Brachymeria podagrica (Fabricius, 1787)

Delvare, Gérard & Huchet, Jean-Bernard, 2017, Brachymeria mochica, a new Neotropical species of Chalcididae (Hymenoptera: Chalcidoidea) discovered on the archaeological site of Huacas de Moche, Peru with a review of related species, Zootaxa 4290 (1), pp. 43-60 : 51-56

Brachymeria podagrica (Fabricius, 1787)

Chalcis podagrica FabriciUS 1787: 272. Original deScription ♂. India, TranqUebar. Lectotype by BoUček 1972: 241.

Brachymeria podagrica (FabriciUS): BoUček 1972: 241.

= Chalcis femorata NeeS 1834: 412. Original deScription. Germany. Synonymy by Dalla Torre 1898: 391. Type apparently deStroyed.

= Chalcis Fonscolombei DUfoUr 1841: 16 – 17. Original deScription ♀. SoUthern France. Lectotype by BUrkS (1936). Synonymy by BoUček 1972: 241.

= Chalcis alphius Walker 1846a: 108 . Original deScription ♂. India, Bombay. Lectotype and Synonymy by BoUček 1972: 241.

= Chalcis xerxena Walker 1846a: 108 – 109. Original deScription ♂. India, Bombay. Lectotype and Synonymy by BoUček 1972: 241.

= Chalcis restituta Walker 1862: 351 – 352. Original deScription Jamaica. Lectotype and Synonymy by BoUček & DelVare 1992: 30.

= Chalcis pulchripes Holmgren 1868: 436 – 437. Original deScription ♂. PhilippineS, Manila. Lectotype and Synonymy by BoUček 1972: 241.

= Chalcis mansueta Walker 1871: 48 – 49. Original deScription ♀. Hong-Kong. Lectotype and Synonymy by BoUček 1972: 241.

= Chalcis callipus Kirby 1883: 75 . Original deScription ♀. Japan. Lectotype and Synonymy by BoUček 1972: 241.

= Chalcis mikado Cameron 1888: 117 . Original deScription ♀. Japan. Lectotype and Synonymy par BoUček 1972: 241.

= Chalcis eccentrica Cameron 1897: 39 . Original deScription ♂. India, Bombay. Lectotype and Synonymy by BoUček 1972: 241.

= Chalcis ferox Kieffer 1905: 263 – 264. Original deScription ♀ ♂. ReUnion ISland [BoUrbon] and MadagaScar, NoSSi-Bé. Syn. nov. Synonymy With B. fonscolombei by Steffan 1959: 42.

= Chalcis ferox Var. coxalis Kieffer 1905: 264 . Original deScription ♀. MadagaScar, NoSSi-Bé. Syn. nov. Synonymy With B. fonscolombei by Steffan 1959: 42.

= Chalcis borneanus Cameron 1905a : Original deScription ' ♀'. Borneo. Lectotype ♂ and Synonymy by BoUček 1972: 241.

= Chalcis spilopus Cameron 1905b: 231 . Original deScription ♀. SoUth Africa , TranSVaal. Syn. nov.

= Chalcis capensis Cameron 1905b: 311 – 312. Original deScription ♀. SoUth Africa , Katberg. Syn. nov.

= Chalcis transvaalensis Cameron 1911: 214 – 215. Original deScription ♀. SoUth Africa . Syn. nov.

= Tumidicoxoides kurandaensis GiraUlt 1913: 86 . Original deScription ♀. AUStralia, QUeenSland. Synonymy by BoUček 1988: 72.

= Chalcis dipterophaga GiraUlt & Dodd in GiraUlt 1915: 320 – 321. Original deScription ♀. AUStralia, QUeenSland. Synonymy by BoUček 1988: 72.

= Tumidicoxoides paucipunctatus GiraUlt 1915: 326 . Original deScription ♀. AUStralia, Northern Territory. Synonymy by BoUček 1988: 72.

= Chalcis neglecta MaSi 1916: 84 . Original deScription ♀. Italia, Giglio. Syn. nov. Synonymy With B. fonscolombei by Steffan 1959: 42.

= Chalcis sodalis MaSi 1917: 128 – 129. Original deScription ♀ ♂. SeychelleS ISlandS: Mahé, SilhoUette. Syn. nov. Synonymy With B. fonscolombei by Steffan 1959: 42.

= Chalcis vegai GiraUlt 1924: 175 . Original deScription ♀. AUStralia, QUeenSland. Synonymy by BoUček, 1988: 72.

= Chalcis garutianus Günther in Hellen &Günther 1936: 73 – 74. Original deScription ♀. JaVa, Kamodjan. Lectotype and Synonymy by BoUček 1972: 241.

= Brachymeria Fonscolombei Var. gananensis MaSi 1938: 210 – 212. Original deScription ♀. Somalia, Belet-Amin and MogadiScio. Syn. nov.

