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Towards a phylogeny of robber flies (Insecta: Diptera: Asilidae) -- evidence from morphology of the imagines

Torsten Dikow

American Museum of Natural History, New York, NY, USA Cornell University, Ithaca, NY, USA

. introduction

Asilidae - robber flies

» predatory flies (larvae & imagines) » 7,000 species world-wide

Efferia sp

Hoplistomerus nobilis

. introduction

Leptogastrinae

» morphologically very distinct » feeding on resting prey, e.g. spiders » 46 species in 7 genera » phylogenetic position within Asilidae ? > sister-group to Asilidae > derived clade related to Asilinae

Lasiocnemus fascipennis

. introduction

At om os iin La i ph ri i n H op i lis to m er Da in m i al is G St ru ic pp ho e po La go si ni op og ni on üb ri g e Da sy po Le go pt ni og na e as te ri n i O m m As atii ilin n i i

Karl 959

?

?

? ?

Dasypogoninae

Asilinae

Bybee et al. 004

outgroups

4

Leptogaster sp Psilonyx annulatus Holcocephala abdominalis Ommatius sp Philonicus arizonensis Machimus sp Promachus bastardii Promachus sp Efferia nemoralis Proctacanthus nearno Diogmites grossus Saropogon fletcheri Andrenosoma fulvicauda Adelodus sp Maira sp Smeryngolaphria sp Laphria sp Holopogon currani Senobasis corsair Laphystia sp Ospriocerus latipennis Stenopogon martini Hypenetes critesi Stichopogon trifasciatus Obelophorus landbecki Creolestes nigribarbis

Bybee, S.M. et al. 2004 Mol. Phyl. & Evo. 30: 789­797. Karl, E. 1959 B. Entomologie 9(5­6): 619­680.

. introduction

Main Objectives

» Higher-level phylogeny » Position of Leptogastrinae within Asilidae » Sister-group of Asilidae » Improve classification

Lasiopogon cinctus - female ovipositor Philonicus albiceps - male genitalia

1 mm

1 mm

Hoplistomerus nobilis - pruinosity on scutum

Lasiocnemus lugens - female spermathecae

1 mm 100 µm

5

.. morphological analysis

Taxon sampling

» 58 Asilidae species » Higher taxa > all subfamily taxa > 9 of the 4 tribal taxa > 9 of the 57 genera » Outgroup taxa > 6 Asiloidea & Nemestrinidae species

··· ··· ·· ·· · · · · ·· ··· · · · · ·· · · · · ·· · · · · · ·· · · · ··· ·· ·· · · ·· · · · ·· ·· · · · · · ·· · · · · · · · ·· · · ·· ·· · · ··· ·· · ··· · · ·· ·

· · ·

· ··· · · · · · · · · ··· · · ·· · ·

· · · · · · · · · · ·· · ·

· ··· ·· · ·

6

.. morphological analysis

Morphological Analysis

» Characters: > 0 parsimony informative characters > 4 binary & 06 multistate > all non-additive (unordered) > few missing or inapplicable data » Analysis: > MacClade & WinClada > TNT (version .0)

search strategy 300 RAS + TBR 1500 ratchet 1500 drift trees found 20 710 720 steps 2761 2760 2760

> Bremer support - 5000 trees, >, above branches > Jackknife - 000 replications, >50, below branches

