Spalangia nigroaenea Curtis, 1839

Gibson, Gary A. P., 2009, 2259, Zootaxa 2259, pp. 1-159 : 122-127

22. Spalangia nigroaenea Curtis, 1839

(Figs 346–365)

Spalangia nigroaenea Curtis, 1839 : folio 740; holotype ♁ (ANIC, not examined). Type data: [ England], in Mr. Shuckard ̕s collection.

Spalangia homalaspis Förster, 1850: 505–507 ; lectotype ♁ (NHMW, not examined) designated by Bouček (1963: 451). Type data: [ Germany], Aachen and Boppard [lectotype: Or. Ex., Boppard Or. Ex. ♁]. Synonymy by Bouček (1963: 450).

Spalangia astuta Förster, 1851: 1–2 ; lectotype ♀ (NHMW, not examined) designated by Bouček (1963: 451). Type data: [ Germany], Aachen [lectotype: ♀ Or. Ex., Collect. G. Mayr]. Synonymy by Bouček (1963: 450).

Spalangia muscidarum Richardson, 1913: 38–39 ; holotype ♁ (USNM; 1♀, 3♁ paratypes examined). Type data: Forest Hills, Mass. [Massachusetts], XII 1912. Synonymy by Bouček (1963: 450).

Prospalangia platensis Brèthes, 1915: 315–317 ; ♀, ♁ syntypes (MLPA, not examined) (2♁ syntypes remain according to Loiácono et al. 2007). Type data: [ Argentina], Général Urquiz, near Buenos Aires, ex Musca domestica L., Stomoxys calcitrans , etc. Synonymy by Bouček (1965: 600).

Spalangia abenabooi Girault, 1932: 1 ; ♀ syntypes (QMBA, not examined). Type data: W. Aust. to Queensland, Johnston and Tiegs, 1921. Tentative synonymy by Bouček (1963: 448) confirmed by Bouček (1988a: 342).

Spalangia sundaica Graham, 1932: 22 ; holotype ♀ (BMNH, examined). Type data: Java, Buitenzorg, 12 th November, 1929, G.L. Windred. Synonymy by Bouček (1963: 450).

Spalangia mors Girault, 1933: 1 ; lectotype ♀ (QMBA, not examined) designated by Bouček (1988a: 342). Type data: [ Australia], Mackay, III, W.A. McDougall. Synonymy with S. abenabooi by Girault (1934: 3); tentative synonymy with S. nigroaenea by Bouček (1963: 448) confirmed by Bouček (1988a: 342).

Description. Female. Length = 2.0– 3.5 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. 346) about 1.2–1.3x as high as wide; in dorsal view about 1.5–1.7x as wide as long; in lateral view (Fig. 347) with malar space about 0.7– 0.8x eye height and about 1.0–1.1x eye width. Head capsule (Figs 346, 347) smooth and shiny except for setiferous punctures as follows: with complete median sulcus extending to elongate-triangular scrobal depression, otherwise upper face and parascrobal region with distinct but variably dense circular punctures, the punctures relatively sparse and separated by about 2–3x own diameter in smaller specimens but in larger specimens mostly only by about own diameter medially and even more crowded laterally toward inner orbit where usually separated by less than own diameter; scrobal depression usually with scrobes punctate-crenulate or in smaller specimens more reticulate-rugulose and sometimes distinct only near torulus, with smooth and shiny interantennal region, and inclined lateral surface of depression punctate similar to parascrobal region; gena without distinct malar sulcus, with distinct circular to oval punctures often separated by almost own diameter but at least by narrow, flat interstices; temple with distinct circular punctures similar to face and gena. Antenna (Fig. 357) with scape about 7.0–9.0x as long as greatest width, the inner surface bare, smooth, and shiny mediolongitudinally over at least basal half, but longitudinally strigose apically (Fig. 361) and outer surface uniformly setose and punctate (Fig. 362); pedicel about 2.4–2.9x as long as apical width and about 1.5– 1.9x as long as fu 1; funicle with fu 1 about 1.3–1.8x as long as wide, fu 2 at least very slightly longer than wide, and apical segments quadrate to slightly longer than wide; clava about 2.0–3.1x as long as wide.

