tdikow mydidae morphological phylogeny esa 2010

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Phylogeny of Mydidae inferred from morphological characters of imagines (Diptera: Asiloidea)

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

2. introduction

Opomydas townsendi

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

Larvae possibly predatory◊

471 species and 66 genera world-wide◊

11 subfamily taxa◊

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

Cretomydas santanensis

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Overview of presentationPrevious Phylogenetic Hypotheses ◊

New Phylogenetic Hypothesis ◊

Character Evolution ◊

Placement of oldest Fossil ◊

Summary + Future◊

Stratiomydas lividus © Charles Melton

Miltinus stenogaster © Shaun Winterton

4

Ectyphinae

Cacatuopyginae

Anom

alomydinae

Diochlistinae

Rhopaliinae

Apiophorinae

Mydinae

Leptom

ydinae

Syllegomydinae

Wilcox & Papavero 1974 Artigas & Papavero 1990› morphology › similarity

.. previous phylogenetic hypotheses

Mitrodetus dentitarsis © Steve Marshall

Diochlistinae remaining Mydidae

› morphology › similarity

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Asilidae

Apiocera (Pyrocera)

Apiocera (Ripidosyrma)

Apiocera (Anypenus)

Apiocera (Apiocera)

Rhaphiomidinae

Megascelinae

Rhopaliinae

Cacatuopyginae

Leptomydinae

Syllegomydinae

Anomalomydinae

Diochlistinae

Ectyphinae

Apiophorinae

Mydinae

Scenopinidae

Therevidae

Apiocera (Anypenus) phillippi

Apiocera (Apiocera) sp 1

Apiocera (Apiocera) sp 2

Apiocera (Pyrocera) haruspex

Apiocera (Pyrocera) sp

Rhaphiomidas terminalis

Megascelus albovillosus

Tongamya stuckenbergi

Diochlistus hackeri

Rhopalia sp

Mitrodetus australis

Nemomydas pantherinus

Arenomydas callosus

Rhaphiomidinae

Megascelinae

Diochlistinae

Rhopaliinae

Diochlistinae

Leptomydinae

Syllegomydinae

Yeates & Irwin 1996 Irwin & Wiegmann 2001› morphology › parsimony (successive weighting)


› 28S rDNA › maximum likelihood

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Afroleptomydas suffusipennis

Apiocera aldrichi

Apiocera painteri

Rhaphiomidas maehleri

Neorhaphiomidas sp (Mandurah)

Opomydas townsendi

Mitrodetus dentitarsis

Mydas clavatus

Pseudonomoneura hirta

Nemomydas brachyrhynchus

Afroleptomydas sp (Clanwilliam)

Asilidae

Apioceridae

Rhaphiomidinae

Diochlistinae

Ectyphinae

Mydinae

Syllegomydinae

Leptomydinae

Megascelinae

...

Dikow 2009› morphology › parsimony


Leptomydas lusitanicus © Valter Jacinto

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Taxon Sampling94 Mydidae species◊ (= 19,9 %)

Higher-level taxa ◊ › 40 genera (= 61,5 %)

› 7 tribal taxa (= 87,5 %) › 11 subfamily taxa (= 100 %)

Outgroup taxa ◊ › 37 Asiloidea species Apioceridae (11), Apsilocephalidae (1), Asilidae (19), Bombyliidae (3), Scenopinidae (1), Therevidae (2) › 1 Acroceridae, 2 Nemestrinidae › 1 Tabanidae

collecting localities of specimens included

... new phylogenetic hypothesis

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Morphological Characters186 discrete, parsimony informative characters◊

tagma # characters remarksHead 42Thorax 83

39 chaetotaxy, sclerites, etc.21 wing23 leg

Abdomen 6116 sclerite shapes, chaetotaxy, etc.25 ? terminalia20 / genitalia


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Phylogenetic AnalysisMorphological data ◊

› MacClade (version 4.08) › WinClada (version 1.0.08)

› all multi-state characters non-additive

Tree Search ◊ › TNT (version 1.1 June 2010) › Ratchet, TreeDrifting, TreeFusing, Sectorial Searches (Goloboff 1999, Nixon 1999)

› regular parsimony › implied weighting (k = 3) (Goloboff 1993)