7

.. morphological analysis

Morphological Hypothesis

Neorhynchocephalus volaticus Bombylius major Poecilognathus sp (el hacha) Prorates sp (escalante) Apsilocephala longistylus Hemigephyra atra Phycus frommeri Apiocera aldrichi Apiocera painteri Rhaphiomidas meahleri Neorhaphiomidas sp (mandurah) Opomydas townsendi Mitrodetus dentitarsis Mydas clavatus Pseudonomoneura hirta Nemomydas brachyrhynchus Afroleptomydas sp (clanwilliam) Perasis transvaalensis Zabrops tagax Protometer sp (el tuparro) Psilocurus modestus Laphystia aegyptiaca Laphystia tolandi Hoplistomerus nobilis Trichardis testacea Laxenecera albicincta Nusa infumata Smeryngolaphria numitor Orthogonis scapularis Goneccalypsis argenteoviridis Loewinella nigripes Aphestia annulipes Atomosia puella Cerotainia albipilosa Maira aenea Lampria clavipes Choerades bella Laphria flava Laphria aktis Lamyra gulo Stiphrolamyra angularis Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris Proagonistes praeceps Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Ommatius senex Ommatius tibialis Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus Lycomya germainii Pogonioefferia pogonias Proctacanthus philadelphicus Dasophrys crenulatus Eichoichemus pyrrhomystax Lochmorhynchus albicans Myaptex brachyptera Neolophonotus bimaculatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Dysmachus trigonus Philonicus albiceps Congomochtherus inachus Heligmonevra leavis Tolmerus atricapillus Zosteria rosevillensis Megaphorus pulchrus Promachus amastrus Apoclea algira Philodicus tenuipes Asilus crabroniformis Asilus sericeus Neomochtherus pallipes Afromochtherus mendax Clephydroneura sundaica Clephydroneura sp (kepong) Colepia rufiventris Neoitamus cyanurus Threnia carbonaria Bathypogon nigrinus Phellus olgae Obelophorus terebratus Tillobroma punctipennis Hypenetes stigmatias Lycostommyia albifacies Coleomyia setigera Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Connomyia varipennis Prolepsis tristis Creolestes nigribarbis Microstylum sp (karkloof) Daspletis stenoura Plesiomma sp (guanacaste) Ospriocerus aeacus Scleropogon subulatus Stenopogon sabaudus Stenopogon rufibarbis Gonioscelis ventralis Ancylorhynchus fulvicollis Scylaticus costalis Acnephalum cylindricum Sisyrnodytes sp (gamka) Ablautus coquilletti Willistonina bilineata Trichoura sp (tierberg) Oligopogon sp (cape recife) Broticosia paramonovi Dicolonus simplex Echthodopa pubera Myelaphus melas Nannodioctria sp (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Acronyches maya Beameromyia bifida Euscelidia pulchra Tipulogaster glabrata Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Damalis sp (palatupana) Damalis monochaetes Damalis annulata Rhipidocephala sp (harold johnson) Trigonomima sp (anamaliensis) Holcocephala abdominalis Holcocephala calva Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Holopgon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster sp (pedion) Heteropgon manicatus Metapogon punctipennis

strict consensus 965 steps Ci=0., Ri=0.65

>50 99

1 mm

Dioctria atricapilla

8

.. morphological analysis

Morphological Hypothesis - simplified

Nemestrinidae Bombyliidae Bombyliidae Scenopinidae Apsilocephalidae Therevidae Mydidae Apioceridae Laphystiini Laphriini Ommatiini Apocleini Asilini Bathypogon nigrinus Phellini Tillobromatini Dasypogonini Coleomyia setigera Stenopogonini Enigmomorphini Ancylorhynchus fulvicollis Scylaticus costalis Willistoninini Oligopogon sp (Cape Recife) Dioctriini Trigonomimini Acronyches maya Leptogastrini Stichopogonini Cyrtopogonini Leptarthrus brevirostris Holopogon priscus Chrysopogonini Brachyrhopalini Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis

> epistomal margin elevated > prothoracic coxa with depression > upper calypter reduced 9

simplified strict consensus of 70 most parsimonious cladograms

.. morphological analysis

Laphriinae + *Laphystiinae

22 95 12 84 11 79 >50 94 >50 89 Perasis transvaalensis Zabrops tagax Protometer sp (el tuparro) Psilocurus modestus Laphystia aegyptiaca Laphystia tolandi Hoplistomerus nobilis Trichardis testacea Laxenecera albicincta Nusa infumata Smeryngolaphria numitor Orthogonis scapularis Goneccalypsis argenteoviridis Loewinella nigripes Aphestia annulipes Atomosia puella Cerotainia albipilosa Maira aenea Lampria clavipes Choerades bella Laphria flava Laphria aktis Lamyra gulo Stiphrolamyra angularis Andrenosoma cyrtoxys Hyperechia nigripennis Pilica formidolosa Dasyllis funebris Proagonistes praeceps

3 >50 19 99 90 11 70 11 93

50

15 84 13 73 15 9 61 21 96 66 30 97 3 50 13 78

Laxenecera albicincta

0

.. morphological analysis

*Apocleinae + *Asilinae + Ommatiinae

10 68 10 54 17 78 Cophinopoda chinensis Michotamia aurata Ommatius incurvatus Ommatius senex Ommatius tibialis Emphysomera conopsoides Emphysomera pallidapex Blepharotes splendidissimus Lycomya germainii Pogonioefferia pogonias Proctacanthus philadelphicus Dasophrys crenulatus Eichoichemus pyrrhomystax Lochmorhynchus albicans Myaptex brachyptera Neolophonotus bimaculatus Nomomyia murina Satanas gigas Machimus chrysitis Machimus occidentalis Dysmachus trigonus Philonicus albiceps Congomochtherus inachus Heligmonevra leavis Tolmerus atricapillus Zosteria rosevillensis Megaphorus pulchrus Promachus amastrus Apoclea algira Philodicus tenuipes Asilus crabroniformis Asilus sericeus Neomochtherus pallipes Afromochtherus mendax Clephydroneura sundaica Clephydroneura sp (kepong) Colepia rufiventris Neoitamus cyanurus Threnia carbonaria