= Chalcis vulcani Schmitz 1946: 63 – 64. Original deScription ♂. RUanda, BUrambi (Volcan MUhaVUra). Syn. nov.

= Brachymeria aligarhensis Husain & Agarwal 1982: 499 – 501. Original description ♀. India, Uttar-Pradesh. Synonymy by Narendran 1989: 260.

Material examinated. Type material. Chalcis Fonscolombei. Lectotype, a pinned ♀ from the Dufour collection, identified by Steffan (MNHN) ( Figs 29, 30 ).

Chalcis capensis . Lectotype, a ♀ on a minutien pin, the designation of which by Bouček (1971) is validated here ( Fig. 33 ) (BMNH).

Chalcis transvaalensis . Lectotype, a ♀ in several pieces, remounted on a rectangular card, the designation of which by Bouček (1971) is validated here ( Fig. 34 ) (BMNH).

Chalcis spilopus . Lectotype, a ♂ on a minutien pin, the designation of which by Bouček (1971) is validated here (BMNH, Hym. type 5.155).

Chalcis ferox . Lectotype, a ♂ on a minutien pin, the designation of which by Rasplus is validated here; it is labelled 'Bourbon [manuscript] / Chalcis ferox Sauss. type ♂' [manuscript, in Kieffer's handwriting] ( Fig. 35 ) (MNHN). The ♀ quoted by Kieffer (1905) from Nossi Bé could not be retrieved.

Chalcis ferox var. coxalis . Lectotype, a pinned ♂ labelled 'Madag.' [manuscript] / ' Ch. ferox Sss. ' [manuscript] / ' Chalcis ferox Ss. var.' [manuscript, in Kieffer's handwriting] ( Fig. 36 ) (MNHN).

Chalcis neglecta . Lectotype, a ♀ glued on a rectangular card, the designation of which by Bouček (1971) is validated here; it is labelled 'Is. Giglio vii 901 G. Doria' / ' Chalcis neglecta Masi 1916 typus!' [manuscript, in Masi's handwriting] / ' ♂ Brachymeria neglecta (Masi) = podagrica (F.) Bouček det. 1971' [manuscript, in Bouček's handwriting] ( Fig. 37 ) (MCSN).

Brachymeria Fonscolombei var. gananensis . Lectotype, the designation of which by Bouček (1971) is validated here, labelled 'EL DUGUL Somalia Italiana E. Confalonieri' / ' Brachymeria Fonscolombei subsp. gananensis Masi ♀ det. L. Masi' [manuscript in Masi's handwriting] / 'co-typus' / 'Lectotype' [in BMNH format] with 'Bčk 1971' in the underside ( Fig. 38 ) (MCSN).

Chalcis vulcani . Holotype ♂, by original designation (MRAC, n° 567).

Other material. PERU. Trujillo , Huaca de la Luna, 08°08′06″S 78°59′29″W, 56 m, 06.v/ 01.vi.2009, in fly trap, J.- B. Huchet and A. Chauchet leg. (1 ♀ GDPC) . FRANCE. Bouches-du-Rhône , Istres, 01.vii.2003, P. Ponel leg. (1 ♂ GDPC) ; Gard , Le-Grau-du-Roi, l'Espiguette beach, 24.v.1997, G. Delvare leg. (1 ♀ GDPC) ; Hérault , Agde, Rochelongue, 29.vii.1979, J.- M. Maldès leg. (1 ♀ GDPC) ; Montpellier , campus ENSA, on mackerel corpse, 15.ix.2000, M. Martinez leg. (3 ♀ GDPC) ; same locality and collector, ex Sarcophaga proxima Rondani [ Sarcophagidae ] on a snake carcass, adult emergence 10/ 13.vi.2003 (2 ♀, 11 ♂ GDPC) ; Lot-et-Garonne , Buzetsur-Baïsse, 30.v.1995, Malaise trap, J.- P. Sarthou leg. (1 ♀ GDPC) . SPAIN. Tarragona , Cambrils-de-Mar, 30.v.1990, H. Tussac leg. (2 ♂ GDPC) .

Comment. Steffan (1959) previously synonymised C. ferox Kieffer and its variety C. ferox coxalis with B. fonscolombei , as well as C. neglecta Masi. Brachymeria fonscolombei was itself synonymised later with B. podagrica by Bouček (1972). The name podagrica was known from Dufour (1841) but because the type was described from India he thought that the European specimens belonged to another species, which he named Chalcis Fonscolombei in honour of Boyer de Fonscolombe. This name was adopted by subsequent authors until Bouček (1972) synonymised it with B. podagrica .