Heteromydas bicolor © Greg Ballmer

Eremomidas arabicus © Drew Gardner

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Limata tenuicornisNeorhynchocephalus volaticusNemestrinus nigrovillosusEulonchus sp. (SixRivers)Prorates sp. (Escalante)Toxophora maximaBombylius majorDipalta serpentinaApsilocephala longistylaHemigephyra atraPhycus frommeriHoplistomerus nobilisPilica formidolosaLaphria aktisNeolophonotus bimaculatusEmphysomera pallidapexOmmatius tibialisMachimus occidentalisPhilonicus albicepsStichopogon punctumLasiopogon cinctusDiogmites grossusOspriocerus aeacusGonioscelis ventralisDamalis monochaetesPegesimallus laticornisLeptogaster cylindricaEuscelidia pulchraDioctria hyalipennisHolcocephala calvaApiocera (Pyrocera) bilineataApiocera (Ripidosyrma) braunsiApiocera (Pyrocera) aldrichiApiocera (Pyrocera) painteriApiocera (Pyrocera) arenaApiocera (Pyrocera) parkeriApiocera (Anypenus) obscuraApiocera (Anypenus) philippiiApiocera (Apiocera) sp. (Wurarga)Apiocera (Apiocera) sp. (Maranoa)Apiocera (Apiocera) sp. (Stuart)Rhaphiomidas hasbrouckiRhaphiomidas parkeriRhaphiomidas maehleriTongamya mirandaNeorhaphiomidas sp. (Mandurah)Neorhaphiomidas setosusMegascelus albovillosusMegascelus nigricornisRhopalia olivieriRhopalia annulataPerissocerus dumontiAnomalomydas mackerrasiCacatuopyga basifasciaCacatuopyga ruficornisMessiasia californicaPhyllomydas parvulusMydas clavatusMydas xanthopterusParamydas igniticornisStratiomydas lividusApiophora quadricinctataApiophora paulseniMiltinus maculipennisMiltinus viduatusMiltinus stenogasterHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus pinguisEctyphus sp. nov. (Amboseli)Haplomydas crassipesEremohaplomydas desertorumEremohaplomydas sp. nov. (Kotwa)Eremomidas arabicusEremomidas sp. nov. (Tiega)Diochlistus gracilisMitrodetus dentitarsisMitrodetus dimidiatusPseudonomoneura micheneriPseudonomoneura hirtaHessemydas parkeriHessemydas seyrigiNemomydas venosusNemomydas brachyrhynchusLeptomydas gruenbergiNemomydas melanopogonLeptomydas corsicanusLeptomydas sardousLeptomydas notos sp. nov.Leptomydas rapti sp. nov.Leptomydas turcicusEremomidas bekLeptomydas lusitanicusMydaselpis ngurumani sp. nov.Vespiodes lepolditertiiVespiodes phaios sp. nov.Mydaselpis sp. nov. (Daan)Mydaselpis sp. nov. (Gamsberg)Namadytes maculiventrisNamadytes prozeskyiCephalocera imitataCephalocera longirostrisCephalocera sp. (Papendorp)Neolaparopsis puncturatusArenomydas caerulescensArenomydas callosusArenomydas namaquensisNomoneura caffraNomoneura paradoxaNomoneuroides natalensisHalterorchis sp. nov. (Cape Recife)gen. nov. + sp. nov. (Panne)gen. nov. + sp. nov. (Ugab)gen. nov. + sp. nov. (Gobabeb)Namibimydas sp. nov. (Agate)Nothomydas sp. nov. (Skerpioenkop)Afroleptomydas A consanguinensisAfroleptomydas A suffusipennisAfroleptomydas C mauriceiAfroleptomydas A sp. (Clanwilliam)Afroleptomydas C sp. (Stampriet)Syllegomydas sp. nov. (Ruacana)Syllegomydas N arnoldiSyllegomydas N disparSyllegomydas S palestinensisSyllegomydas N heothinos sp. nov.Syllegomydas S algiricusSyllegomydas S elachys sp. nov.Syllegomydas N astrictus sp. nov.

Apioceridae

Asilidae

Mydidae

TabanidaeNemestrinidaeAcroceridaeScenopinidaeBombyliidaeApsilocephalidaeTherevidae



Results & Sister-groupParsimony search ◊

› 421 trees, 1485 steps › strict consensus with large polytomy in derived ingroup

