sp Colnett maleApiocera Ap imminutaApiocera Ap tonnoiriApiocera An obscuraApiocera An philippiiApiocera Ap sp WurargaApiocera Ap asilicaApiocera Ap brevicornisApiocera Ap sp Stuart maleApiocera Ap hardyiApiocera Ap newmaniApiocera Ap sp LloydApiocera Ap maritimaApiocera Ap similisRhaphiomidas hasbrouckiRhaphiomidas maehleriRhaphiomidas parkeriTongamya mirandaMegascelus albovillosusMegascelus nigricornisNeorhaphiomidas setosa maleNeorhaphiomidas sp MandurahCretomydas santanensisDiochlistus auripennis maleDiochlistus gracilisDiochlistus melleipennisAnomalomydas mackerrasiAnomalomydas australicus femaleParamydas igniticornisPlyomydas peruviensis malePlyomydas sp nov TandilMessiasia californica maleMessiasia painteriMidacritus stuardoanus maleMidacritus wagenknechtiCacatuopyga basifasciaCacatuopyga ruficornis maleHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus armipesEctyphus amboseli maleEctyphus pinguisApiophora paulseniApiophora quadricinctataMiltinus stenogasterMiltinus maculipennisMiltinus viduatusMydas clavatusMydas xanthopterusStratiomydas lividusDolichogaster brevicornisPhyllomydas bruesiiPhyllomydas parvulusPseudorhopalia mirandaiRhopalia annulataRhopalia cinctaRhopalia olivieriPerissocerus dumonti femalePerissocerus transcaspicusHessemydas parkeri maleHessemydas seyrigiNomoneura caffraNomoneura paradoxaNomoneuroides natalensisMydaselpis ngurumaniArenomydas partitusArenomydas caerulescensArenomydas callosusArenomydas namaquensis maleNeolaparopsis puncturatusHaplomydas crassipesEremohaplomydas desertorumLachnocorynus sp nov Kotwa maleMydaselpis sp nov DaanMydaselpis sp nov Gamsberg maleVespiodes leopolditertiiVespiodes phaiosNemomydas brachyrhynchusNemomydas venosusLeptomydas corsicanusNemomydas gruenbergi maleLeptomydas lusitanicusEremomidas bekLeptomydas rapti maleLeptomydas turcicusLeptomydas sardousLeptomydas notosNemomydas pantherinusPseudonomoneura hirtaPseudonomoneura micheneriNemomydas lamiaNotosyllegomydas brinckiNamibimydas stuckenbergi sp novNothomydas picketti sp novCephalocera longirostrisCephalocera sp PapendorpCephalocera sp Sors SorsCephalocera imitata maleCephalocera sp RatelfonteinNemomydas melanopogonNamadytes sp Gobabeb maleNamadytes vansoniAfroleptomydas C mauriceiAfroleptomydas C sp GobabebAfroleptomydas C sp StamprietAfroleptomydas A inhacaeAfroleptomydas A consanguinensisAfroleptomydas A opacicinctusAfroleptomydas A paganusAfroleptomydas A suffusipennisAfroleptomydas A sp ClanwilliamAfroleptomydas A sp ObibAfroleptomydas C sp KoeroegabAfroleptomydas A omeriAfroleptomydas A rufithoraxSyllegomydas S lineatusSyllegomydas S vittatusSyllegomydas S palestinensisSyllegomydas N astrictusSyllegomydas N heothinosSyllegomydas sp nov RuacanaSyllegomydas N arnoldi maleSyllegomydas N disparHispanomydas hispanicusSyllegomydas S algiricusSyllegomydas S elachysSyllegomydas S proximusHalterorchis karooensis femaleEremomidas arabicusEremomidas sp nov Tiega maleMitrodetus sp nov Argentina femaleMitrodetus nanoglossa maleMitrodetus dentitarsisMitrodetus dimidiatus male

Apiocera subgenera

South Africa

western North America

Chile

Australian Apiocera (Apiocera)

