1

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

Torsten DikowDepartment of Entomology National Museum of Natural History

Apioceridae: Apiocera sp. © J. Hort Mydidae: Miltinus minutus © J. Hort

SmithsonianNational Museum of Natural History

2

Diptera = 159,294 valid species (Pape et al. 2011)

phylogeny from Wiegmann et al. 2011 (simplified), images Apiocera alleni, Leptomydas lusitanicus, Cyrtopogon lutatius, Echthodopa formosa © G. Ballmer, V. Jacinto, M. Thomas

PipunculidaeAcalyptratae I

Acroceridae

Ptychopteridae

Apystomyiidae

Nymphomyiidae

Perissommatidae

Syrphidae

Calyptratae

Tipulomorpha

Psychodomorpha

Tabanomorpha

Empidoidea

Stratiomyomorpha

Platypezoidea

NemestrinidaeXylophagidae

Hilarimorphidae

Deuterophlebiidae

Acalyptratae II

Culicomorpha

Bibionomorpha

Asiloidea

Ephydroidea

Asilidae

Apsilocephalidae

Mydidae

Apioceridae

Evocoidae

Bombyliidae

Therevidae

Scenopinidae

> 15,500 species

Brachycera

Eremoneura

Cyclorrhapha

Schizophora

Neodiptera

Orthorrhapha

> 8,100 species

. introduction

3

Apiocera (Apiocera) pica © J. Hort

Apioceridae – “flower-loving flies”◊ adult flies flower-feeders

◊ size = 15 – 30 mm

◊ 138 species in 1 genus (4 subgenera)

◊ occur only in › Argentina and Chile –> 4 species › Australia –> 67 species › South Africa –> 3 species › western North America (Canada, Mexico, USA) –> 64 species

Apiocera (Apiocera) sp. © J. Hort

. introduction

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Opomydas townsendi

Mydidae – “mydas flies”◊ adult flies flower-feeders

› some adults do not feed at all

◊ size = 9 – 60 mm (individual wing length up to 52 mm)

◊ 473 species in 66 genera

◊ world-wide primarily in arid regions › southern Africa most speciose

◊ rarely collected

◊ mimetic coloration, morphology, and behavior present in several clades › aculeate Hymenoptera

◊ Cretaceous fossil

Eremomidas arabicus © D. Gardner

. introduction

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Yeates & Irwin 1996 – study of Apioceridae relationshipsApiocera (Pyrocera) western North America

Apiocera (Apiocera) Australia

Apiocera (Ripidosyrma) South Africa

Apiocera (Anypenus) Argentina / Chile

Apioceridae area relationships

. introduction

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Apiocera (Pyrocera) western North America

Apiocera (Apiocera) Australia

Apiocera (Ripidosyrma) South Africa

Apiocera (Anypenus) Argentina / Chile

Tongamya

Megascelus

Neorhaphiomidas

RhaphiomidasApioceridaeMydidae

remaining Mydidae

area relationships

Yeates & Irwin 1996 – study of Apioceridae relationships

. introduction

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Taxon Sampling◊ 134 Mydidae species (= 28.4 %) › 46 genera (= 69.7 %)

› 7 tribal taxa (= 87.5 %) › 11 subfamily taxa (= 100.0 %)

◊ 27 Apiocera species (= 19.6 %)

◊ Outgroup taxa › 19 Asilidae species › 7 Asiloidea species › 3 Nemestrinoidea › 1 Tabanomorpha

collecting localities of specimens included

Apioceridae

Mydidae

.. phylogenetic analysis

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Characters◊ 214 discrete, parsimony informative characters

◊ 36 non-genitalic features sexually dimorphictagma # characters remarksHead 44 10 sexually dimorphicThorax 103 22 sexually dimorphic

47 chaetotaxy, sclerites etc.23 wing33 leg

Abdomen 67 4 sexually dimorphic11 chaetotaxy, sclerites etc.30 ? terminalia26 / terminalia

◊ DNA data are being gathered

.. phylogenetic analysis

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Characters – sexual dimorphismSyllegomydas vittatus

? /› 30 char. ? terminalia › 19 char. / terminalia

36 characters

.. phylogenetic analysis

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Phylogenetic analysis◊ TNT 1.1 (July 2012) Goloboff et al. 2008

◊ exhaustive search of tree space (MacBook Pro 2.2 GHz Intel Core i7)

◊ Bremer support Bremer 1988

◊ DIVA 1.2 Ronquist 1994, Nylander Mac version

hit best length 20x

random sectorial searches after hitting best length, find additional trees by fusing suboptimal with optimal trees

consense until strict consensus is stabilized 10x

command(s) time (min) # re-arrangements examinedmult300 01:04 8,383,595,488xmult=rss hits 20 fuse 2 drift 3 ratchet 10 multiply consense 10 12:31 79,691,144,513

