Valery Korneyev. 2008. Genera of the Old World Pyrgotidae

Genera of the Old World Pyrgotidae

(Diptera: Tephritoidea)with the special review of the fauna of the Oriental Region

Valery A. KorneyevI.I.Schmalhausen Institute of Zoology

National Academy of Sciences of Ukraine, Kiev

Catalogof the Family Pyrgotidae

of the World •Prepared in 2001-2004•592 records in species database•187 records in genera database•330 valid / 381 available species names•46 valid / 83 available genus names•ca. 330-340 species recognized in collections; many names are possible synonyms

The Catalog:Problems

•Species were poorly described (no essential characters)

•Most species unrecognizable w/o examination of types

•Concepts of genera extremely confused

•Generic position of almost all Old World species is obscure

The Catalog:Tasks

•Study of all available type material on the Old World Pyrgotidae (except Australian)

•Redescription. Figures. New characters

•Testing monophyly of genera

•Revision of diagnoses of genera

•and generic position of species

What the Pyrgotids are?

Photo by Mike Boone,with permission

What the Pyrgotids are?


•Pyrgotidae are the sister-group of the Tephritidae

•All pyrgotids are believed to be parasites of adult scarabs (Melolonthidae & Cetoniidae). Almost no data on biology

•No comprehensive diagnosis of the family. Monophyly?

Morphological charactersand their taxonomic significance


•Pyrgotis are the flies active in the dusk.

•Females are highly specialized for ovipositing into and for emergence of adults from adult scarab bodies:

•Diverse frictional structures (brushes, suckers) for attaching them firmly to beetle bodies during oviposition and to pierce abdominal sternites


• Head shape is believed to be an adaptation for emerging from the crusty beetle body (very large ptilinum => soft gena => movable face).

• Additionally sclerotized head skeleton: orbital plate=> parafacialium=> subocular plate

• Epiclypeal (epistomal) sclerite =secondarily sclerotized oral cavity membrane

• Strong occiput and deep occipital slit • Synapomorphy of the Pyrgotidae


• Occipital slit is present in all species, including the genera Descoleia, Nosferatumyia, Teretrura, Pyrgotosoma and Toxopyrgota

• Pyrgotids are rather uniform in coloration and wing pattern (mostly dusk activity)

• ~90% have no ocelli • R2+3 vein with stump vein (in ~60%)

• Adapsilia coarctata (Europe)

• Gelomyia sp. n. (Namibia)


• Euphya tripunctata. (Moluccas)

• Tephritopyrgota tripunctata. (Madagascal)


• Females in certain species have so-called femoral organ, a bare area on anterior surface of the midfemur, a structure of unknown function:

• either a sort of timpanal organ (Malloch 1931; Chen 1947)

• or merely a “gall” for better grip around the beetle body

• It is common in the subtribe Adapsilina; closely related species often differ in position, shape or absence of the f.o. Non-applicable in phylogenetic analysis (high homoplasy)

• Rare, but present in the subtribe Pyrgotina, but here it is small and probably non-homological?

•E

upyr

go

ta c

aff

ra (

Afr

ica

)

• Trichopeltia scutellina (Sri Lanka)


• Oviscape is strong, conical and non-compressible dorso-laterally(in ~97%)

• Rarely almost flat and soft (Teretrurinae, Descoleia, Toxopyrgota)

• Bulbous oviscape without eversible membrane ovipositor in Epice



• Pyrgotosoma flavida (Chile)

• Teretrura tinctipennis (Chile)

Epice sp. (Australia)


• Oviscape apex is different from that in Tephritidae

• In Pyrgotinae, eversible membrane is thick

• It bears 1 pair of lateral lobes with the field of trichoid and campaniform sensilla at base

• Medioventral lobe is taenia-like, unpaired, similar to that in the Tephritidae





• Mediodorsal lobe is sometimes very long (Adapsilia)

• often strongly reduced or absent• Adapsilia

coarctata (Europe)


