Tijdschrift voor Entomologie 157 (2014) 95–103 brill.com/tve

Tachinidae (Diptera) reared from Ropalidia nests (Hymenoptera: Vespidae) from Madagascar, with two new species of Anacamptomyia Theo Zeegers

This paper deals with the Tachinidae reared from Ropalidia nests collected on Madagascar. The material belongs to two genera, Anacamptomyia and Parapales,each represented by two species. Both species of Anacamptomyia are described as new: A. aurifrons sp. n. and A. blommersi sp. n. Their relation to the species from the Afrotropical mainland is discussed. The host records for Parapales are the first for this genus, supporting their placement in the Anacamptomyia group. Keywords: Diptera, Tachinidae, hosts, Ropalidia, Madagascar. Theo Zeegers, Eikenlaan 24, 3768 EV Soest, the Netherlands. [email protected]

Introduction Symmorphus Wesmael, 1836. However Hamanishi Tachinidae is one of the families of Diptera with the (1996) proved the actual host to be not Symmorphus, highest number of genera and species (Ziegler 2003, but beetle larvae preyed by Symmorphus. O’Hara 2012, 2013). Tachinid flies are well known Crosskey (1976) placed the genera Anacampto- for their biology. The larvae develop as parasitoids myia, Koralliomyia and Euvespivora in a separate tribe in insects and in a few other arthropods, caterpillars Anacamptomyiini, apparently considering them to of Lepidoptera being the most numerous (Herting constitute a monophyletic group. Mesnil (1977) 1960, Ferrar 1987). The ecology and evolutionary added Parapales to this tribe. Because the monophyly history of the family haven been reviewed by Stire- of this tribe needs substantiation, I here refer to this man et al. (2006). Our knowledge of the phylogeny group as the Anacamptomyia group. Mesnil (1977) of the family has been improved by the usage of added the genus Parapales Mesnil, 1950 to this group recent molecular techniques (Cerretti et al. 2014). without knowledge of its host. Some Tachinidae are of economic importance, since During his visits to Madagascar in the years 1970– their hosts are considered pests (O’Hara 2007). 1973, Leo Blommers made an extensive study of Vespidae, or more general Hymenoptera Aculeata, the nests of vespid wasps belonging to the genus are exceptional as host for Tachinidae (Ferrar 1987). Ropalidia Guérin-Méneville (Blommers 2012). In Vespidae have been recorded as host for only five of this process, he reared several Tachinidae belonging the 1519 currently recognized genera of Tachinidae to two genera: Anacamptomyia and Parapales.This (O’Hara 2012): Anacamptomyia Bischof, 1904, Eu- contribution aims to describe the Tachinidae reared vespivora Baranov, 1942, Koralliomyia Mesnil, 1950 by Blommers from Ropalidia nests from Madagascar. in the Old World (Crosskey 1973, 1976, 1984, In the genus Anacamptomyia, two different species Wood & Zumbado 2010) and Lixophaga Townsend, have been found, both described as new here. Also in 1908 and Ophirion Townsend, 1911 in the New the genus Parapales, two species have been found. World (Curran 1937, Wood 1985). Mesnil & Shima (1977) reared Symmorphomyia from the nests of

Tijdschrift voor Entomologie 157: 95–103, Figs 1–8. [ISSN 0040-7496]. brill.com/tve © Nederlandse Entomologische Vereniging. Published by Koninklijke Brill NV, Leiden. Published 20 November 2014. DOI 10.1163/22119434-00002041

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Material and methods the flat occiput without black hairs posterior to pos- Backgrounds on hosts, nests and collecting tech- tocular row, the large calypter with angular inner niques have been extensively described by Blom- margin, the crossed erect apical scutellar setae and mers (2012). In summary, he collected many nests the presence in the male of a pair of patches of spe- of Ropalidia on Madagascar in the period 1970– cialized setae on the ventral side of both abdominal 1975. He killed the wasps with a short exposure of tergites 3 and 4 and the presence of the regular comb ethyl acetate, thus deliberately leaving the present of anterodorsal setae on the hind tibia. With the ex- pupae alive. He kept the nest to see if any more ception of the males of Parapales, the frontal setae are wasps or parasitoids would emerge. In this process, mostly reclinate and thus not well separated from the he reared some Tachinidae. Blommers (2012) men- reclinate orbital setae. tioned those belonging to the genus Other material, Mesnil (1977) provides a key to separate the two belonging to the genus Parapales, became available genera. Based on the material studied here, his key after Blommers’ manuscript was accepted for pub- needs