45 99

11 55

> maxillae with blunt tip > setae on anepimeron

4 6

20 92

39 91 5 5 >50 99 6 55

Neolophonotus sp

.. morphological analysis

*Dasypogoninae + *Stenopogoninae

>50 94 6 3 6 60 6 6 5 11 85 6 6 64 70 >50 99 3 3 7 72 6 60 26 87 Bathypogon nigrinus Phellus olgae Obelophorus terebratus Tillobroma punctipennis Hypenetes stigmatias Lycostommyia albifacies Coleomyia setigera Lestomyia fraudiger Dasypogon diadema Saropogon dispar Saropogon luteus Thereutria amaraca Archilestris magnificus Neodiogmites melanogaster Diogmites grossus Blepharepium cajennensis Molobratia teutonus Pegesimallus laticornis Senobasis corsair Cyrtophrys attenuatus Deromyia fuscipennis Megapoda labiata Pseudorus distendens Connomyia varipennis Prolepsis tristis Creolestes nigribarbis Microstylum sp (karkloof) Daspletis stenoura Plesiomma sp (guanacaste) Ospriocerus aeacus Scleropogon subulatus Stenopogon sabaudus Stenopogon rufibarbis Gonioscelis ventralis Ancylorhynchus fulvicollis Scylaticus costalis ...

Leptarthrus brevirostris Dasypogon diadema prothoracic tibiae in lateral view, not to scale

.. morphological analysis

Dioctriinae + Leptogastrinae + *Stenopogoninae + Trigonomiminae

3 11 85 3 Acnephalum cylindricum Sisyrnodytes sp (gamka) Ablautus coquilletti Willistonina bilineata Trichoura sp (tierberg) Oligopogon sp (cape recife) Broticosia paramonovi Dicolonus simplex Echthodopa pubera Myelaphus melas Nannodioctria sp (formicaphaga) Eudioctria albius Dioctria atricapillus Dioctria hyalipennis Dioctria rufipes Acronyches maya Beameromyia bifida Euscelidia pulchra Tipulogaster glabrata Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Damalis sp (palatupana) Damalis monochaetes Damalis annulata Rhipidocephala sp (harold johnson) Trigonomima sp (anamaliensis) Holcocephala abdominalis Holcocephala calva ...

7

13 81 7

7 7 >50 93

>50 99 >50 3

3

100 4 53 7 53 7 62 >50 99 18 10 75 75 7 72 11 63 >50 99

14 84

Damalis speciosa

.. morphological analysis

Leptogastrinae + Acronyches maya

» Autapomorphies for entire taxon > hypopharynx `teeth' far apart > pulvilli reduced & claws not angled > abdominal segment » Autapomorphies of Leptogastrinae > postpronotal lobe development > metathoracic coxae directed anteriorly > male genitalia

Beameromyia disfascia

4

© Michael Thomas 2006

.. morphological analysis

*Dasypogoninae + *Stenopogoninae + Stichopogoninae

14 93 >50 99 10 89 >50 99 50 82

5

>50 99 62 98

Lasiopogon aldrichii Lasiopogon cinctus Townsendia albomacula Stichopogon elegantulus Stichopogon trifasciatus Stichopogon punctum Lissoteles aquilonius Cyrtopogon lateralis Cyrtopogon rattus Afroholopogon peregrinus Dasycyrton gibbosus Leptarthrus brevirostris Holopgon priscus Chrysopogon pilosifacies Codula limbipennis Austrosaropogon nigrinus Brachyrhopala ruficornis Cabasa pulchella Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster sp (pedion) Heteropgon manicatus Metapogon punctipennis

Stichopogon elegantulus (l) Diogmites grossus (r) compound eyes in lateral view not to scale

5

.. morphological analysis

Changes to classification

Apioceridae + Mydidae Laphystiini Laphriini Ommatiini Apocleini Asilini Bathypogon nigrinus Phellini Tillobromatini Dasypogonini Coleomyia setigera Stenopogonini Enigmomorphini Ancylorhynchus fulvicollis Scylaticus costalis Willistoninini Oligopogon sp (Cape Recife) Dioctriini Trigonomimini Acronyches maya Leptogastrini Stichopogonini Cyrtopogonini Leptarthrus brevirostris Holopogon priscus Chrysopogonini Brachyrhopalini Ceraturgus fasciatus Nannocyrtopogon lestomyiformis Nicocles politus Cophura brevicornis Rhabdogaster pedion Heteropogon manicatus Metapogon punctipennis

6

simplified strict consensus of 70 most parsimonious cladograms

... preliminary molecular study

Preliminary Molecular Analysis » Taxon sampling > 5 Asilidae species + Asiloidea species » Genes > CAD > EF-1 > 8S rDNA > 8S rDNA 900 bp ,00 bp ,00 bp ,000 bp