Apiocera (Pyrocera) alleni © Greg Ballmer

Apiocera (Apiocera) sp. © Farhan Bokhari

... new phylogenetic hypothesis

12

Rhaphiomidas hasbrouckiRhaphiomidas maehleriRhaphiomidas parkeriTongamya mirandaMegascelus albovillosusMegascelus nigricornisNeorhaphiomidas setosa maleNeorhaphiomidas sp MandurahCretomydas santanensisDiochlistus auripennis maleDiochlistus gracilisDiochlistus melleipennisAnomalomydas mackerrasiAnomalomydas australicus femaleParamydas igniticornisPlyomydas peruviensis malePlyomydas sp nov TandilMessiasia californica maleMessiasia painteriMidacritus stuardoanus maleMidacritus wagenknechtiCacatuopyga basifasciaCacatuopyga ruficornis maleHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus armipesEctyphus amboseli maleEctyphus pinguisApiophora paulseniApiophora quadricinctataMiltinus stenogasterMiltinus maculipennisMiltinus viduatusMydas clavatusMydas xanthopterusStratiomydas lividusDolichogaster brevicornisPhyllomydas bruesiiPhyllomydas parvulusPseudorhopalia mirandaiRhopalia annulataRhopalia cinctaRhopalia olivieriPerissocerus dumonti femalePerissocerus transcaspicusHessemydas parkeri maleHessemydas seyrigiNomoneura caffraNomoneura paradoxaNomoneuroides natalensisMydaselpis ngurumaniArenomydas partitusArenomydas caerulescensArenomydas callosusArenomydas namaquensis maleNeolaparopsis puncturatusHaplomydas crassipesEremohaplomydas desertorumLachnocorynus sp nov Kotwa maleMydaselpis sp nov DaanMydaselpis sp nov Gamsberg maleVespiodes leopolditertiiVespiodes phaiosNemomydas brachyrhynchusNemomydas venosusLeptomydas corsicanusNemomydas gruenbergi maleLeptomydas lusitanicusEremomidas bekLeptomydas rapti maleLeptomydas turcicusLeptomydas sardousLeptomydas notosNemomydas pantherinusPseudonomoneura hirtaPseudonomoneura micheneriNemomydas lamiaNotosyllegomydas brinckiNamibimydas stuckenbergi sp novNothomydas picketti sp novCephalocera longirostrisCephalocera sp PapendorpCephalocera sp Sors SorsCephalocera imitata maleCephalocera sp RatelfonteinNemomydas melanopogonNamadytes sp Gobabeb maleNamadytes vansoniAfroleptomydas C mauriceiAfroleptomydas C sp GobabebAfroleptomydas C sp StamprietAfroleptomydas A inhacaeAfroleptomydas A consanguinensisAfroleptomydas A opacicinctusAfroleptomydas A paganusAfroleptomydas A suffusipennisAfroleptomydas A sp ClanwilliamAfroleptomydas A sp ObibAfroleptomydas C sp KoeroegabAfroleptomydas A omeriAfroleptomydas A rufithoraxSyllegomydas S lineatusSyllegomydas S vittatusSyllegomydas S palestinensisSyllegomydas N astrictusSyllegomydas N heothinosSyllegomydas sp nov RuacanaSyllegomydas N arnoldi maleSyllegomydas N disparHispanomydas hispanicusSyllegomydas S algiricusSyllegomydas S elachysSyllegomydas S proximusHalterorchis karooensis femaleEremomidas arabicusEremomidas sp nov Tiega maleMitrodetus sp nov Argentina femaleMitrodetus nanoglossa maleMitrodetus dentitarsisMitrodetus dimidiatus male

