1. 1
Phylogenetic relationships within Mydidae based on
morphology of imagines (Diptera: Asiloidea)
Torsten Dikow
Biodiversity Synthesis Center
Field Museum of Natural History

2. . introduction 2
Mydidae – “mydas flies”
◊ Imagines flower-feeders, some do not feed at all
◊ Larvae possibly predatory
◊ 472* species and 66 genera world-wide
◊ 11 subfamily taxa
◊ Imagines 9 – 60 mm (wing length up to 52 mm)
Opomydas townsendi
◊ World-wide primarily in arid regions
◊ Oldest definitive fossil Cretomydas santanensis
› Crato Formation, Brazil
› ≈ 110 Million years
* includes species currently being described Cretomydas santanensis

3. . introduction 3
Taxa overview
Taxon Known Diversity Known Distribution
Genera Species Afr Aus Nea Neo Ori Pal
Anomalomydinae 1 2 ×
Apiophorinae 5 29 × ×
Cacatuopyginae 2 6 ×
Diochlistinae 2 20 × ×
Ectyphinae 4 10* × ×
Anomalomydas mackerrasi
Leptomydinae 6 54* × × × × ×
Megascelinae 3 15* × × ×
Mydinae 12 84 × ×
Rhaphiomidinae 1 23* ×
Rhopaliinae 4 21 × × ×
Syllegomydinae 25 206* × × ×
unplaced 1 2
* includes species currently being described
Cacatuopyga basifascia

4. 4
Overview of presentation
◊ Previous Phylogenetic Hypotheses
◊ New Phylogenetic Hypothesis
◊ Character Evolution
Miltinus stenogaster © Shaun Winterton
◊ Summary + Future
Stratiomydas lividus © Charles Melton

5. .. previous phylogenetic hypotheses 5
Wilcox & Papavero 1974 Artigas & Papavero 1990
› morphology › morphology
› similarity ae › similarity
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Diochlistinae remaining Mydidae
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Mitrodetus dentitarsis © Steve Marshall

6. .. previous phylogenetic hypotheses 6
Yeates & Irwin 1996 Irwin & Wiegmann 2001
› morphology › 28S rDNA
› parsimony (successive weighting) › maximum likelihood
Asilidae Scenopinidae
Apiocera (Pyrocera) Therevidae
Apiocera (Ripidosyrma) Apiocera (Anypenus) phillippi
Apiocera (Anypenus) Apiocera (Apiocera) sp 1
Apiocera (Apiocera) Apiocera (Apiocera) sp 2
Rhaphiomidinae
Apiocera (Pyrocera) haruspex
Megascelinae
Apiocera (Pyrocera) sp
Rhopaliinae
Rhaphiomidas terminalis Rhaphiomidinae
Cacatuopyginae
Megascelus albovillosus
Leptomydinae Megascelinae
Tongamya stuckenbergi
Syllegomydinae
Diochlistus hackeri Diochlistinae
Anomalomydinae
Rhopalia sp Rhopaliinae
Diochlistinae
Ectyphinae Mitrodetus australis Diochlistinae
Apiophorinae Nemomydas pantherinus Leptomydinae
Mydinae Arenomydas callosus Syllegomydinae

7. .. previous phylogenetic hypotheses 7
Dikow 2009 Dikow unpublished presented at ICD6 in Fukuoka
› morphology › 18S + 28S rDNA, CAD
› parsimony › parsimony (POY)
Bombylius major
Apsilocephala longistylus
Prorates sp (Esacalante)
Phycus frommeri
... Asilidae Hemigephyra atra
Apiocera (Apiocera) sp (Barakula)
Apiocera aldrichi Apiocera (Pyrocera) haruspex Apioceridae
Apioceridae Apiocera (Pyrocera) painteri
Apiocera painteri Stichopogon punctum
Saropogon luteus
Rhaphiomidas maehleri Rhaphiomidinae Laxenecera albicincta
Hoplistomerus nobilis Asilidae
Neorhaphiomidas sp (Mandurah) Megascelinae Gonioscelis ventralis
Leptogaster cylindrica
Opomydas townsendi Ectyphinae Asilus crabroniformis
Promachus amastrus
Mitrodetus dentitarsis Diochlistinae Emphysomera pallidapex
Rhaphiomidas xanthos Rhaphiomidinae
Mitrodetus dentitarsis Diochlistinae
Mydas clavatus Mydinae Opomydas townsendi Ectyphinae
Miltinus cardinalis Apiophorinae
Pseudonomoneura hirta Mydas clavatus Mydinae
Leptomydinae
Mahafalymydas wiegmanni
Nemomydas brachyrhynchus Syllegomydas sp (Nizzanem)
Afroleptomydas sp (Clanwilliam) Syllegomydinae
Afroleptomydas sp (Clanwilliam) Syllegomydinae Afroleptomydas sp (RSA)
Cephalocera imitata
Cephalocera sp (Montegu)
Nemomydas brachyrhynchus
Nemomydas sp (Tesopaco)
Leptomydinae

8. ... new phylogenetic hypothesis 8
Taxon Sampling
◊ 94 Mydidae species (= 19,9 %)
◊ Higher-level taxa
› 40 genera (= 61,5 %)
› 7 tribal taxa (= 87,5 %)
› 11 subfamily taxa (= 100 %)
collecting localities of specimens included
◊ Outgroup taxa
› 37 Asiloidea species
Apioceridae (11), Apsilocephalidae (1), Asilidae (19), Bombyliidae (3), Scenopinidae (1), Therevidae (2)
› 1 Acroceridae, 2 Nemestrinidae
› 1 Tabanidae

9. ... new phylogenetic hypothesis 9
Morphological Characters
◊ 186 discrete, parsimony informative characters
tagma # characters remarks
Head 42
Thorax 83
39 chaetotaxy, sclerites, etc.
21 wing
23 leg
Abdomen 61
16 sclerite shapes, chaetotaxy, etc.
25 ? terminalia
20 / genitalia

10. ... new phylogenetic hypothesis 10
Phylogenetic Analysis
◊ Morphological data
› MacClade (version 4.08)
› WinClada (version 1.0.08)
› all characters non-additive
◊ Tree Search
› TNT (version 1.1 June 2010)
Eremomidas arabicus © Drew Gardner
› Ratchet, TreeDrifting, TreeFusing, Sectorial Searches
(Goloboff 1999, Nixon 1999)
› regular parsimony
› implied weighting (k = 3) (Goloboff 1993)
Heteromydas bicolor © Greg Ballmer