• Pyrgotina tephritoides (Africa)

• Pyrgotomyia ciliata (Africa)


• Abdominal sternites 1 and 2 fused (except in Teretrura, Pyrgotosoma, Descoleia, and Nosferatumyia)

= Synapomorphy of Pyrgotinae?• In Toxopyrgota, sternites 1+2 are fused with visible seam• In Teretrura, Pyrgotosoma, Descoleia, and

Nosferatumyia sternites 1 & 2 are fusedwithout a seam

• Teretrura tinctipennis (Chile)

• Toxopyrgota inclinata (Africa)

• Tylotrypes immsi (Nepal)

• Females sometimes have densely spinulose abdominal tergites


• Eupyrgota echinata (Congo)

• Tephritopyrgota echinogaster (Madagascar)

• Campylocera kenyana (Africa)

• Trichopewltiascutellina (Sri Lanca)

• Or on the contrary, bare sterniteswith cellular surface


• Non- abdominal friction structures

• Foretrochanter with spinulae (Eupyrgota) • Mid– and hindcoxae with brushes of thick

setae (some Tephritopyrgota, Siridapha, and all Trichopeltia)

Trichopeltia mindanensis (Philippines)

Eupyrgota caffra (Aftica)


• Aculeus• Acute, stiletto-like, with sternite 8 reduced,

membranous• Setae reduced

(except in Prodalmanniapresent on tergite 8)

• Highly modified ovipositorin Epice spp.

Teretrura flaveola (Chile)

Eupyrgota latipennis(Africa)

Adapsilia coarctata (Europe)

Prodalmanniavariabilis (Australia)Epice sp. (Australia)


• Ventral receptacle • Not known in Descoleia and Nosferatumyia• With spherical apical chamber in ALL other

Pyrgotidae• Morula-like or snake-like petal in Teretrura

Teretrura flaveola (Chile)

Prodalmanniavariabilis (Australia)


Teretruratinctipennis (Chile)


• Male genitalia

• Epandrium without setae (most Pyrgotina, all Adapsilina) or with setae (Pyrgotinae: Toxurini, some Pyrgotini, Teretrurinae, Descoleia, Nosferatumyia)

• Surstyli without prensisetae (except in Descoleia, Nosferatumyia and Toxopyrgota)

Eupyrgota echinata (Africa)

Nosferatumyia no (Madeira)

Geloemyia trifasciata (Madagascar)

Descoleiateretrura (Argentina)

Pyrgotosoma flavida (Chile)

Tephritopyrgota microceras (Far East Palaearctics)


• Male genitalia

• Hypandrium with the phallic guide (absent in Descoleia, Nosferatumyia and Toxopyrgota)

• A synapomorphy of the Pyrgotinae?

Geloemyia trifasciata (Madagascar)

Classification of the Pyrgotidae

• Aczél (1956): Toxurinae Pyrgotinae (Teretrurini + Pyrgotini)

• D.K.McAlpine (1990):Teretrurinae(2 genera:Teretrura and

Pyrgotosoma)Pyrgotinae(Toxurini + Pyrgotini)

Classification of the Pyrgotidae

• Basal lineages:

DescoleiaNosferatumyiaTeretruraPyrgotosomaToxopyrgota

Pyrgotinae

Teretrurinae

Classification of the Pyrgotinae

• Basal lineages:

Prodalmanniini

Toxurini

Pyrgotini (Pyrgotina and Adapsilina)

Pyrgotinae

• Monophyly is well supported:

1. Abdominal sternites 1+2 fused2. Surstyli without prensisetae3. Phallic guide present4. Oviscape rigid 5. Face with sclerotized pentagonal area dorsal of

clypeus (epiclypeal sclerite or epistomal plate)6. Stump vein present (secondarily lost in some

species)

Pyrgotinae

• Monophyletic groups recognized:

1. Prodalmanniini (Australia)(Sc at acute angle P; ocelli present P; aculeus with long setulae P) 2 or more genera: Adapsona, Prodalmannia

