TDikow Mydidae morphological phylogeny ESA 2010

1

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

Torsten Dikow
Biodiversity Synthesis Center
Field Museum of Natural History, Chicago, IL

. introduction 2

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)
Opomydas townsendi
◊ 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

3

Overview of presentation
◊ Previous Phylogenetic Hypotheses

◊ New Phylogenetic Hypothesis

◊ Character Evolution
Miltinus stenogaster © Shaun Winterton
◊ Placement of oldest Fossil

◊ Summary + Future

Stratiomydas lividus © Charles Melton

.. previous phylogenetic hypotheses 4

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

ae
in

m e
in

in
e
yd

go na
ae
g

yd rina
ae

yd
e

py

om

lle di
tin
na

iin

Sy my
Le ae
uo

o
is
al
hi

ph
al

in
hl
at

om

o
yp

op

io

pt
oc

Diochlistinae remaining Mydidae
ac
t

Ap
Rh
An
Ec

M
Di
C

Mitrodetus dentitarsis © Steve Marshall


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


Dikow 2009
› morphology
› parsimony

... Asilidae
Apiocera aldrichi
Apioceridae
Apiocera painteri

Rhaphiomidas maehleri Rhaphiomidinae

Neorhaphiomidas sp (Mandurah) Megascelinae Afroleptomydas suffusipennis
Opomydas townsendi Ectyphinae

Mitrodetus dentitarsis Diochlistinae

Mydas clavatus Mydinae

Pseudonomoneura hirta
Leptomydinae
Nemomydas brachyrhynchus

Afroleptomydas sp (Clanwilliam) Syllegomydinae

Leptomydas lusitanicus © Valter Jacinto

... new phylogenetic hypothesis 7

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


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


Phylogenetic Analysis
◊ Morphological data
› MacClade (version 4.08)
› WinClada (version 1.0.08)
› all multi-state 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

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 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
Hessemydas parkeri
Hessemydas seyrigi
Nemomydas venosus
Mydidae
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.
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
Pilica formidolosa
Laphria aktis bimaculatus
Namadytes maculiventris
Neolophonotus
Ommatius tibialis
Namadytes prozeskyi
Philonicus albiceps
Stichopogoncinctus
Lasiopogon punctum
Diogmites grossus
Cephalocera imitata
Ospriocerus aeacus
Damalis monochaetes
Pegesimalluscylindrica
Cephalocera longirostris
Leptogaster laticornis
Euscelidia pulchra Cephalocera sp. (Papendorp)
Holcocephala calva bilineata
Apiocera (Pyrocera)
Apiocera (Pyrocera) aldrichi Neolaparopsis puncturatus
Apiocera (Pyrocera) obscura
Apiocera (Anypenus)parkeri
Arenomydas caerulescens
Apiocera (Anypenus)sp. (Wurarga)
philippii
Apiocera (Apiocera)
Arenomydas callosus
Tongamya miranda (Mandurah)
Arenomydas namaquensis
Neorhaphiomidas sp.
Nomoneura caffra
Rhopalia olivieri
Rhopalia annulata
Nomoneura paradoxa
Nomoneuroides natalensis
Mydas clavatus
Mydas xanthopterus
Halterorchis sp. nov. (Cape Recife)
Apiophora paulseni
Miltinus viduatus
gen. nov. + sp. nov. (Panne)
Heteromydas bicolor
Opomydas townsendi
gen. nov. + sp. nov. (Ugab)
Ectyphus pinguis (Amboseli)
Ectyphus sp. crassipes
Haplomydas nov.
gen. nov. + sp. nov. (Gobabeb)
Eremomidas arabicus(Tiega)
Eremomidas sp. nov. Namibimydas sp. nov. (Agate)
Nothomydas sp. nov. (Skerpioenkop)
Pseudonomoneura
Hessemydas seyrigihirta
parkeri
Hessemydas venosus
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)
Afroleptomydas C sp. (Stampriet)
Namadytes prozeskyi
Cephalocera imitata
Cephalocera sp. (Papendorp)
Syllegomydas sp. nov. (Ruacana)
Neolaparopsis puncturatus
Arenomydas callosus
Arenomydascaffra
Syllegomydas N arnoldi
Nomoneura namaquensis
Nomoneura paradoxa Syllegomydas N dispar
Nomoneuroides nov. (Cape
sp. natalensis
Halterorchissp. nov. (Panne)Recife)
gen. nov. +
gen. nov. + sp. nov. (Gobabeb) Syllegomydas S palestinensis

strict consensus tree from implied weights analysis
Namibimydassp. nov. (Skerpioenkop)
Nothomydas sp. nov. (Agate)
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.

