Brachymeria zygaenae Delvare & Shaw, 2022

Delvare, Gérard & Shaw, Mark R., 2022, Characterisation of some species groups of Brachymeria (Hymenoptera: Chalcididae), with a review of the B. tibialis-group and description of a new species parasitizing Zygaena pupae (Lepidoptera: Zygaenidae), Zootaxa 5178 (5), pp. 401-432 : 404-412

Brachymeria zygaenae Delvare & Shaw , sp. n.

( Figs 1 , 2A–I , 3A–D , 8E–H , 9D–F , 10A–I )

Material examined. TYPE MATERIAL. Holotype ♀ (in NMS). FRANCE: Alpes-de-Haute-Provence: Lac de Sainte-Croix , ex pupa of Zygaena sarpedon , collect 23.vi.1996, emergence 06.vii.1996 (Hofmann A.) . Paratypes (all NMS unless stated otherwise). EUROPE. FRANCE: Alpes-Maritimes: Saint-Martin-Vésubie, La Colmiane, ex cocoon Charops sp. from cocoon of Zygaena filipendulae , collect 02.viii.1987, emergence 22.viii.1987 (Shaw M. R.) (1♂) ; Saint-Martin-Vésubie, Valdeblore , ex pupa of Zygaena filipendulae , collect 01.viii.1997, emergence 04.viii.1987 (Shaw M. R.) (1♂) . Charente-Maritime : Benon, Forêt de Benon, ex pupa of Zygaena transalpina , 07.vi.1976 (Rouch A.) (1♂) . Deux-Sèvres : Availles-Thouarsais, ex pupa of Zygaena filipendulae , collect 19.viii.1979, emergence 22.viii.1979 (Drouet E.) (1♀) . Dordogne : Brantôme ( NW), NW Mareuil, 100 m, ex pupa of Zygaena transalpina , collect 16.ix.1999, emergence 16.ix.1999 (Tremewan W.G. & M.A.) (1♂) ; Saint-Alvère , ex pupa of Zygaena transalpina , collect 18.v.1996, emergence vii.1996 (Shaw M. R.) (1♀) . Haute-Marne : Fayl-Billot (towards Langres), ex pupa of Zygaena transalpina , collect 10.vii.1996, emergence vii.1996 (Shaw M. R.) (1♂) ; same data but collect 24.vii.1996, emergence on 28.vii.1996 (4♀; 1 in CNC, 1 in MNHN, 1 in NHMUK), on 29.vii.1996 (1♀) , on 06.viii.1996 (1♀), on 08.viii.1996 (1♀), on 19.x.1996 (1♀). Hautes-Pyrénées : Gèdre, Héas, 1450-1500 m, ex pupa of Zygaena filipendulae , collect 24.vii.1978, emergence 24.viii.1998 (Tremewan W.G. & S.M.) (1♀) . Hérault : Coursan, ex pupa of Zygaena sarpedon , 01.vi.1993 (Hofmann A.) (1♂) . Indre : Pouligny-Saint-Pierre, Les Veillons, ex pupa of Zygaena filipendulae , collect 05.viii.1979, emergence 23.viii.1979 (Drouet E.) (1♂) , emergence on 24.viii.1978 (1♂); same data but collect on 04.viii.1978, emergence 05.ix.1978 (Drouet E.) (1♀) . Isère : Le Quichat, Mont Rachais, ex pupa of Zygaena transalpina alpina , 12.ix.1983 (Drouet E.) (2♀ 1♂) . Landes : Dax, Labenne-Océan (N.), 5-10 m, ex pupa of Zygaena filipendulae , collect 20.ix.1999, emergence 22.ix.1999 (1♀) (Tremewan W.G. & S.M.) ; on 29.ix.1999 (1♀), on 30.ix.1999 (1♀), on 02.x.1999 (2♀). Loire-Atlantique : Guérande, Kervin, ex pupa of Zygaena filipendulae , 08.vii.1991 (Rouch A.) (1♀) ; Guérande , Sandun, ex pupa of Zygaena filipendulae , collect 04.vi.1980, emergence 29.vi.1980 (Drouet E.) (1♀) ; same locality, ex pupa of Zygaena trifolii , 28.vi.1992 (Rouch A.) (1♂; in MNHN) ; Le Grave, Forêt du Grave , ex pupa of Zygaena trifolii , collect 16.v.1976 (Rouch A.) (1♀) ; Saint-André-des-Eaux, Le Gros Chêne , ex pupa of Zygaena trifolii , collect 02.vii.1991 (Rouch A.) (1♂) . Lot-et-Garonne : Villeneuve-sur-Lot, Castelnaud-de-Gratecambe, 200 m, ex pupa of Zygaena transalpina , collect 31.v.1998, emergence 20.vi.1998 (Tremewan W.G. & S.M.) (1♀) . Manche : Coutances, Pirou Plage (4 km S.), 5 m, ex pupa of Zygaena filipendulae , collect 17.viii.1995, emergence viii.1995 (Tremewan W.G.) (2♀) ; same locality, ex pupa of Zygaena trifolii , collect 08.vi.1998, emergence 24.vi.1998 (Tremewan W.G. & S.M.) (1♀ 1♂) ; same data but collect on 08.vi.1998, emergence on 28.vi.1998 (1♀) , on 22.vi.1998 (1♂; in CNC), on 23.vi.1998 (1♀) ; Coutances , Pointe d’Agon, 0-5 m, ex pupa of Zygaena trifolii , collect 08.vi.1998, emergence 24.vi.1998 (Tremewan W.G. & S.M.) (1♂, in NHMUK), same data but emergence on 25.vi.1998 (1♀) , on 27.vi.1998 (1♀). Puy-de-Dôme : Issoire ( NW), 4 km S Champeix, 725 m, ex pupa of Zygaena transalpina , collect 12-13.vii.1998, emergence 28.vii.1998 (Tremewan W.G.) (1♂) . Sarthe : vic. Vibraye, ex pupa of Zygaena transalpina , collect 05.vi.1998, emergence 30.vi.1998 (Nicolle M.) (1♂) . Seine-Maritime : Sauveur Mesnil, Lieubray, ex pupa of Zygaena filipendulae , 09.viii.1990 (Drouet E.) (1♀) ; same data but emergence on 15.viii.1990 (1♀) . Var : Taradeau, ex pupa of Zygaena filipendulae , collect on 28.v.2014, emergence 09.vi.2014 (Kan P. & B.) (1♀, GDEL0388 , in CBGP) . GERMANY: Baden-Wüttemberg : Schwäbische-Alb., Thaxhaim-Onst., ex pupa of Zygaena angelicae elegans , collect 31.vii.1996, emergence viii.1996 (Hofmann A.) (1♀) . ITALY: Potenza : Monte Pollino (N.), Serra Capellino, Voscari (3 km W), 1050-1100 m, ex pupa of Zygaena lonicerae , emergence 14.vi.2003 (1♂) (Tremewan W.G.) ; same date but emergence on 15.vi.2003 (1♂), on 19.vi.2003 (1♀), on 26.vi.2003 (2♀), on 01.vii.2003 (1♀).