11. ... new phylogenetic hypothesis 11
Limata tenuicornis Tabanidae
Results & Sister-group
Neorhynchocephalus volaticus
Nemestrinus nigrovillosus
Nemestrinidae
Eulonchus sp. (SixRivers) Acroceridae
Prorates sp. (Escalante) Scenopinidae
Toxophora maxima
Bombylius major Bombyliidae
Dipalta serpentina
Apsilocephala longistyla Apsilocephalidae
Hemigephyra atra
Phycus frommeri Therevidae
◊ Parsimony search
Hoplistomerus nobilis
Pilica formidolosa
Laphria aktis
Neolophonotus bimaculatus
Emphysomera pallidapex
Ommatius tibialis
Machimus occidentalis
Asilidae
Philonicus albiceps
› 421 trees, 1485 steps
Stichopogon punctum
Lasiopogon cinctus
Diogmites grossus
Ospriocerus aeacus
Gonioscelis ventralis
Damalis monochaetes
Pegesimallus laticornis
Leptogaster cylindrica
› strict consensus with large polytomy
Euscelidia pulchra
Dioctria hyalipennis
Holcocephala calva
Apiocera (Pyrocera) bilineata
Apiocera (Ripidosyrma) braunsi
Apioceridae
Apiocera (Pyrocera) aldrichi
Apiocera (Pyrocera) painteri
Apiocera (Pyrocera) arena
Apiocera (Pyrocera) parkeri
in derived ingroup
Apiocera (Anypenus) obscura
Apiocera (Anypenus) philippii
Apiocera (Apiocera) sp. (Wurarga)
Apiocera (Apiocera) sp. (Maranoa)
Apiocera (Apiocera) sp. (Stuart)
Rhaphiomidas hasbroucki
Rhaphiomidas parkeri
Rhaphiomidas maehleri
Tongamya miranda
Neorhaphiomidas sp. (Mandurah)
Neorhaphiomidas setosus
◊ Implied weights
Megascelus albovillosus
Megascelus nigricornis
Rhopalia olivieri
Rhopalia annulata
Perissocerus dumonti
Anomalomydas mackerrasi
Cacatuopyga basifascia
Cacatuopyga ruficornis
Messiasia californica
› 6 trees, fit 95.58644
Phyllomydas parvulus
Mydas clavatus
Mydas xanthopterus
Paramydas igniticornis
Stratiomydas lividus
Apiophora quadricinctata
Apiophora paulseni
Miltinus maculipennis
› = 1556 steps
Miltinus viduatus
Miltinus stenogaster
Heteromydas bicolor
Opomydas townsendi
Parectyphus namibiensis
Ectyphus pinguis
Ectyphus sp. nov. (Amboseli)
Haplomydas crassipes
› strict consensus well-resolved
Eremohaplomydas desertorum
Eremohaplomydas sp. nov. (Kotwa)
Eremomidas arabicus
Eremomidas sp. nov. (Tiega)
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Pseudonomoneura micheneri
Pseudonomoneura hirta
Hessemydas parkeri
Hessemydas seyrigi
Nemomydas venosus
Mydidae
Nemomydas brachyrhynchus
Leptomydas gruenbergi
Nemomydas melanopogon
Leptomydas corsicanus
Leptomydas sardous
Leptomydas notos sp. nov.
Leptomydas rapti sp. nov.
Leptomydas turcicus
Eremomidas bek
Leptomydas lusitanicus
Mydaselpis ngurumani sp. nov.
Vespiodes lepolditertii
Limata tenuicornis volaticus
Neorhynchocephalus
Vespiodes phaios sp. nov.
Nemestrinus nigrovillosus
Eulonchus sp. (SixRivers)
Prorates sp. (Escalante)
Toxophora maxima
Mydaselpis sp. nov. (Daan)
Bombylius major
Dipalta serpentina
Apsilocephalaatra
Hemigephyra longistyla
Mydaselpis sp. nov. (Gamsberg)
Phycus frommeri
Hoplistomerus nobilis
Pilica formidolosa
Laphria aktis bimaculatus
Namadytes maculiventris
Neolophonotus
Emphysomera pallidapex
Ommatius tibialis
Machimus occidentalis
Namadytes prozeskyi
Philonicus albiceps
Stichopogoncinctus
Lasiopogon punctum
Diogmites grossus
Cephalocera imitata
Ospriocerus aeacus
Gonioscelis ventralis
Damalis monochaetes
Pegesimalluscylindrica
Cephalocera longirostris
Leptogaster laticornis
Euscelidia pulchra Cephalocera sp. (Papendorp)
Dioctria hyalipennis
Holcocephala calva bilineata
Apiocera (Pyrocera)
Apiocera (Ripidosyrma) braunsi
Apiocera (Pyrocera) aldrichi Neolaparopsis puncturatus
Apiocera (Pyrocera) painteri
Apiocera (Pyrocera) arena
Apiocera (Pyrocera) obscura
Apiocera (Anypenus)parkeri
Arenomydas caerulescens
Apiocera (Anypenus)sp. (Wurarga)
philippii
Apiocera (Apiocera) sp. (Maranoa)
Apiocera (Apiocera)
Apiocera (Apiocera) sp. (Stuart)
Arenomydas callosus
Rhaphiomidas hasbroucki
Rhaphiomidas parkeri
Rhaphiomidas maehleri
Tongamya miranda (Mandurah)
Arenomydas namaquensis
Neorhaphiomidas sp.
Neorhaphiomidas setosus
Megascelus albovillosus
Megascelus nigricornis
Nomoneura caffra
Rhopalia olivieri
Rhopalia annulata
Perissocerus dumonti
Anomalomydas mackerrasi
Nomoneura paradoxa
Cacatuopyga basifascia
Cacatuopyga ruficornis
Messiasia californica
Phyllomydas parvulus
Nomoneuroides natalensis
Mydas clavatus
Mydas xanthopterus
Paramydas igniticornis
Stratiomydas lividus
Halterorchis sp. nov. (Cape Recife)
Apiophora quadricinctata
Apiophora paulseni
Miltinus maculipennis
Miltinus viduatus
gen. nov. + sp. nov. (Panne)
Miltinus stenogaster
Heteromydas bicolor
Opomydas townsendi
Parectyphus namibiensis
gen. nov. + sp. nov. (Ugab)
Ectyphus pinguis (Amboseli)
Ectyphus sp. crassipes
Haplomydas nov.
Eremohaplomydas desertorum
gen. nov. + sp. nov. (Gobabeb)
Eremohaplomydas sp. nov. (Kotwa)
Eremomidas arabicus(Tiega)
Eremomidas sp. nov. Namibimydas sp. nov. (Agate)
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Pseudonomoneura micheneri
Nothomydas sp. nov. (Skerpioenkop)
Pseudonomoneura
Hessemydas seyrigihirta
parkeri
Hessemydas venosus
Nemomydas brachyrhynchus
Afroleptomydas A consanguinensis
Nemomydas gruenbergi
Leptomydas
Nemomydas corsicanus
Leptomydas melanopogon
Afroleptomydas A suffusipennis
Leptomydas sardous nov.
Leptomydas notos sp.
Leptomydas rapti sp.
Leptomydas turcicus nov.
Afroleptomydas C mauricei
Eremomidas lusitanicus
Leptomydas bek
Mydaselpislepolditertii sp. nov.
Vespiodes ngurumani
Afroleptomydas A sp. (Clanwilliam)
Vespiodes phaios sp. nov.
Mydaselpis sp. nov. (Daan)
Mydaselpis sp. nov. (Gamsberg)
Namadytes maculiventris
Afroleptomydas C sp. (Stampriet)
Namadytes prozeskyi
Cephalocera imitata
Cephalocera longirostris
Cephalocera sp. (Papendorp)
Syllegomydas sp. nov. (Ruacana)
Neolaparopsis puncturatus
Arenomydas caerulescens
Arenomydas callosus
Arenomydascaffra
Syllegomydas N arnoldi
Nomoneura namaquensis
Nomoneura paradoxa Syllegomydas N dispar
Nomoneuroides nov. (Cape
sp. natalensis
Halterorchissp. nov. (Panne)Recife)
gen. nov. + sp. nov. (Ugab)
gen. nov. +
gen. nov. + sp. nov. (Gobabeb) Syllegomydas S palestinensis
strict consensus tree from implied weights analysis
Namibimydassp. nov. (Skerpioenkop)
Nothomydas sp. nov. (Agate)
Afroleptomydas A consanguinensis
Afroleptomydas A suffusipennis
Syllegomydas N heothinos sp. nov.
Afroleptomydas C sp. (Clanwilliam)
Afroleptomydas A mauricei
Afroleptomydas C sp.
Syllegomydas sp. nov.(Stampriet)
Syllegomydas S algiricus
Syllegomydas N dispar(Ruacana)
arnoldi
Syllegomydas N palestinensis
Syllegomydas S
Syllegomydas N heothinos
Syllegomydas S elachys sp. nov.
Syllegomydas S algiricus sp. nov.
Syllegomydas S astrictus sp. nov.
Syllegomydas N elachys sp. nov. Syllegomydas N astrictus sp. nov.