to be revised, given the variability of some lication, therefore, it was not mentioned in Blom- features. To separate both genera, I propose the mers (2012). The nests from which the Parapales following key. have been reared, though, are discussed. Strictly speaking, Blommers (2012) did not estab- lish Ropalidia as host; he merely reared the tachinid Key to genera of Afrotropical flies from the hosts nests. Anacamptomyia group The terminology for the body parts used follows 1. Frontal setae and reclinate orbital setae Tschorsnig & Richter (1998) and Merz & Haenni irregularly placed in several rows. Genal (1998). Photos of the habitus of flies have been dilation reduced, consisting of 1–3 rows taken with a Nikon 105 mm macro lens with macro of hairs. Proepimeral seta absent. Synter- flashes; the results have been digitally enhanced. gite 1 + 2 with a pair of strong marginal Photos of heads have been made by using a separate setae. Male: Outer vertical seta present . . . . phototube on the stereomicroscope. Focal depth has ...... Anacamptomyia been enhanced by stacking several images using the – Frontal setae and reclinate orbital setae software program CombineZ (Hadley). placed in one row. Genal dilation well de- The acronyms for collections follow Evenhuis veloped, with about 6 rows of hairs present. (2012), repeated here for convenience. Proepimeral seta present, though small. CNC = Canada, Ontario, Ottawa, Canadian Syntergite 1 + 2 without differentiated National Collection of Insects; marginal setae. Male: Outer vertical seta ab- CTZS = Private collection Th. Zeegers, Soest, sentorhair-like...... Parapales the Netherlands; MNHN = France, Paris, Muséum National d’Histoire Naturelle; Genus Anacamptomyia Bischof, 1904 MZUR = Italy, Roma, Museo di Zoologia, Uni- Anacamptomyia Bischof, 1904: 79. Type species: versità degli Studi di Roma “La Anacamptomyia africana Bischof, 1904, by mono- Sapienza”; typy; Mesnil 1950b: 22–24 [key to Afrotropical NMW = Austria, Wien, Naturhistorisches species]. Museum Wien; Roubaudia Villeneuve, 1910: 249. Type species: RMNH = Netherlands, Leiden, Naturalis Bio- Roubaudia rufescens Villeneuve, 1910, by origi- diversity Center. nal designation; Mesnil 1950b: 22 [as synonym]; Holotypes and some paratypes of both sexes of the Mesnil 1977: 190 [as valid genus]. new species have been deposited in RMNH. Pararoubaudia Roubaud & Villeneuve, 1914: 124; as On the variant names and spellings for cities on subgenus of Roubaudia. Type species: Roubaudia Madagascar, consult Blommers (2012): 135. (Pararoubaudia) bisetosa Roubaud & Villeneuve, 1914, by monotypy; Mesnil 1950b: 22 [as syn- onym]. Results The material found belongs to two genera: Anacamp- Remarks. The genus Anacamptomyia differs from tomyia and Parapales. These genera are the members most other Tachinidae by its broad fronto-orbital of the Anacamptomyia group in the Afrotropical re- plate and very narrow frontal vitta. These features gion (Mesnil 1977). They share many features, such are seen in other genera of the Anacamptomyia group as the reduced or absent ocellar seta, a row of strong as well. The genus occurs in the Afrotropical and setae covering the facial ridge for most of its length, the Australian region (O’Hara 2012). Seven species

Downloaded from Brill.com09/29/2021 10:10:07PM via free access Zeegers: Tachinidae reared from Ropalidia nests from Madagascar 97 have been described from the Afrotropical region frontal orbital plate descending downwards (Crosskey 1980, Zeegers 2007). The latest full re- beyond on parafacial below lowest frontal view is by Mesnil (1950b). From Madagascar, only seta. Wing: base of vein R4+5 above with A. africana has previously been recorded (Crosskey only 1 setula. Male: proclinate orbital seta 1980). This study reports two new species from absent.[Afrotropicalmainland]...... Madagascar, both closely related to A. africana.The ...... africana Bischof, 1904 old records of A. africana from Madagascar might – Mid tibia with 1 very strong anterodor- refer to one of these species. Therefore, the literature sal seta only. Hairs on frontal orbital records need to be reevaluated. plate scarce, not descending downwards on Mesnil (1977) defines the genus Roubaudia based parafacial below lowest frontal seta. Wing: on the presence of median discal setae on abdomi- base of vein R4+5 above usually with 2–3 nal tergites, though Mesnil (1950b) cited it in syn- setulae. Male: at least 2 pairs of strong pro- onymy with Anacamptomyia. This synonymy is fol- clinate orbital setae present [Madagascar] lowed by Crosskey (1980, 1984) and O’Hara (2012)...... → 4 Zeegers (2007) described