NORTH AFRICA. MOROCCO: Fès-Boulmane, Boulemane , Moyen-Atlas, M.F. Tirhboula, 1600 m, ex pupa of Zygaena favonia ssp. thevestis , collect 28.v.1980, emergence 01.vi.1980 (Hofmann A. & Reiss G.) (1♂) . DrâaTafilalet: Ouarzazate, vic. Skoura ex pupa of Zygaena maroccana , emergence vi.1993 (Weiss J.-C.) (4♀ 4♂) .

NEAR EAST. IRAN: Fars: Eqlid (W.), via Timur Gun , 2650 m, ex pupa of Zygaena nocturna , 28.vi.2001 (Tremewan W.G.) (1♀) . Kerman: 50 km N Kerman, Gardaneh-ye Khorasani , 2450-2600 m, ex pupa of Zygaena rubricollis , collect 30-31.v.2002, emergence 21.vi.2002 (Tremewan W.G.) (1♀) ; Mazandarân, Gardaneh-ye Lavashm , 2800-2950 m, ex pupa of Zygaena haberhaueri , collect 29-30.vii.1997, emergence 06.viii.1997 (Tremewan W.G.) (1♀) ; Goulujett , ex pupa of Zygaena tamara (Hofmann A.) (1♀) . TURKEY: Ankara: vic. Beynam , 1200-1250 m, ex pupa of Zygaena laeta , collect 10-13.vii.1999, emergence 24.vii.1999 (Tremewan W.G.) (1♀) . AlaDagh , ex pupa of Zygaena lydia , vii.1995 (Eckweiler W.) (1♂) .