Rhaphiomidinae
Monophyly & Rhaphiomidinae
Tongamya miranda
Megascelinae
Rhopalia olivieri
Rhopalia annulata Rhopaliinae
Anomalomydas mackerrasi Anomalomydinae
Cacatuopyga ruficornis Cacatuopyginae
Mydas clavatus
Mydas xanthopterus Mydinae
Apiophora paulseni
Miltinus viduatus
Apiophorinae
Heteromydas bicolor
Opomydas townsendi
Ectyphus pinguis
Ectyphinae
Syllegomydinae
Eremomidas arabicus
Eremomidas sp. nov. (Tiega) Leptomydinae
Diochlistinae
Hessemydas parkeri
Hessemydas seyrigi
Nemomydas venosus
Rhaphiomidas auratus © Greg Ballmer Leptomydas gruenbergi
Leptomydinae
Leptomydas sardous
Leptomydas turcicus
Eremomidas bek
Namadytes prozeskyi
Cephalocera imitata
Arenomydas callosus
Neorhynchocephalus
Nomoneura caffra
Toxophora maxima
Bombylius major
Dipalta serpentina
Apsilocephalaatra

Nomoneura paradoxa
Phycus frommeri
Pilica formidolosa
Neolophonotus

Ommatius tibialis

Syllegomydinae
Philonicus albiceps
Stichopogoncinctus
Lasiopogon punctum
Diogmites grossus

Ospriocerus aeacus
Damalis monochaetes
Euscelidia pulchra
Apiocera (Pyrocera)
Apiocera (Pyrocera) aldrichi gen. nov. + sp. nov. (Panne)
philippii

Apiocera (Apiocera)

monophyletic
Neorhaphiomidas sp.
Rhopalia olivieri Namibimydas sp. nov. (Agate)
Rhopalia annulata

Mydas clavatus
Mydas xanthopterus
Apiophora paulseni
Miltinus viduatus
Heteromydas bicolor

Opomydas townsendi
Haplomydas nov.


non-monophyletic
Eremomidas sp. nov.

Pseudonomoneura
parkeri
Hessemydas venosus

Leptomydas

Leptomydas bek
Vespiodes ngurumani

Syllegomydas N dispar
Namadytes prozeskyi

Syllegomydas S palestinensis
Cephalocera imitata
Arenomydas callosus

Arenomydascaffra
Nomoneura paradoxa
sp. natalensis

gen. nov. +
arnoldi Syllegomydas S elachys sp. nov.
Syllegomydas N astrictus sp. nov.
Syllegomydas S
Syllegomydas N elachys sp. nov.


Syllegomydinae non-monophyly
Eremomidas arabicus
Hessemydas parkeri
Hessemydas seyrigi
Nemomydas venosus
Leptomydas sardous
Leptomydas turcicus
Eremomidas bek
Namadytes prozeskyi
Cephalocera imitata
Arenomydas callosus
Nomoneura caffra
Nomoneura paradoxa
Limata tenuicornis volaticus gen. nov. + sp. nov. (Panne)
Neorhynchocephalus
Toxophora maxima
Bombylius major
Dipalta serpentina

Apsilocephalaatra
Phycus frommeri
Pilica formidolosa
Neolophonotus

Namibimydas sp. nov. (Agate)
Ommatius tibialis
Philonicus albiceps
Stichopogoncinctus
Lasiopogon punctum
Diogmites grossus
Ospriocerus aeacus

Damalis monochaetes
Euscelidia pulchra
Apiocera (Pyrocera)

philippii

Apiocera (Apiocera)
Neorhaphiomidas sp.