a species of Anacamptomyia 4. Fronto-orbital plate ochraceously yellow. in which median discal setae are either present or ab- First flagellomere yellow, distinctly dark- sent. Therefore, Roubaudia is treated as a synonym ened at tip; arista darkened. Abdominal ter- of Anacamptomyia. gites with silvery pruinescence restricted to The new material from Madagascar clearly con- anterior half. Eye with very long hairs (each sists of two species. Both lack median discal setae hairabout5–6timesaslongasthediameter on abdominal tergites, thus resembling A. africana of an ommatidium). Male: hairs on abdom- (the only described species with this feature and hairy inal tergites 3 and 4 on central axis much eyes). Comparison with the type of africana showed longer and more erect than those as sides. neither of them to be conspecific: the males are easily Lowest proclinate orbital seta about twice separated by the presence of proclinate orbital setae. as strong as the one above...... aurifrons sp. n. This is in line with the general experience that en- – Fronto-orbital plate silvery white. First flag- demism is rule rather than exception on Madagascar ellomere yellow, tip not darkened; arista (Irwin et al. 2003, Paulian & Viette 2003). brown or yellow. Abdominal tergites uni- As a consequence, both species are described here formly covered with silvery white pruines- as new. A diagnostic key to the species of Madagascar cence. Eye with shorter hairs (each hair with reference to the other Afrotropical species is about3timesaslongasthediameterof presented, followed by a description of the new an ommatidium). Male: hairs on abdominal species. A short redescription of A. africana,basedon tergites3and4adpressed,onlyonthecen- a reexamination of the male paralectotypes, is given. tral axis of abdominal tergite 4 before the marginal setae slightly erect. Lowest procli- nate orbital seta about as long as the one Key to species of Anacamptomyia from above...... blommersi sp. n. Madagascar with reference to Afrotropical species Anacamptomyia aurifrons sp. n. 1. At least abdominal tergite 3 without me- Anacamptomyia spec.: Blommers 2012: 143, 150, 188. dian discal setae. Afrotropical mainland, Figs 1–4 YemenorMadagascar...... → 2  – Abdominal tergite 3 with distinct median Type material. Holotype : Madagascar [Rép. Mal- discal setae. Five species from Afrotropical gache]: Ambatolampy, Ankaratra Mts., Manjaka- mainlandandonefromYemen...... tompo, 2000 m, 21.iii.1973, reared ex nest 73.22 ...... nottreated Ropalidia variabilis (de Saussure, 1890), leg. R. & here; see Mesnil (1950b) & Zeegers (2007) L. Blommers (RMNH).  2.Eyebare(atcloseinspectionsparselyand Paratypes. Madagascar [Rép. Malgache]: :as short-haired). Fore and mid tibiae with 2 holotype, but reared ex nest 73.22/24, same host  posterior setae . . . gymnops Zeegers, 2007 [part] (CTZS); 2 : as holotype, but reared ex nest 73.24,    – Eye distinctly hairy. Fore and usually mid same host (CTZS); 1 :same(RMNH);1 &1 : tibiae with 3–4 posterior setae ...... → 3 same, but altitude 1750 m, 5.xi.1972, reared ex nest  3. Mid tibia with 1–2 weaker anterodorsal 72.41, same host (RMNH); 1 with pupa: Tana- setae additional to the strong anterodorsal narive, 1300 m, 22.xi.1970, reared ex nest 70.11 one, which is about twice as long. Hairs on Ropalidia grandidieri (de Saussure, 1890), leg. L. & R. Blommers (CTZS).

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Figs 1–8. Anacamptomyia aurifrons sp. n., male (Madagascar): (1) habitus, dorsal (holotype); (2) head and thorax, lateral; (3) head, dorsal; (4) abdomen, latero-dorsal. Anacamptomyia blommersi sp. n. male (Madagascar): (5) habitus, dorsal (holotype); (6) head and thorax, lateral; (7) head, dorsal; (8) abdomen, latero-dorsal.

Description. Length 5.5–7.5 mm. est frontal seta. Antenna yellow, apical third of first Male. Colour. General colouration yellow and red- flagellomere contrastingly blackish; arista dark. Pal- dish, as usual in the genus Anacamptomyia.Fronto- pus yellow. Thoracic dorsum reddish brown with 4 orbital plate, frontal vitta and lunula ochraceous yel- narrow black vittae; postpronotum yellow. Scutellum low. Colour of parafacial changing with angle of view yellow. Legs yellow. Wings hyaline; tegula and basi- from ochraceous to silvery white: seen obliquely from costa yellow. Abdomen red, abdominal tergites with front white (sharply contrasting with fronto-orbital silvery diffuse pruinescence on anterior half, some- plate), in frontal view ochraceous as well. Border what more intense to lateral margin of abdominal between the two areas sharp, just below the low- tergite.