12. ... new phylogenetic hypothesis 12
Rhaphiomidas hasbroucki
Rhaphiomidinae
Monophyly & Rhaphiomidinae
Rhaphiomidas parkeri
Rhaphiomidas maehleri
Tongamya miranda
Neorhaphiomidas sp. (Mandurah)
Neorhaphiomidas setosus
Megascelus albovillosus
Megascelinae
Megascelus nigricornis
Rhopalia olivieri
Rhopalia annulata Rhopaliinae
Perissocerus dumonti
Anomalomydas mackerrasi Anomalomydinae
Cacatuopyga basifascia
Cacatuopyga ruficornis Cacatuopyginae
Messiasia californica
Phyllomydas parvulus
Mydas clavatus
Mydas xanthopterus Mydinae
Paramydas igniticornis
Stratiomydas lividus
Apiophora quadricinctata
Apiophora paulseni
Miltinus maculipennis
Miltinus viduatus
Apiophorinae
Miltinus stenogaster
Heteromydas bicolor
Opomydas townsendi
Parectyphus namibiensis
Ectyphus pinguis
Ectyphinae
Ectyphus sp. nov. (Amboseli)
Haplomydas crassipes
Eremohaplomydas desertorum
Eremohaplomydas sp. nov. (Kotwa)
Syllegomydinae
Eremomidas arabicus
Eremomidas sp. nov. (Tiega) Leptomydinae
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Diochlistinae
Pseudonomoneura micheneri
Pseudonomoneura hirta
Hessemydas parkeri
Hessemydas seyrigi
Nemomydas venosus
Nemomydas brachyrhynchus
Rhaphiomidas auratus © Greg Ballmer Leptomydas gruenbergi
Nemomydas melanopogon
Leptomydas corsicanus
Leptomydinae
Leptomydas sardous
Leptomydas notos sp. nov.
Leptomydas rapti sp. nov.
Leptomydas turcicus
Eremomidas bek
Leptomydas lusitanicus
Mydaselpis ngurumani sp. nov.
Vespiodes lepolditertii
Vespiodes phaios sp. nov.
Mydaselpis sp. nov. (Daan)
Mydaselpis sp. nov. (Gamsberg)
Namadytes maculiventris
Namadytes prozeskyi
Cephalocera imitata
Cephalocera longirostris
Cephalocera sp. (Papendorp)
Neolaparopsis puncturatus
Arenomydas caerulescens
Arenomydas callosus
Limata tenuicornis volaticus
Neorhynchocephalus
Nemestrinus nigrovillosus
Eulonchus sp. (SixRivers)
Prorates sp. (Escalante)
Arenomydas namaquensis
Nomoneura caffra
Toxophora maxima
Bombylius major
Dipalta serpentina
Apsilocephalaatra
Hemigephyra longistyla
Nomoneura paradoxa
Phycus frommeri
Hoplistomerus nobilis
Pilica formidolosa
Laphria aktis bimaculatus
Neolophonotus
Emphysomera pallidapex
Nomoneuroides natalensis
Ommatius tibialis
Syllegomydinae
Machimus occidentalis
Philonicus albiceps
Stichopogoncinctus
Lasiopogon punctum
Diogmites grossus
Halterorchis sp. nov. (Cape Recife)
Ospriocerus aeacus
Gonioscelis ventralis
Damalis monochaetes
Pegesimalluscylindrica
Leptogaster laticornis
Euscelidia pulchra
Dioctria hyalipennis
Holcocephala calva bilineata
Apiocera (Pyrocera)
Apiocera (Ripidosyrma) braunsi
Apiocera (Pyrocera) aldrichi gen. nov. + sp. nov. (Panne)
gen. nov. + sp. nov. (Ugab)
Apiocera (Pyrocera) painteri
Apiocera (Pyrocera) arena
Apiocera (Pyrocera) obscura
Apiocera (Anypenus)parkeri
Apiocera (Anypenus)sp. (Wurarga)
philippii
Apiocera (Apiocera) sp. (Maranoa)
gen. nov. + sp. nov. (Gobabeb)
Apiocera (Apiocera)
Apiocera (Apiocera) sp. (Stuart)
Rhaphiomidas hasbroucki
Rhaphiomidas parkeri
monophyletic
Rhaphiomidas maehleri
Tongamya miranda (Mandurah)
Neorhaphiomidas sp.
Neorhaphiomidas setosus
Megascelus albovillosus
Megascelus nigricornis
Rhopalia olivieri Namibimydas sp. nov. (Agate)
Nothomydas sp. nov. (Skerpioenkop)
Rhopalia annulata
Perissocerus dumonti
Anomalomydas mackerrasi
Cacatuopyga basifascia
Cacatuopyga ruficornis
Messiasia californica
Afroleptomydas A consanguinensis
Phyllomydas parvulus
Mydas clavatus
Mydas xanthopterus
Paramydas igniticornis
Stratiomydas lividus
Apiophora quadricinctata
Afroleptomydas A suffusipennis
Apiophora paulseni
Miltinus maculipennis
Miltinus viduatus
Miltinus stenogaster
Heteromydas bicolor
Afroleptomydas C mauricei
Opomydas townsendi
Parectyphus namibiensis
Ectyphus pinguis (Amboseli)
Ectyphus sp. crassipes
Haplomydas nov.
Eremohaplomydas desertorum
Afroleptomydas A sp. (Clanwilliam)
Eremohaplomydas sp. nov. (Kotwa)
Eremomidas arabicus(Tiega)
non-monophyletic
Eremomidas sp. nov.
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Afroleptomydas C sp. (Stampriet)
Pseudonomoneura micheneri
Pseudonomoneura
Hessemydas seyrigihirta
parkeri
Hessemydas venosus
Nemomydas brachyrhynchus
Syllegomydas sp. nov. (Ruacana)
Nemomydas gruenbergi
Leptomydas
Nemomydas corsicanus
Leptomydas melanopogon
Leptomydas sardous nov.
Leptomydas notos sp.
Syllegomydas N arnoldi
Leptomydas rapti sp.
Leptomydas turcicus nov.
Eremomidas lusitanicus
Leptomydas bek
Mydaselpislepolditertii sp. nov.
Vespiodes ngurumani
Syllegomydas N dispar
Vespiodes phaios sp. nov.
Mydaselpis sp. nov. (Daan)
Mydaselpis sp. nov. (Gamsberg)
Namadytes maculiventris
Namadytes prozeskyi
Syllegomydas S palestinensis
Cephalocera imitata
Cephalocera longirostris
Cephalocera sp. (Papendorp)
Neolaparopsis puncturatus
Arenomydas caerulescens
Arenomydas callosus
Syllegomydas N heothinos sp. nov.
Arenomydascaffra
Nomoneura namaquensis
Nomoneura paradoxa
Nomoneuroides nov. (Cape
sp. natalensis
Halterorchissp. nov. (Panne)Recife)
gen. nov. + sp. nov. (Ugab)
Syllegomydas S algiricus
gen. nov. +
gen. nov. + sp. nov. (Gobabeb)
Namibimydassp. nov. (Skerpioenkop)
Nothomydas sp. nov. (Agate)
Afroleptomydas A consanguinensis
Afroleptomydas A suffusipennis
Afroleptomydas C sp. (Clanwilliam)
Afroleptomydas A mauricei
Afroleptomydas C sp.
Syllegomydas sp. nov.(Stampriet)
Syllegomydas N dispar(Ruacana)
arnoldi Syllegomydas S elachys sp. nov.
Syllegomydas N astrictus sp. nov.
Syllegomydas N palestinensis
Syllegomydas S
Syllegomydas N heothinos
Syllegomydas S algiricus sp. nov.
Syllegomydas S astrictus sp. nov.
Syllegomydas N elachys sp. nov.