Australia

Argentina + Chile

† Cretomydas santanensis, Brazil

≈ 112 myoPlacement of † Cretomydas santanensis

Limata tenuicornisEulonchus sp Six RiversNeorhynchocephalus volaticusNemestrinus nigrovillosusBombylius majorDipalta serpentinaToxophora maximaProrates sp EscalanteApsilocephala longistylaHemigephyra atraPhycus frommeriDiogmites grossusOspriocerus aeacusGonioscelis ventralisHoplistomerus nobilisPilica formidolosaLaphria aktisEmphysomera pallidapexOmmatius tibialisNeolophonotus bimaculatusMachimus occidentalisPhilonicus albicepsPegesimallus laticornisDioctria hyalipennisLeptogaster cylindricaEuscelidia pulchraStichopogon punctumLasiopogon cinctusDamalis monochaetesHolcocephala calvaApiocera R alastor maleApiocera R braunsiApiocera P bilineataApiocera P mexicana maleApiocera P parahydraApiocera P barriApiocera P martinorumApiocera P haruspexApiocera P aldrichiApiocera P painteriApiocera P arenaApiocera P parkeriApiocera P sp Colnett maleApiocera Ap imminutaApiocera Ap tonnoiriApiocera An obscuraApiocera An philippiiApiocera Ap sp WurargaApiocera Ap asilicaApiocera Ap brevicornisApiocera Ap sp Stuart maleApiocera Ap hardyiApiocera Ap newmaniApiocera Ap sp LloydApiocera Ap maritimaApiocera Ap similisRhaphiomidas hasbrouckiRhaphiomidas maehleriRhaphiomidas parkeriTongamya mirandaMegascelus albovillosusMegascelus nigricornisNeorhaphiomidas setosa maleNeorhaphiomidas sp MandurahCretomydas santanensisDiochlistus auripennis maleDiochlistus gracilisDiochlistus melleipennisAnomalomydas mackerrasiAnomalomydas australicus femaleParamydas igniticornisPlyomydas peruviensis malePlyomydas sp nov TandilMessiasia californica maleMessiasia painteriMidacritus stuardoanus maleMidacritus wagenknechtiCacatuopyga basifasciaCacatuopyga ruficornis maleHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus armipesEctyphus amboseli maleEctyphus pinguisApiophora paulseniApiophora quadricinctataMiltinus stenogasterMiltinus maculipennisMiltinus viduatusMydas clavatusMydas xanthopterusStratiomydas lividusDolichogaster brevicornisPhyllomydas bruesiiPhyllomydas parvulusPseudorhopalia mirandaiRhopalia annulataRhopalia cinctaRhopalia olivieriPerissocerus dumonti femalePerissocerus transcaspicusHessemydas parkeri maleHessemydas seyrigiNomoneura caffraNomoneura paradoxaNomoneuroides natalensisMydaselpis ngurumaniArenomydas partitusArenomydas caerulescensArenomydas callosusArenomydas namaquensis maleNeolaparopsis puncturatusHaplomydas crassipesEremohaplomydas desertorumLachnocorynus sp nov Kotwa maleMydaselpis sp nov DaanMydaselpis sp nov Gamsberg maleVespiodes leopolditertiiVespiodes phaiosNemomydas brachyrhynchusNemomydas venosusLeptomydas corsicanusNemomydas gruenbergi maleLeptomydas lusitanicusEremomidas bekLeptomydas rapti maleLeptomydas turcicusLeptomydas sardousLeptomydas notosNemomydas pantherinusPseudonomoneura hirtaPseudonomoneura micheneriNemomydas lamiaNotosyllegomydas brinckiNamibimydas stuckenbergi sp novNothomydas picketti sp novCephalocera longirostrisCephalocera sp PapendorpCephalocera sp Sors SorsCephalocera imitata maleCephalocera sp RatelfonteinNemomydas melanopogonNamadytes sp Gobabeb maleNamadytes vansoniAfroleptomydas C mauriceiAfroleptomydas C sp GobabebAfroleptomydas C sp StamprietAfroleptomydas A inhacaeAfroleptomydas A consanguinensisAfroleptomydas A opacicinctusAfroleptomydas A paganusAfroleptomydas A suffusipennisAfroleptomydas A sp ClanwilliamAfroleptomydas A sp ObibAfroleptomydas C sp KoeroegabAfroleptomydas A omeriAfroleptomydas A rufithoraxSyllegomydas S lineatusSyllegomydas S vittatusSyllegomydas S palestinensisSyllegomydas N astrictusSyllegomydas N heothinosSyllegomydas sp nov RuacanaSyllegomydas N arnoldi maleSyllegomydas N disparHispanomydas hispanicusSyllegomydas S algiricusSyllegomydas S elachysSyllegomydas S proximusHalterorchis karooensis femaleEremomidas arabicusEremomidas sp nov Tiega maleMitrodetus sp nov Argentina femaleMitrodetus nanoglossa maleMitrodetus dentitarsisMitrodetus dimidiatus male