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Head. Eye with very long hairs, about 5–6 times as nal tergites, however, hairs on central axis longer and long as the diameter of an ommatidium. Vertex at its sub-erect, seta-like especially on abdominal tergite 4. narrowest point about 0.8 times width of one eye. Abdominal tergite 5 with a row of marginal and dis- Fronto-orbital plate very broad, frontal vitta narrow, cal setae. Ventral surface of abdominal tergites 3 and about as wide as anterior ocellus. Fronto-orbital plate 4 with a pair of more or less circular patches of spe- with 2–3 irregular rows of reclinate setae (frontal cialized setae (‘Sturmia-spot’), approaching but not setae and reclinate orbital setae), outwards of these 2 reaching anterior and posterior margins of abdomi- pairs of proclinate orbital setae, the lowest one about nal tergites. twice as strong as the one above. 1–2 pairs of frontal Female. Very similar to the male, sexual dimorphism setae above the antennal base crossed. Ocellar seta very small. Female differs in the following points: absent, inner vertical seta strong, outer vertical seta proclinate orbital setae of the same size. Comb of about half as strong as inner; one pair of postocellar anterodorsal setae on hind tibia less dense. Hairs setulae. Facial ridge nearly completely covered with on abdominal tergite 3 and 4 shorter than in male a row of strong setae. Parafacial narrow, at most and adpressed. Ventral surface of abdominal tergites half as wide as first flagellomere, bare below lowest without patches of specialized setae. frontal seta. Gena narrow, narrower than width of parafacial at level of antennal base. Genal dilation Variability reduced, with one row of strong setae and 1–2 rows Very little. of hairs only. Occiput with only white hairs behind postocular row. First flagellomere about 4 times as long as broad and as long as pedicel. Arista thickened Remarks at basal 1/4–1/5. Anacamptomyia aurifrons is characterized by ochra- ceously yellow fronto-orbital plate and the distinc- Thorax. Scutum with 3 + 3 acrostichal setae, 3 + 4 dorsocentral setae, 1 + 3 intra-alar setae, 3 supra- tive black tip of postpedicel. The pruinescence on alar setae, the anterior one strong. Postpronotum abdominal tergites 3–4 is restricted to the anterior with 4 setae, the basal 3 placed in a straight line. part. The hairs on the eye are long, as in A. africana. Notopleuron with 2 setae. Proepimeral seta absent, Male with 2 pairs of proclinate orbital setae, of proepisternal seta strong, often doubled by a smaller which the lower one is much larger than the upper one. Anepisternum with, in addition to the row of one and the hairs on central axis of abdominal anepisternal setae, a seta in the anterodorsal corner, tergites 3–4 are sub-erect. just below the anterior notopleural seta. One strong anepimeral seta present; katepimeron bare. Proster- Biology num with a few strong setulae. Scutellum with 4 Reared from nests of Ropalidia variabilis and R. pairs of marginal setae, basal and subapical strongest, grandidieri.TherecordofR. galimatia as a host apical erect and crossed. (Blommers 2012: 188) is incorrect: this refers to the Wing with cell r4+5 open, ending close to wing next species. tip (sixth section of costa half as long as fourth); apical section of vein M concave, shortest distance Etymology from bend in vein M to hind margin of wing half The name aurifrons is derived from Latin, meaning as long as the distance between bend and crossvein ‘golden frons’. It should be treated as a noun. dm-cu. Base of vein R4+5 with 2–3 setulae. Lower calypter large with its lateral margin convex and bent Anacamptomyia blommersi sp. n. ventrally. Anacamptomyia spec.: Blommers 2012: 141, 156, 188. Frontal tarsus with short claws and pulvilli. Fore tibia with 3 posterior setae; mid tibia with 1 very Figs 5–8 strong anterodorsal seta, a strong ventral seta and 3–4 Type material. Holotype : Madagascar [Rép. Mal- smaller posterior setae; hind tibia with a regular row gache]: Tananarive, 1300 m, 20.iv.1972, reared ex of anterodorsal setae and a nearly regular posterodor- nest 72.15 Ropalidia merina Blommers, 2012, leg. L. sal row, 3–4 smaller ventral setae; 2 dorsal praeapical & R. Blommers (RMNH). setae present (anterodorsal and posterodorsal). Hind Paratypes. Madagascar [Rép. Malgache]:3 as coxa bare on posterior surface. holotype (RMNH); 2  as holotype (CTZS); 2  & Abdomen. Excavation of abdominal syntergite 1 + 2 : Tuléar, 18.v.1971, reared ex nest 71.24 Ropalidia reaching hind margin. Syntergite 1 + 2andabdom- galimatia (de Saussure, 1890), leg. L. & R. Blom- inal tergite 3 each with a pair of median marginal mers (CTZS); 1  &1: Tananarive, 1300 m, setae; abdominal tergite 4 with a row of marginal 13.ii.1971, reared ex nest 71.9 Ropalidia shestakowi setae; median discal setae absent on these abdomi- (von Schulthess, 1931), leg. L. & R. Blommers