13. ... new phylogenetic hypothesis 13
Apiophorinae non-monophyly
Rhaphiomidas hasbroucki
Rhaphiomidas parkeri
Rhaphiomidas maehleri
Tongamya miranda
Neorhaphiomidas sp. (Mandurah)
Neorhaphiomidas setosus
Megascelus albovillosus
Megascelus nigricornis
Rhopalia olivieri
Rhopalia annulata
Perissocerus dumonti
Anomalomydas mackerrasi
Cacatuopyga basifascia
Cacatuopyga ruficornis
Messiasia californica
Phyllomydas parvulus
Mydas clavatus
Mydas xanthopterus
Paramydas igniticornis
Stratiomydas lividus
Apiophora quadricinctata
Apiophora paulseni
Miltinus maculipennis
Miltinus viduatus
Miltinus stenogaster
Paramydas igniticornis
Heteromydas bicolor
Opomydas townsendi
Parectyphus namibiensis
Ectyphus pinguis
Ectyphus sp. nov. (Amboseli)
...
Limata tenuicornis volaticus
Neorhynchocephalus
Nemestrinus nigrovillosus
Eulonchus sp. (SixRivers)
Prorates sp. (Escalante)
Toxophora maxima
Bombylius major
Dipalta serpentina
Apsilocephalaatra
Hemigephyra longistyla
Phycus frommeri
Hoplistomerus nobilis
Pilica formidolosa
Laphria aktis bimaculatus
Neolophonotus
Emphysomera pallidapex
Ommatius tibialis
Machimus occidentalis
Philonicus albiceps
Stichopogoncinctus
Lasiopogon punctum
Diogmites grossus
Ospriocerus aeacus
Gonioscelis ventralis
Damalis monochaetes
Pegesimalluscylindrica
Leptogaster laticornis
Euscelidia pulchra
Dioctria hyalipennis
Holcocephala calva bilineata
Apiocera (Pyrocera)
Apiocera (Ripidosyrma) braunsi
Apiocera (Pyrocera) aldrichi
Apiocera (Pyrocera) painteri
Apiocera (Pyrocera) arena
Apiocera (Pyrocera) obscura
Apiocera (Anypenus)parkeri
Apiocera (Anypenus)sp. (Wurarga)
philippii
Apiocera (Apiocera) sp. (Maranoa)
Apiocera (Apiocera)
Apiocera (Apiocera) sp. (Stuart)
Rhaphiomidas hasbroucki
Rhaphiomidas parkeri
Apiophorinae
Rhaphiomidas maehleri
Tongamya miranda (Mandurah)
Neorhaphiomidas sp.
Neorhaphiomidas setosus
Megascelus albovillosus
Megascelus nigricornis
Rhopalia olivieri
Rhopalia annulata
Perissocerus dumonti
Anomalomydas mackerrasi
Cacatuopyga basifascia
Cacatuopyga ruficornis
Messiasia californica
Phyllomydas parvulus
Mydas clavatus
Mydas xanthopterus
Paramydas igniticornis
Stratiomydas lividus
Apiophora quadricinctata
Apiophora paulseni
Miltinus maculipennis
Miltinus viduatus
Miltinus stenogaster
Heteromydas bicolor
Opomydas townsendi
Parectyphus namibiensis
Ectyphus pinguis (Amboseli)
Ectyphus sp. crassipes
Haplomydas nov.
Eremohaplomydas desertorum
Eremohaplomydas sp. nov. (Kotwa)
Eremomidas arabicus(Tiega)
Mydinae
Eremomidas sp. nov.
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Pseudonomoneura micheneri
Pseudonomoneura
Hessemydas seyrigihirta
parkeri
Hessemydas venosus
Nemomydas brachyrhynchus
Nemomydas gruenbergi
Leptomydas
Nemomydas corsicanus
Leptomydas melanopogon
Leptomydas sardous nov.
Leptomydas notos sp.
Leptomydas rapti sp.
Leptomydas turcicus nov.
Eremomidas lusitanicus
Leptomydas bek
Mydaselpislepolditertii sp. nov.
Vespiodes ngurumani
Vespiodes phaios sp. nov.
Mydaselpis sp. nov. (Daan)
Mydaselpis sp. nov. (Gamsberg)
Namadytes maculiventris
Namadytes prozeskyi
Cephalocera imitata
Cephalocera longirostris
Cephalocera sp. (Papendorp)
Neolaparopsis puncturatus
Arenomydas caerulescens
Arenomydas callosus
Arenomydascaffra
Nomoneura namaquensis
Nomoneura paradoxa
Nomoneuroides nov. (Cape
sp. natalensis
Halterorchissp. nov. (Panne)Recife)
gen. nov. + sp. nov. (Ugab)
Miltinus viduatus © Gerhard Weber
gen. nov. +
gen. nov. + sp. nov. (Gobabeb)
Namibimydassp. nov. (Skerpioenkop)
Nothomydas sp. nov. (Agate)
Afroleptomydas A consanguinensis
Afroleptomydas A suffusipennis
Afroleptomydas C sp. (Clanwilliam)
Afroleptomydas A mauricei
Afroleptomydas C sp.
Syllegomydas sp. nov.(Stampriet)
Syllegomydas N dispar(Ruacana)
arnoldi
Syllegomydas N palestinensis
Syllegomydas S
Syllegomydas N heothinos
Syllegomydas S algiricus sp. nov.
Syllegomydas S astrictus sp. nov.
Syllegomydas N elachys sp. nov.

