Brachymeria mochica, a new Neotropical species of

Zootaxa 4290 (1): 043–060 http://www.mapress.com/j/zt/ Copyright © 2017 Magnolia Press

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https://doi.org/10.11646/zootaxa.4290.1.2 http://zoobank.org/urn:lsid:zoobank.org:pub:8172109F-E545-4EB7-9E9D-D91E7DD1FA42

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 GÉRARD DELVARE1 & JEAN-BERNARD HUCHET2 1

Cirad UMR CBGP, 755 avenue du Campus Agropolis, CS 30016, 34988 Montferrier-sur-Lez Cedex. E-mail: [email protected] Institut de Systématique, Evolution, Biodiversité (ISYEB), UMR 7205 CNRS MNHN UPMC EPHE, Muséum national d'Histoire naturelle, CP 50, 45 rue Buffon, 75005 Paris France. E-mail: [email protected] 2

Abstract Two parasitoid wasp species of the Brachymeria minuta species group are studied. The specimens were collected in Peru at the archaeological site of Huacas de Moche, the former capital of the Mochica culture between 400–600 A.D. Brachymeria mochica Delvare sp. nov. (Chalcididae) is described and illustrated while B. podagrica (Fabricius, 1787) is reviewed with an update of the nomenclature and a summary of knowledge regarding its hosts and biology. The morphology and biology of both species are newly defined and characterized. Brachymeria brevicornis (Klug, 1834) stat. rev., previously synonymized with B. minuta (Linnaeus, 1767), is re-established as a valid species and a lectotype is designated. The following names are synonymised with B. podagrica: Chalcis ferox Kieffer, 1905 and its variety C. ferox coxalis Kieffer, 1905, Chalcis spilopus Cameron, 1905, Chalcis capensis Cameron, 1905, Chalcis transvaalensis Cameron, 1911, Chalcis neglecta Masi, 1916, Chalcis sodalis Masi, 1917, Brachymeria Fonscolombei var. gananensis Masi, 1938, Chalcis vulcani Schmitz, 1946, and Brachymeria fonscolombei f. neglecta Masi, 1951 new synonymies. Brachymeria amenocles (Walker, 1846) stat. rev. is re-established as a valid species with Chalcis varipes Walker, 1871 and Brachymeria Beccarii Masi, 1929 as junior synonyms syn. rev. Lectotypes are designated for the above names except for C. sodalis and C. vulcani. Key words: taxonomy, description, Sarcophagidae, parasitoids

Résumé Les auteurs présentent les deux Brachymeria collectés sur le site archéologique de las Huacas de Moche, ancienne capitale de la culture Mochica (400–600 A.D.), au Pérou. Toutes deux appartiennent au groupe minuta, nouvellement défini et caractérisé, au niveau de sa morphologie et de sa biologie. Brachymeria mochica Delvare sp. nov. (Chalcididae) est décrit et illustré tandis que B. podagrica (Fabricius, 1787) fait l'objet d'une synthèse avec actualisation de la nomenclature et un résumé des connaissances sur ses hôtes et sa biologie. Brachymeria brevicornis (Klug, 1834), préalablement mis en synonymie avec B. minuta est revalidé et un lectotype désigné. Les noms suivants sont mis en synonymie avec B. podagrica: Chalcis ferox Kieffer, 1905 et sa variété C. ferox coxalis Kieffer, 1905, Chalcis spilopus Cameron, 1905, Chalcis capensis Cameron, 1905, Chalcis transvaalensis Cameron, 1911, Chalcis neglecta Masi, 1916, Chalcis sodalis Masi, 1917, Brachymeria Fonscolombei var. gananensis Masi, 1938, Chalcis vulcani Schmitz, 1946, et Brachymeria fonscolombei f. neglecta Masi, 1951 nouvelles synonymies. Brachymeria amenocles (Walker, 1846) est revalidé stat. rev., avec pour synonymes Chalcis varipes Walker, 1871 et Brachymeria Beccarii Masi, 1929 syn. rev. Des lectotypes ont été désignés ou validés pour l'ensemble de ces noms à l'exception de C. sodalis et C. vulcani.