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(RMNH); 1 : same, but date 16.ii.1972 (CTZS); second, smaller one; katepimeron bare. Prosternum  (form with yellow arista): same, but date 9.ii.1972 with a few strong setulae. Scutellum with 4–5 pairs (RMNH); 1  (form with yellow arista): same, but of marginal setae, basal and subapical strongest, api- date 11.ii.1972 (RMNH); 1  (formwithyellow cal erect and crossed, lateral ones usually double. arista): Nosy Bé, dunes, Angorombalo, 8.i.1972, Wing with cell r4+5 open, ending close to wing ex nest 72.4 Ropalidia shestakowi (von Schulthess, tip (sixth section of costa half as long as fourth); 1931), leg. R. & L. Blommers (CTZS). apical section of vein M concave, shortest distance Description. Length 5.5–7.5 mm. from bend in vein M to hind margin of wing half Male. Colour. General colouration yellow and red- as long as the distance between bend and crossvein dish, as usual in the genus Anacamptomyia.Fronto- dm-cu. Base of vein R4+5 with 1–3 setulae. Lower orbital plate and lunula silvery white, frontal vitta calypter large with its lateral margin convex and bent reddish or brownish. Colour of parafacial changing ventrally. from light yellow to silvery white with angle of view: Frontal tarsus with short claws and pulvilli. Fore seen obliquely from front silvery white (as fronto- tibia with 3 posterior setae; mid tibia with 1 very orbital plate), in frontal view light yellow as well. An- strong anterodorsal seta, a strong ventral seta and 2– tenna yellow, arista brown; but in 3 specimens (see at 4 smaller posterior setae; hind tibia with a regular ‘Variation’), the arista is light yellow. Palpus yellow. but little dense row of anterodorsal setae and a nearly Thoracic dorsum reddish brown with silvery pru- regular posterodorsal row, 3–4 smaller ventral setae; inescence leaving 4 narrow black vittae; postprono- 2 dorsal praeapical setae present (anterodorsal and tum yellow. Scutellum yellow, with more or less sil- posterodorsal). Hind coxa bare on posterior surface. very pruinescence. Legs yellow. Wings hyaline; tegula Abdomen. Excavation of abdominal syntergite 1 + 2 and basicosta yellow. Abdomen red, abdominal ter- reaching hind margin. Syntergite 1 + 2andabdom- gites uniformly covered with silvery pruinescence. inal tergite 3 each with a pair of median marginal Head. Eye with shorter hairs, about 3 times as long setae; abdominal tergite 4 with a row of marginal se- as long as the diameter of an ommatidium. Vertex tae; median discal setae absent on these abdominal at its narrowest point about 0.8 times as broad as tergites. Abdominal tergite 5 with a row of marginal width of one eye. Fronto-orbital plate very broad, and discal setae. Ventral surface of abdominal tergites leaving a narrow frontal vitta, its width at narrow- 3 and 4 with a pair of rectangular patches of spe- est point only slightly wider than anterior ocellus. cialized hairs (‘Sturmia-spot’), reaching both anterior Fronto-orbital plate with 2–3 irregular rows of recli- and posterior margin of abdominal tergite. nate setae (frontal setae and reclinate orbital setae), Female. Very similar to the male, sexual dimorphism outwards of these 2–3 pairs of proclinate orbital se- very small. Female differs in the following points: tae, the lowest one about as long as the one above. always two pairs of proclinate orbital setae (of similar 1–2 pairs of frontal setae above the antennal base size). Comb of anterodorsal setae on hind tibia less crossed. Ocellar seta absent, inner vertical seta strong, dense. Ventral surface of abdominal tergites without outer vertical seta about half as strong as inner; one patches of specialized setae. pair of postocellar setulae. Facial ridge nearly com- pletely covered with a row of strong setae. Parafacial Variability narrow, at most half as broad as width of first flagel- In some specimens, there is only 1 setula at the base lomere, bare below lowest frontal seta. Gena narrow, of vein R4+5. Three specimens represent a form with narrower than width of parafacial at level of antennal light yellow arista. The shape of the Sturmia-spot base. Genal dilation reduced, with one row of strong in the male is slightly varying. Some males have an setae and 1–2 rows of hairs only. Occiput with only additional smaller third upper proclinate orbital seta white hairs behind postocular row. First flagellomere present. about4timesaslongasbroadandaslongaspedicel. Arista thickened at basal 1/4–1/5. Thorax. Scutum with 3 + 3 acrostichal setae, 3 + 4 Remarks dorsocentral setae, 1 + 3 intra-alar setae, 3 supra- Anacamptomyia blommersi is similar to A. africana in alar setae, the anterior one strong. Postpronotum the colouration of the fronto-orbital plate and the with 4 setae, the basal 3 placed in a straight line. abdomen. The hairs on eye are shorter than in the Notopleuron with 2 setae. Proepimeral seta absent, other species treated. The male is characterized by proepisternal seta strong, often doubled by a smaller the presence of 2–3 pairs of proclinate orbitals setae, one. Anepisternum with, in addition to the row of of which the lower 2 are similar in size. The female anepisternal setae, a seta in the anterodorsal cor- differs from A. africana bythepresenceofonly1 ner, just below the anterior notopleural seta. One anterodorsal seta on the mid tibia and from aurifrons strong anepimeral seta present, often doubled by a by the yellow first flagellomere.