.. phylogenetic analysis

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Nemomydas venosus

Halterorchis karooensis F

Apiocera Ap brevicornis

Dipalta serpentina

Namadytes cimbebasiensis

Rhopalia annulata

Pseudonomoneura micheneri

Diochlistus gracilis

Apiophora paulseni

Eremomidas bek

Mydaselpis sp nov Gamsberg M

Pegesimallus laticornis

Anomalomydas australicus F

Apiocera R alastor M

Arenomydas namaquensis M

Apiocera Ap imminuta

Laphria aktis

Megascelus albovillosus

Afroleptomydas C sp Stampriet

Leptogaster cylindrica

Diogmites grossus

Dolichogaster brevicornis

Halterorchis sp nov Cape Recife M

Paramydas igniticornis

Nemomydas brachyrhynchus

Miltinus maculipennis

Rhopalia cincta

Hessemydas seyrigi

Cacatuopyga ruficornis M

Nemomydas pantherinus

Nothomydas picketti

Eremomidas arabicus

Syllegomydas N astrictus

Bombylius major

Stichopogon punctum

Rhaphiomidas hasbroucki

Phycus frommeri

Messiasia painteri

Cretomydas santanensis

Apiophora quadricinctata

Afroleptomydas C sp Koeroegab

Neorhaphiomidas hardyi

Mydaselpis sp nov Daan

Notosyllegomydas brincki

Mitrodetus nanoglossa M

Syllegomydas sp nov Ruacana

Cephalocera sp Papendorp

Leptomydas corsicanus

Plyomydas sp nov Tandil

Hemigephyra atra

Neolaparopsis puncturatus

Apiocera P sp Colnett M

Nemomydas lamia

Namibimydas psamminos

Nemestrinus nigrovillosus

Syll gen nov sp nov minute proboscis F

Midacritus wagenknechti

Apiocera P mexicana M

Limata tenuicornis F

Emphysomera pallidapex

Mydas clavatus

Leptomydas sardous

Cephalocera longirostris

Apiocera P arena

Syllegomydas N dispar

Apiocera Ap asilica

Afroleptomydas C sp Gobabeb

Anomalomydas mackerrasi

Nemomydas gruenbergi M

Neorhaphiomidas sp Mandurah

Phyllomydas parvulus

Vespiodes phaios

Rhaphiomidas parkeri

Prorates sp Escalante

Tongamya miranda

Midacritus stuardoanus M

Rhopalia olivieri

Cephalocera sp Sors Sors

Apsilocephala longistyla

Apiocera Ap sp Wurarga

Cephalocera imitata M

Perissocerus transcaspicus

Megascelus melanoproctus

Syllegomydas S lineatus

Apiocera P parkeri

Syllegomydas S algiricus

Apiocera Ap minor

Miltinus stenogaster

Afroleptomydas A opacicinctus

Cleamydas gen nov sp nov Ugab F

Haplomydas crassipes

Stenomydas gen nov sp nov Panne M

Mitrodetus dentitarsis

Syllegomydas S vittatus

Neorhaphiomidas sp Yanchep

Arenomydas callosus

Hoplistomerus nobilis

Syllegomydas S elachys

Megascelus nigricornis

Messiasia californica M

Apiocera P painteri

Leptomydas turcicus

Perissocerus dumonti F

Afroleptomydas A sp Clanwilliam

Damalis monochaetes

Arenomydas partitus

Neolophonotus bimaculatusMachimus occidentalis

Diochlistus melleipennis

Euscelidia pulchra

Miltinus viduatus

Apiocera P bilineata

Pseudonomoneura bajaensis

Eulonchus sp Six Rivers M

Diochlistus auripennis M

Syllegomydas S proximus

Pseudonomoneura hirta

Vespiodes leopolditertii

Apiocera An philippii

Apiocera Ap hardyi

Heteromydas bicolor

Nemomydas melanopogon

Apiocera P martinorum

Dioctria hyalipennis

Apiocera P barri

Phyllomydas bruesii

Ospriocerus aeacus

Nomoneura paradoxa

Apiocera P haruspex

Apiocera Ap pica

Afroleptomydas A suffusipennis

Cacatuopyga basifascia

Afroleptomydas A paganus

Toxophora maxima

Cephalocera sp Ratelfontein

Gonioscelis ventralis

Leptomydas notos

Afroleptomydas A rufithorax

Arenomydas caerulescens

Apiocera R braunsi

Namibimydas stuckenbergi

Mydaselpis ngurumani

Syllegomydas N arnoldi M

Opomydas townsendi

Ectyphus pinguis

Apiocera Ap maritima

Plyomydas peruviensis M

Mydas tricinctus

Apiocera Ap sp Lloyd

Eremomidas sp nov Tiega M

Syll gen nov sp nov Gobabeb

Syllegomydas N heothinos

Mydas xanthopterus

Lachnocorynus sp nov Kotwa M

Afroleptomydas A omeri

Apiocera Ap sp Stuart M

Pilica formidolosa

Afroleptomydas C mauricei

Miltinus minutus

Nomoneura caffra

Ommatius tibialis

Pseudorhopalia mirandai

Nomoneuroides natalensis

Hessemydas parkeri M

Neorhynchocephalus volaticus

Afroleptomydas A consanguinensis

Apiocera P aldrichi

Syllegomydas S palestinensis

Namadytes vansoni

Apiocera An obscura

Ectyphus armipes

Hessemydas daugeroni

Hispanomydas hispanicus

Eremohaplomydas desertorum

Leptomydas lusitanicus

Lasiopogon cinctus

Afroleptomydas A sp Obib

Parectyphus namibiensis

Philonicus albiceps

Afroleptomydas A inhacae

Stratiomydas lividus

Holcocephala calva

Leptomydas rapti M

Rhaphiomidas maehleri

Ectyphus amboseli M

Apiocera Ap similis

Mitrodetus dimidiatus

Diochlistinae Argentina F

Mitrodetus microglossa

Apiocera P parahydra

Apiocera Ap tonnoiri

Apiocera Ap newmani

Neorhaphiomidas sp Mingenew M

Mydidae

Asilidae

Apioceridae

Nemomydas venosus

Halterorchis karooensis F