Diochlistus auripennis, Australia © Peter Chew

... new phylogenetic hypothesis

13... new phylogenetic hypothesis

Limata tenuicornisEulonchus sp Six RiversNeorhynchocephalus volaticusNemestrinus nigrovillosusBombylius majorDipalta serpentinaToxophora maximaProrates sp EscalanteApsilocephala longistylaHemigephyra atraPhycus frommeriDiogmites grossusOspriocerus aeacusGonioscelis ventralisHoplistomerus nobilisPilica formidolosaLaphria aktisEmphysomera pallidapexOmmatius tibialisNeolophonotus bimaculatusMachimus occidentalisPhilonicus albicepsPegesimallus laticornisDioctria hyalipennisLeptogaster cylindricaEuscelidia pulchraStichopogon punctumLasiopogon cinctusDamalis monochaetesHolcocephala calvaApiocera R alastor maleApiocera R braunsiApiocera P bilineataApiocera P mexicana maleApiocera P parahydraApiocera P barriApiocera P martinorumApiocera P haruspexApiocera P aldrichiApiocera P painteriApiocera P arenaApiocera P parkeriApiocera P sp Colnett maleApiocera Ap imminutaApiocera Ap tonnoiriApiocera An obscuraApiocera An philippiiApiocera Ap sp WurargaApiocera Ap asilicaApiocera Ap brevicornisApiocera Ap sp Stuart maleApiocera Ap hardyiApiocera Ap newmaniApiocera Ap sp LloydApiocera Ap maritimaApiocera Ap similisRhaphiomidas hasbrouckiRhaphiomidas maehleriRhaphiomidas parkeriTongamya mirandaMegascelus albovillosusMegascelus nigricornisNeorhaphiomidas setosa maleNeorhaphiomidas sp MandurahCretomydas santanensisDiochlistus auripennis maleDiochlistus gracilisDiochlistus melleipennisAnomalomydas mackerrasiAnomalomydas australicus femaleParamydas igniticornisPlyomydas peruviensis malePlyomydas sp nov TandilMessiasia californica maleMessiasia painteriMidacritus stuardoanus maleMidacritus wagenknechtiCacatuopyga basifasciaCacatuopyga ruficornis maleHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus armipesEctyphus amboseli maleEctyphus pinguisApiophora paulseniApiophora quadricinctataMiltinus stenogasterMiltinus maculipennisMiltinus viduatusMydas clavatusMydas xanthopterusStratiomydas lividusDolichogaster brevicornisPhyllomydas bruesiiPhyllomydas parvulusPseudorhopalia mirandaiRhopalia annulataRhopalia cinctaRhopalia olivieriPerissocerus dumonti femalePerissocerus transcaspicusHessemydas parkeri maleHessemydas seyrigiNomoneura caffraNomoneura paradoxaNomoneuroides natalensisMydaselpis ngurumaniArenomydas partitusArenomydas caerulescensArenomydas callosusArenomydas namaquensis maleNeolaparopsis puncturatusHaplomydas crassipesEremohaplomydas desertorumLachnocorynus sp nov Kotwa maleMydaselpis sp nov DaanMydaselpis sp nov Gamsberg maleVespiodes leopolditertiiVespiodes phaiosNemomydas brachyrhynchusNemomydas venosusLeptomydas corsicanusNemomydas gruenbergi maleLeptomydas lusitanicusEremomidas bekLeptomydas rapti maleLeptomydas turcicusLeptomydas sardousLeptomydas notosNemomydas pantherinusPseudonomoneura hirtaPseudonomoneura micheneriNemomydas lamiaNotosyllegomydas brinckiNamibimydas stuckenbergi sp novNothomydas picketti sp novCephalocera longirostrisCephalocera sp PapendorpCephalocera sp Sors SorsCephalocera imitata maleCephalocera sp RatelfonteinNemomydas melanopogonNamadytes sp Gobabeb maleNamadytes vansoniAfroleptomydas C mauriceiAfroleptomydas C sp GobabebAfroleptomydas C sp StamprietAfroleptomydas A inhacaeAfroleptomydas A consanguinensisAfroleptomydas A opacicinctusAfroleptomydas A paganusAfroleptomydas A suffusipennisAfroleptomydas A sp ClanwilliamAfroleptomydas A sp ObibAfroleptomydas C sp KoeroegabAfroleptomydas A omeriAfroleptomydas A rufithoraxSyllegomydas S lineatusSyllegomydas S vittatusSyllegomydas S palestinensisSyllegomydas N astrictusSyllegomydas N heothinosSyllegomydas sp nov RuacanaSyllegomydas N arnoldi maleSyllegomydas N disparHispanomydas hispanicusSyllegomydas S algiricusSyllegomydas S elachysSyllegomydas S proximusHalterorchis karooensis femaleEremomidas arabicusEremomidas sp nov Tiega maleMitrodetus sp nov Argentina femaleMitrodetus nanoglossa maleMitrodetus dentitarsisMitrodetus dimidiatus male