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Biology to generic level]; Mesnil (1977) [raise to generic Reared from nests of Ropalidia galimatia, R. merina, level; key to species]; Evenhuis & O’Hara (2008). R. shestakowi. Parapales was described by Mesnil (1950a) as sub- genus of Ctenophorocera [currently: Pales Robineau- Etymology Desvoidy, 1830 (O’Hara, 2012)]. In the original de- This species is dedicated to the collector of this new scription of the subgenus Parapales, a second species species, Leo Blommers. The name should be treated was included, P. s t u r m i o i d e s from the Oriental re- as the genitive of a noun. gion (Mesnil 1950a). Mesnil (1963) raised Para- pales to generic level and added erebiae Mesnil, Anacamptomyia africana Bischof, 1904 1963 from the eastern Palaearctic and laevis Vil- Anacamptomyia africana: Bischof, 1904: 81. Lecto- leneuve, 1932, pilosa Villeneuve, 1942 and pacta type female, South Africa (NMW) [not seen]. Villeneuve, 1932 from the Afrotropical region to Paralectotypes 2 : South Africa: Capland, Algoa this genus. Later, Mesnil (1977) associated Parapales bay, 6.ii.[18]96, leg. Dr. Brauns (NMW) [seen]. with Anacamptomyiini, removed sturmioides from Anacamptomyia africana: Townsend (1940) [designa- Parapales and transferred erebiae to Pseudoperichaeta tion female as lectotype], Mesnil 1950b [key to Brauer & Bergenstamm, 1889. Crosskey (1980) Afrotropical species]. placed the three Afrotropical species mentioned in Pseudoperichaeta as well. We concur with the latter views. Sturmioides is still unplaced but is not closely Redescription related to either Parapales or Pales. As a consequence, To the description by Mesnil (1950b), I add the the genus Parapales is endemic to Madagascar, cur- following. First flagellomere yellow with distinctly or rently represented by six known species. indistinctly darkened apex. Arista brown. Parafacial Diagnosis. The genus Parapales as currently under- and frontal orbital plate white, slightly more yellow stood resembles Anacamptomyia in many aspects, in some angles of view. Abdomen reddish with a such as the reduced or absent ocellar seta, the fa- distinct central dark grey vitta, as wide as the distance cial ridge covered with a row of setae for most of its between median marginal setae on abdominal tergite length, the flat occiput without black hairs behind 3, dorsal surface of abdominal tergites completely postocular row, the large calypter with angular inner covered with non-dense silvery pruinescence, only margin, the crossed erect apical scutellar setae and denser at the anterior margin of abdominal tergites. the presence in the male of a pair of patches of spe- This pruinescence is lacking on ventral surface of the cialized setae on the ventral side of both abdominal abdominal tergites. tergites 3 and 4 and the presence of the regular comb Eye with long hairs, length of one hair about 6 of anterodorsal setae on the hind tibia. The orienta- times as long as the diameter of an ommatidium. tion of the frontal setae is sexually dimorphic. In the Frontal orbital plate densely haired, these hairs de- male, those are crossed; in the female, reclinate as in scending on parafacial below lower frontal seta. Two Anacamptomyia. pairs of proclinate orbital setae present, the lower one Remarks. According to Mesnil (1977), Parapales dif- less or equal in size than the upper one. Scutellum fers from Anacamptomyia among others by the nar- with 4 pairs of marginal setae. Front tibia with 4 pos- rower vertex and by the broader frontal vitta. Both terior setae, mid tibia with 1–2 weaker anterodor- features, however, display quite some variation in sal setae additional to the strong anterodorsal one. Parapales and one could imagine a gradual transition The distance of vein M between crossvein dm-cu towards Anacamptomyia. Therefore, I introduce sev- and bend only slightly longer than the distance from eral new features in the key above to define Parapales. bend to hind margin of wing. Vein R4+5 reaching wing margin far from apex of wing, therefore the Parapales pallidula (Mesnil, 1950) th th combined length of 5 and 6 costal section hardly Ctenophorocera (Parapales) pallidula Mesnil, 1950a: less than length of 4th. Hairs on abdominal tergites 122–123. Holotype : Madagascar, Bekily, iii. 3and4shortandadpressed. 1930, [leg.] A. Seyrig (CNC) [not seen]. Parapales pallidula: Mesnil, 1977. Material. Madagascar [Rép. Malgache]:1,Tsara- Genus Parapales Mesnil, 1950 mandroso, Ampijoroa, 1.v.1972, leg. L. & R. Blom- Parapales Mesnil, 1950a: 122 [as subgenus of Cteno- mers, ex nest 72.26 Ropalidia flavoviridis Kojima, phorocera Brauer & Bergenstamm, 1891]. Type 1988 (CTZS). species: Parapales pallidula Mesnil, 1950, by orig- inal designation; Mesnil (1963) [implicit raise