Apiocera Ap brevicornis

Dipalta serpentina

Namadytes cimbebasiensis

Rhopalia annulata

Pseudonomoneura micheneri

Diochlistus gracilis

Apiophora paulseni

Eremomidas bek

Mydaselpis sp nov Gamsberg M

Pegesimallus laticornis

Anomalomydas australicus F

Apiocera R alastor M

Arenomydas namaquensis M

Apiocera Ap imminuta

Laphria aktis

Megascelus albovillosus

Afroleptomydas C sp Stampriet

Leptogaster cylindrica

Diogmites grossus

Dolichogaster brevicornis

Halterorchis sp nov Cape Recife M

Paramydas igniticornis

Nemomydas brachyrhynchus

Miltinus maculipennis

Rhopalia cincta

Hessemydas seyrigi

Cacatuopyga ruficornis M

Nemomydas pantherinus

Nothomydas picketti

Eremomidas arabicus

Syllegomydas N astrictus

Bombylius major

Stichopogon punctum

Rhaphiomidas hasbroucki

Phycus frommeri

Messiasia painteri

Cretomydas santanensis

Apiophora quadricinctata

Afroleptomydas C sp Koeroegab

Neorhaphiomidas hardyi

Mydaselpis sp nov Daan

Notosyllegomydas brincki

Mitrodetus nanoglossa M

Syllegomydas sp nov Ruacana

Cephalocera sp Papendorp

Leptomydas corsicanus

Plyomydas sp nov Tandil

Hemigephyra atra

Neolaparopsis puncturatus

Apiocera P sp Colnett M

Nemomydas lamia

Namibimydas psamminos

Nemestrinus nigrovillosus

Syll gen nov sp nov minute proboscis F

Midacritus wagenknechti

Apiocera P mexicana M

Limata tenuicornis F

Emphysomera pallidapex

Mydas clavatus

Leptomydas sardous

Cephalocera longirostris

Apiocera P arena

Syllegomydas N dispar

Apiocera Ap asilica

Afroleptomydas C sp Gobabeb

Anomalomydas mackerrasi

Nemomydas gruenbergi M

Neorhaphiomidas sp Mandurah

Phyllomydas parvulus

Vespiodes phaios

Rhaphiomidas parkeri

Prorates sp Escalante

Tongamya miranda

Midacritus stuardoanus M

Rhopalia olivieri

Cephalocera sp Sors Sors

Apsilocephala longistyla

Apiocera Ap sp Wurarga

Cephalocera imitata M

Perissocerus transcaspicus

Megascelus melanoproctus

Syllegomydas S lineatus

Apiocera P parkeri

Syllegomydas S algiricus

Apiocera Ap minor

Miltinus stenogaster

Afroleptomydas A opacicinctus

Cleamydas gen nov sp nov Ugab F

Haplomydas crassipes

Stenomydas gen nov sp nov Panne M

Mitrodetus dentitarsis

Syllegomydas S vittatus

Neorhaphiomidas sp Yanchep

Arenomydas callosus

Hoplistomerus nobilis

Syllegomydas S elachys

Megascelus nigricornis

Messiasia californica M

Apiocera P painteri

Leptomydas turcicus

Perissocerus dumonti F

Afroleptomydas A sp Clanwilliam

Damalis monochaetes

Arenomydas partitus

Neolophonotus bimaculatusMachimus occidentalis

Diochlistus melleipennis

Euscelidia pulchra

Miltinus viduatus

Apiocera P bilineata

Pseudonomoneura bajaensis

Eulonchus sp Six Rivers M

Diochlistus auripennis M

Syllegomydas S proximus

Pseudonomoneura hirta

Vespiodes leopolditertii

Apiocera An philippii

Apiocera Ap hardyi

Heteromydas bicolor

Nemomydas melanopogon

Apiocera P martinorum

Dioctria hyalipennis

Apiocera P barri

Phyllomydas bruesii

Ospriocerus aeacus

Nomoneura paradoxa

Apiocera P haruspex

Apiocera Ap pica

Afroleptomydas A suffusipennis

Cacatuopyga basifascia

Afroleptomydas A paganus

Toxophora maxima

Cephalocera sp Ratelfontein

Gonioscelis ventralis

Leptomydas notos

Afroleptomydas A rufithorax

Arenomydas caerulescens

Apiocera R braunsi

Namibimydas stuckenbergi

Mydaselpis ngurumani

Syllegomydas N arnoldi M

Opomydas townsendi

Ectyphus pinguis

Apiocera Ap maritima

Plyomydas peruviensis M

Mydas tricinctus

Apiocera Ap sp Lloyd

Eremomidas sp nov Tiega M

Syll gen nov sp nov Gobabeb

Syllegomydas N heothinos

Mydas xanthopterus

Lachnocorynus sp nov Kotwa M

Afroleptomydas A omeri

Apiocera Ap sp Stuart M

Pilica formidolosa

Afroleptomydas C mauricei

Miltinus minutus

Nomoneura caffra

Ommatius tibialis

Pseudorhopalia mirandai

Nomoneuroides natalensis

Hessemydas parkeri M

Neorhynchocephalus volaticus

Afroleptomydas A consanguinensis

Apiocera P aldrichi

Syllegomydas S palestinensis

Namadytes vansoni

Apiocera An obscura

Ectyphus armipes

Hessemydas daugeroni

Hispanomydas hispanicus

Eremohaplomydas desertorum

Leptomydas lusitanicus

Lasiopogon cinctus

Afroleptomydas A sp Obib

Parectyphus namibiensis

Philonicus albiceps

Afroleptomydas A inhacae

Stratiomydas lividus

Holcocephala calva

Leptomydas rapti M

Rhaphiomidas maehleri

Ectyphus amboseli M

Apiocera Ap similis

Mitrodetus dimidiatus

Diochlistinae Argentina F

Mitrodetus microglossa

Apiocera P parahydra

Apiocera Ap tonnoiri

Apiocera Ap newmani

Neorhaphiomidas sp Mingenew M

Results◊ 440 most parsimonious trees

◊ 1837 steps

◊ strict consensus › several polytomies › not well-resolved are: › earliest branchings within Apioceridae › derived Mydidae ingroup