NON TYPE MATERIAL. EUROPE. CZECH REPUBLIC: Moravia, Mer. Or. Poštornǎ. BV. sans by ‘ Františkŭv’ pond, 02.vi.2003 (Kment P.) (1♂, GDPC) . FRANCE: Bouches-du-Rhône, Aix-en-Provence, Europole de l’Arbois , 27.viii.2007 (Ponel P.) (1♀, GDEL0471 , CBGP) . Hérault: Grabels ; 13.vii.1989 (Tussac H.) (1♀, GDPC) ; Lauroux, Cirque de Labeil , 25.v.1983 (Delvare G.) (1♀, GDPC) ; Mireval, Mas d’Andos , 19.vi.1994 (Delvare G.) (1 m, GDPC) ; Montpellier , 03.iv.1989 (Maldès J.-M) (1♀, GDPC) ; Saint-Geniès-de-Varensal, Albès , 785 m, 01.ix.1989 (Delvare G.) (7♀, GDPC) ; same locality and collector, 15.viii.2004 (1♀) ; Viols-le-Fort , 13.v.1995 (Cocquempot C.) (1♀) (both GDPC) . Lot: Comiac , 03.vii.1977 (Tussac H.) (1♂, GDPC) ; Saint-Géry , 28.vii.1989 (Tussac H.) (1♀, GDPC) .

NORTH AFRICA. MOROCCO: Drâa-Tafilalet: Ouarzazate, Anti-Atlas, 1 km N Tizi n’ Tiniffifft , 29.v.1992 (Delvare G.) (7♂, GDPC) . Souss-Massa: Tasserirt , emergence v.1985 [no collector name but handwriting identical with that of J.-C. Weiss] (2♂, NMS) .

NEAR EAST. IRAN: [no locality] ex pupa of Zygaena cuvieri , collect 25.vii.2009, emergence 10.viii.2009 (Hofmann A.) (1♀, NMS) ; [no locality] ex pupa of Zygaena tamara , collect 14.viii.2009, emergence 25.ix.2009 (Hofmann A.) (1 ♂, NMS) ; [no locality] ex pupa of Zygaena tenhagenova , viii.2009 (Hofmann A.) (1♂, NMS) . TURKEY [no locality] ex pupa of Zygaena filipendulae, 1996 (Hofmann A.) (1♀, NMS) . Bingöl: Genç, Karhova, Merkez , 1046-1885 m, 05.v-03.vi.2019, on flowers of various plants (Kaplan E.) (6♀ 1♂, BU) ; Diyarbakar: Ҫermik, Hani, Lice, Silvan , 777-1188 m, on flowers of various plants (Kaplan E.) (5♀, BU) .

Description. FEMALE (holotype). See relevant parts above relative to tibialis species-group characters ( Figs 2A–I, 2A – 3D ). Characters that distinguish females of B. zygaenae from B. tibialis include: mesepisternum with adscrobal area entirely areolate ( Fig. 8E ); metacoxa without or with vestigial inner ventral tooth ( Fig. 8F ); metafemur broad, 1.55× as long as wide and densely punctulate, the interspaces about as large as the diameter of the punctures, with dorsal and ventral margins of femur similarly curved and teeth on ventral margin with same appearance throughout, appearing as equilateral triangles; metatibia with edge of apical truncation somewhat concave ( Fig. 8G ); hind tarsus slender, 4.35× as long as width of metatibia ( Fig. 8G ); hind pretarsus with falcate seta narrow, not broadening from base ( Fig. 8H ). Additional characters are as follows:

Colour. Body black, including mandibles, antenna, coxae, trochanters, basal two thirds of profemur, meso- and metafemora except apices, outer ventral spots on pro- and mesotibiae, ventral strip broadening mesally on metatibia; tegula and rest of legs bright yellow; venation dark brown; setation of wings black ( Fig. 1 ).

Setation. Setae thin, filiform and suberect on frons and dorsum of mesosoma, not visible from above because orientated subvertically ( Fig. 2H ); setae longer on mesoscutellum, especially in front of frenal carina where adpressed ( Fig. 2I ), setae on propodeal callus and postspiracular region, thicker, also longer on metepimeron ventrally ( Figs 10D, E ). Gt2‒Gt5 with moderately thick and adpressed setation, distributed in 2‒3 intricate rows on each tergite ( Fig. 3D ).