14. ... new phylogenetic hypothesis 14
Leptomydinae non-monophyly
Haplomydas crassipes
Eremohaplomydas desertorum
Eremohaplomydas sp. nov. (Kotwa)
Eremomidas arabicus
Eremomidas sp. nov. (Tiega)
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Pseudonomoneura micheneri
Pseudonomoneura hirta
Hessemydas parkeri
Hessemydas seyrigi
Nemomydas venosus
Nemomydas brachyrhynchus
Leptomydas gruenbergi
Nemomydas melanopogon
Leptomydas corsicanus
Leptomydas sardous
Leptomydas notos sp. nov.
Leptomydas rapti sp. nov.
Leptomydas turcicus
Eremomidas bek
Leptomydas lusitanicus
Mydaselpis ngurumani sp. nov.
Vespiodes lepolditertii
Vespiodes phaios sp. nov.
Mydaselpis sp. nov. (Daan)
Mydaselpis sp. nov. (Gamsberg)
Eremomidas arabicus © Drew Gardner
Namadytes maculiventris
Namadytes prozeskyi
Cephalocera imitata
Cephalocera longirostris
Cephalocera sp. (Papendorp)
Neolaparopsis puncturatus
Arenomydas caerulescens
Arenomydas callosus
Arenomydas namaquensis
Nomoneura caffra
Nomoneura paradoxa
Nomoneuroides natalensis
Halterorchis sp. nov. (Cape Recife)
Limata tenuicornis volaticus gen. nov. + sp. nov. (Panne)
gen. nov. + sp. nov. (Ugab)
Neorhynchocephalus
Nemestrinus nigrovillosus
Eulonchus sp. (SixRivers)
Prorates sp. (Escalante)
Toxophora maxima
Bombylius major
Dipalta serpentina
gen. nov. + sp. nov. (Gobabeb)
Apsilocephalaatra
Hemigephyra longistyla
Phycus frommeri
Hoplistomerus nobilis
Pilica formidolosa
Laphria aktis bimaculatus
Neolophonotus
Emphysomera pallidapex
Namibimydas sp. nov. (Agate)
Ommatius tibialis
Machimus occidentalis
Philonicus albiceps
Stichopogoncinctus
Lasiopogon punctum
Diogmites grossus
Ospriocerus aeacus
Nothomydas sp. nov. (Skerpioenkop)
Gonioscelis ventralis
Damalis monochaetes
Pegesimalluscylindrica
Leptogaster laticornis
Euscelidia pulchra
Dioctria hyalipennis
Holcocephala calva bilineata
Apiocera (Pyrocera)
Afroleptomydas A consanguinensis
Apiocera (Ripidosyrma) braunsi
Apiocera (Pyrocera) aldrichi
Apiocera (Pyrocera) painteri
Apiocera (Pyrocera) arena
Apiocera (Pyrocera) obscura
Apiocera (Anypenus)parkeri
Apiocera (Anypenus)sp. (Wurarga)
philippii
Afroleptomydas A suffusipennis
Apiocera (Apiocera) sp. (Maranoa)
Apiocera (Apiocera)
Apiocera (Apiocera) sp. (Stuart)
Rhaphiomidas hasbroucki
Rhaphiomidas parkeri
Rhaphiomidas maehleri
Tongamya miranda (Mandurah)
Neorhaphiomidas sp.
Afroleptomydas C mauricei
Neorhaphiomidas setosus
Megascelus albovillosus
Megascelus nigricornis
Rhopalia olivieri
Rhopalia annulata
Perissocerus dumonti
Anomalomydas mackerrasi
Afroleptomydas A sp. (Clanwilliam)
Cacatuopyga basifascia
Cacatuopyga ruficornis
Messiasia californica
Phyllomydas parvulus
Mydas clavatus
Mydas xanthopterus
Paramydas igniticornis
Stratiomydas lividus
Afroleptomydas C sp. (Stampriet)
Apiophora quadricinctata
Apiophora paulseni
Miltinus maculipennis
Miltinus viduatus
Miltinus stenogaster
Heteromydas bicolor
Opomydas townsendi
Parectyphus namibiensis
Syllegomydas sp. nov. (Ruacana)
Ectyphus pinguis (Amboseli)
Ectyphus sp. crassipes
Haplomydas nov.
Eremohaplomydas desertorum
Eremohaplomydas sp. nov. (Kotwa)
Eremomidas arabicus(Tiega)
Eremomidas sp. nov.
Syllegomydas N arnoldi
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Pseudonomoneura micheneri
Pseudonomoneura
Hessemydas seyrigihirta
parkeri
Hessemydas venosus
Nemomydas brachyrhynchus
Syllegomydas N dispar
Nemomydas gruenbergi
Leptomydas
Nemomydas corsicanus
Leptomydas melanopogon
Leptomydas sardous nov.
Leptomydas notos sp.
Leptomydas rapti sp.
Leptomydas turcicus nov.
Syllegomydas S palestinensis
Eremomidas lusitanicus
Leptomydas bek
Mydaselpislepolditertii sp. nov.
Vespiodes ngurumani
Vespiodes phaios sp. nov.
Mydaselpis sp. nov. (Daan)
Mydaselpis sp. nov. (Gamsberg)
Syllegomydas N heothinos sp. nov.
Namadytes maculiventris
Namadytes prozeskyi
Cephalocera imitata
Cephalocera longirostris
Cephalocera sp. (Papendorp)
Neolaparopsis puncturatus
Arenomydas caerulescens
Syllegomydas S algiricus
Arenomydas callosus
Arenomydascaffra
Nomoneura namaquensis
Nomoneura paradoxa
Nomoneuroides nov. (Cape
sp. natalensis
Halterorchissp. nov. (Panne)Recife)
gen. nov. + sp. nov. (Ugab)
Leptomydas lusitanicus © Valter Jacinto
gen. nov. +
Syllegomydas S elachys sp. nov.
gen. nov. + sp. nov. (Gobabeb)
Namibimydassp. nov. (Skerpioenkop)
Nothomydas sp. nov. (Agate)
Afroleptomydas A consanguinensis
Afroleptomydas A suffusipennis
Afroleptomydas C sp. (Clanwilliam)
Afroleptomydas A mauricei
Afroleptomydas C sp.
Syllegomydas sp. nov.(Stampriet)
Syllegomydas N astrictus sp. nov.
Syllegomydas N dispar(Ruacana)
arnoldi
Syllegomydas N palestinensis
Syllegomydas S
Syllegomydas N heothinos
Syllegomydas S algiricus sp. nov.
Syllegomydas S astrictus sp. nov.
Syllegomydas N elachys sp. nov.