strict consensus topology (2121 steps)Bremer ≥ 3

... results

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Nemomydas lamia

Phyllomydas parvulus

Heteromydas bicolor

Megascelus albovillosus

Leptomydas corsicanus

Mitrodetus dimidiatus

Nemomydas venosus

Syllegomydas S vittatus

Cacatuopyga ruficornis M

Phyllomydas bruesii

Nomoneura paradoxa

Megascelus nigricornis

Miltinus viduatus

Nemomydas melanopogon

Syllegomydas S algiricus

Mydas clavatus

Plyomydas peruviensis M

Hessemydas daugeroni

Diochlistus auripennis M

Pseudorhopalia mirandai

Syllegomydas S elachys

Syll gen nov sp nov minute proboscis F

Perissocerus transcaspicus

Anomalomydas australicus F

Rhopalia olivieri

Stratiomydas lividus

Afroleptomydas A consanguinensis

Midacritus wagenknechti

Halterorchis karooensis F

Diochlistus melleipennis

Vespiodes phaios

Leptomydas turcicus

Arenomydas caerulescens

Apiophora quadricinctata

Hessemydas parkeri M

Plyomydas sp nov Tandil

Syllegomydas N astrictus

Neolaparopsis puncturatus

Syllegomydas N dispar

Arenomydas callosus

Neorhaphiomidas sp Mandurah

Namadytes cimbebasiensis

Afroleptomydas A suffusipennis

Syllegomydas S palestinensis

Arenomydas namaquensis M

Diochlistinae Argentina F

Afroleptomydas A sp Clanwilliam

Leptomydas sardous

Arenomydas partitus

Mydas xanthopterus

Namibimydas stuckenbergi

Afroleptomydas A paganus

Opomydas townsendi

Cephalocera sp Sors Sors

Mydaselpis ngurumani

Mydaselpis sp nov Gamsberg M

Afroleptomydas C sp Stampriet

Eremomidas sp nov Tiega M

Neorhaphiomidas hardyi

Pseudonomoneura bajaensis

Ectyphus armipes

Mitrodetus dentitarsis

Nomoneura caffra

Neorhaphiomidas sp Yanchep

Diochlistus gracilis

Messiasia californica M

Dolichogaster brevicornis

Rhaphiomidas parkeri

Afroleptomydas C sp Koeroegab

Pseudonomoneura hirta

Leptomydas notos

Leptomydas lusitanicus

Rhopalia annulata

Nemomydas brachyrhynchus

Cacatuopyga basifascia

Cretomydas santanensis

Paramydas igniticornis

Neorhaphiomidas sp Mingenew M

Miltinus minutus

Syllegomydas S lineatus

Halterorchis sp nov Cape Recife M

Megascelus melanoproctus

Afroleptomydas C mauricei

Namadytes vansoni

Syllegomydas N arnoldi M

Rhopalia cincta

Messiasia painteri

Eremohaplomydas desertorum

Syllegomydas S proximus

Nemomydas pantherinus

Afroleptomydas A omeri

Lachnocorynus sp nov Kotwa M

Afroleptomydas A opacicinctus

Rhaphiomidas hasbroucki

Namibimydas psamminos

Pseudonomoneura micheneri

Nothomydas picketti

Ectyphus pinguis

Afroleptomydas A rufithorax

Leptomydas rapti M

Afroleptomydas C sp Gobabeb

Syll gen nov sp nov Gobabeb

Miltinus stenogaster

Miltinus maculipennis

Midacritus stuardoanus M

Parectyphus namibiensis

Afroleptomydas A sp Obib

Hispanomydas hispanicus

Perissocerus dumonti F

Mitrodetus nanoglossa M

Cephalocera sp Papendorp

Vespiodes leopolditertii

Haplomydas crassipes

Nemomydas gruenbergi M

Anomalomydas mackerrasi

Mydaselpis sp nov Daan

Syllegomydas sp nov Ruacana

Cephalocera sp Ratelfontein

Cleamydas gen nov sp nov Ugab F

Syllegomydas N heothinos

Hessemydas seyrigi

Nomoneuroides natalensis

Mitrodetus microglossa

Stenomydas gen nov sp nov Panne M

Tongamya miranda

Cephalocera longirostris

Eremomidas arabicus

Apiophora paulseni

Afroleptomydas A inhacae

Notosyllegomydas brincki

Ectyphus amboseli M

Rhaphiomidas maehleri

Eremomidas bek

Mydas tricinctus

Cephalocera imitata M

Mydidae

Nemomydas venosus

Halterorchis karooensis F

Apiocera Ap brevicornis

Dipalta serpentina

Namadytes cimbebasiensis

Rhopalia annulata

Pseudonomoneura micheneri

Diochlistus gracilis

Apiophora paulseni

Eremomidas bek

Mydaselpis sp nov Gamsberg M

Pegesimallus laticornis

Anomalomydas australicus F

Apiocera R alastor M

Arenomydas namaquensis M

Apiocera Ap imminuta

Laphria aktis

Megascelus albovillosus

Afroleptomydas C sp Stampriet

Leptogaster cylindrica

Diogmites grossus

Dolichogaster brevicornis

Halterorchis sp nov Cape Recife M

Paramydas igniticornis

Nemomydas brachyrhynchus

Miltinus maculipennis

Rhopalia cincta

Hessemydas seyrigi

Cacatuopyga ruficornis M

Nemomydas pantherinus

Nothomydas picketti

Eremomidas arabicus

Syllegomydas N astrictus

Bombylius major

Stichopogon punctum

Rhaphiomidas hasbroucki

Phycus frommeri

Messiasia painteri

Cretomydas santanensis

Apiophora quadricinctata

Afroleptomydas C sp Koeroegab

Neorhaphiomidas hardyi

Mydaselpis sp nov Daan

Notosyllegomydas brincki

Mitrodetus nanoglossa M

Syllegomydas sp nov Ruacana

Cephalocera sp Papendorp

Leptomydas corsicanus

Plyomydas sp nov Tandil

Hemigephyra atra

Neolaparopsis puncturatus

Apiocera P sp Colnett M

Nemomydas lamia

Namibimydas psamminos

Nemestrinus nigrovillosus

Syll gen nov sp nov minute proboscis F

Midacritus wagenknechti

Apiocera P mexicana M

Limata tenuicornis F

Emphysomera pallidapex

Mydas clavatus

Leptomydas sardous

Cephalocera longirostris

Apiocera P arena

Syllegomydas N dispar

Apiocera Ap asilica

Afroleptomydas C sp Gobabeb

Anomalomydas mackerrasi

Nemomydas gruenbergi M

Neorhaphiomidas sp Mandurah

Phyllomydas parvulus

Vespiodes phaios

Rhaphiomidas parkeri

Prorates sp Escalante

Tongamya miranda

Midacritus stuardoanus M

Rhopalia olivieri

Cephalocera sp Sors Sors

Apsilocephala longistyla

Apiocera Ap sp Wurarga

Cephalocera imitata M

Perissocerus transcaspicus

Megascelus melanoproctus

Syllegomydas S lineatus

Apiocera P parkeri

Syllegomydas S algiricus

Apiocera Ap minor

Miltinus stenogaster

Afroleptomydas A opacicinctus

Cleamydas gen nov sp nov Ugab F

Haplomydas crassipes

Stenomydas gen nov sp nov Panne M

Mitrodetus dentitarsis

Syllegomydas S vittatus

Neorhaphiomidas sp Yanchep

Arenomydas callosus

Hoplistomerus nobilis

Syllegomydas S elachys

Megascelus nigricornis

Messiasia californica M

Apiocera P painteri

Leptomydas turcicus

Perissocerus dumonti F

Afroleptomydas A sp Clanwilliam

Damalis monochaetes

Arenomydas partitus

Neolophonotus bimaculatusMachimus occidentalis

Diochlistus melleipennis

Euscelidia pulchra

Miltinus viduatus

Apiocera P bilineata

Pseudonomoneura bajaensis

Eulonchus sp Six Rivers M

Diochlistus auripennis M

Syllegomydas S proximus

Pseudonomoneura hirta

Vespiodes leopolditertii

Apiocera An philippii

Apiocera Ap hardyi

Heteromydas bicolor

Nemomydas melanopogon

Apiocera P martinorum

Dioctria hyalipennis

Apiocera P barri

Phyllomydas bruesii

Ospriocerus aeacus

Nomoneura paradoxa

Apiocera P haruspex

Apiocera Ap pica

Afroleptomydas A suffusipennis

Cacatuopyga basifascia

Afroleptomydas A paganus

Toxophora maxima

Cephalocera sp Ratelfontein

Gonioscelis ventralis

Leptomydas notos

Afroleptomydas A rufithorax

Arenomydas caerulescens

Apiocera R braunsi

Namibimydas stuckenbergi

Mydaselpis ngurumani

Syllegomydas N arnoldi M

Opomydas townsendi

Ectyphus pinguis

Apiocera Ap maritima

Plyomydas peruviensis M

Mydas tricinctus

Apiocera Ap sp Lloyd

Eremomidas sp nov Tiega M

Syll gen nov sp nov Gobabeb

Syllegomydas N heothinos

Mydas xanthopterus

Lachnocorynus sp nov Kotwa M

Afroleptomydas A omeri

Apiocera Ap sp Stuart M

Pilica formidolosa

Afroleptomydas C mauricei

Miltinus minutus

Nomoneura caffra

Ommatius tibialis

Pseudorhopalia mirandai

Nomoneuroides natalensis

Hessemydas parkeri M

Neorhynchocephalus volaticus

Afroleptomydas A consanguinensis

Apiocera P aldrichi

Syllegomydas S palestinensis

Namadytes vansoni

Apiocera An obscura

Ectyphus armipes

Hessemydas daugeroni

Hispanomydas hispanicus

Eremohaplomydas desertorum

Leptomydas lusitanicus

Lasiopogon cinctus

Afroleptomydas A sp Obib

Parectyphus namibiensis

Philonicus albiceps

Afroleptomydas A inhacae

Stratiomydas lividus

Holcocephala calva

Leptomydas rapti M

Rhaphiomidas maehleri

Ectyphus amboseli M

Apiocera Ap similis

Mitrodetus dimidiatus

Diochlistinae Argentina F

Mitrodetus microglossa

Apiocera P parahydra

Apiocera Ap tonnoiri

Apiocera Ap newmani

Neorhaphiomidas sp Mingenew M † Cretomydas santanensis, Crato Formation, Brazil

≈ 112 myo

Placement of † Cretomydas santanensis

1 of the mp trees

◊ explicitly included in matrix

◊ 80% missing data

◊ wing venation very similar to Diochlistus (Australia) and Rhaphiomidas (wNA)