2. Toxurini (Australia, New Guinea)(Sc bowed at 90° SA; ocelli present P; aculeus non-setulose A) 11 genera, > 60 species, ~10 undescribed (New Guinea)

Pyrgotinae


3. Pyrgotini (Sc at acute angle P; ocelli lacking A; aculeus non-setulose A; ocelli lacking A)

3.1. Pyrgotina (Old World, except Australia; New World)(no subocular sclerite, gena soft P; medioventral lobe (unpaired taenia) ending with hook, or ovipositungus (Steyskal 1978) SA; glans very simple, no neck, ring-like sclerite around gonopore SA) >60 species of 13 genera in Americas; >20 species in Madagascar; >35 in mainland Africa; >5 in Asia (Oriental and Palaearctic)

Madagascan fauna is represented only by species assigned to Tephritopyrgota, but these species are almost as diverse as American Pyrgotina.

Pyrgotinae


3.2. Adapsiliina (Old World, including Australia, not in Madagascar)(subocular sclerite strong A; medioventral lobe without a hook P; glans of more diverse and complex morphology, not as in Pyrgotina) >60 species of 13 genera in Americas; >20 species in Madagascar; >35 in mainland Africa; >5 in Asia (Oriental and Palaearctic)

PyrgotinaePyrgotinaeProdalmanniini and ToxuriniProdalmanniini and Toxurini

are strictly Australasian and not considered hereare strictly Australasian and not considered here

Subtribe Adapsiliina

• Geloemyia group of genera:3-4 dc, facial ridge reduced, R4+5 setulose, surstyli posteriorly produced, oviscape apex without lobes except ventral, aculeus spear-like

1. Geloemyia Hendel 1908= Trichempodia Malloch 1930 new synonym= Parageloemyia Hendel 1934 new synonym

= Dicranostira Enderlein 1942 new synonym

1.1. Geloemyia stylata Hendel 1908 Vietnam, Thailand, Malaysia = Trichempodia cockerelli Malloch 1930 new synonym

1.2. Geloemyia nigrofasciata (Hendel 1933) new combinationChina, Korea, Russia = Parageloemyia ornata Hering 1940

1.3. Geloemyia quadriseta (Hendel 1933) new combination China, Korea

1.4. Geloemyia trifasciata (Enderlein 1942) new combination Madagascar

1.5. Geloemyia globa (Shi 1996) new combination China

1.6. Geloemyia wonjuensis (Kim & Han 2001) new combination Korea, Russia

1.7. Geloemyia sp. 1 new species Namibia

Geloemyiatrifasciata (Madagascar)

Geloemyiasp. n. (Namibia)

Trichempodiacockerelli (Thailand)

Geloemyia stylata (Vietnam)

Subtribe Adapsiliina • Geloemyia group of genera:

2. Pyrgotomyia Hendel 1934= Metropina Enderlein 1942 new synonym

2.1. Pyrgotomyia ciliata Hendel 1934 Africa =Metropina richteri Steyskal 1977 new synonym

2.2. Pyrgotomyia temporalis (Enderlein 1942) new combination Africa

2.3. Pyrgotomyia sp. n. 1 Africa

• Hendelpyrgota Vanschuytbroek 19633.1. Hendelpyrgota longicornis Vanschuytbroek 1963 Africa

4. Porpomastix Enderlein 1942= Paradapsilia Chen 1947

4.1. Porpomastix foveolata Enderlein 1942 Far East Palaearctics, Indochina

4.2. Porpomastix sp. n. 1 India


• Adapsilia group of genera:1-2 dc, facial ridge usually well-developed, surstyli not posteriorly produced, oviscape apex with 4 pairs of lobes except ventral, aculeus widened at base

