“Parasitic Fruit Flies”:Pyrgotidae, Ctenostylidae, Tachiniscidae

“Parasitic Fruit Flies”:Pyrgotidae, Ctenostylidae,

Tachiniscidae

Valery A. Korneyev

I.I.Schmalhausen Institute of Zoology

National Academy of SciencesKyiv, Ukraine

Background

• The tephritid lineage of the Higher Tephritoidea

• Family-group taxa of unresolved relationships and limits

• Pyrgotidae, Tachniniscidae, Ctenostylidae and Tephritidae.

Campylocera angustigenis

Ramuliseta sp.n.

Tachiniscini gen. sp. (photo: T.Saigusa)

Rioxa discoidalis (photo: orionmystery diptera info; J. Smit det.)

Background

• Ctenostylidae: • a subfamily of Pyrgotidae (as

Lochmostyliinae).

• D.K.McAlpine (1990):Separate family beyond Tephritoidea.

• V.A.Korneyev (2001; 2006; 2010): Separate family of uncertain position of Higher Tephritoidea.

Background

• Tachiniscidae:• Hennig (1958) – sister-

group of Tephritidae.• J.F.McAlpine (1989):

sister-group of Pyrgotidae

• V.A.Korneyev (1999); Korneyev & Norrbom (2006): subfamily of Tephritidae, sister-group of Tephritidae.

• Han & Ro (2009): sister-group of Tephritidae (poor bootstrap support)

Background

• Tachiniscidae:

• Ortalotrypetini are similar to Tephritidae Phytalmiinae and poorly distinguishable.

• For practical identification reasons, they were retained in Tephritidae (Korneyev & Ovchinnikova, 2006).

• Tachiniscini are similar to Pyrgotidae and poorly distinguishable.

Background

• Pyrgotidae:• Subfamily Pyrgotinae

is monophyletic (fused sternites 1 & 2)

• Five basal taxa:Teretrurinae,Prodalmannia, Maenomenus, Descoleia, Nosferatumyia are of uncertain relationships.

Background

Rioxa discoidalis (photo: orionmystery diptera info; J. Smit det.)

• Tephrtidae:• some subfamilies/

tribes with saprophagous larvae (Phytalmiinae, Gastrozonini, possibly some Adramini)

• phytophagy evolved many times in the family.

• Basal lineages of Pyrgotidae, Tachiniscidae and Tephritidae are represented by the genera, sharing many diagnostic characters.

Problem

Problem

• Must we lump Pyrgotidae and Tachiniscidae into Tephrtidae to form one family?

• Must we adopt an oxymoron “parasitic fruit flies”?

Basal lineages of Pyrgotidae / Tachiniscidae

• Ortalotrypetini • Tephritid-like appearance, incl.

presence of ocelli, frontal setae, 3 pairs of scutellar setae, apex of Sc strongly angled; sternites 1 and 2 separate.

• Female: oviscape opened dorsally. Eversible membrane ventrally dentate. Aculeus short, acute. (SA with Tachiniscini).

• Male: surstyli present; glans and hypandrium as in Blepharoneurini (Tephritidae).

Korneyev & Norrbom 2006

Neortalotrypeta Norrbom


• Descoleia Aczél 1956• Partly tephritid-like

appearance, incl. presence of ocelli, apex of Sc strongly angled; sternites 1 and 2 separate. No frontal setae, 2 pairs of scutellar setae.

• Female: oviscape apparently not opened dorsally. Eversible membrane possibly dentate. Aculeus acute. (Aczél, 1956).

• Male: prensisetae present; glans and hypandrium as in Blepharoneurini (Tephritidae).

• Other details see Korneyev & Norrbom (2006).


• Nosferatumyia Korneyev 2006• Partly tephritid-like appearance,

incl. presence of ocelli, apex of Sc strongly angled; sternites 1 and 2 separate. No frontal setae, 2 pairs of scutellar setae.

• Female unknown.• Male: prensisetae present; glans

and hypandrium as in Blepharoneurini (Tephritidae).

• Other details see Korneyev & Norrbom (2006).

• Larval feeding unknown.• Distribution: dubious data

(Palaearctic Region?)


• Maenomenus Bezzi 1929• Partly tephritid-like appearance,

incl. presence of ocelli, 3 pairs of scutellar setae sternites 1 and 2 separate. No frontal setae. Apex of Sc strongly angled at 75°.

Female: oviscape opened posteriorly. Eversible membrane bare. Aculeus long and acute.

Male: prensisetae present; glans and hypandrium as in Blepharoneurini (Tephritidae).

Larvae in adult Melolontinae (Scarabaeidae).

Distribution: Australia.


• Teretrurini• Partly tephritid-like appearance,

incl. ocelli present, 3 pairs of scutellar setae, sternites 1 and 2 separate. No frontal setae. Apex of Sc strongly angled at 75°.

Female: oviscape opened posteriorly. Eversible membrane bare. Aculeus long and narrow, with cercal unit acute, entirely integrated.

Male: prensisetae absent; glans partly simplified.


Distribution: Argentina, Chile.


• Prodalmannia Bezzi 1929• Partly tephritid-like appearance,

incl. 3 pairs of scutellar setae; sternites 1 and 2 separate. No frontal setae, no ocelli. Apex of Sc strongly angled at 75°.

