Spalangia nigra Latreille, 1805

Gibson, Gary A. P., 2009, 2259, Zootaxa 2259, pp. 1-159 : 109-115

20. Spalangia nigra Latreille, 1805

(Figs 298–317)

Spalangia nigra Latreille, 1805: 228 ; lectotype ♀ (MRSN, not examined) designated by Bouček (1963: 445). Type data: [ France] Paris.

Spalangia hirta Haliday, 1833: 334 ; lectotype ♀ (OXUM, not examined) designated by Graham (1969: 52). Type data: England. Synonymy by Bouček (1963: 443).

Spalangia rugosicollis Ashmead, 1894: 35 , 36; holotype ♀ (USNM, examined). Type data: Missouri. Synonymy by Bouček (1963: 443).

Description. Female. Length = 2.1–3.6 mm. Legs dark except at least basal 3 tarsal segments yellow and sometimes knees and apex of tibiae narrowly yellow. Head in anterior view (Fig. 299) about 1.1–1.2x as high as wide; in dorsal view (Fig. 304) about 1.6–1.9x as wide as long; in lateral view (Fig. 300) with malar space about 0.9–1.0x eye height and about 1.2–1.4x eye width. Head capsule smooth and shiny except for setiferous punctures as follows: with complete median sulcus extending ventrally to elongate-triangular scrobal depression, otherwise upper face and parascrobal region densely punctate, the punctures sometimes separated by about own diameter but more often by less than own diameter and sometimes so crowded as to be more or less multisided with ridge-like interstices, at least laterally; scrobal depression with scrobes broadly, transversely to obliquely carinate-crenulate at least ventrally above torulus (Fig. 299), the inclined lateral surface of depression otherwise punctate similar to parascrobal region, but interantennal region smooth and shiny; gena without distinct malar sulcus, with crowded punctures separated by linear ridges; temple variably densely punctate similar to upper face. Antenna (Fig. 312) with scape about 8.3–9.2x as long as greatest width, the outer surface (Fig. 314) uniformly setose and longitudinally strigose, but inner surface (Fig. 313) bare mediolongitudinally where usually more distinctly longitudinally striate; pedicel about 2.2–2.7x as long as apical width and about 1.4–1.8x as long as fu 1; funicle with fu 1 about 1.3–1.9x as long as wide, fu 2 at least quadrate and usually at least slight longer than wide, and apical segments quadrate to up to about 1.4x as wide as long; clava about 2.3–2.6x as long as wide.

Fig. 298. Distribution of Spalangia masneri Gibson and S. nigra Latreille.

Pronotal collar in lateral view convexly arched behind neck and anterolaterally with a short smooth bar interrupting circumpronotal furrow, but anteriorly smoothly rounded to neck; usually completely or almost completely punctate without distinctly differentiated cross-line posteriorly, except often medially where collar smooth or at least less densely punctate anterior to cross-line (Fig. 306), and dorsally with punctures becoming increasingly crowded so as to be more or less multisided or rugose anteriorly and/or laterally, but laterally sculpture smoothly rounded to circumpronotal furrow above lateral panel. Mesoscutal median lobe (Fig. 308) with anterior convex region mostly smooth and shiny, though sometimes finely coriaceous posteriorly; internotaular region completely punctate-rugose except often for irregular median carina. Axillae (Fig. 308) usually with distinct punctures but sometimes setae originating from only pinprick-like pores. Scutellum (Fig. 308) more or less uniformly covered by distinct, circular setiferous punctures lateral to median smooth band; frenum (Fig. 308) differentiated by complete crenulate frenal line. Mesopleuron (Fig. 310) smooth and shiny except as follows: pectal region crenulate along anterior margin and bare except for 1 posteroventral seta; acropleuron longitudinally carinate, the carinae extending posteriorly onto alar shelf and variably extensively, sometimes completely, over upper mesepimeron; subalar scrobe a large, reticulate-rugose, triangular region widened posteroventrally along transepisternal line; episternal scrobe an arcuate, crenulate-reticulate furrow connected to subalar scrobe by punctate-crenulate episternal line and to precoxal scrobe by punctate precoxal

Figs 299–307. Spalangia nigra Latreille. 299–305, head: 299, anterior view ♀, 300, lateral view ♀, 301, lateral view ♁, 302, anterior view ♁, 303, anterior view ♁, 304, dorsal view ♀, 305, dorsal view ♁; 306 & 307, pronotum, dorsolateral view: 306, ♀, 307 ♁.