Measurements. Head respectively 2.3× and 1.34× as wide as long and high; POL and OOL respectively 1.9× and 0.67× as long as diameter of lateral ocellus; temple 0.26× as long as eye; eyes separated by 0.94× their height; malar space 0.34× as long as eye height and 0.43× as oral fossa width; antennal scrobes 1.22× as long as wide; distance from ventral edge of antennal toruli to lower ocular line 0.18× as long as antennal scrobes. Antenna with scape, pedicel, F1 and F7, respectively, 4.15×, 0.83×, 0.90× and 0.68× as long as wide; F1 1.18× as long as pedicel; clava 1.73× as long as wide. Mesosoma 1.38× as long as wide, mesoscutellum as long as mesoscutum and 0.97× as long as wide. Hind leg with metacoxa and metafemur respectively 1.87× and 1.55× as long as wide; hind tarsus 4.35× as long as metatibia width. Fore wing 2.87× as long as wide, costal cell 1.11× as long as wing width, marginal vein half as long as costal cell; postmarginal vein respectively 0.55× and 2.55× as long as marginal and stigmal veins. Metasoma with gaster 0.94× as long as mesosoma and 1.4× as long as wide; Gt1 0.34× as long as gaster.

Structure. Labrum semicircular, its surface hardly concave, setose on ventral half. Clypeus as low triangle, with step-like upper margin and 3 rows of setiferous points ( Fig. 2F ). Frons, lower face and temples rugulose-areolate; gena below postgenal groove with finer sculpture than rest of head, with small setiferous points behind malar groove and areolate-rugulose near genal carina ( Figs 2F , 10B ); vertex coriaceous on interspaces between punctures ( Fig. 10A ); genal carina faintly continuing on occiput. Antenna with scape somewhat narrowing subapically, with a row of setae along either side of ventral margin, the scape apex reaching median ocellus; anellus discoidal ( Fig. 2G ); flagellomeres with several intricate rows of short MPS and numerous short, suberect setae ( Fig. 10C ).

Pronotum with pronotal carina restricted to sides. Pro- and mesonotum regularly punctured ( Fig. 2H ). Notauli as impressed punctured lines, their apices facing axillar grooves on transscutal line, the latter also appearing as punctured impressions. Mesoscutellum strongly sloping subapically, its punctuation similar to that of mesoscutum. Edge of frenal carina uniformly curved, preceded by a row of areolae. Propodeum sloping at an angle of about 70° with dorsum of mesonotum, entirely areolate, with a median areola and a sublateral projection at base of a setose postspiracular region ( Figs 10D, E ). Femoral depression of mesepisternum entirely strigose dorsally ( Fig. 8E ). Mesodiscrimen with projection separating its dorsal and ventral sections moderately expanded ( Fig. 3B ). Metepisternum with bulging coriaceous adpetiolar region ( Fig. 3B ). Metepimeron coarsely areolate ( Fig. 1 ).

Fore wing ( Fig. 1 ) with basal cell mostly bare except a few setae apically; speculum visible as a narrow strip behind parastigma; costal cell entirely setose ventrally; marginal fringe absent apically; parastigma with 3 sensilla, uncus with 4 aligned sensilla.

Metasomal petiole with exposed ventral part longitudinally strigose ( Fig. 3C ). Gaster shortly acute ( Fig. 3D ). Gt2 with 3 rows of setiferous points basally on anterior half of mesal surface, bare and delicately punctulate beyond. Gt3‒Gt6 entirely setose dorsally and laterally. Syntergum very short. Outer plate with 2‒3 rows of coarse setiferous punctures. Tip of hypopygium level with hind margin of Gt5.

MALE. Characters separating males of B. zygaenae from B. tibialis include special, spatulate setae present on F1‒F5, with F1 having 8‒10 and their number decreasing from 12‒15 on F2 to 4‒6 on F5 ( Fig. 9D ). Otherwise, the male is mostly similar to the female except all parts of the hind leg are more slender ( Figs 9E, 9F ).

Morphological variation. Variation is evident mainly in the intensity of sculpture of various body parts: gena sometimes with more delicate sculpture, with only small setiferous punctures; adscrobal area of mesepisternum occasionally not completely areolate; femoral groove also not entirely strigose; propodeum without postspiracular projection. Such variation is exhibited by specimens reared from the same host at the same time and thus represent individual variation among the same populations. Different variation, apparently of a regional nature, occurs in both sexes of specimens reared or collected in Morocco in the Northern slope of the Anti-Atlas. One series was reared near Skoura in the Vallée des Roses, the other was collected independently near the pass separating the northern from the southern slope of the Anti-Atlas with a reduced distance between the two sites. In these specimens the hind tarsus is shorter and thicker in both sexes, tending to the habitus of B. tibialis ( Figs 10F–I ). All other characters of the legs and of the male flagellum agree with those described above for B. zygaenae ; in addition, the two sexes were reared from Zygaena maroccana .