Introduction Between 2008 and 2009, archaeo-entomological investigations were conducted by one of the authors (JBH) on a funerary platform at the foot of the Huaca de la Luna (archaeological site of Huacas de Moche, the former capital Accepted by G. Gibson: 24 Apr. 2017; published: 6 Jul. 2017

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of the Mochica civilization in Peru) (Fig. 1). The aim was to study the insect remains present in the mochica tombs from an 'archaeo-forensic' perspective. Within the archaeological material, largely consisting of puparia from the main Diptera families of forensic interest (i.e. Calliphoridae, Muscidae and Sarcophagidae), the Sarcophagidae puparia showed distinct exit holes, indicating that they had been parasitized (Huchet & Greenberg 2011) (Fig. 2). In order to collect information on the biology and colonization stages of the necrophagous flies present in the mochica graves, and to identify the parasitoid Hymenoptera, a selective trapping protocol, involving the measurement of daily temperature and humidity, was conducted in situ (Tsurikov 2006; Moretti & Godoy 2013; Farinha et al. 2014). Two distinct hymenopteran species belonging to the genus Brachymeria (Chalcididae) were collected in the traps among a crawling mass of Synthesiomyia nudiseta Van der Wulp, 1883 larvae (Muscidae) and unidentified Sarcophagidae species. Their examination revealed the presence of a single specimen of B. podagrica (Fabricius) and four specimens of a new species of the Brachymeria minuta species group, described herein as B. mochica sp. nov. The genus Brachymeria currently includes about 350 described species worldwide (Noyes 2016) and 37 species in the Neotropical Region, if excluding those presently included in Ceyxia Girault, which contains an additional 25 described species (Andrade & Tavares 2009). Currently, there is no key available to identify Brachymeria species; furthermore, species groups remain relatively undefined and little is known about their infrageneric relationships. We describe here the new species, place it within the newly diagnosed minuta species group, and review the known species from the New World.

Material and methods The traps (aerial and partially buried in the soil) were baited with pieces of pork (fig. 3). All the Brachymeria specimens were collected in traps baited with pork meat placed in the vicinity of the Huaca de la Luna temple (Fig. 3a–b). These Chalcididae were found among a huge number of 2nd and 3rd instar larvae of necrophagous Diptera including the muscid fly Synthesiomyia nudiseta Van der Wulp 1883 as well as unidentified sarcophagid flies (Fig. 3c). Morphological characterization. Following desiccation, specimens were immediately prepared for morphological examination as recommended by Noyes (1982). Specimens were labelled according to the information provided by JBH. Terms for morphology follow Bouček (1988; 1992), Delvare (1992) and Delvare et al. (2011) except 'scutellum' and 'dorsellum', replaced here by 'mesoscutellum' and 'metascutellum' in accordance with Seltmann et al. (2012). Specimens were mostly photographed with a JVC KY-75U 3CCD digital camera attached to an EntoVision microscope and the stacked, serial images obtained were combined using Cartograph 5.6.0 (Microvision, Evry, France) software. The photos were then digitally optimized (artefacts removal, background standardized) using the Photoshop® V program. Some images were further made using Keyence VHH 5000 equipment. At BMN a Multifocus-Station Leica was used consisting of a Z16Apo Stereomicroscope linked to a Leica DFC 495 digital camera and managed with the Leica Application Suite 4.5. The program Helicon Focus 6 was used for stacking of the serial images. Collection acronyms used are: BMN, Berlin Museum für Naturkunde, Berlin, Germany; BMNH, Natural History Museum, London, UK; CIRAD, Cirad collection in Centre de Biologie pour la Gestion des Populations (CBGP), Montferrier-sur-Lez, France; CJBH, Jean-Bernard Huchet personal collection, Bordeaux, France; GDPC, Gérard Delvare personal collection, Montpellier, France; MCSN, Museo Civico di Storia Naturale, Genova, Italia, MNHN, Muséum National d'Histoire Naturelle, Paris, France; MRAC, Musée Royal d'Afrique Centrale, Tervuren, Belgium; USNM, United States National Museum of Natural History, Washington DC, USA. Species identification. GD examined specimens from approximately 300 species of Brachymeria, including most of the available types of species described from the New World and selected specimens from other regions. Abbreviations. F1 to F7 correspond to the rank of the funicular segments, F1 being the second flagellomere; GT1 to GT6 rely to the rank of the gastral tergites.