» Analysis > POY (version .0.a-550477.78) > cluster computer

data-set complete Morph+18S+28S search strategy 27 RAS + TBR 50 RAS + TBR steps 19776 9463

7

... preliminary molecular study

Morphology, CAD, EF-1, 8S, 8S

Bombylius major Hemigephyra atra Apsilocephala longistylus Phycus frommeri Prorates sp Leptarthrus brevirostris Apiocera painteri Opomydas townsendi Mydas clavatus Mitrodetus dentitarsis Afrleptomydas sp (Clanwilliam) Nemomydas brachyrhynchus Stichopogon punctum Stichopogon elegantulus Atomosia puella Laphria aktis Choerades bella Laphystia tolandi Hoplistomerus nobilis Trichardis testacea Laxenecera albicincta Perasis transvaalensis Stiphrolamyra angularis Damalis annulata Damalis monochaetes Pegesimallus laticornis Diogmites grossus Lestomyia fraudiger Saropogon luteus Dasypogon diadema Stenopogon rufibarbis Molobratia teutonus Cyrtopogon rattus Ablautus coquilletti Connomyia varipennis Lycostommyia albifacies Gonioscelis ventralis Acnephalum cylindricum Eudioctria albius Dioctria rufipes Dioctria hyalipennis Rhipidocephala sp (Harold Johnson) Holcocephala calva Holcocephala abdominalis Lasiopogon cinctus Leptogaster arida Leptogaster cylindrica Lasiocnemus lugens Tipulogaster glabrata Beameromyia bifida Euscelidia pulchra Megaphorus pulchrus Promachus amastrus Philonicus albiceps Tolmerus atricapillus Asilus sericeus Asilus crabroniformis Dasophrys crenulatus Neolophonotus bimaculatus Emphysomera pallidapex Ommatius senex Ommatius tibialis

8

most parsimonious cladogram from 7 RAS+TBR, 9776 steps

... preliminary molecular study

Morphology, 8S, 8S

Bombylius major Hemigephyra atra Apsilocephala longistylus Phycus frommeri Prorates sp Apiocera painteri Opomydas townsendi Mydas clavatus Mitrodetus dentitarsis Nemomydas brachyrhynchus Afrleptomydas sp (Clanwilliam) Acnephalum cylindricum Stenopogon rufibarbis Gonioscelis ventralis Hoplistomerus nobilis Tricahrdis testacea Laxenecera albicincta Stiphrolamyra angularis Perasis transvaalensis Choerades bella Laphria aktis Atomosia puella Laphystia tolandi Lycostommyia albifacies Connomyia varipennis Cyrtopogon rattus Leptarthrus brevirostris Ablautus coquilletti Molobratia teutonus Pegesimallus laticornis Lestomyia fraudiger Diogmites grossus Dasypogon diadema Saropogon luteus Lasiopogon cinctus Stichopogon punctum Stichopogon elegantulus Rhipidocephala sp (Harold Johnson) Holcocephala calva Holcocephala abdominalis Damalis monochaetes Damalis annulata Eudioctria albius Dioctria rufipes Dioctria hyalipennis Beameromyia bifida Euscelidia pulchra Leptogaster arida Leptogaster cylindrica Tipulogaster glabrata Lasiocnemus lugens Dasophrys crenulatus Neolophonotus bimaculatus Megaphorus pulchrus Promachus amastrus Emphysomera pallidapex Ommatius senex Ommatius tibialis Philonicus albiceps Tolmerus atricapillus Asilus crabroniformis Asilus sericeus

9

most parsimonious cladogram from 50 RAS+TBR, 946 steps

.... conclusions

Summary & Conclusion

» Asilidae is not monophyletic in all analyses » Leptogastrinae is always a derived clade » Apioceridae + Mydidae sister-group

Taxon Asilidae Apocleinae Asilinae Dasypogoninae Dioctriinae Laphriinae Laphystiinae Leptogastrinae Ommatiinae Stenopogoninae Stichopogoninae Trigonomiminae Morph Complete Morph, 18S, 28S

+ -- -- -- + + -- + + -- + +

-- -- + -- + -- -- + + -- -- --

+ -- + -- + -- -- + + -- + --

0

+ --

= monophyletic = non-monophyletic

Acknowledgements

» Specimens: museum curators, Jason Londt, Martin Hauser, Shaun Winterton, Jean-Michel Maldes, Eric Fisher, David Clements, Fritz Geller-Grimm » Laboratory: Ward Wheeler & lab, Brian Wiegmann & lab » Analysis: Taran Grant, Kurt Pickett » Jim Liebherr, David Grimaldi, Roberto Keller

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