15. ... new phylogenetic hypothesis 15
Syllegomydinae non-monophyly + tribal taxa
Haplomydas crassipes
Eremohaplomydas desertorum
Eremohaplomydas sp. nov. (Kotwa)
Eremomidas arabicus
Eremomidas sp. nov. (Tiega)
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Pseudonomoneura micheneri
Pseudonomoneura hirta
Hessemydas parkeri
Hessemydas seyrigi
Nemomydas venosus
Nemomydas brachyrhynchus
Leptomydas gruenbergi
Nemomydas melanopogon
Leptomydas corsicanus
Leptomydas sardous
Leptomydas notos sp. nov.
Leptomydas rapti sp. nov.
Leptomydas turcicus
Eremomidas bek
Leptomydas lusitanicus
Mydaselpis ngurumani sp. nov.
Vespiodes lepolditertii
Vespiodes phaios sp. nov.
Mydaselpis sp. nov. (Daan)
Mydaselpis sp. nov. (Gamsberg)
Haplomydas crassipes
Mydaselpidini Namadytes maculiventris
Namadytes prozeskyi
Cephalocera imitata
Cephalocera longirostris
Cephalocera sp. (Papendorp)
Neolaparopsis puncturatus
Arenomydas caerulescens
Arenomydas callosus
Arenomydas namaquensis
Nomoneura caffra
Nomoneura paradoxa
Nomoneuroides natalensis
Halterorchis sp. nov. (Cape Recife)
Limata tenuicornis volaticus
Cephalocerini gen. nov. + sp. nov. (Panne)
gen. nov. + sp. nov. (Ugab)
Neorhynchocephalus
Nemestrinus nigrovillosus
Eulonchus sp. (SixRivers)
Prorates sp. (Escalante)
Toxophora maxima
Bombylius major
Dipalta serpentina
gen. nov. + sp. nov. (Gobabeb)
Apsilocephalaatra
Hemigephyra longistyla
Phycus frommeri
Hoplistomerus nobilis
Pilica formidolosa
Laphria aktis bimaculatus
Neolophonotus
Emphysomera pallidapex
Namibimydas sp. nov. (Agate)
Ommatius tibialis
Machimus occidentalis
Philonicus albiceps
Stichopogoncinctus
Lasiopogon punctum
Diogmites grossus
Ospriocerus aeacus
Nothomydas sp. nov. (Skerpioenkop)
Gonioscelis ventralis
Damalis monochaetes
Pegesimalluscylindrica
Leptogaster laticornis
Euscelidia pulchra
Dioctria hyalipennis
Holcocephala calva bilineata
Apiocera (Pyrocera)
Afroleptomydas A consanguinensis
Apiocera (Ripidosyrma) braunsi
Apiocera (Pyrocera) aldrichi
Apiocera (Pyrocera) painteri
Apiocera (Pyrocera) arena
Apiocera (Pyrocera) obscura
Apiocera (Anypenus)parkeri
Apiocera (Anypenus)sp. (Wurarga)
philippii
Afroleptomydas A suffusipennis
Apiocera (Apiocera) sp. (Maranoa)
Apiocera (Apiocera)
Apiocera (Apiocera) sp. (Stuart)
Rhaphiomidas hasbroucki
Rhaphiomidas parkeri
Rhaphiomidas maehleri
Tongamya miranda (Mandurah)
Neorhaphiomidas sp.
Afroleptomydas C mauricei
Neorhaphiomidas setosus
Megascelus albovillosus
Megascelus nigricornis
Rhopalia olivieri
Rhopalia annulata
Perissocerus dumonti
Anomalomydas mackerrasi
Afroleptomydas A sp. (Clanwilliam)
Cacatuopyga basifascia
Cacatuopyga ruficornis
Messiasia californica
Phyllomydas parvulus
Mydas clavatus
Mydas xanthopterus
Paramydas igniticornis
Stratiomydas lividus
Afroleptomydas C sp. (Stampriet)
Apiophora quadricinctata
Apiophora paulseni
Miltinus maculipennis
Miltinus viduatus
Miltinus stenogaster
Heteromydas bicolor
Opomydas townsendi
Parectyphus namibiensis
Halterorchini Syllegomydas sp. nov. (Ruacana)
Ectyphus pinguis (Amboseli)
Ectyphus sp. crassipes
Haplomydas nov.
Eremohaplomydas desertorum
Eremohaplomydas sp. nov. (Kotwa)
Eremomidas arabicus(Tiega)
Eremomidas sp. nov.
Syllegomydas N arnoldi
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Pseudonomoneura micheneri
Pseudonomoneura
Hessemydas seyrigihirta
parkeri
Hessemydas venosus
Nemomydas brachyrhynchus
Syllegomydas N dispar
Nemomydas gruenbergi
Leptomydas
Nemomydas corsicanus
Leptomydas melanopogon
Leptomydas sardous nov.
Leptomydas notos sp.
Leptomydas rapti sp.
Leptomydas turcicus nov.
Syllegomydas S palestinensis
Eremomidas lusitanicus
Leptomydas bek
Mydaselpislepolditertii sp. nov.
Vespiodes ngurumani
Vespiodes phaios sp. nov.
Mydaselpis sp. nov. (Daan)
Mydaselpis sp. nov. (Gamsberg)
Syllegomydas N heothinos sp. nov.
Namadytes maculiventris
Namadytes prozeskyi
Syllegomydini
Cephalocera imitata
Cephalocera longirostris
Cephalocera sp. (Papendorp)
Neolaparopsis puncturatus
Arenomydas caerulescens
Syllegomydas S algiricus
Arenomydas callosus
Arenomydascaffra
Nomoneura namaquensis
Nomoneura paradoxa
Nomoneuroides nov. (Cape
sp. natalensis
Halterorchissp. nov. (Panne)Recife)
gen. nov. + sp. nov. (Ugab)
Cephalocera imitata
gen. nov. +
Syllegomydas S elachys sp. nov.
gen. nov. + sp. nov. (Gobabeb)
Namibimydassp. nov. (Skerpioenkop)
Nothomydas sp. nov. (Agate)
Afroleptomydas A consanguinensis
Afroleptomydas A suffusipennis
Afroleptomydas C sp. (Clanwilliam)
Afroleptomydas A mauricei
Afroleptomydas C sp.
Syllegomydas sp. nov.(Stampriet)
Syllegomydas N astrictus sp. nov.
Syllegomydas N dispar(Ruacana)
arnoldi
Syllegomydas N palestinensis
Syllegomydas S
Syllegomydas N heothinos
Syllegomydas S algiricus sp. nov.
Syllegomydas S astrictus sp. nov.
Syllegomydas N elachys sp. nov.