1. Adapsilia Waga 1843

1.1. Adapsila coarctata Waga 1843 Europe, Far East Asia = Adapsilia alini Hering 1940

1.2. Adapsila verrucifer Hendel 1933 China, Korea, Russia; Oriental Region= A. cornugaster Kim & Han 2001

1.3. Adapsila trypetoides Chen 1947 China, Oriental Region


• Adapsilia group of genera:

2. [Genus nova 1] (Adapsilia hirtoscutellata group)2.1. Campylocera myopina Wulp 1880 Indonesia: Java

2.2. Adapsilia magnicornis Hendel 1908 Sri Lanka

2.3. Adapsila hirtoscutellata Hendel 1933 China, Oriental Region

2.4. Adapsila myopoides Chen 1947 China, Oriental Region

2.4. Euthioza latigenis Enderlein 1942 Indonesia: Sumatra

2.4. Euthioza sumatrana Enderlein 1942 Indonesia: Sumatra

Euthioza latigenis (Sumatra)

Campylocera myopina (Java)

Adapsilia magnicornis (Sri Lanka)

Euthioza sumatrana (Sumatra)

Adapsilia hirtoscutellata (Sumatra)



3. Plectrobrachis Enderlein 19423.1. Plectrobrachis filigena Enderlein 1942 Indonesia: Sumatra

3.2. Plectrobrachis hispida (Kim & Han 2001) new combination Korea

3.3. sp. n. 1 Sri Lanka

3.4. sp. n. 2 Thailand

3.5. sp. n. 3 Philippines

3.6. sp. n. 4 Indonesia

Plectrobrachis filigena (Java)



4. Trichopeltia Enderlein 19424.1. Trichopeltia squalida (Walker 1861) comb. n. India; Turkey

= Adapsilia scutellata Hendel 1914

4.2. Trichopeltia longicornis (Wulp 1885) comb. n. Indonesia: Java

4.3. Trichopeltia scutellina (Hendel 1934) comb. n. India; Sri Lanka

4.4. Trichopeltia mindanensis (Hennig 1936) comb. n. Philippines

4.5. Trichopeltia penicillaticoxis Enderlein 1942 Indonesia: Sumatra; Malaysia (Peninsular)


Trichopeltia penicillaticoxis (Sumatra)

Trichopeltia longicornis (Java)




5. Campylocera Macquart 1843

• The largest genus in the group (>40 nominal species)

• Mostly in the Afrotropical Region

• Identity, including redescription of the type species

• Diagnostic characters: costa reduced between R4+5 and M; facial carina high; epiclypeal sclerite very low; male cerci usually fused

• Apparently monophyletic, though monophyly has not been rigorously tested

Campyloceraferruginea (Africa)



5. Campylocera Macquart 1843= Prosyrogaster Rondani 1875 new synonym = Hypotyphla Hendel 1908 new synonym= Prohypotyphla Hendel 1934 new synonym= Clemaxia Enderlein 1942 = Dicrostira Enderlein 1942 = Congopyrgota Vanschuytbroeck 1963 new synonym

Hypotyphla loewi (Africa)

Campylocera angustigennis (Africa)

Campyloceraferruginea (Africa)

Clemaxia angustigenis (Africa)

Congopyrgotahyalipennis (Africa)



5. Campylocera Macquart 1843Oriental species= Prosyrogaster Rondani 1875 new synonym = Teliophleps Enderlein 1942 new synonym

5.1. Campylocera chelionota (Rondani 1875) comb. n.

= Campylocera robusta Wulp 1880 new synonym

= Campylocera hirsuta Aldrich 1928 new synonym

= Teliophleps apicalis Enderlein 1942 new synonym

5.2. Campylocera thoracalis Hendel 1914

5.3. Campylocera rufina Bezzi 1916



6. Diasteneura Hendel 1908

6.1. Diasteneura laticeps Hendel 1908

6.2. Diasteneura variceps Curran 1928

6.3. Diasteneura similis Steyskal 1963

6.4. Diasteneura basilewskyi Vanschuytbroeck 1963

6.5. Diasteneura obscura Vanschuytbroeck 1963

6.6. Diasteneura sp. n. 1

6.7. Diasteneura sp. n. 2

Apparently also a synonym of Campylocera

Most species are known from holotype (either males or females only) and not dissected.