Brachymeria . podagrica was sometimes mixed with B. parvula (Walker) under its junior synonym Chalcis Dalmanni Thomson , especially by Stefani (1889) because the females of both species have the metafemur mostly reddish. It is nevertheless evident that Stefani dealt with B. podagrica because he reared his specimens from flies emerging from carrion while B. parvula is a secondary parasitoid of locusts.

Diagnosis. Apex of scape reaching base of median ocellus. Antennal flagellum mostly linear, hardly fusiform or clavate and bearing short erect setae only ( Fig. 21 ). Mesonotum with moderately dense puncturation, the interspaces between punctures narrower that the points themselves ( Fig. 22 ). Propodeum without spiracular callosity. Metacoxa and metafemur reddish, the latter with a small ivory spot apically. Metafemur slender 1.85 – 1.95× as long as broad, with an acute inner tooth basally; its outer surface with relatively dense puncturation, the basal piliferous points hardly more distant from each other than their own diameter ( Fig. 24 ). Metatibia reddish, respectively with a subbasal and dorsoapical ivory spots. Gaster acuminate, 1.8 – 2.0× as long as broad ( Fig. 23 ).

Hosts and biology. Table 1 lists the known hosts of the species and the authorships of the observations. Dubious host records are not included in the table, especially those for the hyperparasitoids of locusts which, as already mentioned by Steffan (1959), result from past confusion with B. parvula. We also omitted the hosts quoted for B. amenocles , which until now was wrongly mixed with B. podagrica following Bouček (1972) (see below).

Sarcophagidae (18 species) are predominant in the list of known hosts, which also includes a number of Calliphoridae (11 species) together with a few Muscidae . Cultures made by several authors (Stefani 1889; Parker 1924; Roberts 1933; Carlos 2014) showed that females emerging from sarcophagid hosts were always much more numerous compared to those that developed at the expense of the two other dipterous families. These flies often feed on corpses of various vertebrates, primarily mammals, but also from snakes (Beaver 1936; Michel Martinez pers. com.) and even fishes, as shown by the trapping of B. podagrica by exposing mackerel corpses (Michel Martinez, pers. com.). Some blowflies, such as Cochliomyia macellaria (Fabricius) (Calliphoridae) , also figure in the spectrum of hosts. The presence of individuals along the Mediterranean seaside (see above, examined material) suggests that B. podagrica is normally resident in this habitat, possibly searching for larval stages of flies developing at the expense of fish corpses washed up on the shore.

HoSt SpecieS HoSt family Reference and diStribUtion

Ophyra aenescens (Wiedemann, 1830) MUScidae Marchiori et al. 2003a and 2003c: Brazil Ophyra Sp. MUScidae CarloS 2014: Brazil

Synthesiomyia nudiseta (WUlp, 1883) MUScidae RobertS 1933: USA

Philornis Sp. MUScidae CoUri et al. 2006 : Brazil

Calliphora coloradensis HoUgh, 1899 Calliphoridae RobertS 1933: USA

Calliphora grahami Aldrich, 1930 Calliphoridae HabU 1960: Japan

Cochliomyia macellaria (FabriciUS, 1775) Calliphoridae RobertS 1933: USA

Chrysomya albiceps (Wiedemann, 1819) Calliphoridae Marchiori et al. 2002b and 2003a; CarloS 2014: Brazil Chrysomya megacephala (FabriciUS, 1794) Calliphoridae CarloS 2014: Brazil

Hemilucilia flavifacies Enderlein, 1931 Calliphoridae Marchiori et al. 2003b ; CarloS 2014: Brazil Lucilia caesar (LinnaeUS, 1758) Calliphoridae BUrkS 1960: USA

Lucilia illustris (Meigen, 1826) Calliphoridae BUrkS 1960: USA

Phaenicia mexicana (MacqUart, 1843) Calliphoridae RobertS 1933: USA

Phaeanicia sericata (Meigen, 1826) Calliphoridae RobertS 1933: USA: OliVa 2008: Argentina Phormia regina (Meigen, 1826) Calliphoridae BUrkS 1960: USA

Kellymyia plinthopyga (Wiedemann, 1830) Sarcophagidae RobertS 1933: USA

Kellymyia impar (Aldrich, 1916) Sarcophagidae RobertS 1933: USA

Kellymyia plinthopyga (Wiedemann, 1830) Sarcophagidae RobertS 1933: USA

Oxysarcodexia thornax (Walker, 1849) Sarcophagidae Marchiori et al. 2002a and 2003a; GonçalVeS et al. 2012;

CarloS 2014: Brazil

Peckia chrysostoma (Wiedemann, 1830) Sarcophagidae Marchiori 2001 ; Marchiori et al. 2003a; CarloS 2014: Brazil Sarcophaga africa (Wiedemann, 1824) Sarcophagidae DUfoUr 1841: France [Under S. haemorrhoidalis ]: RobertS