16. .... character evolution 16
Rhaphiomidas hasbroucki
Character evolution: head + female
Rhaphiomidas parkeri
Rhaphiomidas maehleri
Tongamya miranda
Neorhaphiomidas sp. (Mandurah)
Neorhaphiomidas setosus
Megascelus albovillosus
Megascelus nigricornis
Rhopalia olivieri
Rhopalia annulata
Perissocerus dumonti
Anomalomydas mackerrasi
Cacatuopyga basifascia
Cacatuopyga ruficornis
Messiasia californica
Phyllomydas parvulus
Mydas clavatus
Mydas xanthopterus
Paramydas igniticornis
Stratiomydas lividus
Apiophora quadricinctata
Apiophora paulseni
Miltinus maculipennis
Miltinus viduatus
Miltinus stenogaster
Heteromydas bicolor
Opomydas townsendi
Parectyphus namibiensis
Ectyphus pinguis
Ectyphus sp. nov. (Amboseli)
Haplomydas crassipes
Eremohaplomydas desertorum
Eremohaplomydas sp. nov. (Kotwa)
Eremomidas arabicus
Eremomidas sp. nov. (Tiega)
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Pseudonomoneura micheneri
Pseudonomoneura hirta
Hessemydas parkeri
Hessemydas seyrigi
Nemomydas venosus
Nemomydas brachyrhynchus
gen. nov. + sp. nov. Gobabeb Leptomydas gruenbergi
Nemomydas melanopogon
Leptomydas corsicanus
Leptomydas sardous
Leptomydas notos sp. nov.
Leptomydas rapti sp. nov.
Leptomydas turcicus
Eremomidas bek
Leptomydas lusitanicus
Mydaselpis ngurumani sp. nov.
Vespiodes lepolditertii
Vespiodes phaios sp. nov.
Mydaselpis sp. nov. (Daan)
Mydaselpis sp. nov. (Gamsberg)
Namadytes maculiventris
Namadytes prozeskyi
Cephalocera imitata
Cephalocera longirostris
Cephalocera sp. (Papendorp)
Neolaparopsis puncturatus
Arenomydas caerulescens
Arenomydas callosus
Limata tenuicornis volaticus
Neorhynchocephalus
Nemestrinus nigrovillosus
Eulonchus sp. (SixRivers)
Prorates sp. (Escalante)
Arenomydas namaquensis
Nomoneura caffra
Toxophora maxima
Bombylius major
Dipalta serpentina
Apsilocephalaatra
Hemigephyra longistyla
Nomoneura paradoxa
Phycus frommeri
acanthophorite spurs absent
Hoplistomerus nobilis
Pilica formidolosa
Laphria aktis bimaculatus
Neolophonotus
Emphysomera pallidapex
Nomoneuroides natalensis
Ommatius tibialis
Machimus occidentalis
Philonicus albiceps
Stichopogoncinctus
Lasiopogon punctum
Diogmites grossus
Halterorchis sp. nov. (Cape Recife)
Ospriocerus aeacus
Gonioscelis ventralis
Damalis monochaetes
Pegesimalluscylindrica
Leptogaster laticornis
Euscelidia pulchra
Dioctria hyalipennis
Holcocephala calva bilineata
Apiocera (Pyrocera)
Apiocera (Ripidosyrma) braunsi
Apiocera (Pyrocera) aldrichi gen. nov. + sp. nov. (Panne)
gen. nov. + sp. nov. (Ugab)
Apiocera (Pyrocera) painteri
Apiocera (Pyrocera) arena
Apiocera (Pyrocera) obscura
Apiocera (Anypenus)parkeri
Apiocera (Anypenus)sp. (Wurarga)
philippii
Apiocera (Apiocera) sp. (Maranoa)
gen. nov. + sp. nov. (Gobabeb)
Apiocera (Apiocera)
Apiocera (Apiocera) sp. (Stuart)
Rhaphiomidas hasbroucki
Rhaphiomidas parkeri
proboscis vestigial
Rhaphiomidas maehleri
Tongamya miranda (Mandurah)
Neorhaphiomidas sp.
Neorhaphiomidas setosus
Megascelus albovillosus
Megascelus nigricornis
Rhopalia olivieri Namibimydas sp. nov. (Agate)
Nothomydas sp. nov. (Skerpioenkop)
Rhopalia annulata
Perissocerus dumonti
Anomalomydas mackerrasi
Cacatuopyga basifascia
Cacatuopyga ruficornis
Messiasia californica
Afroleptomydas A consanguinensis
Phyllomydas parvulus
Mydas clavatus
Mydas xanthopterus
Paramydas igniticornis
Stratiomydas lividus
Apiophora quadricinctata
Afroleptomydas A suffusipennis
Apiophora paulseni
Miltinus maculipennis
Miltinus viduatus
Miltinus stenogaster
Heteromydas bicolor
Afroleptomydas C mauricei
Opomydas townsendi
Parectyphus namibiensis
Ectyphus pinguis (Amboseli)
Ectyphus sp. crassipes
Haplomydas nov.
Eremohaplomydas desertorum
Afroleptomydas A sp. (Clanwilliam)
Eremohaplomydas sp. nov. (Kotwa)
Eremomidas arabicus(Tiega)
proboscis and acanthophorite spurs
Eremomidas sp. nov.
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Afroleptomydas C sp. (Stampriet)
Pseudonomoneura micheneri
Pseudonomoneura
Hessemydas seyrigihirta
parkeri
Hessemydas venosus
Nemomydas brachyrhynchus
Syllegomydas sp. nov. (Ruacana)
Nemomydas gruenbergi
Leptomydas
Nemomydas corsicanus
Leptomydas melanopogon
Leptomydas sardous nov.
Leptomydas notos sp.
Syllegomydas N arnoldi
Leptomydas rapti sp.
Leptomydas turcicus nov.
Eremomidas lusitanicus
Leptomydas bek
Mydaselpislepolditertii sp. nov.
Vespiodes ngurumani
Syllegomydas N dispar
Vespiodes phaios sp. nov.
Mydaselpis sp. nov. (Daan)
Mydaselpis sp. nov. (Gamsberg)
Namadytes maculiventris
Namadytes prozeskyi
Syllegomydas S palestinensis
Cephalocera imitata
Cephalocera longirostris
Cephalocera sp. (Papendorp)
Neolaparopsis puncturatus
Arenomydas caerulescens
Arenomydas callosus
well-developed
Syllegomydas N heothinos sp. nov.
Arenomydascaffra
Nomoneura namaquensis
Nomoneura paradoxa
Nomoneuroides nov. (Cape
sp. natalensis
Halterorchissp. nov. (Panne)Recife)
gen. nov. + sp. nov. (Ugab)
Syllegomydas S algiricus
gen. nov. +
gen. nov. + sp. nov. (Gobabeb)
Namibimydassp. nov. (Skerpioenkop)
Nothomydas sp. nov. (Agate)
Afroleptomydas A consanguinensis
Afroleptomydas A suffusipennis
Afroleptomydas C sp. (Clanwilliam)
Afroleptomydas A mauricei
Afroleptomydas C sp.
Syllegomydas sp. nov.(Stampriet)
Syllegomydas N dispar(Ruacana)
arnoldi Syllegomydas S elachys sp. nov.
Syllegomydas N astrictus sp. nov.
Syllegomydas N palestinensis
Syllegomydas S
Syllegomydas N heothinos
Syllegomydas S algiricus sp. nov.
Syllegomydas S astrictus sp. nov.
Syllegomydas N elachys sp. nov.