monophyletic* non-monophyletic

Syllegomydinae: Cephalocera imitata

Monophyly of subfamily taxa Rhaphiomidas hasbrouckiRhaphiomidas maehleriRhaphiomidas parkeriTongamya mirandaMegascelus albovillosusMegascelus nigricornisNeorhaphiomidas setosa maleNeorhaphiomidas sp MandurahCretomydas santanensisDiochlistus auripennis maleDiochlistus gracilisDiochlistus melleipennisAnomalomydas mackerrasiAnomalomydas australicus femaleParamydas igniticornisPlyomydas peruviensis malePlyomydas sp nov TandilMessiasia californica maleMessiasia painteriMidacritus stuardoanus maleMidacritus wagenknechtiCacatuopyga basifasciaCacatuopyga ruficornis maleHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus armipesEctyphus amboseli maleEctyphus pinguisApiophora paulseniApiophora quadricinctataMiltinus stenogasterMiltinus maculipennisMiltinus viduatusMydas clavatusMydas xanthopterusStratiomydas lividusDolichogaster brevicornisPhyllomydas bruesiiPhyllomydas parvulusPseudorhopalia mirandaiRhopalia annulataRhopalia cinctaRhopalia olivieriPerissocerus dumonti femalePerissocerus transcaspicusHessemydas parkeri maleHessemydas seyrigiNomoneura caffraNomoneura paradoxaNomoneuroides natalensisMydaselpis ngurumaniArenomydas partitusArenomydas caerulescensArenomydas callosusArenomydas namaquensis maleNeolaparopsis puncturatusHaplomydas crassipesEremohaplomydas desertorumLachnocorynus sp nov Kotwa maleMydaselpis sp nov DaanMydaselpis sp nov Gamsberg maleVespiodes leopolditertiiVespiodes phaiosNemomydas brachyrhynchusNemomydas venosusLeptomydas corsicanusNemomydas gruenbergi maleLeptomydas lusitanicusEremomidas bekLeptomydas rapti maleLeptomydas turcicusLeptomydas sardousLeptomydas notosNemomydas pantherinusPseudonomoneura hirtaPseudonomoneura micheneriNemomydas lamiaNotosyllegomydas brinckiNamibimydas stuckenbergi sp novNothomydas picketti sp novCephalocera longirostrisCephalocera sp PapendorpCephalocera sp Sors SorsCephalocera imitata maleCephalocera sp RatelfonteinNemomydas melanopogonNamadytes sp Gobabeb maleNamadytes vansoniAfroleptomydas C mauriceiAfroleptomydas C sp GobabebAfroleptomydas C sp StamprietAfroleptomydas A inhacaeAfroleptomydas A consanguinensisAfroleptomydas A opacicinctusAfroleptomydas A paganusAfroleptomydas A suffusipennisAfroleptomydas A sp ClanwilliamAfroleptomydas A sp ObibAfroleptomydas C sp KoeroegabAfroleptomydas A omeriAfroleptomydas A rufithoraxSyllegomydas S lineatusSyllegomydas S vittatusSyllegomydas S palestinensisSyllegomydas N astrictusSyllegomydas N heothinosSyllegomydas sp nov RuacanaSyllegomydas N arnoldi maleSyllegomydas N disparHispanomydas hispanicusSyllegomydas S algiricusSyllegomydas S elachysSyllegomydas S proximusHalterorchis karooensis femaleEremomidas arabicusEremomidas sp nov Tiega maleMitrodetus sp nov Argentina femaleMitrodetus nanoglossa maleMitrodetus dentitarsisMitrodetus dimidiatus male