Implied weights ◊ › 6 trees, fit 95.58644 › = 1556 steps

› strict consensus well-resolved

strict consensus tree from implied weights analysis

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Rhaphiomidas hasbrouckiRhaphiomidas parkeriRhaphiomidas maehleriTongamya mirandaNeorhaphiomidas sp. (Mandurah)Neorhaphiomidas setosusMegascelus albovillosusMegascelus nigricornisRhopalia olivieriRhopalia annulataPerissocerus dumontiAnomalomydas mackerrasiCacatuopyga basifasciaCacatuopyga ruficornisMessiasia californicaPhyllomydas parvulusMydas clavatusMydas xanthopterusParamydas igniticornisStratiomydas lividusApiophora quadricinctataApiophora paulseniMiltinus maculipennisMiltinus viduatusMiltinus stenogasterHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus pinguisEctyphus sp. nov. (Amboseli)Haplomydas crassipesEremohaplomydas desertorumEremohaplomydas sp. nov. (Kotwa)Eremomidas arabicusEremomidas sp. nov. (Tiega)Diochlistus gracilisMitrodetus dentitarsisMitrodetus dimidiatusPseudonomoneura micheneriPseudonomoneura hirtaHessemydas parkeriHessemydas seyrigiNemomydas venosusNemomydas brachyrhynchusLeptomydas gruenbergiNemomydas melanopogonLeptomydas corsicanusLeptomydas sardousLeptomydas notos sp. nov.Leptomydas rapti sp. nov.Leptomydas turcicusEremomidas bekLeptomydas lusitanicusMydaselpis ngurumani sp. nov.Vespiodes lepolditertiiVespiodes phaios sp. nov.Mydaselpis sp. nov. (Daan)Mydaselpis sp. nov. (Gamsberg)Namadytes maculiventrisNamadytes prozeskyiCephalocera imitataCephalocera longirostrisCephalocera sp. (Papendorp)Neolaparopsis puncturatusArenomydas caerulescensArenomydas callosusArenomydas namaquensisNomoneura caffraNomoneura paradoxaNomoneuroides natalensisHalterorchis sp. nov. (Cape Recife)gen. nov. + sp. nov. (Panne)gen. nov. + sp. nov. (Ugab)gen. nov. + sp. nov. (Gobabeb)Namibimydas sp. nov. (Agate)Nothomydas sp. nov. (Skerpioenkop)Afroleptomydas A consanguinensisAfroleptomydas A suffusipennisAfroleptomydas C mauriceiAfroleptomydas A sp. (Clanwilliam)Afroleptomydas C sp. (Stampriet)Syllegomydas sp. nov. (Ruacana)Syllegomydas N arnoldiSyllegomydas N disparSyllegomydas S palestinensisSyllegomydas N heothinos sp. nov.Syllegomydas S algiricusSyllegomydas S elachys sp. nov.Syllegomydas N astrictus sp. nov.

Rhaphiomidinae

Megascelinae

RhopaliinaeAnomalomydinaeCacatuopyginae

Mydinae

Apiophorinae

Ectyphinae

SyllegomydinaeLeptomydinaeDiochlistinae

Leptomydinae

Syllegomydinae



monophyleticnon-monophyletic

Rhaphiomidas auratus © Greg Ballmer

Monophyly & Rhaphiomidinae

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Haplomydas crassipesEremohaplomydas desertorumEremohaplomydas sp. nov. (Kotwa)Eremomidas arabicusEremomidas sp. nov. (Tiega)Diochlistus gracilisMitrodetus dentitarsisMitrodetus dimidiatusPseudonomoneura micheneriPseudonomoneura hirtaHessemydas parkeriHessemydas seyrigiNemomydas venosusNemomydas brachyrhynchusLeptomydas gruenbergiNemomydas melanopogonLeptomydas corsicanusLeptomydas sardousLeptomydas notos sp. nov.Leptomydas rapti sp. nov.Leptomydas turcicusEremomidas bekLeptomydas lusitanicusMydaselpis ngurumani sp. nov.Vespiodes lepolditertiiVespiodes phaios sp. nov.Mydaselpis sp. nov. (Daan)Mydaselpis sp. nov. (Gamsberg)Namadytes maculiventrisNamadytes prozeskyiCephalocera imitataCephalocera longirostrisCephalocera sp. (Papendorp)Neolaparopsis puncturatusArenomydas caerulescensArenomydas callosusArenomydas namaquensisNomoneura caffraNomoneura paradoxaNomoneuroides natalensisHalterorchis sp. nov. (Cape Recife)gen. nov. + sp. nov. (Panne)gen. nov. + sp. nov. (Ugab)gen. nov. + sp. nov. (Gobabeb)Namibimydas sp. nov. (Agate)Nothomydas sp. nov. (Skerpioenkop)Afroleptomydas A consanguinensisAfroleptomydas A suffusipennisAfroleptomydas C mauriceiAfroleptomydas A sp. (Clanwilliam)Afroleptomydas C sp. (Stampriet)Syllegomydas sp. nov. (Ruacana)Syllegomydas N arnoldiSyllegomydas N disparSyllegomydas S palestinensisSyllegomydas N heothinos sp. nov.Syllegomydas S algiricusSyllegomydas S elachys sp. nov.Syllegomydas N astrictus sp. nov.