1933: USA

Sarcophaga argyrostoma

(RobineaU-DeSVoidy, 1830) Sarcophagidae UrSU & TUdor 1975; AndrieScU 1988: Romania; OliVa 2008:

Argentina

Sarcophaga carnaria (LinnaeUS, 1758) Sarcophagidae Stefani 1889 : Italy: Sicily; BUrkS 1960: USA Sarcophaga crassipalpis MacqUart, 1839 Sarcophagidae OliVa 2008 : Argentina

Sarcophaga dux ThompSon, 1869 Sarcophagidae ChoWdhUry & HoWlader 1975: BangladeSh Sarcophaga exuberans Pandellé, 1896 Sarcophagidae ChoWdhUry & HoWlader 1975 and 1978: BangladeSh Sarcodexia lambens (Wiedemann, 1830) Sarcophagidae Marchiori et al. 2003a ; GonçalVeS et al. 2012; CarloS 2014:

Brazil

Sarcophaga musitasi CUrran, 1934 Sarcophagidae Fry 1989 : PakiStan

Sarcophaga peregrina

RobineaU-DeSVoidy, 1830 Sarcophagidae HabU 1960: Japan

Sarcophaga proxima Rondani, 1860 Sarcophagidae Martinez perS. comm.: France (preSent pUblication) Sarcophaga Sp. Sarcophagidae Parker 1924 : France (cUltUre) Sarcophaga Sp. ex corpSe Achatina fulica Sarcophagidae BeaVerS 1986: Thailand

Sarcophaga tuberosa (Pandellé, 1896) Sarcophagidae FabritiUS & AndrieScU 1987; AndrieScU 1988: Romania As is the case for the other species belonging to the minuta group, B. podagrica is a solitary larval-pupal endoparasitoid ( Dufour 1841; Stefani 1889; Parker 1924; Roberts 1933). The latter author studied in detail the biology, ethology and phenology of B. podagrica , collecting copious samples and conducting large scale rearing of flies with the aim of evaluating the potential of the chalcidid to regulate the populations of the blowfly. All the above authors found a clear preference of B. podagrica for the pre-imaginal stages of Sarcophagidae relative to those of Calliphoridae , often rejecting the latter family. This might be explained by the fact that the adults of the Brachymeria , having emerged from the puparia of Calliphoridae , are much smaller in size—sometimes only half the size—than those emerging from Sarcophagidae , apparently relating to differences in the size of the hosts themselves ( Roberts 1933).

Females of B. podagrica oviposit within the body of the last larval stage of the host, which survives until pupation. Oviposition is especially induced by the presence of fluids issuing from the animals attacked by the flies, whether from living tissues or decaying corpses. The female uses her posterior legs to hold the larva, which promptly reacts to her attack. Reproduction is of the preovigenic type; the single mating occurs very soon after emergence of the imagos and is quickly followed by oviposition. This might be an adaptation to the presence of a large number of host larvae, grouped into clusters on decaying carrion. In temperate regions B. podagrica diapauses during winter in the last larval or prepupal stage, within the host puparium. Several generations occur during summertime ( Roberts 1933).

Distribution. Brachymeria podagrica is a cosmopolitan species, recorded from 45 countries (Noyes 2016), distributed from Australia to Sweden and Argentina. In the Neotropical Region the chalcidid is recorded from Mexico, Haiti, Jamaica, Peru, Brazil and Argentina. Its region of origin is not known and it is possible that is was introduced by Europeans when colonising the Americas.

Chalcis amenocles Walker and Brachymeria Beccarii Masi were synonymised with B. podagrica by Bouček (1972). However, examination of their types show that they belong to another distinct and valid species which is revalidated here.

Figures

FIGURES 29 – 39. 29, type SerieS of Brachymeria Fonscolombei DUfoUr (MNHN). 30 – 39, labelS of primary typeS. 30, Brachymeria Fonscolombei. 31, Chalcis varipes Walker. 32, Chalcis amenocles Walker. 33, Chalcis capensis Cameron. 34, Chalcis transvaalensis Cameron. 35, Chalcis ferox Kieffer. 36, Chalcis ferox Var. Kieffer. 37, Chalcis neglecta MaSi. 38, Brachymeria Fonscolombei Var. gananensis MaSi. 39, Brachymeria Beccarii MaSi.

FIGURES 21 – 28. 21 ‒ 24, Brachymeria podagric a, ♀. 21, antenna exclUding Scape. 22, meSoSoma in dorSal VieW. 23, metaSoma in dorSal VieW. 24, metafemUr. 25 ‒ 28, Brachymeria amenocles, ♀. 25, antenna exclUding Scape. 26, meSoSoma in dorSal VieW. 27, metaSoma in dorSal VieW. 28, metafemUr.