RhaphiomidinaeMegascelinae

* Rhopaliinae

Anomalomydinae

Cacatuopyginae

* Mydinae

* Apiophorinae

* Ectyphinae

* Leptomydinae* Diochlistinae

* Leptomydinae

* Syllegomydinae

* Diochlistinae

* Mydinae

* Leptomydinae

* Syllegomydinae

* Apiophorinae

Syllegomydinae (43 % species diversity)

14

Rhaphiomidas hasbrouckiRhaphiomidas maehleriRhaphiomidas parkeriTongamya mirandaMegascelus albovillosusMegascelus nigricornisNeorhaphiomidas setosa maleNeorhaphiomidas sp MandurahCretomydas santanensisDiochlistus auripennis maleDiochlistus gracilisDiochlistus melleipennisAnomalomydas mackerrasiAnomalomydas australicus female

Rhaphiomidinae

Megascelinae

Anomalomydinae

* Diochlistinae

western North America

Namibia + South AfricaChileAustralia

BrazilAustralia

Australia

remaining Mydidae

Mydidae: Rhaphiomidas auratus © Greg Ballmer

Mydidae: Anomalomydas mackerrasi © Jean Hort

Limata tenuicornisEulonchus sp Six RiversNeorhynchocephalus volaticusNemestrinus nigrovillosusBombylius majorDipalta serpentinaToxophora maximaProrates sp EscalanteApsilocephala longistylaHemigephyra atraPhycus frommeriDiogmites grossusOspriocerus aeacusGonioscelis ventralisHoplistomerus nobilisPilica formidolosaLaphria aktisEmphysomera pallidapexOmmatius tibialisNeolophonotus bimaculatusMachimus occidentalisPhilonicus albicepsPegesimallus laticornisDioctria hyalipennisLeptogaster cylindricaEuscelidia pulchraStichopogon punctumLasiopogon cinctusDamalis monochaetesHolcocephala calvaApiocera R alastor maleApiocera R braunsiApiocera P bilineataApiocera P mexicana maleApiocera P parahydraApiocera P barriApiocera P martinorumApiocera P haruspexApiocera P aldrichiApiocera P painteriApiocera P arenaApiocera P parkeriApiocera P sp Colnett maleApiocera Ap imminutaApiocera Ap tonnoiriApiocera An obscuraApiocera An philippiiApiocera Ap sp WurargaApiocera Ap asilicaApiocera Ap brevicornisApiocera Ap sp Stuart maleApiocera Ap hardyiApiocera Ap newmaniApiocera Ap sp LloydApiocera Ap maritimaApiocera Ap similisRhaphiomidas hasbrouckiRhaphiomidas maehleriRhaphiomidas parkeriTongamya mirandaMegascelus albovillosusMegascelus nigricornisNeorhaphiomidas setosa maleNeorhaphiomidas sp MandurahCretomydas santanensisDiochlistus auripennis maleDiochlistus gracilisDiochlistus melleipennisAnomalomydas mackerrasiAnomalomydas australicus femaleParamydas igniticornisPlyomydas peruviensis