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FIGURES 1–3. 1, Collection site. 2, Sarcophagidae puparia collected from an ancient grave and showing emergence holes of adult parasitoids (Photo C. Chauchat and J.-B. Huchet). 3a, design of the trap used for collecting flies; 3b, installed trap ready to collect; 3c, details of the inside of the trap with 2nd and 3rd instars larvae of Muscidae and Sarcophagidae—the arrow points to an adult Brachymeria (Photos J.-B. Huchet).

Results The Brachymeria minuta species group Diagnosis. Head conspicuously transverse in frontal view (Fig. 5). Both mandibles 2-toothed, the teeth blunt at apex, with lower tooth longer than upper tooth (Fig. 8). Postorbital carinae present and complete, reaching posteriorly genal carina (Fig. 7). Lower surface of gena trapezoidal, delimited, respectively, by postorbital carina, genal carina, and lateral margin of oral fossa. Pedicel squat, at most as long as broad in dorsal view and basally ‘bottle necked’ (Fig. 11). Flagellum weakly fusiform, flagellomeres short and often strongly transverse and frequently bearing long, sparse and erect setae (Fig. 10). Ventral section of the epicnemial carina never raised into submedian lobes. Apical truncation of metatibia clearly emarginate (Fig. 15). Spatulate seta (Steffan 1954) of posterior pretarsus slender and not curved very much, generally surpassing apex of pretarsus (Fig. 16). BRACHYMERIA MOCHICA IN PERU

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Subdivision. The minuta group can be subdivided into two distinct sections. The first one, here defined as the minuta section, includes primary parasitoids of synanthropic flies (mainly Sarcophagidae and Calliphoridae) developing within carrion and dung, while the parvula section, comprises secondary parasitoids, mainly of Lepidoptera through Tachinidae (Blanchard 1935; Burks 1960; Sychevskaya 1966; Louranção & Sabino 1994; Tavares et al. 2006), more rarely of Coleoptera or Orthoptera through Sarcophagidae or Anthomyiidae (Steffan 1959; Burks 1960; Léonide & Léonide 1969). The proposed subdivision is otherwise corroborated by several morphological features. The minuta section is characterized by: 1) outer surface of metafemur with moderately to distinctly sparse piliferous points only, with engraved network extremely superficial, visible at high magnification only [over 300×] (Figs 24, 28); 2) metafemur with an inner basal tooth or tubercle; 3) posterior tarsus slender; 4) antennal scrobes never carinate dorsally (Fig. 5). Additionally in most species, the mesonotum is completely punctured (Figs 12, 22, 26), at most areolate on anterior third in some species of this section. Conversely, species of the parvula section do not show the combination of characters stated above for the minuta section: outer surface of metafemur generally with an evident engraved, alutaceous network in addition to piliferous points and mesonotum areolate. Other characters of the parvula section are variable: the metafemur usually has no inner basal tooth and in some species the antennal scrobes are completely carinate and/or the posterior tarsus markedly broadened towards the apex. The minuta section is represented in the New World by B. podagrica (Fabricius), a species also widespread throughout the world. Otherwise, only species of the parvula section have been described from the Americas: B. aeca Burks, B. carinatifrons Gahan, B. compacta (Walker), B. compsilurae (Crawford), B. koehleri Blanchard, B. nigritibialis Tavares & Navarro-Tavares, B. parvula (Walker)—this latter species with a Holarctic distribution—B. tegularis (Cresson), B. truncatella (Burks) and B. weemsi Burks. Species recognition. The Brachymeria minuta species group exhibits numerous variable characters that can be used for separating species: setation generally silvery but sometimes golden; sculpture of the labrum and clypeus; variable development of the preorbital carinae; density of the puncturation of the mesonotum; habitus of the posterior margin of the scutellum, resulting from the more or less deep emargination of the frenal carina; punctulation of the metafemur; habitus of the gaster which may be squat or acuminate, etc. Diversity. The group includes at least 28 species already described from the Old World and 8 from the Americas, but the examination of diverse collections showed that many species remain undescribed. Hosts and biology. The species for which the hosts and development are known are solitary endoparasitoids of Diptera, showing a koinobiont strategy, with the female ovipositing within the larval stages of the host while the adult emerges from the pupae (Stefani 1889; Roberts 1933; Dowden 1935; Léonide & Léonide 1969). The biology of B. compsilurae (Crawford) was studied by Dowden (1935) in North America. The species develops within tachinids that are primary parasitoids of several species of Lepidoptera. In the Holarctic Region B. moerens (Ruschka) and B. parvula (Walker) parasitize sarcophagids and anthomyiides that are primary parasitoids of Orthoptera (Léonide & Léonide 1969). The Brachymeria that develop at the expense of coprophagous or necrophagous Calliphoridae and Sarcophagidae are apparently attracted by chemicals given off by the organic tissues in which the flies feed and consequently can be trapped using meat or dead bodies (Roberts 1933; Horenstein & Salvo 2012; Michel Martinez pers. com.; present paper).