17. .... character evolution 17
Rhaphiomidas hasbroucki
Character evolution: met tibia
Rhaphiomidas parkeri
Rhaphiomidas maehleri
Tongamya miranda
Neorhaphiomidas sp. (Mandurah)
Neorhaphiomidas setosus
Megascelus albovillosus
Megascelus nigricornis
Rhopalia olivieri
Rhopalia annulata
Perissocerus dumonti
Anomalomydas mackerrasi
Cacatuopyga basifascia
Cacatuopyga ruficornis
Messiasia californica
Phyllomydas parvulus
Mydas clavatus
Mydas xanthopterus
Paramydas igniticornis
Stratiomydas lividus
Apiophora quadricinctata
Apiophora paulseni
Miltinus maculipennis
Miltinus viduatus
Miltinus stenogaster
Heteromydas bicolor
Opomydas townsendi
Parectyphus namibiensis
Ectyphus pinguis
Ectyphus sp. nov. (Amboseli)
Haplomydas crassipes
Eremohaplomydas desertorum
Eremohaplomydas sp. nov. (Kotwa)
Eremomidas arabicus
Eremomidas sp. nov. (Tiega)
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Pseudonomoneura micheneri
Pseudonomoneura hirta
Hessemydas parkeri
Hessemydas seyrigi
Nemomydas venosus
Nemomydas brachyrhynchus
Diochlistus cardinalis Leptomydas gruenbergi
Nemomydas melanopogon
Leptomydas corsicanus
Leptomydas sardous
Leptomydas notos sp. nov.
Leptomydas rapti sp. nov.
Leptomydas turcicus
Eremomidas bek
Leptomydas lusitanicus
Mydaselpis ngurumani sp. nov.
Vespiodes lepolditertii
Vespiodes phaios sp. nov.
Mydaselpis sp. nov. (Daan)
Mydaselpis sp. nov. (Gamsberg)
Namadytes maculiventris
Namadytes prozeskyi
Cephalocera imitata
Cephalocera longirostris
Cephalocera sp. (Papendorp)
Neolaparopsis puncturatus
Arenomydas caerulescens
Arenomydas callosus
Limata tenuicornis volaticus
Neorhynchocephalus
Nemestrinus nigrovillosus
Eulonchus sp. (SixRivers)
Prorates sp. (Escalante)
Arenomydas namaquensis
Nomoneura caffra
Toxophora maxima
Bombylius major
Dipalta serpentina
Apsilocephalaatra
Hemigephyra longistyla
Nomoneura paradoxa
Phycus frommeri
ventral keel absent
Hoplistomerus nobilis
Pilica formidolosa
Laphria aktis bimaculatus
Neolophonotus
Emphysomera pallidapex
Nomoneuroides natalensis
Ommatius tibialis
Machimus occidentalis
Philonicus albiceps
Stichopogoncinctus
Lasiopogon punctum
Diogmites grossus
Halterorchis sp. nov. (Cape Recife)
Ospriocerus aeacus
Gonioscelis ventralis
Damalis monochaetes
Pegesimalluscylindrica
Leptogaster laticornis
Euscelidia pulchra
Dioctria hyalipennis
Holcocephala calva bilineata
Apiocera (Pyrocera)
Apiocera (Ripidosyrma) braunsi
Apiocera (Pyrocera) aldrichi gen. nov. + sp. nov. (Panne)
gen. nov. + sp. nov. (Ugab)
Apiocera (Pyrocera) painteri
Apiocera (Pyrocera) arena
Apiocera (Pyrocera) obscura
Apiocera (Anypenus)parkeri
Apiocera (Anypenus)sp. (Wurarga)
philippii
Apiocera (Apiocera) sp. (Maranoa)
gen. nov. + sp. nov. (Gobabeb)
Apiocera (Apiocera)
Apiocera (Apiocera) sp. (Stuart)
Rhaphiomidas hasbroucki
Rhaphiomidas parkeri
keel at least proximally
Rhaphiomidas maehleri
Tongamya miranda (Mandurah)
Neorhaphiomidas sp.
Neorhaphiomidas setosus
Megascelus albovillosus
Megascelus nigricornis
Rhopalia olivieri Namibimydas sp. nov. (Agate)
Nothomydas sp. nov. (Skerpioenkop)
Rhopalia annulata
Perissocerus dumonti
Anomalomydas mackerrasi
Cacatuopyga basifascia
Cacatuopyga ruficornis
Messiasia californica
Afroleptomydas A consanguinensis
Phyllomydas parvulus
Mydas clavatus
Mydas xanthopterus
Paramydas igniticornis
Stratiomydas lividus
Apiophora quadricinctata
Afroleptomydas A suffusipennis
Apiophora paulseni
Miltinus maculipennis
Miltinus viduatus
Miltinus stenogaster
Heteromydas bicolor
Afroleptomydas C mauricei
Opomydas townsendi
Parectyphus namibiensis
Ectyphus pinguis (Amboseli)
Ectyphus sp. crassipes
Haplomydas nov.
Eremohaplomydas desertorum
Afroleptomydas A sp. (Clanwilliam)
Eremohaplomydas sp. nov. (Kotwa)
Eremomidas arabicus(Tiega)
keel well-developed with spur
Eremomidas sp. nov.
Diochlistus gracilis
Mitrodetus dentitarsis
Mitrodetus dimidiatus
Afroleptomydas C sp. (Stampriet)
Pseudonomoneura micheneri
Pseudonomoneura
Hessemydas seyrigihirta
parkeri
Hessemydas venosus
Nemomydas brachyrhynchus
Syllegomydas sp. nov. (Ruacana)
Nemomydas gruenbergi
Leptomydas
Nemomydas corsicanus
Leptomydas melanopogon
Leptomydas sardous nov.
Leptomydas notos sp.
Syllegomydas N arnoldi
Leptomydas rapti sp.
Leptomydas turcicus nov.
Eremomidas lusitanicus
Leptomydas bek
Mydaselpislepolditertii sp. nov.
Vespiodes ngurumani
Syllegomydas N dispar
Vespiodes phaios sp. nov.
Mydaselpis sp. nov. (Daan)
Mydaselpis sp. nov. (Gamsberg)
Namadytes maculiventris
Namadytes prozeskyi
Syllegomydas S palestinensis
Cephalocera imitata
Cephalocera longirostris
Cephalocera sp. (Papendorp)
Neolaparopsis puncturatus
Arenomydas caerulescens
Arenomydas callosus
Syllegomydas N heothinos sp. nov.
Arenomydascaffra
Nomoneura namaquensis
Nomoneura paradoxa
Nomoneuroides nov. (Cape
sp. natalensis
Halterorchissp. nov. (Panne)Recife)
gen. nov. + sp. nov. (Ugab)
Syllegomydas S algiricus
gen. nov. +
gen. nov. + sp. nov. (Gobabeb)
Namibimydassp. nov. (Skerpioenkop)
Nothomydas sp. nov. (Agate)
Afroleptomydas A consanguinensis
Afroleptomydas A suffusipennis
Afroleptomydas C sp. (Clanwilliam)
Afroleptomydas A mauricei
Afroleptomydas C sp.
Syllegomydas sp. nov.(Stampriet)
Syllegomydas N dispar(Ruacana)
arnoldi Syllegomydas S elachys sp. nov.
Syllegomydas N astrictus sp. nov.
Syllegomydas N palestinensis
Syllegomydas S
Syllegomydas N heothinos
Syllegomydas S algiricus sp. nov.
Syllegomydas S astrictus sp. nov.
Syllegomydas N elachys sp. nov.

18. .... summary + future 18
Summary
◊ Mydidae monophyletic
◊ (Mydidae + Apioceridae) + Asilidae
◊ Afrotropical Eremomidas in Leptomydinae?
◊ Eremohaplomydas + Haplomydas in Syllegomydinae?
◊ Chilean Paramydas to Mydinae – Apiophorinae monophyly
◊ Future Research:
› additional exemplar species
› internal head characters
› inclusion of †Cretomydas, †Mydas miocenicus, †Protapiocera
Phyllomydas parvulus © Lyn Atherton

19. 19
Acknowledgements
◊ Museum curators and collectors who made specimens available
◊ U.S. National Science Foundation (REVSYS DEB-0919333)
◊ I would be happy to study additional specimens from your collection!
www.mydidae.tdvia.de