malePlyomydas sp nov TandilMessiasia californica maleMessiasia painteriMidacritus stuardoanus maleMidacritus wagenknechtiCacatuopyga basifasciaCacatuopyga ruficornis maleHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus armipesEctyphus amboseli maleEctyphus pinguisApiophora paulseniApiophora quadricinctataMiltinus stenogasterMiltinus maculipennisMiltinus viduatusMydas clavatusMydas xanthopterusStratiomydas lividusDolichogaster brevicornisPhyllomydas bruesiiPhyllomydas parvulusPseudorhopalia mirandaiRhopalia annulataRhopalia cinctaRhopalia olivieriPerissocerus dumonti femalePerissocerus transcaspicusHessemydas parkeri maleHessemydas seyrigiNomoneura caffraNomoneura paradoxaNomoneuroides natalensisMydaselpis ngurumaniArenomydas partitusArenomydas caerulescensArenomydas callosusArenomydas namaquensis maleNeolaparopsis puncturatusHaplomydas crassipesEremohaplomydas desertorumLachnocorynus sp nov Kotwa maleMydaselpis sp nov DaanMydaselpis sp nov Gamsberg maleVespiodes leopolditertiiVespiodes phaiosNemomydas brachyrhynchusNemomydas venosusLeptomydas corsicanusNemomydas gruenbergi maleLeptomydas lusitanicusEremomidas bekLeptomydas rapti maleLeptomydas turcicusLeptomydas sardousLeptomydas notosNemomydas pantherinusPseudonomoneura hirtaPseudonomoneura micheneriNemomydas lamiaNotosyllegomydas brinckiNamibimydas stuckenbergi sp novNothomydas picketti sp novCephalocera longirostrisCephalocera sp PapendorpCephalocera sp Sors SorsCephalocera imitata maleCephalocera sp RatelfonteinNemomydas melanopogonNamadytes sp Gobabeb maleNamadytes vansoniAfroleptomydas C mauriceiAfroleptomydas C sp GobabebAfroleptomydas C sp StamprietAfroleptomydas A inhacaeAfroleptomydas A consanguinensisAfroleptomydas A opacicinctusAfroleptomydas A paganusAfroleptomydas A suffusipennisAfroleptomydas A sp ClanwilliamAfroleptomydas A sp ObibAfroleptomydas C sp KoeroegabAfroleptomydas A omeriAfroleptomydas A rufithoraxSyllegomydas S lineatusSyllegomydas S vittatusSyllegomydas S palestinensisSyllegomydas N astrictusSyllegomydas N heothinosSyllegomydas sp nov RuacanaSyllegomydas N arnoldi maleSyllegomydas N disparHispanomydas hispanicusSyllegomydas S algiricusSyllegomydas S elachysSyllegomydas S proximusHalterorchis karooensis femaleEremomidas arabicusEremomidas sp nov Tiega maleMitrodetus sp nov Argentina femaleMitrodetus nanoglossa maleMitrodetus dentitarsisMitrodetus dimidiatus male