Distribution ( Table 1 ). The species is present in North Africa ( Morocco), southern Europe ( France, Italy, Spain), Central Europe ( Czech Republic and Germany) and Near East ( Turkey). It is also known from England ( Shaw et al. 2010, as B. tibialis ) but its presence here appears to be the result of recent establishment. The most easterly known distribution is Iran; it occurs up to 2950 m in Iran and 1600 m in Morocco. The known distribution of B. zygaenae indicates it might be better adapted to prolonged winter conditions than B. tibialis .

Hosts and biology. When only a single date is given in the above list of reared specimens it is usually a date of collection (although ambiguous) and interpreted as such here. All the reared specimens examined were from cocoons of various Zygaena species ( Table 1 ), almost always as a primary parasitoid of the Zygaena pupa. The preponderance of host records from Zygaena species whose cocoons are formed high up in field layer vegetation, such as Z. filipendulae , Z. lonicerae , Z. transalpina and Z. trifolii , is likely to be no more than a sampling artefact and we suspect that cocooned pupae of all Zygaena species would be potentially susceptible to being parasitized by this species. Only a single male was reared from a cocoon of Charops sp. [undoubtedly C. cantator (DeGeer) ] ( Ichneumonidae : Campopleginae) which had parasitized the larva of Zygaena filipendulae and made its cocoon within that of the Zygaena host. Thus, secondary parasitism can occur but is probably exceptional for this species. The adults emerge, always in the year of host collection, from June to early October. Host cocoon collection dates range from May to September, and in some localities (e.g. in southern Europe) bivoltine Zygaena species are found. From the rearing data it is clear that B. zygaenae is plurivoltine under suitable conditions. One female collected at the beginning of April, and others in early May, further strongly suggest that hibernation is as an adult.

Figures

FIGURE 1. Brachymeria zygaenae Delvare & Shaw sp. n., holotype ♀. Habitus in lateral view.

FIGURE 2A–I. Characters of Brachymeria: 2A–C, characters of Brachymeriinae; 2D–I, characters of the Brachymeria tibialis species group. 2A–C and 2E–I, Brachymeria zygaenae, holotype ♀: 2A, antennal clava; 2B, hind pretarsus; 2C, hamuli; 2E, head in lateroventral view; 2F, head in frontal view; 2G, pedicel and base of flagellum; 2H, mesosoma in dorsal view; 2I, mesoscutellum and axillae in dorsal view. 2D, B. zygaenae, paratype ♂, flagellum in ventral view. Abbreviations: fc, falcate seta; poc, postorbital carina.

FIGURES 3A–D. Characters of the Brachymeria tibialis species group, Brachymeria zygaenae, holotype ♀: 3A, mesopleuron in anterior view; 3B and 3C, ventral side of mesosoma, respectively in lateroventral and ventral views; 3D, metasoma in dorsal view. Abbreviations: cx3-f, metacoxal foramen; dm2-ds, mesodiscrimen, dorsal section; dm2-vs, mesodiscrimen, ventral section; dm3-as, metadiscrimen, anterior section; dm3-is, metadiscrimen, intermediate section; dm3-ps, metadiscrimen, posterior section; epcm, epicnemium; epcm-c, epicnemial carina; epcm-ce, submedian elevations of epicnemial carina; epst2- vsf, ventral shelf of mesepisternum; epst3, metepisternum; pet, petiole; ppt-vb, ventral belt of prepectus; smd, submedian depressions.

FIGURES 8A–H. Comparative characters of B. tibialis and B. zygaenae. 8A–D, B. tibialis, ♀. 8E–H, Brachymeria zygaenae, holotype ♀. 8A and 8E, mesopleuron in lateral view with adscrobal area enlarged; 8B and 8F, hind leg with sculpture of metafemur enlarged; 8C and 8G, apex of metatibia and hind tarsus; 8D and 8H, hind pretarsus. Abbreviation: fc, falcate seta.

FIGURES 9A–F. Comparative characters of B. tibialis and B. zygaenae. 9A–C, B. tibialis, ♂. 9D–F, Brachymeria zygaenae, paratype ♂. 8A and 8D, antennal flagellum with base enlarged; 8B and 8E, hind leg with sculpture of metafemur enlarged; 8C and 8F, apex of metatibia and hind tarsus.

FIGURES 10A–I. Characters of B. zygaenae. 10A–E, Brachymeria zygaenae, holotype ♀. 10F–I, Brachymeria zygaenae, special phenotype from southern Morocco: 10F–G, ♀, 10H–I, ♂. 10A and 10B, head, respectively in dorsal and lateral views; 10D and 10E,. propodeum, respectively in posterior and laterodorsal views; 10F and 10H, hind leg with sculpture of metafemur enlarged; 10G and 10I, apex of metatibia and hind tarsus.