Brachymeria collected on the Huaca de la Luna site Brachymeria mochica Delvare sp. nov. (Figs 4–20) Material examined. Holotype ♀. PERU. Trujillo, Huaca de la Luna, Libertad Region, 8°08′06″S 78°59′29″W, 56 m, 06.V/01.VI.2009, in fly trap with 2nd and 3rd instar larvae of Muscidae and Sarcophagidae, J.-B. Huchet leg. (MNHN). Paratypes. 1 ♀, same references as holotype (GDPC). 2 ♀, same references but J.-B. Huchet and A. Chauchat leg. (CJBH), with one ♀ dated 11.V.2009 (BMNH). Description (female holotype). Body length 6.5 mm. Colour (Fig. 4). Body and most of coxae, basal half of metafemur on both sides, and ventral margin of metatibia black. Mandible with basal half dark brown, then with a reddish ring and black teeth (Fig. 8). Inner side of mandible and remaining mouth parts, radicle and pedicel of


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antenna, and apices of anterior and middle trochanters reddish brown. Labrum and basal halves of pro- and mesofemora dark brown to reddish. Flagellum dark brown except clava yellowish. Tegula, rest of pro- and mesofemora, and a broad spot on metafemur apically bright yellow. Pro- and mesotibia with a reddish ring, this wider on mesotibia. Metatibia with entire inner side reddish and outer side with reddish ring broadening towards its ventral margin; basodorsal and apicoventral spots yellow. Tarsi and scape of antenna mostly testaceous but scape darker within diffuse region dorsobasally. Submarginal vein of fore and hind wings testaceous at base, then becoming brown, the same colour as rest of venation. Setation silvery on body and dark on fore wing.

FIGURE 4. Brachymeria mochica Delvare, holotype ♀. Habitus in lateral view.

Head. Head transverse in dorsal and frontal views, 2.5× as wide as long (Fig. 6) and 1.42× as wide as high (Fig. 5), hardly wider than mesosoma (1.06×). Distance between lateral ocellus and compound eye 0.88× diameter of median ocellus. Eyes separated by 1.21× their own height. Oral fossa 2.5× as long as length of malar space, which is consequently very short. Mandible elongate, 2-toothed, with lower tooth longer than upper tooth (Fig. 8); disk densely punctured basally and bearing fine setae, the longest setae inserted at base of the teeth and as long as width of mandible. Labrum subtriangular, with a marginal fringe of setae and 12 irregularly distributed piliferous points (Fig. 9). Clypeus conspicuously transverse, with a line of 8 piliferous points. Lower face regularly punctured (Fig. 9). Postorbital carina present, complete, diverging markedly relatively to posterior margin of eye, delimiting dorsally a ventral trapezoidal area on gena (Fig. 7). Gena with puncturation coarse but relatively sparse, with