Mydidae: Neorhaphiomidas sp. © Fred Hort

Earliest branchings within Mydidae

... new phylogenetic hypothesis

15

Rhaphiomidas hasbrouckiRhaphiomidas maehleriRhaphiomidas parkeriTongamya mirandaMegascelus albovillosusMegascelus nigricornisNeorhaphiomidas setosa maleNeorhaphiomidas sp MandurahCretomydas santanensisDiochlistus auripennis maleDiochlistus gracilisDiochlistus melleipennisAnomalomydas mackerrasiAnomalomydas australicus femaleParamydas igniticornisPlyomydas peruviensis malePlyomydas sp nov TandilMessiasia californica maleMessiasia painteriMidacritus stuardoanus maleMidacritus wagenknechtiCacatuopyga basifasciaCacatuopyga ruficornis maleHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus armipesEctyphus amboseli maleEctyphus pinguisApiophora paulseniApiophora quadricinctataMiltinus stenogasterMiltinus maculipennisMiltinus viduatusMydas clavatusMydas xanthopterusStratiomydas lividusDolichogaster brevicornisPhyllomydas bruesiiPhyllomydas parvulusPseudorhopalia mirandaiRhopalia annulataRhopalia cinctaRhopalia olivieriPerissocerus dumonti femalePerissocerus transcaspicusHessemydas parkeri maleHessemydas seyrigiNomoneura caffraNomoneura paradoxaNomoneuroides natalensisMydaselpis ngurumaniArenomydas partitusArenomydas caerulescensArenomydas callosusArenomydas namaquensis maleNeolaparopsis puncturatusHaplomydas crassipesEremohaplomydas desertorumLachnocorynus sp nov Kotwa maleMydaselpis sp nov DaanMydaselpis sp nov Gamsberg maleVespiodes leopolditertiiVespiodes phaiosNemomydas brachyrhynchusNemomydas venosusLeptomydas corsicanusNemomydas gruenbergi maleLeptomydas lusitanicusEremomidas bekLeptomydas rapti maleLeptomydas turcicusLeptomydas sardousLeptomydas notosNemomydas pantherinusPseudonomoneura hirtaPseudonomoneura micheneriNemomydas lamiaNotosyllegomydas brinckiNamibimydas stuckenbergi sp novNothomydas picketti sp novCephalocera longirostrisCephalocera sp PapendorpCephalocera sp Sors SorsCephalocera imitata maleCephalocera sp RatelfonteinNemomydas melanopogonNamadytes sp Gobabeb maleNamadytes vansoniAfroleptomydas C mauriceiAfroleptomydas C sp GobabebAfroleptomydas C sp StamprietAfroleptomydas A inhacaeAfroleptomydas A consanguinensisAfroleptomydas A opacicinctusAfroleptomydas A paganusAfroleptomydas A suffusipennisAfroleptomydas A sp ClanwilliamAfroleptomydas A sp ObibAfroleptomydas C sp KoeroegabAfroleptomydas A omeriAfroleptomydas A rufithoraxSyllegomydas S lineatusSyllegomydas S vittatusSyllegomydas S palestinensisSyllegomydas N astrictusSyllegomydas N heothinosSyllegomydas sp nov RuacanaSyllegomydas N arnoldi maleSyllegomydas N disparHispanomydas hispanicusSyllegomydas S algiricusSyllegomydas S elachysSyllegomydas S proximusHalterorchis karooensis femaleEremomidas arabicusEremomidas sp nov Tiega maleMitrodetus sp nov Argentina femaleMitrodetus nanoglossa maleMitrodetus dentitarsisMitrodetus dimidiatus male

* Rhopaliinae

* Apiophorinae

* Ectyphinae

* Apiophorinae

Mydidae: Apiophorinae

Mydidae: Ectyphinae

Mydidae: Rhopaliinae

Biogeographically interesting taxa

.... biogeographically interesting taxa

16.... summary + future

Summary

Mydidae: Phyllomydas parvulus © Lyn Atherton

Apioceridae + Mydidae monophyletic◊

(Apioceridae + Mydidae) + Asilidae◊

Apiocera (Apiocera)◊ not monophyletic

earliest branchings within Mydidae well-supported◊

subfamily taxa relationships still not resolved ◊

Future Research: ◊ › field work in Australia (Dec 2011) and Namibia (Feb 2012) › internal head characters › addition of molecular data › adding yet additional taxa?

17

AcknowledgementsMuseum curators and collectors who made specimens available◊

Photographers◊

U.S. National Science Foundation (REVSYS DEB-0919333)◊

www.mydidae.tdvia.de