... results

13

Nemomydas lamia

Phyllomydas parvulus

Heteromydas bicolor

Megascelus albovillosus

Leptomydas corsicanus

Mitrodetus dimidiatus

Nemomydas venosus

Syllegomydas S vittatus

Cacatuopyga ruficornis M

Phyllomydas bruesii

Nomoneura paradoxa

Megascelus nigricornis

Miltinus viduatus

Nemomydas melanopogon

Syllegomydas S algiricus

Mydas clavatus

Plyomydas peruviensis M

Hessemydas daugeroni

Diochlistus auripennis M

Pseudorhopalia mirandai

Syllegomydas S elachys

Syll gen nov sp nov minute proboscis F

Perissocerus transcaspicus

Anomalomydas australicus F

Rhopalia olivieri

Stratiomydas lividus

Afroleptomydas A consanguinensis

Midacritus wagenknechti

Halterorchis karooensis F

Diochlistus melleipennis

Vespiodes phaios

Leptomydas turcicus

Arenomydas caerulescens

Apiophora quadricinctata

Hessemydas parkeri M

Plyomydas sp nov Tandil

Syllegomydas N astrictus

Neolaparopsis puncturatus

Syllegomydas N dispar

Arenomydas callosus

Neorhaphiomidas sp Mandurah

Namadytes cimbebasiensis

Afroleptomydas A suffusipennis

Syllegomydas S palestinensis

Arenomydas namaquensis M

Diochlistinae Argentina F

Afroleptomydas A sp Clanwilliam

Leptomydas sardous

Arenomydas partitus

Mydas xanthopterus

Namibimydas stuckenbergi

Afroleptomydas A paganus

Opomydas townsendi

Cephalocera sp Sors Sors

Mydaselpis ngurumani

Mydaselpis sp nov Gamsberg M

Afroleptomydas C sp Stampriet

Eremomidas sp nov Tiega M

Neorhaphiomidas hardyi

Pseudonomoneura bajaensis

Ectyphus armipes

Mitrodetus dentitarsis

Nomoneura caffra

Neorhaphiomidas sp Yanchep

Diochlistus gracilis

Messiasia californica M

Dolichogaster brevicornis

Rhaphiomidas parkeri

Afroleptomydas C sp Koeroegab

Pseudonomoneura hirta

Leptomydas notos

Leptomydas lusitanicus

Rhopalia annulata

Nemomydas brachyrhynchus

Cacatuopyga basifascia

Cretomydas santanensis

Paramydas igniticornis

Neorhaphiomidas sp Mingenew M

Miltinus minutus

Syllegomydas S lineatus

Halterorchis sp nov Cape Recife M

Megascelus melanoproctus

Afroleptomydas C mauricei

Namadytes vansoni

Syllegomydas N arnoldi M

Rhopalia cincta

Messiasia painteri

Eremohaplomydas desertorum

Syllegomydas S proximus

Nemomydas pantherinus

Afroleptomydas A omeri

Lachnocorynus sp nov Kotwa M

Afroleptomydas A opacicinctus

Rhaphiomidas hasbroucki

Namibimydas psamminos

Pseudonomoneura micheneri

Nothomydas picketti

Ectyphus pinguis

Afroleptomydas A rufithorax

Leptomydas rapti M

Afroleptomydas C sp Gobabeb

Syll gen nov sp nov Gobabeb

Miltinus stenogaster

Miltinus maculipennis

Midacritus stuardoanus M

Parectyphus namibiensis

Afroleptomydas A sp Obib

Hispanomydas hispanicus

Perissocerus dumonti F

Mitrodetus nanoglossa M

Cephalocera sp Papendorp

Vespiodes leopolditertii

Haplomydas crassipes

Nemomydas gruenbergi M

Anomalomydas mackerrasi

Mydaselpis sp nov Daan

Syllegomydas sp nov Ruacana

Cephalocera sp Ratelfontein

Cleamydas gen nov sp nov Ugab F

Syllegomydas N heothinos

Hessemydas seyrigi

Nomoneuroides natalensis

Mitrodetus microglossa

Stenomydas gen nov sp nov Panne M

Tongamya miranda

Cephalocera longirostris

Eremomidas arabicus

Apiophora paulseni

Afroleptomydas A inhacae

Notosyllegomydas brincki

Ectyphus amboseli M

Rhaphiomidas maehleri

Eremomidas bek

Mydas tricinctus

Cephalocera imitata M

Mydidae

Australia

Argentina/Chile

Diochlistinae

Diochlistinae

≈ 112 myo

Placement of † Cretomydas santanensis

Diochlistus auripennis, Australia © P. Chew

Non-monophyly of Diochlistinae

1 of the mp treesNemomydas venosus

Halterorchis karooensis F

Apiocera Ap brevicornis

Dipalta serpentina

Namadytes cimbebasiensis

Rhopalia annulata

Pseudonomoneura micheneri

Diochlistus gracilis

Apiophora paulseni

Eremomidas bek

Mydaselpis sp nov Gamsberg M

Pegesimallus laticornis

Anomalomydas australicus F

Apiocera R alastor M

Arenomydas namaquensis M

Apiocera Ap imminuta

Laphria aktis

Megascelus albovillosus

Afroleptomydas C sp Stampriet

Leptogaster cylindrica

Diogmites grossus

Dolichogaster brevicornis

Halterorchis sp nov Cape Recife M

Paramydas igniticornis

Nemomydas brachyrhynchus

Miltinus maculipennis

Rhopalia cincta

Hessemydas seyrigi

Cacatuopyga ruficornis M

Nemomydas pantherinus

Nothomydas picketti

Eremomidas arabicus

Syllegomydas N astrictus

Bombylius major

Stichopogon punctum

Rhaphiomidas hasbroucki

Phycus frommeri

Messiasia painteri

Cretomydas santanensis

Apiophora quadricinctata

Afroleptomydas C sp Koeroegab

Neorhaphiomidas hardyi

Mydaselpis sp nov Daan

Notosyllegomydas brincki

Mitrodetus nanoglossa M

Syllegomydas sp nov Ruacana

Cephalocera sp Papendorp

Leptomydas corsicanus

Plyomydas sp nov Tandil

Hemigephyra atra

Neolaparopsis puncturatus

Apiocera P sp Colnett M

Nemomydas lamia

Namibimydas psamminos

Nemestrinus nigrovillosus

Syll gen nov sp nov minute proboscis F

Midacritus wagenknechti

Apiocera P mexicana M

Limata tenuicornis F

Emphysomera pallidapex

Mydas clavatus

Leptomydas sardous

Cephalocera longirostris

Apiocera P arena

Syllegomydas N dispar

Apiocera Ap asilica

Afroleptomydas C sp Gobabeb

Anomalomydas mackerrasi

Nemomydas gruenbergi M

Neorhaphiomidas sp Mandurah

Phyllomydas parvulus

Vespiodes phaios

Rhaphiomidas parkeri

Prorates sp Escalante

Tongamya miranda

Midacritus stuardoanus M

Rhopalia olivieri

Cephalocera sp Sors Sors

Apsilocephala longistyla

Apiocera Ap sp Wurarga

Cephalocera imitata M

Perissocerus transcaspicus

Megascelus melanoproctus

Syllegomydas S lineatus

Apiocera P parkeri

Syllegomydas S algiricus

Apiocera Ap minor

Miltinus stenogaster

Afroleptomydas A opacicinctus

Cleamydas gen nov sp nov Ugab F

Haplomydas crassipes

Stenomydas gen nov sp nov Panne M

Mitrodetus dentitarsis

Syllegomydas S vittatus

Neorhaphiomidas sp Yanchep

Arenomydas callosus

Hoplistomerus nobilis

Syllegomydas S elachys

Megascelus nigricornis

Messiasia californica M

Apiocera P painteri

Leptomydas turcicus

Perissocerus dumonti F

Afroleptomydas A sp Clanwilliam

Damalis monochaetes