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smooth interspaces between points; area above gena rugose-punctured (Fig. 7). Frons with blunt preorbital carina visible in its ventral half (Fig. 5). Eyes not very large, with dorsal margin of clypeus clearly below lower ocular line and lower margins of antennal scrobes somewhat above ocular line (Fig. 5). Antennal scrobes 1.25× as high as broad, extending to median ocellus and acarinate dorsally. Interantennal projection not strongly compressed laterally, with two lines of piliferous points dorsally. Antenna. Scape not reaching median ocellus, 3.6× as long as wide. Pedicel transverse, conspicuously ‘bottle necked’ dorsobasally (Fig. 11). Flagellum weakly fusiform, its maximum width at F3–F5 (Fig. 10); with long and sparse erect setae, the longest setae ventrally where two-thirds length of F7. Funiculars transverse and progressively decreasing in length, with F1 and F7 respectively 0.90× and 0.54× as long as broad. Clava 1.47× as long as broad, tapering from base to apex, the last segment hemispherical, hardly sclerotized, and with numerous sensilla. Mesosoma short and squat, only 1.37× as long as broad (Fig. 12); setae on dorsum relatively long, about 2.6× as long as width of a point, adpressed on pronotum, and suberect on mesonotum; puncturation sparse with interspaces between points smooth, as wide as diameter of a point (65 µm on centre of mesoscutum). Pronotal carina vestigial, hardly visible laterally; pronotal collar widening posteriorly (Fig. 12), its sides convex in dorsal view, and its lateral surface well delimited relatively to dorsal surface and with a narrow oblique grove and a small depression ventrally. Mesoscutum and mesoscutellum (Fig. 12) respectively 1.23× and 0.88× as long as broad; puncturation sparse and superficial on inner half of scapula, dense on axillae and apical part of mesoscutellum. Mesoscutellum with edge of the frenal carina forming two submedian rounded lobes, separated by a moderately wide emargination; frenal area [not visible from above] coriaceous with vertical rugulae, the latter more closely set in posterior part (Fig. 13). Metascutellum areolate. Propodeum steeply sloped, entirely areolate, with a few sublateral setae on each side at mid length. Mesepisternum with dorsal stripe of adscrobal area broadly smooth, but with an incomplete line of piliferous points (Fig. 14). Mesepimeron and femoral scrobe mostly smooth though the latter with a few transverse crests, and both crossed by a narrow, crenulate transepimeral sulcus. Epicnemial carina not raised on the ventral section. Metapleuron areolate with a wide areola anterobasally just above supracoxal stripe of punctures (Fig. 14). Legs. Metacoxa moderately slender, 1.7× as long as broad, with ventral setation long and adpressed (Fig. 4). Metafemur 1.8× as long as broad, with disc devoid of engraved network, having only piliferous points separated from each other by 2–3× diameter of a point (Fig. 4); with longest setae basoventrally, 1.5× as long as the basal tooth; ventral margin with 12 teeth progressively closer to each other; basal inner tooth present but blunt. Metatibia with apical truncation emarginate and forming a small acute projection (Fig. 15). Tarsi slender; falciform seta of posterior pretarsus slender, not very curved, surpassing apex of tarsus (Fig. 16). Wings. Fore wing 2.8× as long as broad (Fig. 4). Costal cell elongate, 1.24× as long as width of wing (Fig. 17). Marginal vein 0.35× as long as costal cell and 2.38× as long as postmarginal vein, which is 2.4× as long as stigmal vein (Fig. 18). Cubital fold bare. Basal fold with eight setae dorsally. Costal cell with two lines of setae ventrally (Fig. 17). Metasoma. Gaster 1.79× as long as broad (Fig. 19). Syntergum short, 0.55× as long as broad (Fig. 20); not carinate dorsally, punctured laterally anterior to cercal plates. First gastral tergite dorsally smooth, its posterior margin hardly concave (Fig. 19); second tergite with piliferous points basally and several lines of setae laterally, the lines broken dorsally; tergites 3‒5 with one row of setae subapically; and tergite 6 completely setose with larger piliferous points. Tip of hypopygium at level of hind margin of penultimate tergite. Male. Unknown. Variation. Variability among the specimens was limited to their length, which ranges from 4.7‒5.5 mm and reflects the degree of telescoping of the gastral tergites. Diagnosis. The species exhibits all the characters of the minuta species group and minuta section, as described above. Otherwise, body entirely black including metacoxa and basal half of metafemur (Fig. 4). Body with silvery setae. Preorbital carinae incomplete (Fig. 5). Antennal scrobes acarinate dorsally. Scape relatively long, 3.6× as long as broad (Fig. 10). Erect setae on flagellum moderately numerous and visible on ventral side only. Puncturation of mesonotum sparse and fine with interspaces between punctures smooth and as broad as diameter of a point (Fig. 12). Tegula and apical half of metafemur bright yellow. Metafemur 1.8× as long as broad, its disk devoid of engraved network and exhibiting spaced piliferous points only (Fig. 4).


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FIGURES 5–13. Brachymeria mochica, holotype ♀. 5–7, head in frontal, dorsal and lateral views, respectively. 8, right mandible. 9, clypeus and mouth parts. 10, antenna excluding scape. 11, pedicel and base of flagellum. 12, mesosoma in dorsal view. 13, mesoscutellum in lateral view.