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Parapales aff. micronychia Mesnil, 1977 the manuscript and provided relevant information Parapales micronychia Mesnil, 1977: 191. Holotype on the lectotype of Anacamptomyia africana and : Madagascar, Joffreville, 11.v.1958 (MNHN) the holotype of Parapales micronychia. Peter Sehnal [not seen]. (NMW) was so kind to loan me the paralectotypes Material. Madagascar [Rép. Malgache]:1 &1, of Anacamptomyia africana. James O’Hara (Ottawa, Mananjary, 14.i.1973, leg. L. & R. Blommers, ex Canada) kindly helped me out with the New World nest 73.1 Ropalidia carinata (de Saussure, 1890) tachinid parasitoids with vespid hosts. (MZUR); 1  &2,same(CTZS). This species resembles, but might be not identi- cal, with P. micronychia. Dr. Cerretti (Rome), who References compared a couple of the Blommers-material with Baranov, N., 1942. Ein neuer Vespidenparasit von Java und the holotype of micronychia, kindly informed me (in eine mit ihm verwante Fliege von den Salomon-Inseln. litt.) that he concurs with this opinion. Therefore, I – Veterinaryski Archiv 12: 161–163. report the species as aff. micronychia. Bischof, J., 1904. Beitrag zur Kenntnis der Mus- caria schizometopa. – Verhandlungen der Kaiserlich- Königlichen Zoologisch-Botanischen gesellschaft in Discussion and conclusions Wien 54: 79–101. This article reports the Tachinidae reared from Ropa- Blommers, L.H.M., 2012. Taxonomy and natural history of 18 Ropalidia species (Hymenoptera, Vespidae) of lidia nests from Madagascar by Blommers. Two Madagascar. – Tijdschrift voor Entomologie 155: 133– species belonging to Anacamptomyia and two belong- 192. ing to Parapales have been found, both genera be- Cerretti, P., J.E. O’Hara, D.M. Wood, H. Shima, D.J. In- longing to the Anacamptomyia group in the tribe clan & J.O. Stireman III, 2014. Signal through the Eryciini. noise? Phylogeny of the Tachinidae (Diptera) as in- The hosts of members of this group are Vespi- ferred from morphological evidence. – Systematic En- dae (Hymenoptera Aculeata), which is unusual in Ta- tomology 39: 335–353. chinidae. This feature is a candidate synapomorphy. Crosskey, R.W., 1973. A conspectus of the Tachinidae In the literature, the genera Belonogaster Saussure, (Diptera) of Australia, including keys to the supraspe- 1845, Euvespivora Baranov, 1942, Polistes Latreille, cific taxa and taxonomic and host catalogues. – Bulletin 1802 and Ropalidia are recorded as hosts (Crosskey of the British Museum (Natural History) Entomology, 1973, 1976, 1984, Shima 2006). All these Vespid Supplement 21: 1–221. genera are social or quasi-social wasps of the sub- Crosskey, R.W., 1976. A taxonomic conspectus of the family Polistinae. The only exception is the record Tachinidae (Diptera) of the Oriental region. – Bulletin by Baranov (1942) of Vespa analis Fabricius, 1775 as of the British Museum (Natural History) Entomology, ahostforEuvespivora orientalis Baranov, 1942 from Supplement 26: 1–357. Crosskey, R.W., 1980. Family Tachinidae. – In: the Oriental region (Crosskey 1976). This study es- R.W. Crosskey (ed.), Catalogue of the Diptera of tablishes the vespid genus Ropalidia as host for Para- the Afrotropical Region. British Museum (Natural pales and thus supports the placement of Parapales in History), London, pp. 822–882. the Anacamptomyia group of Erycini. Crosskey, R.W., 1984. Annotated keys to the genera of In this study, only Ropalidia nests have been in- Tachinidae (Diptera) found in tropical and southern vestigated. Therefore, no conclusions can be drawn Africa. – Annals of the Natal Museum 26: 189–337. on other genera of Vespidae as possible hosts. No Curran, C.H., 1937. Two new Tachinidae (Diptera) par- overlap has been found in hosts between the differ- asitic on Polybia species (Hymenoptera). – American ent species: Anacamptomyia aurifrons was reared from Museum Novitates 923: 1–4. Ropalidia grandidieri and R. variabilis; A. blommersi Evenhuis, N.L. & J.E. O’Hara, 2008. The status of Mes- from R. galimatia, R. merina and R. shestakowi; Para- nil’s 1949 Die Fliegen genus-group names (Diptera: Ta- pales pallidula from R. flavoviridis and P. aff. microny- chinidae). – Zootaxa 1827: 65–68. chia from R. carinata. Due to the small numbers of Evenhuis, N.L., 2012. The insect and spider collections Tachinidae found, no conclusions can be drawn on of the world website. http://hbs.bishopmuseum.org/ the exclusiveness of these relationships. codens [assessed on: 8 November 2012]. Ferrar, P., 1987. A Guide to the Breeding Habits and Im- mature Stages of Diptera Cyclorrhapha. Entomograph Acknowledgements 8. – E.J. Brill, Leiden, 907 pp. Hadley, A. CombineZ [software program]. http://www. First and foremost, I would like to thank Leo Blom- hadleyweb.pwp.blueyonder.co.uk/index.htm [assessed mers introducing me into his Ropalidia project and on: 8 November 2012]. for the pleasant cooperation. Pierfilippo Cerretti made valuable comments to an earlier version of