Eupyrgota vespiformis (Africa)

Eupyrgota sublatipennis (Africa)

Eupyrgota vespiformis (Africa) Eupyrgota

spinifemur (Africa)

Eupyrgota spinifemur (Africa)


7. Eupyrgota Coquillett 1900= Peltodasia Enderlein 1942 = Apyrgota Hendel 1908 new synonym = Acropyrgota Hendel 1914 new synonym

= Taeniomastix Enderlein 1942 new synonym



Eupyrgota Coquillett 1900• Two subgenera: Eupyrgota s. str. & Taeniomastix

with several groups of species

• Palaearctic and Afrotropical species have been revised recently(Kim & Han 2000; Korneyev 2006 a & b)

• 20 described and 5 undescribed species in Oriental andAustralasian Regions

Eupyrgota nocturna (India)

Eupyrgota sumatrana (Sumatra)

Eupyrgota (=Apyrgota) scioida (Moluccas)Eupyrgota

fusca (Japan)



8. Euphya Wulp 18858.1. Euphya tripunctata (Doleschall 1857) Indonesia: Moluccas

8.2. Euphya bistrigata Enderlein 1942 Indonesia: Java; Philippines= Euphya davisi Steyskal 1973 syn. n.

8.3. sp. n. 1 New Guinea

Euphya bistrigata (Java)

Euphya bistrigata (Java)

Euphya tripunctata (Moluccas)



• Siridapha Enderlein 19429.1. Siridapha longina (Hendel 1914) Africa: Tanzania

9.2. Siridapha filipalpis Enderlein 1942 Africa: Cameroun

9.3. Siridapha ophionea Enderlein 1942 Africa: Cameroun

9.4. Siridapha muluguensis Vanschuytbroeck 1963 Africa: Zaire

= Peltodasia mageraensis Vanschuytbroeck 1963 Africa: Zaire syn.n.

9.5. Siridapha sp. n. 1 Africa: Zaire

9.5. Siridapha sp. n. 2 Africa: Tanzania

Siridapha filipalpis (Cameroun)

Siridapha ophionea (Fernando Po)

Siridapha sp. n. (Tanzania)

Siridapha muluguensis (Zaire)

1. Tephritopyrgota Hendel 1914= Tephritocampyclocera Hendel 1934 new synonym

= Stypina Enderlein 1942

= Euthioza Enderlein 1942 new synonym

> 40 described species, most diverse fauna in Madagascar

Tephritopyrgota belzebuth (Africa)

Tephritopyrgota passerina (Africa)

Tephritopyrgota gowdei (Africa)

Tephritopyrgota carbonaria (Africa)

Subtribe Pyrgotina

Subtribe Pyrgotina

• Tephritopyrgota Hendel 1914 Madagascan species are almost as diverse as the rest of the Pyrgotina, but at least some were improperly placed

Tephritoprgota spinosa (Madagascar)

Tephritoprgota rutricauda (Madagascar)

Tephritoprgota rufonigra (Madagascar)

Tephritoprgota laevigaster (Madagascar)

Tephritopyrgota haltericauda (Africa)

Subtribe Pyrgotina

• Tephritopyrgota Hendel 1914 Oriental and East Palaearctic fauna are less diverse

9.1. Tephritopyrgota ferruginea (Walker 1914) India, Indochina, Korea= Adapsilia longicaudata Kim & Han 2001

9.2. Tephritopyrgota microceras (Portschinsky 1892) comb. n. Far East Russia, Korea

= Adapsilia tenebrosa Kim & Han 2001

9.3. Tephritopyrgota miliaria Hendel 1933 China: Szechwan

9.4. Tephritopyrgota yunnanensis Shi 1996 China: Yunnan

9.5. Tephritopyrgota sp. n.1 China: Taiwan

Tephritoprgota ferruginea (India)