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FIGURES 14–20. Brachymeria mochica, holotype ♀. 14, meso- and metapleuron. 15, hind tarsus. 16, hind pretarsus. 17, base of fore wing. 18, stigmal and postmarginal vein. 19, metasoma in dorsal view. 20, apex of metasoma.

Recognition. Brachymeria mochica, some specimens of B. brevicornis (Klug, 1834), and B. minuta have a similar colouration of the metafemur, being black with a bright yellow spot apically (Figs 4, 43). In contrast, the other species of the minuta section have the femur mainly or broadly reddish with an ivory spot apically. Brachymeria mochica and B. brevicornis are separated from B. minuta by the scape, mostly light coloured in B. mochica (Fig. 10) but dorsally dark in B. minuta, by the more delicate and sparser puncturation of its mesonotum (Fig. 12), by the dorsal outline of the mesoscutellum, regularly convex when examined in lateral view (Fig. 13) versus subhorizontal and then sloping apically in B. minuta, by its relatively broader mesoscutellum, by the absence of spiracular callosity/tooth on the propodeum, by the more expanded apical yellow spot on the metafemur, and by the colour of the dark parts of the tibiae, being reddish in B. mochica (Fig. 4) versus brown to black in B. minuta. Brachymeria mochica is extremely close to B. brevicornis and all characters quoted above for separating it from B. minuta, except the puncturation of the mesonotum, are also present in B. brevicornis. This species differs from B. mochica by its fusiform flagellum bearing numerous erect setae on the dorsal side only (Fig. 40) and the mesonotum having golden setation and larger punctures (Fig. 41). Distribution. The species is known only from its type locality, in Peru. Hosts. Basically unknown but a suspected host is the unidentified sarcophagid species or even the muscid fly Synthesiomyia nudiseta, both species occurring simultaneously in the traps.


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Update of the nomenclature Brachymeria brevicornis (Klug, 1834) stat. rev. Chalcis brevicornis Klug, 1834: 4, plate 37, fig. 10. Original description. Egypt, Anjun. Brachymeria brevicornis (Klug): Lucas, 1849: 318. = Vespa minuta Linnaeus, 1767: Bouček, 1956: 232.

Type material. Three conspecific syntypes (2 ♀ 1 ♂) are labelled as such in the BMN collection (Fig. 43). The female in the best condition is selected as lectotype. It has the infuscate parts of the legs dark brown to black (Fig. 42); the other female has the infuscate parts of the legs reddish.

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.



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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).


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FIGURES 21–28. 21‒24, Brachymeria podagrica, ♀. 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.



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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.

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.


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TABLE 1. Known hosts of Brachymeria podagrica. With a very few exceptions, only data from original publications are included, hence excluding second-hand citations in catalogues or data. Host species

Host family

Reference and distribution

Ophyra aenescens (Wiedemann, 1830) Ophyra sp. Synthesiomyia nudiseta (Wulp, 1883) Philornis sp. Calliphora coloradensis Hough, 1899 Calliphora grahami Aldrich, 1930 Cochliomyia macellaria (Fabricius, 1775) Chrysomya albiceps (Wiedemann, 1819) Chrysomya megacephala (Fabricius, 1794) Hemilucilia flavifacies Enderlein, 1931 Lucilia caesar (Linnaeus, 1758) Lucilia illustris (Meigen, 1826) Phaenicia mexicana (Macquart, 1843) Phaeanicia sericata (Meigen, 1826) Phormia regina (Meigen, 1826) Kellymyia plinthopyga (Wiedemann, 1830) Kellymyia impar (Aldrich, 1916) Kellymyia plinthopyga (Wiedemann, 1830) Oxysarcodexia thornax (Walker, 1849)

Muscidae Muscidae Muscidae Muscidae Calliphoridae Calliphoridae Calliphoridae Calliphoridae Calliphoridae Calliphoridae Calliphoridae Calliphoridae Calliphoridae Calliphoridae Calliphoridae Sarcophagidae Sarcophagidae Sarcophagidae Sarcophagidae

Peckia chrysostoma (Wiedemann, 1830) Sarcophaga africa (Wiedemann, 1824)