Arenomydas partitus

Neolophonotus bimaculatusMachimus occidentalis

Diochlistus melleipennis

Euscelidia pulchra

Miltinus viduatus

Apiocera P bilineata

Pseudonomoneura bajaensis

Eulonchus sp Six Rivers M

Diochlistus auripennis M

Syllegomydas S proximus

Pseudonomoneura hirta

Vespiodes leopolditertii

Apiocera An philippii

Apiocera Ap hardyi

Heteromydas bicolor

Nemomydas melanopogon

Apiocera P martinorum

Dioctria hyalipennis

Apiocera P barri

Phyllomydas bruesii

Ospriocerus aeacus

Nomoneura paradoxa

Apiocera P haruspex

Apiocera Ap pica

Afroleptomydas A suffusipennis

Cacatuopyga basifascia

Afroleptomydas A paganus

Toxophora maxima

Cephalocera sp Ratelfontein

Gonioscelis ventralis

Leptomydas notos

Afroleptomydas A rufithorax

Arenomydas caerulescens

Apiocera R braunsi

Namibimydas stuckenbergi

Mydaselpis ngurumani

Syllegomydas N arnoldi M

Opomydas townsendi

Ectyphus pinguis

Apiocera Ap maritima

Plyomydas peruviensis M

Mydas tricinctus

Apiocera Ap sp Lloyd

Eremomidas sp nov Tiega M

Syll gen nov sp nov Gobabeb

Syllegomydas N heothinos

Mydas xanthopterus

Lachnocorynus sp nov Kotwa M

Afroleptomydas A omeri

Apiocera Ap sp Stuart M

Pilica formidolosa

Afroleptomydas C mauricei

Miltinus minutus

Nomoneura caffra

Ommatius tibialis

Pseudorhopalia mirandai

Nomoneuroides natalensis

Hessemydas parkeri M

Neorhynchocephalus volaticus

Afroleptomydas A consanguinensis

Apiocera P aldrichi

Syllegomydas S palestinensis

Namadytes vansoni

Apiocera An obscura

Ectyphus armipes

Hessemydas daugeroni

Hispanomydas hispanicus

Eremohaplomydas desertorum

Leptomydas lusitanicus

Lasiopogon cinctus

Afroleptomydas A sp Obib

Parectyphus namibiensis

Philonicus albiceps

Afroleptomydas A inhacae

Stratiomydas lividus

Holcocephala calva

Leptomydas rapti M

Rhaphiomidas maehleri

Ectyphus amboseli M

Apiocera Ap similis

Mitrodetus dimidiatus

Diochlistinae Argentina F

Mitrodetus microglossa

Apiocera P parahydra

Apiocera Ap tonnoiri

Apiocera Ap newmani

Neorhaphiomidas sp Mingenew M

Mitdrodetus dentitarsis, Chile © S. Marshall

† Cretomydas santanensis, Crato Formation, Brazil

... results

14

Apiocera alleni © G. Balmer

Relationships within Apioceridae

Nemomydas venosus

Halterorchis karooensis F

Apiocera Ap brevicornis

Dipalta serpentina

Namadytes cimbebasiensis

Rhopalia annulata

Pseudonomoneura micheneri

Diochlistus gracilis

Apiophora paulseni

Eremomidas bek

Mydaselpis sp nov Gamsberg M

Pegesimallus laticornis

Anomalomydas australicus F

Apiocera R alastor M

Arenomydas namaquensis M

Apiocera Ap imminuta

Laphria aktis

Megascelus albovillosus

Afroleptomydas C sp Stampriet

Leptogaster cylindrica

Diogmites grossus

Dolichogaster brevicornis

Halterorchis sp nov Cape Recife M

Paramydas igniticornis

Nemomydas brachyrhynchus

Miltinus maculipennis

Rhopalia cincta

Hessemydas seyrigi

Cacatuopyga ruficornis M

Nemomydas pantherinus

Nothomydas picketti

Eremomidas arabicus

Syllegomydas N astrictus

Bombylius major

Stichopogon punctum

Rhaphiomidas hasbroucki

Phycus frommeri

Messiasia painteri

Cretomydas santanensis

Apiophora quadricinctata

Afroleptomydas C sp Koeroegab

Neorhaphiomidas hardyi

Mydaselpis sp nov Daan

Notosyllegomydas brincki

Mitrodetus nanoglossa M

Syllegomydas sp nov Ruacana

Cephalocera sp Papendorp

Leptomydas corsicanus

Plyomydas sp nov Tandil

Hemigephyra atra

Neolaparopsis puncturatus

Apiocera P sp Colnett M

Nemomydas lamia

Namibimydas psamminos

Nemestrinus nigrovillosus

Syll gen nov sp nov minute proboscis F

Midacritus wagenknechti

Apiocera P mexicana M

Limata tenuicornis F

Emphysomera pallidapex

Mydas clavatus

Leptomydas sardous

Cephalocera longirostris

Apiocera P arena

Syllegomydas N dispar

Apiocera Ap asilica

Afroleptomydas C sp Gobabeb

Anomalomydas mackerrasi

Nemomydas gruenbergi M

Neorhaphiomidas sp Mandurah

Phyllomydas parvulus

Vespiodes phaios

Rhaphiomidas parkeri

Prorates sp Escalante

Tongamya miranda

Midacritus stuardoanus M

Rhopalia olivieri

Cephalocera sp Sors Sors

Apsilocephala longistyla

Apiocera Ap sp Wurarga

Cephalocera imitata M

Perissocerus transcaspicus

Megascelus melanoproctus

Syllegomydas S lineatus

Apiocera P parkeri

Syllegomydas S algiricus

Apiocera Ap minor

Miltinus stenogaster

Afroleptomydas A opacicinctus

Cleamydas gen nov sp nov Ugab F

Haplomydas crassipes

Stenomydas gen nov sp nov Panne M

Mitrodetus dentitarsis

Syllegomydas S vittatus

Neorhaphiomidas sp Yanchep

Arenomydas callosus

Hoplistomerus nobilis

Syllegomydas S elachys

Megascelus nigricornis

Messiasia californica M

Apiocera P painteri

Leptomydas turcicus

Perissocerus dumonti F

Afroleptomydas A sp Clanwilliam

Damalis monochaetes

Arenomydas partitus

Neolophonotus bimaculatusMachimus occidentalis

Diochlistus melleipennis

Euscelidia pulchra

Miltinus viduatus

Apiocera P bilineata

Pseudonomoneura bajaensis

Eulonchus sp Six Rivers M

Diochlistus auripennis M

Syllegomydas S proximus

Pseudonomoneura hirta

Vespiodes leopolditertii

Apiocera An philippii

Apiocera Ap hardyi

Heteromydas bicolor

Nemomydas melanopogon

Apiocera P martinorum

Dioctria hyalipennis

Apiocera P barri

Phyllomydas bruesii

Ospriocerus aeacus

Nomoneura paradoxa

Apiocera P haruspex

Apiocera Ap pica

Afroleptomydas A suffusipennis

Cacatuopyga basifascia

Afroleptomydas A paganus

Toxophora maxima

Cephalocera sp Ratelfontein

Gonioscelis ventralis

Leptomydas notos

Afroleptomydas A rufithorax

Arenomydas caerulescens

Apiocera R braunsi

Namibimydas stuckenbergi

Mydaselpis ngurumani

Syllegomydas N arnoldi M

Opomydas townsendi

Ectyphus pinguis

Apiocera Ap maritima

Plyomydas peruviensis M

Mydas tricinctus

Apiocera Ap sp Lloyd

Eremomidas sp nov Tiega M

Syll gen nov sp nov Gobabeb

Syllegomydas N heothinos

Mydas xanthopterus

Lachnocorynus sp nov Kotwa M

Afroleptomydas A omeri

Apiocera Ap sp Stuart M

Pilica formidolosa

Afroleptomydas C mauricei

Miltinus minutus

Nomoneura caffra

Ommatius tibialis

Pseudorhopalia mirandai