Subtribe Pyrgotina

• Rhagostira Enderlein 1942= Tephritocampylocera Hendel pro parte major

= Tephritohypotyphla Vanschuytbroeck 1963

10.1. Rhagostira irrorata (Malloch 1929) comb.n. Kenya

10.2. Rhagostira breviseta (Hendel 1934) comb.n. Sudan

10.3. Rhagostira abessinica (Hennig 1937) comb.n. Ethiopia

10.4. Rhagostira erlangeri Enderlein 1942 Ethiopia

10.5. Rhagostira latigena Enderlein 1942 Tanzania

10.6. Rhagostira nana Enderlein 1942 Tanzania

10.7. Rhagostira biseta (Vanschuytbroeck 1963) comb.n. Zaire

10.8. Rhagostira masaiensis (Vanschuytbroeck 1963) comb.n. Tanzania

10.9. Rhagostira brincki (Vanschuytbroeck 1967) comb.n. RSARhagostyra latigena (Africa)

Rhagostira breviseta (Africa)

Rhagostyra erlangeri (Africa)

Subtribe Pyrgotina

• Tylotrypes Bezzi 1914

11.1. Tylotrypes immsi Bezzi 1914 India

11.2. Tylotrypes biseta (Shi 1996) comb. n. China: Beijing

11.3. Tylotrypes sp. n. 1 China: Taiwan

11.4. Tylotrypes sp. n. 2 China: Fujian

11.5. Tylotrypes sp. n. 3 Nepal

11.6. Tylotrypes sp. n. 4 Indonesia: Borneo

11.7. Tylotrypes sp. n. 5 Thailand

Tylotrypesimmsi (India)



Subtribe Pyrgotina


Pyrgotina tephritoides (South Africa)

Pyrgotina tephritoides (South Africa)

Pyrgotina turbata

(South Africa)

• Pyrgotina Malloch 1929= Platynostira Enderlein 1942 syn. n.

12.1. Pyrgotina muiri (Hendel 1914) RSA

12.2. Pyrgotina tephritoides (Hendel 1914) RSA

12.3. Pyrgotina tipuloides (Hendel 1914) RSA

12.4. Pyrgotina turbata (Enderlein 1942) comb. n. RSA

12.5. Pyrgotina antidorcas Bruggen 1961 RSA

12.6. Pyrgotina sp. n. 1 Tanzania

12.7. Pyrgotina sp. n. 2 Kenia

12.8. Pyrgotina sp. n. 3 Namibia

Pyrgotina tipuloides (South Africa)

Subfamily Toxopyrgotinae subfam. n.

• Toxopyrgota Hendel 1914

Toxopyrgota inclinata Hendel 1914 Tropical Africa

Aknowledgements


To all the curators of museums who kindly borrowed almost all the known types and material:

A.L.Norrbom, A.Kirk-Spriggs, D. Grimaldi, D. Notton, E. Marais, J.Cummings, N. Evenhuis, M. von Tschirnhaus, P.Limbourg, B. Merz, M.De Meyer, M. Mostovsky, K. van Achterberg, B.Brugge, J.Ziegler, L.Papp, R.Urban, H.-Y.Han, H.-P.Tschorsnig, P.Sehnal, M.Kotrba, W. Schacht, C.Daugerone, J.Stevington

Aknowledgements


•Work was supported by the Belgian Federal Office for Scientific, Technical and Cultural Affairs stipend in 2005-2006 and Short-Term Visitor Grant (Smithsonian Institution) in 2008 and by the National Academy of Sciences of Ukraine Program on Biodiversity in 2006-2010

Aknowledgements


•Special thanks :Alan Norrbom, Lucrecia Rodriguez and Gary Steck for organization of the meeting

Thank you Thank you for for

attention!attention!

Education

Valery Korneyev. 2008. Genera of the Old World Pyrgotidae