Sarcophagidae Sarcophagidae

Marchiori et al. 2003a and 2003c: Brazil Carlos 2014: Brazil Roberts 1933: USA Couri et al. 2006: Brazil Roberts 1933: USA Habu 1960: Japan Roberts 1933: USA Marchiori et al. 2002b and 2003a; Carlos 2014: Brazil Carlos 2014: Brazil Marchiori et al. 2003b; Carlos 2014: Brazil Burks 1960: USA Burks 1960: USA Roberts 1933: USA Roberts 1933: USA: Oliva 2008: Argentina Burks 1960: USA Roberts 1933: USA Roberts 1933: USA Roberts 1933: USA Marchiori et al. 2002a and 2003a; Gonçalves et al. 2012; Carlos 2014: Brazil Marchiori 2001; Marchiori et al. 2003a; Carlos 2014: Brazil Dufour 1841: France [under S. haemorrhoidalis]: Roberts 1933: USA

Sarcophaga argyrostoma (Robineau-Desvoidy, 1830)

Sarcophagidae

Sarcophaga carnaria (Linnaeus, 1758) Sarcophaga crassipalpis Macquart, 1839 Sarcophaga dux Thompson, 1869 Sarcophaga exuberans Pandellé, 1896 Sarcodexia lambens (Wiedemann, 1830)

Sarcophagidae Sarcophagidae Sarcophagidae Sarcophagidae Sarcophagidae

Sarcophaga musitasi Curran, 1934 Sarcophaga peregrina Robineau-Desvoidy, 1830 Sarcophaga proxima Rondani, 1860 Sarcophaga sp. Sarcophaga sp. ex corpse Achatina fulica Sarcophaga tuberosa (Pandellé, 1896)

Sarcophagidae

Ursu & Tudor 1975; Andriescu 1988: Romania; Oliva 2008: Argentina Stefani 1889: Italy: Sicily; Burks 1960: USA Oliva 2008: Argentina Chowdhury & Howlader 1975: Bangladesh Chowdhury & Howlader 1975 and 1978: Bangladesh Marchiori et al. 2003a; Gonçalves et al. 2012; Carlos 2014: Brazil Fry 1989: Pakistan

Sarcophagidae Sarcophagidae Sarcophagidae Sarcophagidae Sarcophagidae

Habu 1960: Japan Martinez pers. comm.: France (present publication) Parker 1924: France (culture) Beavers 1986: Thailand 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 BRACHYMERIA MOCHICA IN PERU


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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 40–43. Brachymeria brevicornis, lectotype ♀. 40, antenna excluding base of scape. 41, mesosnotum. 42, hind leg. 43, labels.

Brachymeria amenocles (Walker, 1846) stat. rev. Chalcis Amenocles Walker 1846b: 84. Original description ♀. Sierra Leone. Lectotype by Bouček 1972: 241 but type locality erroneously quoted as Hong-Kong by this author. Brachymeria amenocles (Walker): Thompson 1955: 195 (earliest known use of combination vide Noyes 2016). = Chalcis varipes Walker 1871: 48. Original description ♀. South Africa. Syn. rev. = Brachymeria Beccarii Masi 1929: 142–144. Original description ♀. Kenya. Syn. rev.

Type material. Chalcis Amenocles. Lectotype, a ♀ on a minutien pin, designated by Bouček (1972), this designation being validated here (Fig. 32) (BMNH, Hym. type 5.440).


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Chalcis varipes. Lectotype, a ♀ on a minutien pin, the designation of which by Bouček (1971) is also validated (Fig. 31) (BMNH, Hym. type 5.441). Brachymeria Beccarii. Holotype ♀, by monotypy (Fig. 39) (MCSN). Other material. IVORY COAST. Bouaké, Malaise trap in rice field, 28.i.1978, P. Cochereau leg. (1 ♀ CIRAD). GUINEA. Koliegbé, 01.iii.1992, J.-F. Vayssières leg. (13 ♀ CIRAD). Diagnosis. Mainly as B. podagrica but separated from it by its flagellum being clearly fusiform and having long erect setae (Fig. 25); the mesonotum having much sparser puncturation with numerous interspaces being broader than the points (Fig. 26); the metafemur having much sparser punctulation, basally the femur having the interspaces between the piliferous points much larger than their own diameter (Fig. 28); and the gaster being lanceolate and relatively longer, at least 2× as long as broad (Fig. 27). Hosts and Biology. In Namibia and Zimbabwe, B. amenocles [as B. varipes or B. podagrica] develops at the expense of Sarcophaga inzi Curran, 1834, infesting corpses of Triaenostreptus heros Porat, 1872 [quoted as Spirostreptus triodus (Attems, 1909)] (Diplopoda, Spirostreptidae) (Curthbertson 1932 & 1938; Kirk-Spriggs 1999). The female oviposits below the cuticle of the third instar larvae before these move within the ground for pupation. Brachymery amenocles may also parasitize the larvae of Glossina morsitans Westwood, 1851 (Glossinidae) following the same behavioural pattern (Curthbertson 1932).