Rhopalia olivieri
Rhopalia annulata

Mydas clavatus
Mydas xanthopterus

Apiophora paulseni
Miltinus viduatus
Heteromydas bicolor
Opomydas townsendi

Haplomydas nov.
Eremomidas sp. nov.

Pseudonomoneura
parkeri
Hessemydas venosus

Leptomydas

Leptomydas bek
Vespiodes ngurumani

Namadytes prozeskyi
Cephalocera imitata

Arenomydas callosus
Arenomydascaffra
Nomoneura paradoxa
sp. natalensis

Cephalocera imitata
gen. nov. +


arnoldi
Syllegomydas S


Monophyly of higher-level taxa & Revised classification
Taxon (sensu Papavero & Wilcox 1975) Monophyly Revised, monophyletic taxon
Anomalomydinae monogeneric
Apiophorinae — Paramydas
Cacatuopyginae √
Diochlistinae √
Ectyphinae √
Leptomydinae — Eremomidas sp.
Megascelinae √
Mydinae √ + Paramydas (Chile)
Rhaphiomidinae √
Rhopaliinae √
Syllegomydinae — Haplomydas, Eremohaplomydas

.... character evolution 14

Character evolution: head + female
Tongamya miranda
Rhopalia olivieri
Rhopalia annulata
Mydas clavatus
Mydas xanthopterus
Apiophora paulseni
Miltinus viduatus
Heteromydas bicolor
Opomydas townsendi
Ectyphus pinguis
Eremomidas arabicus
Hessemydas parkeri
Hessemydas seyrigi
Nemomydas venosus
gen. nov. + sp. nov. Gobabeb Leptomydas gruenbergi
Leptomydas sardous
Leptomydas turcicus
Eremomidas bek
Namadytes prozeskyi
Cephalocera imitata
Arenomydas callosus
Neorhynchocephalus
Nomoneura caffra
Toxophora maxima
Bombylius major
Dipalta serpentina
Apsilocephalaatra

Nomoneura paradoxa
Phycus frommeri

acanthophorite spurs absent
Pilica formidolosa
Neolophonotus

Ommatius tibialis
Philonicus albiceps
Stichopogoncinctus
Lasiopogon punctum
Diogmites grossus

Ospriocerus aeacus
Damalis monochaetes
Euscelidia pulchra
Apiocera (Pyrocera)
Apiocera (Pyrocera) aldrichi gen. nov. + sp. nov. (Panne)
philippii

Apiocera (Apiocera)

proboscis vestigial
Neorhaphiomidas sp.
Rhopalia olivieri Namibimydas sp. nov. (Agate)
Rhopalia annulata

Mydas clavatus
Mydas xanthopterus
Apiophora paulseni
Miltinus viduatus
Heteromydas bicolor

Opomydas townsendi
Haplomydas nov.


proboscis and acanthophorite spurs
Eremomidas sp. nov.

Pseudonomoneura
parkeri
Hessemydas venosus

Leptomydas

Leptomydas bek
Vespiodes ngurumani

Namadytes prozeskyi

Cephalocera imitata
Arenomydas callosus

well-developed
Arenomydascaffra
Nomoneura paradoxa
sp. natalensis

gen. nov. +
arnoldi Syllegomydas S elachys sp. nov.
Syllegomydas S

..... oldest fossil 15

Placement of oldest fossil Mydidae
Limata tenuicornis Tabanidae
Neorhynchocephalus volaticus
Nemestrinus nigrovillosus Nemestrinidae
Eulonchus sp. Acroceridae
Prorates sp. (Escalante) Scenopinidae
Toxophora maxima
Bombylius major Bombyliidae
Dipalta serpentina
Apsilocephala longistyla
Hemigephyra atra
Apsilocephalidae
Phycus frommeri Therevidae

› † Cretomydas santanensis
Pilica formidolosa
Laphria aktis
Neolophonotus bimaculatus
Philonicus albiceps
Ommatius tibialis

Asilidae
› coded for 23.1 % of characters
Stichopogon punctum
Lasiopogon cinctus
Diogmites grossus
Ospriocerus aeacus
Damalis monochaetes
Pegesimallus laticornis
Holcocephala calva
Leptogaster cylindrica
Euscelidia pulchra
Apiocera P bilineata
Apiocera R braunsi
Apiocera P aldrichi