Syllegomydinae non-monophyly

Haplomydas crassipes

Cephalocera imitata

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Taxon (sensu Papavero & Wilcox 1975) MonophylyAnomalomydinae monogeneric

Apiophorinae —Cacatuopyginae √Diochlistinae √Ectyphinae √Leptomydinae —Megascelinae √Mydinae √Rhaphiomidinae √Rhopaliinae √Syllegomydinae —

Revised, monophyletic taxon

Paramydas

Eremomidas sp.

+ Paramydas (Chile)

Haplomydas, Eremohaplomydas

Monophyly of higher-level taxa & Revised classification


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Rhaphiomidas hasbrouckiRhaphiomidas parkeriRhaphiomidas maehleriTongamya mirandaNeorhaphiomidas sp. (Mandurah)Neorhaphiomidas setosusMegascelus albovillosusMegascelus nigricornisRhopalia olivieriRhopalia annulataPerissocerus dumontiAnomalomydas mackerrasiCacatuopyga basifasciaCacatuopyga ruficornisMessiasia californicaPhyllomydas parvulusMydas clavatusMydas xanthopterusParamydas igniticornisStratiomydas lividusApiophora quadricinctataApiophora paulseniMiltinus maculipennisMiltinus viduatusMiltinus stenogasterHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus pinguisEctyphus sp. nov. (Amboseli)Haplomydas crassipesEremohaplomydas desertorumEremohaplomydas sp. nov. (Kotwa)Eremomidas arabicusEremomidas sp. nov. (Tiega)Diochlistus gracilisMitrodetus dentitarsisMitrodetus dimidiatusPseudonomoneura micheneriPseudonomoneura hirtaHessemydas parkeriHessemydas seyrigiNemomydas venosusNemomydas brachyrhynchusLeptomydas gruenbergiNemomydas melanopogonLeptomydas corsicanusLeptomydas sardousLeptomydas notos sp. nov.Leptomydas rapti sp. nov.Leptomydas turcicusEremomidas bekLeptomydas lusitanicusMydaselpis ngurumani sp. nov.Vespiodes lepolditertiiVespiodes phaios sp. nov.Mydaselpis sp. nov. (Daan)Mydaselpis sp. nov. (Gamsberg)Namadytes maculiventrisNamadytes prozeskyiCephalocera imitataCephalocera longirostrisCephalocera sp. (Papendorp)Neolaparopsis puncturatusArenomydas caerulescensArenomydas callosusArenomydas namaquensisNomoneura caffraNomoneura paradoxaNomoneuroides natalensisHalterorchis sp. nov. (Cape Recife)gen. nov. + sp. nov. (Panne)gen. nov. + sp. nov. (Ugab)gen. nov. + sp. nov. (Gobabeb)Namibimydas sp. nov. (Agate)Nothomydas sp. nov. (Skerpioenkop)Afroleptomydas A consanguinensisAfroleptomydas A suffusipennisAfroleptomydas C mauriceiAfroleptomydas A sp. (Clanwilliam)Afroleptomydas C sp. (Stampriet)Syllegomydas sp. nov. (Ruacana)Syllegomydas N arnoldiSyllegomydas N disparSyllegomydas S palestinensisSyllegomydas N heothinos sp. nov.Syllegomydas S algiricusSyllegomydas S elachys sp. nov.Syllegomydas N astrictus sp. nov.