Discussion Specialisation of the Brachymeria spp. belonging to the minuta group. In all references (see Table 1) the hosts are Diptera developing at the expense of vertebrates, whether they are alive or dead (Sarcophagidae, Calliphoridae, Glossinidae and Muscidae) or are parasitoids of other insects (Tachinidae and Anthomyiidae). Furthermore, available observations on the behaviour of these Brachymeria (Dufour 1841; Stefani 1889; Parker 1924; Roberts 1933; Dowden 1935; Léonide & Léonide 1969) show a narrow host specialisation, with the discovery of the host, oviposition and phenology of the parasitoid closely related to the development of the host. The koinobiont strategy, with oviposition at the larval stages of the host which survives until pupation, and the emergence of the adult parasitoid from the puparia, may be considered an adaptation to flies that pupate within the ground and thus are inaccessible to the chalcidid females. The puparium of the host also, provides a shelter for the parasitoid pupation and possible hibernation. This strategy also allows these Brachymeria to gain access to food resources particularly important when the fly larvae are clustered together. The chalcidid adults emerging from pupae within the ground need to dig their way to the surface. The special morphology of the mandibles, especially long and robust and with blunt teeth, is the only known synapomorphy of the group. The mandibular structure might also prove to be a morphological adaptation to emergence from a host within the soil, allowing the wasp adult to escape towards the surface. Previous catalogues or databases (Noyes 2016) include a number of errors probably resulting from misidentifications, especially records listing Brachymeria of the minuta species group as primary parasitoids of Lepidoptera. This contradicts the apparent specialisation of those Brachymeria on Diptera. Here, when searching for their hosts or during oviposition, the females are attracted by semiochemicals, either originating from decaying dead bodies (Horenstein & Salvo 2012; Michel Martinez pers. com.), or resulting from injuries inflicted by the larvae (Roberts 1933). Furthermore, as shown by Léonide & Léonide (op. cit.) for B. moerens, Brachymeria that are secondary parasitoids look for primary hosts that are already parasitized, probably exploiting physiological modifications induced by the parasitism of the tachinid or sarcophagid flies. Lastly, the life cycle of B. moerens is perfectly synchronized with that of the pair primary host and its primary parasitoid. Region of origin and potential hosts for B. mochica. The morphological characters of the new species B. mochica support, as far as is presently known for the species group, a putative sister group relation to B. minuta. However, the huge gap in the respective distributions of the two species—B. minuta is only quoted from Eurasia— challenges this relationship. Because of the great antiquity of the separation of the two continents, it remains open if the current distribution is the result of a very ancient allopatric speciation event or that of an unintentional introduction following the colonisation of the Americas by humans. It presently is impossible to answer this question until a sound phylogeographic study is performed.



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Acknowledgements The authors are sincerely obliged to Nathalie Dale-Skey (BMNH), Roberto Poggi and Maria Tavano (MCSN), Claire Villemant and Agnièle Touret-Alby (MNHN), Eliane De Coninck (MRAC) and finally Lukas Kirschey (BMN) for the loans of types and their kind and helpful welcome during the visits of GD. We are grateful to Michel Martinez for the specimens he collected and generously loaned to us and for information on the behaviour of B. podagrica. We also thank very much Claude Chauchat (UMR 8096, "Archéologie des Amériques” (ArchAm)), Head of the international project Moché, who provided the opportunity to JBH to undertake entomological studies on the Huaca de la Luna site. We especially thank the late Andrée Chauchat, for her kind assistance during the research carried out on the sites. At last we are much obliged to Robert Copeland, from ICIPE, Nairobi, Kenya who kindly corrected the English. We thank the two reviewers for their constructive comments and corrections which improved the text.

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