Apioceridae
Apiocera P painteri
Apiocera P arena
Apiocera P parkeri
Apiocera An obscura
Apiocera An philippii
Apiocera Ap sp. Wurarga
Apiocera Ap sp. Maranoa
Apiocera Ap sp. Stuart
Tongamya miranda
Rhopalia olivieri
Rhopalia annulata
Mydas clavatus
Mydas xanthopterus
Cretomydas santanensis
Apiophora paulseni

≈ 112 myo
Miltinus viduatus
Heteromydas bicolor
Opomydas townsendi
Ectyphus pinguis
Eremomidas arabicus
Hessemydas parkeri
Hessemydas seyrigi
Nemomydas venosus
Mydidae
Eremomidas bek
Leptomydas rapti
Leptomydas turcicus
Leptomydas sardous
Leptomydas notos
Mydaselpis ngurumani
Neorhynchocephalus
Toxophora major
Bombylius maxima
Dipalta serpentina
Apsilocephala longistyla
Hemigephyra atra
Phycus frommeri
Vespiodes phaios
Pilica formidolosa
Neolophonotus Namadytes maculiventris
Ommatius tibialis
Philonicus albiceps Namadytes prozeskyi
Stichopogoncinctus
Lasiopogon punctum
Diogmites grossus
Ospriocerus aeacus Cephalocera imitata
Damalis monochaetes
Euscelidia pulchra
Dioctria hyalipennis Cephalocera sp. (Papendorp)

4 trees, fit 95.86467
Apiocera (Pyrocera)

† Cretomydas santanensis, Crato Formation
Apiocera (Apiocera) philippii
Rhaphiomidas hasbroucki Arenomydas callosus
Neorhaphiomidas sp. Arenomydas namaquensis
Rhopalia olivieri Nomoneura caffra
Rhopalia annulata
Nomoneura paradoxa

= 1563 steps
Mydas clavatus
Mydas xanthopterus
Apiophora paulseni
Miltinus viduatus
Stenomydas gen. nov. sp. nov. (Panne)
Heteromydas bicolor
Opomydas townsendi
Parectyphus namibiensis Namibimydas sp. nov. (Agate)
Ectyphus pinguis
Haplomydas nov. (Amboseli)
Eremohaplomydas desertorum Nothomydas sp. nov. (Skerpioenkop)

CI=0.23
Eremohaplomydas sp.
Eremomidas arabicus nov. (Kotwa)
Diochlistus gracilis Cleamydas gen. nov. sp. nov. (Ugab)
Syll gen. nov. sp. nov. (Gobabeb)
Hessemydas parkeri
Hessemydas seyrigi
Nemomydas venosus
Leptomydas brachyrhynchus
Leptomydas notossp. nov.
Leptomydas turcicus
bek
Leptomydasngurumani sp. nov.

RI=0.75
Mydaselpislepolditertii
Vespiodes
Namadytes maculiventris Afroleptomydas C sp. (Stampriet)
Namadytes prozeskyi
Cephalocera imitata
Cephalocera sp. (Papendorp) Syllegomydas sp. nov. (Ruacana)
callosus
Arenomydascaffra Syllegomydas N arnoldi
Nomoneura
Nomoneura paradoxa
Nomoneuroides nov. (Cape Recife)
natalensis
Halterorchissp. nov. (Panne)
gen. nov. + sp.

strict consensus shown (1564 steps)
Afroleptomydas C sp. (Ruacana)
Syllegomydas S dispar
Syllegomydas N algiricus sp. nov.
Syllegomydas S heothinos
Syllegomydas S elachys
Syllegomydas N astrictus sp. nov. Syllegomydas N astrictus

.... summary + future 16

Summary
◊ 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

Phyllomydas parvulus © Lyn Atherton

17

Acknowledgements
◊ Museum 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

Recommended

Recommended

More Related Content

Viewers also liked

Viewers also liked (10)

TDikow Mydidae morphological phylogeny ESA 2010