proboscis vestigialproboscis and acanthophorite spurs well-developed

acanthophorite spurs absent

Character evolution: head + female

gen. nov. + sp. nov. Gobabeb

.... character evolution

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Limata tenuicornisNeorhynchocephalus volaticusNemestrinus nigrovillosusEulonchus sp.Prorates sp. (Escalante)Toxophora maximaBombylius majorDipalta serpentinaApsilocephala longistylaHemigephyra atraPhycus frommeriHoplistomerus nobilisPilica formidolosaLaphria aktisNeolophonotus bimaculatusMachimus occidentalisPhilonicus albicepsEmphysomera pallidapexOmmatius tibialisStichopogon punctumLasiopogon cinctusDiogmites grossusOspriocerus aeacusGonioscelis ventralisDamalis monochaetesPegesimallus laticornisDioctria hyalipennisHolcocephala calvaLeptogaster cylindricaEuscelidia pulchraApiocera P bilineataApiocera R braunsiApiocera P aldrichiApiocera P painteriApiocera P arenaApiocera P parkeriApiocera An obscuraApiocera An philippiiApiocera Ap sp. WurargaApiocera Ap sp. MaranoaApiocera Ap sp. StuartRhaphiomidas hasbrouckiRhaphiomidas parkeriRhaphiomidas maehleriTongamya mirandaMegascelus albovillosusMegascelus nigricornisNeorhaphiomidas sp. (Mandurah)Neorhaphiomidas setosusRhopalia olivieriRhopalia annulataPerissocerus dumontiAnomalomydas mackerrasiCacatuopyga basifasciaCacatuopyga ruficornisMessiasia californicaPhyllomydas parvulusMydas clavatusMydas xanthopterusStratiomydas lividusParamydas igniticornisCretomydas santanensisApiophora quadricinctataApiophora paulseniMiltinus maculipennisMiltinus viduatusMiltinus stenogasterHeteromydas bicolorOpomydas townsendiParectyphus namibiensisEctyphus pinguisEctyphus sp. nov. (Amboseli)Haplomydas crassipesEremohaplomydas desertorumEremohaplomydas sp. nov. (Kotwa)Eremomidas arabicusEremomidas sp. nov. (Tiega)Diochlistus gracilisMitrodetus dentitarsisMitrodetus dimidiatusHessemydas parkeriHessemydas seyrigiPseudonomoneura micheneriPseudonomoneura hirtaNemomydas venosusNemomydas brachyrhynchusLeptomydas gruenbergiNemomydas melanopogonLeptomydas corsicanusEremomidas bekLeptomydas lusitanicusLeptomydas raptiLeptomydas turcicusLeptomydas sardousLeptomydas notosMydaselpis ngurumaniMydaselpis sp. nov. (Daan)Mydaselpis sp. nov. (Gamsberg)Vespiodes lepolditertiiVespiodes phaiosNamadytes maculiventrisNamadytes prozeskyiCephalocera imitataCephalocera longirostrisCephalocera sp. (Papendorp)Neolaparopsis puncturatusArenomydas caerulescensArenomydas callosusArenomydas namaquensisNomoneura caffraNomoneura paradoxaNomoneuroides natalensisHalterorchis sp. nov. (Cape Recife)Stenomydas gen. nov. sp. nov. (Panne)Namibimydas sp. nov. (Agate)Nothomydas sp. nov. (Skerpioenkop)Cleamydas gen. nov. sp. nov. (Ugab)Syll gen. nov. sp. nov. (Gobabeb)Afroleptomydas A consanguinensisAfroleptomydas A suffusipennisAfroleptomydas C mauriceiAfroleptomydas A sp. (Clanwilliam)Afroleptomydas C sp. (Stampriet)Syllegomydas sp. nov. (Ruacana)Syllegomydas N arnoldiSyllegomydas N disparSyllegomydas S palestinensisSyllegomydas N heothinosSyllegomydas S algiricusSyllegomydas S elachysSyllegomydas N astrictus

Apioceridae

Asilidae

Mydidae

TabanidaeNemestrinidaeAcroceridaeScenopinidaeBombyliidaeApsilocephalidaeTherevidae

† Cretomydas santanensis, Crato Formation

≈ 112 myo

Placement of oldest fossil Mydidae› † Cretomydas santanensis › coded for 23.1 % of characters

4 trees, fit 95.86467= 1563 stepsCI=0.23RI=0.75strict consensus shown (1564 steps)


..... oldest fossil

16.... summary + future

Summary

Phyllomydas parvulus © Lyn Atherton

Mydidae monophyletic◊

(Mydidae + Apioceridae) + Asilidae◊

Afrotropical ◊ Eremomidas in Leptomydinae or Diochlistinae?

Eremohaplomydas ◊ + Haplomydas in Syllegomydinae?

Chilean ◊ Paramydas to Mydinae – Apiophorinae monophyly

Future Research: ◊ › additional exemplar species (8 species) › internal head characters › regular parsimony

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AcknowledgementsMuseum curators and collectors who made specimens available◊

Photographers◊

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

www.mydidae.tdvia.de

tdikow mydidae morphological phylogeny esa 2010

Documents

maranoaapiocera apiocera

wurargaapiocera apiocera

subfamily taxa

outgroup taxa

higherlevel taxa

tribal taxa

characters remarkshead

winclada version