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Hamanishi, Y., 1996. Cleptoparasitic life of the tachinid Paulian, R. & P.Viette, 2003. An introduction to terrestrial fly, Symmorphomyia katayamai Mesnil et Shima, in the and freshwater invertebrates. – In: S.M. Goodman & nest of host wasps hunting the chrysomelid-larva prey. J.P.Benstead (eds), The Natural History of Madagascar. – Japanese Journal of Entomology 64: 843–860. University of Chicago Press, Chicago, pp. 503–511. Herting, B., 1960. Biologie der westpaläarktischen Rau- Roubaud, E. & J. Villeneuve, 1914. Contribution à l’étude penfliegen Dipt., Tachinidae. – Monographien zur des espèces du genre Anacamptomyia Bischof (Dipt.). – Angewandten Entomologie 16: 1–188. Revue Zoologique Africaine 4: 121–128. Irwin, M.E., E.I. Schlinger & F.C. Thompson, 2003. Shima, H., 2006. A host-parasite catalog of Tachinidae Diptera, true flies. – In: S.M. Goodman & J.P.Benstead (Diptera) of Japan. – Contribution from the Biosystem- (eds), The Natural History of Madagascar. University atic Laboratory, Graduate School of Social and Cultural of Chicago Press, Chicago, pp. 692–702. Studies, Kyushu University, Fukuoka 104: 1–171. Merz, B. & J.P. Haenni, 1998. 1.1. Morphology and Stireman, J.O. III, J.E. O’Hara & D.M. Wood, 2006. Ta- terminology of adult Diptera (other than genitalia). – chinidae: Evolution, behaviour and ecology. – Annual In: L. Papp & B. Darvas (eds), Contributions to a Review of Entomology 51: 525–555. Manual of Palaearctic Diptera 1: General and Applied Townsend, C.H.T., 1940. Manual of Myiology, Part X: Dipterology, 21–50 pp. Anacamptomyiini to Frontinini. – Charles Townsend Mesnil, L.P., 1950a. Larvaevorinae (Tachininae). – In: and Filhos, Itaquaquecetuba, Sao Paulo. E. Lindner (ed.), Die Fliegen der Paläarktischen Re- Tschorsnig, H.-P. & V.A. Richter, 1998. 3.54. Family Ta- gion 64g. E. Schweitzerbart’sche Verlagsbuchhandlung, chinidae.–In:L.Papp&B.Darvas(eds),Contri- Stuttgart, 105–160 pp. butions to a Manual of Palaearctic Diptera 3: Higher Mesnil, L.P., 1950b. Notes sur les Carceliina (Dipt. Ta- Brachycera, pp. 691–827. chinidae) et révision des espèces d’Afrique. – Revue de Villeneuve, J., 1910. Description de nouvelles espèces Zoologie et de Botanique Africaines 43: 1–24. de Tachinaires provenant de l’Afrique occidentale. – Mesnil, L.P., 1963. Nouveaux Tachinaires de la région Wiener Entomologische Zeitung 29: 249–254. Paléarctique principalement de l’USSR et du Japon. – Wood, D.M., 1985. A taxonomic conspectus of the Blon- Bulletin de l’Institut Royal des Sciences Naturelles de deliini of north and central America and the West In- Belgique 39(24): 1–56. dies (Diptera: Tachinidae). – Memoirs of the Entomo- Mesnil, L.P., 1977. Nouveaux Tachinaires de Madagascar, logical Society of Canada 132: 1–130. 2e partie (Dipt. Tachinidae). – Verhandlungen der Wood, D.M. & M.A. Zumbado, 2010. Tachinidae Naturforschenden Gesellschaft in Basel 86: 171–192. (tachinid flies, parasitic flies). – In: B.V. Brown, Mesnil, L.P. & H. Shima, 1977. A new genus and species A. Borkent, J.M. Cumming, D.M. Wood, N.E. Wood- of the Japanese Tachinidae (Diptera) reared from the ley & M.A. Zumbado (eds), Manual of Central Amer- nest of a solitary wasp Symmorphus sp. (Hymenoptera, ican Diptera, Vol. 2. NRC Research Press, Ottawa, Vespidae). – Kontyû 45: 36–42. pp. 1343–1417. O’Hara, J.E., 2007. Tachinid flies (Diptera: Tachinidae). Zeegers, Th., 2007. A first account of the Tachinidae – In: J.L. Capinera (ed.), Encyclopedia of Entomology (Insecta: Diptera) of Yemen. – Fauna of Arabia 23: [2nd Edition], Vol. 4 (S–Z), pp. 3675–3686. 369–419. O’Hara, J.E., 2012. World Genera of the Tachinidae Ziegler, J., 2003. Ordnung Diptera, Zweiflügler (Fliegen (Diptera) and their Regional Occurrence. http://www. und Mücken). – In: H.H. Dathe (ed.), Lehrbuch der nadsdiptera.org/Tach/WorldTachs/Genera/Gentach_ Speziellen Zoologie, Band I: Wirbellose Tiere, 5. Teil: ver7.pdf Insecta, zweiter Auflage, pp. 756–860. O’Hara, J.E., 2013. History of tachinid classification (Diptera, Tachinidae). – Zookeys 316: 1–34. Received: March 24, 2014 Accepted: September 9, 2014

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