Nomoneuroides natalensis

Hessemydas parkeri M

Neorhynchocephalus volaticus

Afroleptomydas A consanguinensis

Apiocera P aldrichi

Syllegomydas S palestinensis

Namadytes vansoni

Apiocera An obscura

Ectyphus armipes

Hessemydas daugeroni

Hispanomydas hispanicus

Eremohaplomydas desertorum

Leptomydas lusitanicus

Lasiopogon cinctus

Afroleptomydas A sp Obib

Parectyphus namibiensis

Philonicus albiceps

Afroleptomydas A inhacae

Stratiomydas lividus

Holcocephala calva

Leptomydas rapti M

Rhaphiomidas maehleri

Ectyphus amboseli M

Apiocera Ap similis

Mitrodetus dimidiatus

Diochlistinae Argentina F

Mitrodetus microglossa

Apiocera P parahydra

Apiocera Ap tonnoiri

Apiocera Ap newmani

Neorhaphiomidas sp Mingenew M

1 of the mp treesunambiguous optimizationBremer below branches

... results

Apiocera (Apiocera) tonnoiri

Apiocera (Apiocera) spp. 11 species

Apiocera (Pyrocera) spp. 11 species

Apiocera (Anypenus) spp. 2 species

Apiocera (Apiocera) imminuta

Apiocera (Ripidosyrma) spp. 2 species

Apioceridae100

33

9975

2

74

0

47

0

11

1

182

1

137

1

231

1

214

11

20953

2

4

2

230

1

113

1

89

12

79

90

1

178

01

113

79

11

77

154

6

100

4

133

4

209

2

6

≥10

8

2

3

31

4 Apiocera (Apiocera) sp. © J. Hort

15

Rhaphiomidas auratus © G. Ballmer

Anomalomydas mackerrasi © J. Hort

Neorhaphiomidas villosa © F. Hort

Earliest branchings within Mydidae

Nemomydas venosus

Halterorchis karooensis F

Apiocera Ap brevicornis

Dipalta serpentina

Namadytes cimbebasiensis

Rhopalia annulata

Pseudonomoneura micheneri

Diochlistus gracilis

Apiophora paulseni

Eremomidas bek

Mydaselpis sp nov Gamsberg M

Pegesimallus laticornis

Anomalomydas australicus F

Apiocera R alastor M

Arenomydas namaquensis M

Apiocera Ap imminuta

Laphria aktis

Megascelus albovillosus

Afroleptomydas C sp Stampriet

Leptogaster cylindrica

Diogmites grossus

Dolichogaster brevicornis

Halterorchis sp nov Cape Recife M

Paramydas igniticornis

Nemomydas brachyrhynchus

Miltinus maculipennis

Rhopalia cincta

Hessemydas seyrigi

Cacatuopyga ruficornis M

Nemomydas pantherinus

Nothomydas picketti

Eremomidas arabicus

Syllegomydas N astrictus

Bombylius major

Stichopogon punctum

Rhaphiomidas hasbroucki

Phycus frommeri

Messiasia painteri

Cretomydas santanensis

Apiophora quadricinctata

Afroleptomydas C sp Koeroegab

Neorhaphiomidas hardyi

Mydaselpis sp nov Daan

Notosyllegomydas brincki

Mitrodetus nanoglossa M

Syllegomydas sp nov Ruacana

Cephalocera sp Papendorp

Leptomydas corsicanus

Plyomydas sp nov Tandil

Hemigephyra atra

Neolaparopsis puncturatus

Apiocera P sp Colnett M

Nemomydas lamia

Namibimydas psamminos

Nemestrinus nigrovillosus

Syll gen nov sp nov minute proboscis F

Midacritus wagenknechti

Apiocera P mexicana M

Limata tenuicornis F

Emphysomera pallidapex

Mydas clavatus

Leptomydas sardous

Cephalocera longirostris

Apiocera P arena

Syllegomydas N dispar

Apiocera Ap asilica

Afroleptomydas C sp Gobabeb

Anomalomydas mackerrasi

Nemomydas gruenbergi M

Neorhaphiomidas sp Mandurah

Phyllomydas parvulus

Vespiodes phaios

Rhaphiomidas parkeri

Prorates sp Escalante

Tongamya miranda

Midacritus stuardoanus M

Rhopalia olivieri

Cephalocera sp Sors Sors

Apsilocephala longistyla

Apiocera Ap sp Wurarga

Cephalocera imitata M

Perissocerus transcaspicus

Megascelus melanoproctus

Syllegomydas S lineatus

Apiocera P parkeri

Syllegomydas S algiricus

Apiocera Ap minor

Miltinus stenogaster

Afroleptomydas A opacicinctus

Cleamydas gen nov sp nov Ugab F

Haplomydas crassipes

Stenomydas gen nov sp nov Panne M

Mitrodetus dentitarsis

Syllegomydas S vittatus

Neorhaphiomidas sp Yanchep

Arenomydas callosus

Hoplistomerus nobilis

Syllegomydas S elachys

Megascelus nigricornis

Messiasia californica M

Apiocera P painteri

Leptomydas turcicus

Perissocerus dumonti F

Afroleptomydas A sp Clanwilliam

Damalis monochaetes

Arenomydas partitus

Neolophonotus bimaculatusMachimus occidentalis

Diochlistus melleipennis

Euscelidia pulchra

Miltinus viduatus

Apiocera P bilineata

Pseudonomoneura bajaensis

Eulonchus sp Six Rivers M

Diochlistus auripennis M

Syllegomydas S proximus

Pseudonomoneura hirta

Vespiodes leopolditertii

Apiocera An philippii

Apiocera Ap hardyi

Heteromydas bicolor

Nemomydas melanopogon

Apiocera P martinorum

Dioctria hyalipennis

Apiocera P barri

Phyllomydas bruesii

Ospriocerus aeacus

Nomoneura paradoxa

Apiocera P haruspex

Apiocera Ap pica

Afroleptomydas A suffusipennis

Cacatuopyga basifascia

Afroleptomydas A paganus

Toxophora maxima

Cephalocera sp Ratelfontein

Gonioscelis ventralis

Leptomydas notos

Afroleptomydas A rufithorax

Arenomydas caerulescens

Apiocera R braunsi

Namibimydas stuckenbergi

Mydaselpis ngurumani

Syllegomydas N arnoldi M

Opomydas townsendi

Ectyphus pinguis

Apiocera Ap maritima

Plyomydas peruviensis M

Mydas tricinctus

Apiocera Ap sp Lloyd

Eremomidas sp nov Tiega M

Syll gen nov sp nov Gobabeb

Syllegomydas N heothinos

Mydas xanthopterus

Lachnocorynus sp nov Kotwa M

Afroleptomydas A omeri

Apiocera Ap sp Stuart M

Pilica formidolosa

Afroleptomydas C mauricei

Miltinus minutus

Nomoneura caffra

Ommatius tibialis

Pseudorhopalia mirandai

Nomoneuroides natalensis

Hessemydas parkeri M

Neorhynchocephalus volaticus

Afroleptomydas A consanguinensis

Apiocera P aldrichi

Syllegomydas S palestinensis

Namadytes vansoni

Apiocera An obscura

Ectyphus armipes

Hessemydas daugeroni

Hispanomydas hispanicus

Eremohaplomydas desertorum

Leptomydas lusitanicus

Lasiopogon cinctus

Afroleptomydas A sp Obib

Parectyphus namibiensis

Philonicus albiceps

Afroleptomydas A inhacae

Stratiomydas lividus

Holcocephala calva

Leptomydas rapti M

Rhaphiomidas maehleri

Ectyphus amboseli M

Apiocera Ap similis

Mitrodetus dimidiatus

Diochlistinae Argentina F

Mitrodetus microglossa

Apiocera P parahydra

Apiocera Ap tonnoiri

Apiocera Ap newmani

Neorhaphiomidas sp Mingenew M

1 of the mp treesunambiguous optimizationBremer below branches* newly delimited taxon