Female: oviscape opened ventrally. Eversible membrane bare. Aculeus short, with cercal unit acute, but not entirely integrated.

Male: prensisetae present; glans and hypandrium as in Blepharoneurini (Tephritidae).



Pyrgotinae

• Toxopyrgota Hendel 1914• Ocelli absent, 2 pairs of

scutellar setae, sternites 1 and 2 fused with seam. Frontal setae present. Sc almost straight, apically obsolete.

Female: oviscape opened ventrally. Eversible membrane bare. Aculeus short, acute, entirely integrated.

Male: prensisetae present; glans simplified.

Larvae unknown.Distribution: Southern Africa.

Pyrgotinae

• Toxurini• Ocelli present or absent, 2 pairs

of scutellar setae, sternites 1 and 2 fused without seam. Frontal setae absent. Apex of Sc strongly angled at 90°.

Female: oviscape opened posteriorly or ventrally. Eversible membrane bare. Aculeus variable, entirely integrated.

Male: prensisetae absent; glans simplified.



Pyrgotinae

• Pyrgotini• Ocelli absent, 2 pairs of

scutellar setae, sternites 1 and 2 fused without seam. Frontal setae absent. Sc straight.




Distribution: circumtropical, rich in Madagascar.

Pyrgotinae

• Adapsiliini• Ocelli absent, 2(1) pairs of

scutellar setae, sternites 1 and 2 fused without seam. Frontal setae absent. Sc straight.




Distribution: palaeotropics, with a few genera in the Palaearctics, Madagascar, and PNG.

Ctenostylidae

• Ocelli absent, 2(1) pairs of scutellar setae, sternites 1 and 2 fused with seam. Frontal setae present or absent. Sc straight, apically obsolete.

Female: oviscape opened posteriorly. Eversible membrane bare. NO ACULEUS.

Male: prensisetae absent; glans simplified or lacking.

Larval habits unknown, some species with barbate apex of egg (as in Stylogaster).

Distribution: circumtropical, incl. PNG.

Preliminary phylogenetic analysis

Tachiniscinae

Descoleia

Nosferatumyia

Maenomenus

Prodalmannia

Teretrurinae

Toxopyrgota

ToxuriniPyrgotiniAdapsiliini

Tephritidae

Parasites Parasites of scarabsof scarabs

Parasites Parasites of scarabsof scarabs

Ctenostylidae

??

??

Parasites of Parasites of moths ?moths ?

11

22

66

33

??

??88

99

55

44

8877

1010

Synapomorphies:• Tephritidae (without Tachiniscinae):

• Greater ampulla present (SA).• Aculeus in basal groups with blunt and

separate cercal unit (SP).• Originally, larvae saprophagous or

saprozoophagous.

• Pyrgotidae (with Tachiniscinae):

• Greater ampulla absent (SP).

• Aculeus acute, with entirely integrated cercal unit (SA).

• Specialized parasites.

Taxonomic implications:

• Including Tachiniscinae (as a subfamily?) into Pyrgotidae does not contradict monophyly of the family.

• Concept of Pyrgotidae: Parasitic Higher Tephritoidea without greater ampulla and with acute aculeus for cutting animal tissues.

• Concept of Tephritidae:Saprophagous (or phytophagous) Higher Tephritoidea with blunt (or acute) aculeus and greater ampulla.

• Tachiniscidae Kertesz 1903 is a junior synonym of Tephritidae Newman 1834.

Molecular data

• Mitochondrial 12S, 16S, and COII (Han & Ro, 2009) – low bootstrap support for Tephritidae+ Tachiniscidae clade.

• Wiegman et al. (2011) 28S, cad, tpi, and aats1 genes – significant bootstrap support for Pyrgotidae+ Tachiniscidae clade.

Conclusions

• Despite the monophyly of Pyrgotidae s. l. (with Tachiniscidae) is supported only by the molecular data and having cutting aculeus, adopting such a concept would solve the problem of familial position of Descoleia and Nosferatumyia.

• Ctenostylidae are provisionally considered a related group of Toxopyrgota, with many morphological aberrations, possibly a sister-group of Pyrgotinae.

Conclusions

• Pyrgotidae, especially basal taxa are rare in collections and hardly available as fresh material. Phylogenies supported by molecular data are to be obtained but hardly will be available in the nearest future.

• No tephritoids with blunt aculeus and without greater ampulla are known. Wide hiatus between two families exists due to entire extinction of ancestral forms.

Acknowledgements

• This study is resulted from and partly supported by the :

• The Belgian Science Policy Office Grant in 2005-2006;

• Geddes Visiting Collection Fellowship (Australia) in 2012;

• DAAD (Germany) stipend in 2012.• National Academy of Sciences of Ukraine in 2011-

2015.

Acknowledgements

• Special thanks are due to Ho-Yeon Han, Allen Norrbom, David McAlpine, Joachim Ziegler, Marc De Meyer, Dan Bickel, Shane McEvey, David Yeates, Ashley Kirk-Spriggs, Robert Copeland and many other colleagues for their kind assistance.

• This study would be impossible without a great impact of Sergei J. Paramonov (1894-1967), who stimulated accumulation of the world largest collection in CSIRO (Australia).

Thank you for attention!

“Parasitic Fruit Flies”:Pyrgotidae, Ctenostylidae, Tachiniscidae

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