Figs 308–317. Spalangia nigra Latreille. 308 & 309, thorax, dorsal view: 308, ♀, 309, ♁; 310, ♀ mesopleuron; 311, ♁ frenum–petiole, dorsal view; 312, ♀ antenna; 313 & 314, ♀ scape: 313, inner view, 314, outer view; 315, ♁ antenna; 316 & 317, ♁ scape: 316,

inner view, 317, outer view.

line; upper and lower mesepisternum differentiated by punctate-crenulate transepisternal line and adjacent line of setae. Fore wing hyaline; variably extensively setose behind submarginal vein but mediocubital fold with at least a few setae. Propodeum (Fig. 311) with crenulate postspiracular sulcus differentiated from callus; callus completely punctate-rugose; plical region with variably distinctly widened paramedian crenulate furrows delineating median carina, but at least anterior-most cell obviously wider and larger than more posterior cells; supracoxal band contiguous with paramedian crenulate furrows; propodeal panels smooth and shiny.

Petiole about 2x as long as medial width; transversely carinate to reticulate-rugulose between longitudinal carinae; with several setae laterally. Gaster with tergites smooth and shiny or with only extremely obscure coriaceous sculpture.

Male. Length = 1.5–2.9 mm. Antenna (Fig. 315) with scape about 5.2–7.1x as long as wide, the sculpture usually less distinctly strigose, particularly outer surface (Fig. 317) often more coriaceous-rugulose and inner surface (Fig. 316) sometimes smooth and shiny medially; pedicel subglobular, at most about 1.3x as long as wide; flagellum with setae much shorter than width of respective segment; funicle with fu 1 about 1.7–2.6x as long as wide and about 1.5–3.6x as long as pedicel, and subsequent funicular segments usually all obviously (about 1.5x) longer than wide, but in small individuals middle or most segments sometimes only slightly longer than wide. Otherwise similar to female except as follows. Head in anterior view (Figs 302, 303) about 0.9x as high as wide; in lateral view (Fig. 301) with malar space only about 0.6–0.7x eye height and about 0.8–1.0x eye width. Head capsule sometimes much more sparsely punctate than female, upper face and parascrobal region sometimes with punctures separated by up to about 3x own diameter (Fig. 303) and gena sometimes smooth and shiny near lower orbit but without malar sulcus. Pronotum often with quite large smooth region posteromedially and therefore with more distinctly differentiated cross-line than most females, but at least rugose-punctate anteriorly and laterally (Figs 307, 309). Scutellum quite commonly with relatively few, often tiny setiferous punctures laterally (Fig. 309). Petiole (Fig. 311) about 2.2–2.8x as long as medial width.

Material examined. Nearctic and Neotropical (884 specimens in AEIC, CASC, CLEV, CNC, CUAC, CUIC, DEBU, DENH, FSCA, IRCW, MCZH, NCSU, OSAC, OSUC, ROME, TAMU, UCDC, UCFC, UCRC, UMRM, UNML, USNM, WSUC). Complete collection records are not provided; those with host data: Nearctic. CANADA: New Brunswick, Moncton, St-Léonard-Parent Beaupré Farm, 26.VI.03, 9, 18, 30.VII.03, T. Levesque, M. domestica . Ontario, Guelph, 18.IX.78, D. Levin, Hylemyia antiqua (DEBU) . Fitzroy Harbour, Weir farm, 45º29.349'N 76º11.938'W, G. Gibson & L. Bartels — 26.VI.01, 7.VIII.01,? Hydrotaea (Ophyra) leucostoma ; 11.IX.01 (dissected 13.XI.01),? Phormia regina . Ottawa, Merivale Rd, 45º18.926'N 75º42.762'W, 23.VIII.01 (1 dissected 26.X.01), G. Gibson & L. Bartels,? Phormia regina . Ottawa, Prince of Wales Dr., McEwen farm, 45º18.763'N 75º42.193'W, 14.VI.01, 9.VIII.01 (dissected 6.XI.01), 16.VIII.01 (dissected 15.X.01), G. Gibson & L. Bartels,? Hydrotaea (Ophyra) leucostoma . Ottawa vicinity, various localities and dates 2000 and 2001, G. Gibson & L. Bartels, M. domestica and S. calcitrans (see Gibson and Floate 2004). Styles Side Rd NW West Carleton, Clarence farm, 45º26.622'N 76º10.296'W, dissected 4.XII.01, G. Gibson & L. Bartels,? Phormia regina . Yorks Corners Road, S Kenmore, Donevelyn farm, 45º 11.758'N 75º 23.547'W, 5.VII.01, G. Gibson & L. Bartels,? Hydrotaea (Ophyra) leucostoma . Quebec, Luskville, Alary farm, 45º31.655'N 76º02.898'W, 28.VIII.01 (dissected 29.X.01), G. Gibson & L. Bartels,? Hydrotaea (Ophyra) leucostoma . St. Jean, 15, 22.IX.53, P. Perron, Hylemyia antiqua . USA: Florida, Alachua Co., Gainesville, 19.VI.75, R.L. Escher, H. irritans (FSCA) . Illinois, Libertyville — 25.VII.68, E. Legner, Stomoxys sp. ( UCRC); reared in lab., CES Riverside, VIII, IX.68, M. domestica (UCRC) . Kansas, Wellington, 2.XI.12, S. calcitrans (USNM) . Mississippi, Starkville, 11.VIII.72, K.J. Watts, Ravinia derelicta (USNM) ․ Missouri, Boone Co., Columbia, D.E. Figg, voucher specimens ( UMRM) — 8, 14, 18, 25, 28.IX.81, Orthellia caesarion ; 23.IX.81, Paregle cinerella ; 14, 17.VIII.81, 2.X.81, Ravinia sp. ; 14, 23, 25.IX.81, Anthomyiidae . Nebraska, Lincoln, 1962, C. McCoy, M. domestica (USNM) . New York, Cayuga Co., 14.V.86, 1, 3, 16.VII.86, L. Smith, M. domestica (USNM) . Tompkins Co., 8.IX.87, C. Henderson, M. domestica (CUIC) . North Carolina, Alamance Co., Sutphin, 18.VIII.78, T.D. Edwards, M. domestica (CUIC) . Chatham Co., Silk Hope, 29.IX.76, D.A. Rutz, M. domestica (CUIC) . Lenoir Co., Deep Run, 24.VIII.78, T.D. Edwards, M. domestica (CUIC) . South Carolina, Oconee Co., South Union, 12.VI.73, M. domestica (CUAC) . Texas, Dallas, 1, 24.XI.12, S. calcitrans (USNM) . Gainesville, 31.V.13, S. calcitrans (USNM) . Virginia, Blacksburg, 10.VIII.67, R.P. Burton, Musca autumnalis (USNM) .