Diochlistus tenebrosus © M. Jefferies

Megascelus albovillosus

Megascelus nigricornis

Diochlistus auripennis M

Anomalomydas australicus F

Diochlistus melleipennis

Neorhaphiomidas sp MandurahNeorhaphiomidas hardyi

Neorhaphiomidas sp Yanchep

Diochlistus gracilis

Rhaphiomidas parkeri

† Cretomydas santanensis

Neorhaphiomidas sp Mingenew M

Megascelus melanoproctus

Rhaphiomidas hasbroucki

Anomalomydas mackerrasi

Tongamya miranda

Rhaphiomidas maehleri

remaining Mydidae117 species

≥10

≥10

4

104

≥10

1

≥10

10

50

20

2019

0

18

1

17

0

8

5

7

5

203

2

176

2

114

2

80

1

149

21

148145

1

143

1

47

0

4

0

3

0

160

2

158

3

154

5

152

4

211

2

157

1

116

1

0

15175

2

56

1

47

2

44

1

12

0

169

1

167

1

132

0

104

1

1

158154

1

104

0

77

0

63

1

15

1

106

11

10294

0

93

0

92

0

91

0

72

1

202

7

107

1

51

13

50

0

45

0

39

2

38

1

5

2

60

1

59

0

58

1

57

1

73

2

46 203

1

101

0

96

0

89

1

1

6966

1

53

4

52

4

10

0

6

1

222

0

217

0

160

0

152

3

Rhaphiomidinae

Anomalomydinae

Megascelinae

Diochlistinae *

Mydidae

... results

16

Center of origin

Nemomydas venosus

Halterorchis karooensis F

Apiocera Ap brevicornis

Dipalta serpentina

Namadytes cimbebasiensis

Rhopalia annulata

Pseudonomoneura micheneri

Diochlistus gracilis

Apiophora paulseni

Eremomidas bek

Mydaselpis sp nov Gamsberg M

Pegesimallus laticornis

Anomalomydas australicus F

Apiocera R alastor M

Arenomydas namaquensis M

Apiocera Ap imminuta

Laphria aktis

Megascelus albovillosus

Afroleptomydas C sp Stampriet

Leptogaster cylindrica

Diogmites grossus

Dolichogaster brevicornis

Halterorchis sp nov Cape Recife M

Paramydas igniticornis

Nemomydas brachyrhynchus

Miltinus maculipennis

Rhopalia cincta

Hessemydas seyrigi

Cacatuopyga ruficornis M

Nemomydas pantherinus

Nothomydas picketti

Eremomidas arabicus

Syllegomydas N astrictus

Bombylius major

Stichopogon punctum

Rhaphiomidas hasbroucki

Phycus frommeri

Messiasia painteri

Cretomydas santanensis

Apiophora quadricinctata

Afroleptomydas C sp Koeroegab

Neorhaphiomidas hardyi

Mydaselpis sp nov Daan

Notosyllegomydas brincki

Mitrodetus nanoglossa M

Syllegomydas sp nov Ruacana

Cephalocera sp Papendorp

Leptomydas corsicanus

Plyomydas sp nov Tandil

Hemigephyra atra

Neolaparopsis puncturatus

Apiocera P sp Colnett M

Nemomydas lamia

Namibimydas psamminos

Nemestrinus nigrovillosus

Syll gen nov sp nov minute proboscis F

Midacritus wagenknechti

Apiocera P mexicana M

Limata tenuicornis F

Emphysomera pallidapex

Mydas clavatus

Leptomydas sardous

Cephalocera longirostris

Apiocera P arena

Syllegomydas N dispar

Apiocera Ap asilica

Afroleptomydas C sp Gobabeb

Anomalomydas mackerrasi

Nemomydas gruenbergi M

Neorhaphiomidas sp Mandurah

Phyllomydas parvulus

Vespiodes phaios

Rhaphiomidas parkeri

Prorates sp Escalante

Tongamya miranda

Midacritus stuardoanus M

Rhopalia olivieri

Cephalocera sp Sors Sors

Apsilocephala longistyla

Apiocera Ap sp Wurarga

Cephalocera imitata M

Perissocerus transcaspicus

Megascelus melanoproctus

Syllegomydas S lineatus

Apiocera P parkeri

Syllegomydas S algiricus

Apiocera Ap minor

Miltinus stenogaster

Afroleptomydas A opacicinctus

Cleamydas gen nov sp nov Ugab F

Haplomydas crassipes

Stenomydas gen nov sp nov Panne M

Mitrodetus dentitarsis

Syllegomydas S vittatus

Neorhaphiomidas sp Yanchep

Arenomydas callosus

Hoplistomerus nobilis

Syllegomydas S elachys

Megascelus nigricornis

Messiasia californica M

Apiocera P painteri

Leptomydas turcicus

Perissocerus dumonti F

Afroleptomydas A sp Clanwilliam

Damalis monochaetes

Arenomydas partitus

Neolophonotus bimaculatusMachimus occidentalis

Diochlistus melleipennis

Euscelidia pulchra

Miltinus viduatus

Apiocera P bilineata

Pseudonomoneura bajaensis

Eulonchus sp Six Rivers M

Diochlistus auripennis M

Syllegomydas S proximus

Pseudonomoneura hirta

Vespiodes leopolditertii

Apiocera An philippii

Apiocera Ap hardyi

Heteromydas bicolor

Nemomydas melanopogon

Apiocera P martinorum

Dioctria hyalipennis

Apiocera P barri

Phyllomydas bruesii

Ospriocerus aeacus

Nomoneura paradoxa

Apiocera P haruspex

Apiocera Ap pica

Afroleptomydas A suffusipennis

Cacatuopyga basifascia

Afroleptomydas A paganus

Toxophora maxima

Cephalocera sp Ratelfontein

Gonioscelis ventralis

Leptomydas notos

Afroleptomydas A rufithorax

Arenomydas caerulescens

Apiocera R braunsi

Namibimydas stuckenbergi

Mydaselpis ngurumani

Syllegomydas N arnoldi M

Opomydas townsendi

Ectyphus pinguis

Apiocera Ap maritima

Plyomydas peruviensis M

Mydas tricinctus

Apiocera Ap sp Lloyd

Eremomidas sp nov Tiega M

Syll gen nov sp nov Gobabeb

Syllegomydas N heothinos

Mydas xanthopterus

Lachnocorynus sp nov Kotwa M

Afroleptomydas A omeri

Apiocera Ap sp Stuart M

Pilica formidolosa

Afroleptomydas C mauricei

Miltinus minutus

Nomoneura caffra

Ommatius tibialis

Pseudorhopalia mirandai

Nomoneuroides natalensis

Hessemydas parkeri M

Neorhynchocephalus volaticus

Afroleptomydas A consanguinensis

Apiocera P aldrichi

Syllegomydas S palestinensis

Namadytes vansoni

Apiocera An obscura

Ectyphus armipes

Hessemydas daugeroni

Hispanomydas hispanicus

Eremohaplomydas desertorum

Leptomydas lusitanicus

Lasiopogon cinctus

Afroleptomydas A sp Obib

Parectyphus namibiensis

Philonicus albiceps

Afroleptomydas A inhacae

Stratiomydas lividus

Holcocephala calva

Leptomydas rapti M

Rhaphiomidas maehleri

Ectyphus amboseli M

Apiocera Ap similis

Mitrodetus dimidiatus

Diochlistinae Argentina F

Mitrodetus microglossa

Apiocera P parahydra

Apiocera Ap tonnoiri

Apiocera Ap newmani

Neorhaphiomidas sp Mingenew M

Rhaphiomidas auratus © G. Ballmer

DIVA analysis based on 1 of the mp treesancestral area above branches wNA = western North America Aus = Australia sAfr = southern Africa ArgChi = Argentina/Chile Neo = Neotropical (E of Andes)

Apiocera (Apiocera) sp. © J. Hort

Megascelus albovillosus

Megascelus nigricornis

Diochlistus auripennis M

Anomalomydas australicus F

Diochlistus melleipennis

Neorhaphiomidas sp MandurahNeorhaphiomidas hardyi

Neorhaphiomidas sp Yanchep

Diochlistus gracilis

Rhaphiomidas parkeri

† Cretomydas santanensis

Neorhaphiomidas sp Mingenew M

Megascelus melanoproctus

Rhaphiomidas hasbroucki

Anomalomydas mackerrasi

Tongamya miranda

Rhaphiomidas maehleri

remaining Mydidae117 species

Apiocera (Apiocera) tonnoiri

Apiocera (Apiocera) spp. 11 species

Apiocera (Pyrocera) spp. 11 species

Apiocera (Anypenus) spp. 2 species

Apiocera (Apiocera) imminuta

Apiocera (Ripidosyrma) spp. 2 species

western North America

Namibia + South Africa

Brazil

Australia

Australia

Chile

Australia

wNA

Aus+Neo

Aus

wNA+Aus

Aus

Aus

85 alternatives

47 alternatives

sAfr+Aus / Aus+ArgChi / sAfr+Aus+ArgChi

Chile

western North America

Australia

Australia

South AfricawNA+sAFRwNA+Aus / sAfr+Aus / wNA+sAfr+Aus

Aus+ArgChiAus

Aus

wNA / Aus

Apioceridae

Mydidae

... results

17

A. aux A. bux sp. nov. B. aux † C. aux D. aux D. bux D. cux sp. nov.

evolutionary time

.... future

18

Acknowledgements◊ museum curators – collectors – photographers

◊ National Science Foundation – REVSYS DEB-0919333

◊ Willi Hennig Society for free access to TNT – Pablo Goloboff

..... acknowledgements