TDikow Mydidae morphological phylogeny ICD 2010

1

Phylogenetic relationships within Mydidae based on morphology of imagines (Diptera: Asiloidea)

Torsten DikowBiodiversity Synthesis Center Field Museum of Natural History

2. introduction

Opomydas townsendi

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)

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

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* × × Leptomydinae 6 54* × × × × ×Megascelinae 3 15* × × × Mydinae 12 84 × × Rhaphiomidinae 1 23* × Rhopaliinae 4 21 × × ×Syllegomydinae 25 206* × × ×unplaced 1 2

. introduction

* includes species currently being describedCacatuopyga basifascia

Anomalomydas mackerrasi

Taxa overview

4

Overview of presentationPrevious Phylogenetic Hypotheses ◊

New Phylogenetic Hypothesis ◊

Character Evolution ◊

Summary + Future◊

Stratiomydas lividus © Charles Melton

Miltinus stenogaster © Shaun Winterton

5

Ectyp

hinae

Cacat

uopyg

inae

Anom

alom

ydina

e

Diochli

stina

e

Rhopali

inae

Apiopho

rinae

Myd

inae

Lepto

myd

inae

Sylleg

omyd

inae

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

.. previous phylogenetic hypotheses

Mitrodetus dentitarsis © Steve Marshall

Diochlistinae remaining Mydidae

› morphology › similarity

6

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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Bombylius majorApsilocephala longistylusProrates sp (Esacalante)Phycus frommeriHemigephyra atraApiocera (Apiocera) sp (Barakula)Apiocera (Pyrocera) haruspexApiocera (Pyrocera) painteriStichopogon punctumSaropogon luteusLaxenecera albicinctaHoplistomerus nobilisGonioscelis ventralisLeptogaster cylindricaAsilus crabroniformisPromachus amastrusEmphysomera pallidapexRhaphiomidas xanthosMitrodetus dentitarsisOpomydas townsendiMiltinus cardinalisMydas clavatusMahafalymydas wiegmanniSyllegomydas sp (Nizzanem)Afroleptomydas sp (Clanwilliam)Afroleptomydas sp (RSA)Cephalocera imitataCephalocera sp (Montegu)Nemomydas brachyrhynchusNemomydas sp (Tesopaco)

RhaphiomidinaeDiochlistinaeEctyphinaeApiophorinaeMydinae

Syllegomydinae

Leptomydinae

Asilidae

ApioceridaeApiocera 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 Dikow unpublished presented at ICD6 in Fukuoka

› morphology › parsimony


› 18S + 28S rDNA, CAD › parsimony (POY)

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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 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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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)

...

Miltinus viduatus © Gerhard Weber



Apiophorinae non-monophyly

Paramydas igniticornis

ApiophorinaeMydinae

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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. Leptomydas lusitanicus © Valter Jacinto

Eremomidas arabicus © Drew Gardner



Leptomydinae non-monophyly

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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.

Mydaselpidini

Cephalocerini

Halterorchini

Syllegomydini



Syllegomydinae non-monophyly + tribal taxa

Haplomydas crassipes

Cephalocera imitata

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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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keel at least proximallykeel well-developed with spur

ventral keel absent

Character evolution: met tibia

Diochlistus cardinalis

.... character evolution

18.... summary + future

Summary

Phyllomydas parvulus © Lyn Atherton

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

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AcknowledgementsMuseum 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

TDikow Mydidae morphological phylogeny ICD 2010

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