Distribution. A cosmopolitan species that Noyes (2003) records from all six biogeographic regions, including West Indies ( Barbados, St. Vincent & Grenadines, Trinidad & Tobago) and South America ( Brazil, Chile, Peru). However, I only saw specimens from North America (Fig. 298), including just a single specimen from south of the USA labelled “ Mexico: Nuevo Leon, Mun. Guadalupe, Rincón de la Sierra, 11.VII.83, A. Gonzalez H.” (UCRC). The absence of specimens from the extensive rearings by Loera-Gallardo et al. (2008) in northeastern Mexico near Guadalupe suggests that S. nigra is at most a very rare naturally occurring species south of the USA.

Biology. Noyes (2003) lists S. nigra as a primary parasitoid of at least 30 species in 10 families of Diptera plus other taxa in Coleoptera , Lepidoptera and Hymenoptera , likely as hyperparasitoids through Tachinidae primary parasitoids. Specimens I saw from the New World indicate S. nigra is a parasitoid of the three principal filth-fly pests of livestock, the house fly, stable fly and horn fly, as well as the face fly, Musca autumnalis De Geer , the onion maggot, Delia (= Hylemia) antiqua (Meigen) , and Neomyia cornicina (Fabricius) (= Orthellia caesarion ), Adia (= Paregle ) cinerella (Fallén) and Ravinia derelicta (Walker) . The tentative records of Hydrotaea (Ophyra) leucostoma (Wiedemann) and Phormia regina (Meigen) by Gibson and Floate (2004) were based on comparison of pupal remains correlated with flies emerged from similar puparia.

Recognition. I include S. nigra along with S. alyxia , S. chontalensis , S. masneri , S. nigroides and S. nigroaenea in the nigra species group. These six New World species share a coarsely sculptured pronotum and a petiole with several setae laterally, in contrast to cameroni -group species, which have a coarsely sculptured pronotum but a bare petiole. Spalangia simplex and S. dozieri also have a laterally setose petiole, but S. simplex is distinguished by its pronotal collar being smooth anterior to the cross-line and by its unique internotaular sculpture pattern (Fig. 412). Spalangia dozieri is more similar to such species as S. drosophilae because it has a median sculptured propodeal band (Fig. 104) and its mesopleuron is extensively sculptured (Fig. 106). Spalangia nigra and the other five species of the group have a median propodeal carina and the mesopleuron is mostly smooth and shiny between the various scrobes and connecting lines of sculpture except sometimes for a longitudinally carinate upper mesepimeron.

Within the group of six species, S. nigra and S. nigroides have the subalar scrobe extending broadly along the transepisternal line so that it is more or less triangular in shape (Figs 310, 370). Other nigra -group species have a more vertical subalar scrobe or one that is widest medially and narrowed anteroventrally so that its posteroventral margin and transepisternal line form an abrupt or acute angle (Figs 34, 35, 87, 88, 290). Males of S. nigroides are unknown, but S. nigra females differ from those of S. nigroides mostly only in the relative features given in the key. The pronotum of S. nigroides in lateral view is also flatter and anteriorly less strongly convex (Fig. 367) than for S. nigra females (Fig. 306). Most females of S. nigra also have the anterior-most cell of the paramedian crenulate furrows larger and more obviously differentiated from the more posterior cells than for the two known females of S. nigroides . If mesopleural sculpture is not visible, S. nigra could be mistaken for S. chontalensis or S. nigroaenea , both of which sometimes have a coarsely punctate scutellum. However, unlike the latter species the pronotal collar of S. nigra lacks a distinct vertical carina anterolaterally and anteriorly is evenly rounded to the neck (Figs 306, 307). Both sexes of S. nigra have at least some setae on the mediocubital fold and the anterior convex region of the median mesoscutal lobe smooth and shiny except sometimes posteriorly adjacent to the sculptured internotaular region (Figs 306, 307), whereas both sexes of S. masneri have the mediocubital fold bare and both sexes of S. alyxia have the mesoscutal median lobe anterior convex region completely sculptured (Figs 31, 33).

Figs 318–326. Figs 318–320, Spalangia rugulosa Förster ♀. 318, head, lateral view; 319, head, frontolateral view; 320, pronotum and mesoscutum, frontodorsal view. Figs 321–324, Spalangia irregularis Bouček ♀. 321, head, frontolateral view; 322, pronotum and mesoscutum, frontodorsal view; 323 & 324, scape: 323, inner view, 324, outer view. Figs 325 & 326, S. rugulosa ♀, scape: 325, inner view, 326, outer view.

Bouček (1963) included in his nigra -group S. irregularis Bouček and S. rugulosa Förster , two other Old World species that are very similar to S. nigra . My concept of S. irregularis is based on the one female and two male paratypes from Cyprus deposited in the BMNH by Bouček (1963), two ZSMC females ( Switzerland: Collonge-Bellerive, 23.VII.66, P.P. Babiv; Corsica: Taravotal, 3 km NE Porto Pollo, 27.II.68), one NMPC female ( Cyprus: Platus, 18.VIII.37, Mavromoustakis) and one CNC male ( France: Montpellier, 14.IV.78, J. Huber), whereas my concept of S. rugulosa is based on one NMPC female ( Serbia: Ruma, Dr. Hensch.), one ZSMC female ( Germany: Munich, 18.V.66, W. Schacht), and one USNM male ( France: Yvelines, Bouafle, 21.VI.79, CHOUX Lab.: 13.VII). As described by Bouček (1963: 447), the “antennal socket [is] abruptly raised” in S. nigra but not in S. rugulosa or S. irregularis . Females of S. nigra have the torulus at the end of a short but quite distinct, obliquely projecting tube formed in part from the lower face (Fig. 300), whereas females of S. rugulosa (Fig. 318) and S. irregularis have the lower face at almost a right angle to the face so that the torulus is only rim-like and non-protuberant. This feature is not as obvious for males because males of S. nigra , like males of most species, do not have the lower face protruding as conspicuously into a lobe as for females (cf. Figs 300, 301). The pronotal collar of S. rugulosa (Fig. 320) and S. irregularis (Fig. 322) is also more coarsely sculptured than that of S. nigra (Fig. 306), being more uniformly reticulate-rugulose with all the setiferous punctures, even posteromedially, irregular and separated by linear ridges. Furthermore, individuals of S. nigra that have crowded punctures on the face similar to the densely punctate to punctate-rugose face of S. irregularis or S. rugulosa have the scrobes and inclined surface of the scrobal depression above the scrobes transversely to obliquely carinate-crenulate (Figs 299, 302), whereas the latter two species have the scrobes only narrowly crenulate ventrally so that the inclined portion of the scrobal depression is reticulate-rugose even above the toruli (Figs 319, 321). However, sculpture of the scape probably most readily distinguishes females of S. nigra from those of S. rugulosa and S. irregularis . Both surfaces of the scape of female S. nigra (Figs 313, 314), but particularly the bare inner surface (Fig. 313), are distinctly longitudinally striate-strigose, whereas the scape is punctate-rugose to reticulate-punctate in S. rugulosa (Figs 325, 326) and S. irregularis (Figs 323, 324) as observed originally by Bouček (1963).