Pronotal collar in lateral view quite abruptly but comparatively low convex behind neck, anterolaterally with vertical carina or ridge extending from circumpronotal furrow onto collar (Figs 351, 352) and, more or less conspicuously, across collar as low, almost uniform, usually crenulate but medially interrupted margin (Figs 351, 352), the margin differentiating a mostly smooth and shiny vertical surface of collar above neck and in dorsal view usually appearing ^-like (Fig. 350); with distinct cross-line posteriorly, sometimes with very shallow mediolongitudinal furrow (Fig. 351), and smooth and shiny medially anterior to cross-line, but otherwise with mostly well separated circular setiferous punctures except laterally where sometimes rugose (Figs 350, 352), but the sculpture smoothly rounded to circumpronotal furrow above lateral panel (Fig. 351). Mesoscutal median lobe (Fig. 353) with anterior convex region and median band of internotaular region smooth and shiny, the internotaular region otherwise with crowded, circular to irregular setiferous punctures.

Figs 346–355. Spalangia nigroaenea Curtis. 346–349, head: 346, anterior view ♀, 347, lateral view ♀, 348, lateral view ♁, 349, frontolateral view ♁; 350–352, pronotum: 350, dorsal view ♀, 351, frontolateral view ♁, 352, dorsolateral view ♀; 353 ♁ thorax, dorsal view; 354, ♁ mesopleuron; 355, ♀ frenum–petiole, dorsal view.

Figs 356–364. Figs 356–362, Spalangia nigroaenea Curtis. 356, ♁ frenum–petiole, dorsal view; 357 & 358, antenna: 357, ♀, 358, ♁; 359 & 360, ♁ scape: 359, inner view, 360, outer view; 361 & 362, ♀ scape: 361, inner view, 362, outer view. Figs 363 & 364, Spalangia slovaca Bouček , ♀ scape: 363, inner view; 364, outer view.

Axillae (Fig. 355) mostly smooth and shiny to extensively covered with shallow setiferous punctures. Scutellum (Fig. 353) usually smooth and shiny except for a few pinprick or shallow setiferous punctures laterally; frenum (Figs 353, 355) differentiated by complete crenulate frenal line. Mesopleuron (Fig. 354) smooth and shiny except as follows: pectal region variably distinctly crenulate along anterior margin and bare except for 1 posteroventral seta; acropleuron longitudinally carinate, the carinae extending posteriorly onto alar shelf and sometimes dorsally onto upper mesepimeron; subalar scrobe reticulate-rugulose and more or less vertical or narrowed ventrally such that its posteroventral margin and transepisternal line form abrupt angle (Fig. 354); episternal scrobe a lunate depression usually differentiated into a horizontal anterior portion and more vertically directed posterior portion, and either separated from subalar and precoxal scrobes or connected by only very shallow, inconspicuous furrow or line of minute punctures; upper and lower mesepisternum distinguished by complete line of setae and variably developed, sometimes almost completely effaced transepisternal line (Fig. 354). Fore wing hyaline; bare behind submarginal vein except sometimes basal cell with a few setae distally. Propodeum (Fig. 355) with crenulate postspiracular sulcus differentiated from callus; callus punctate-rugose anteriorly and posteriorly but centrally with effaced sculpture, sometimes almost smooth; plical region with subparallel or narrowly widened paramedian crenulate furrows delineating median carina, and with anterior-most cell either transverse or if elongate then of same width as more posterior cells; supracoxal band contiguous with paramedian crenulate furrows; propodeal panels smooth and shiny or only rarely slightly wrinkled-roughened.

Petiole (Fig. 355) about 2x as long as medial width; transversely carinate to reticulate-rugose between longitudinal carinae; with several setae laterally. Gaster shiny and with Gt 1 smooth but at least Gt 3 very finely coriaceous.

Male. Length = 1.4–3.7 mm. Antenna (Fig. 358) with scape about 6x as long as wide, the inner surface smooth and shiny and either bare or with single medial line of setae over at least basal half (Fig. 359) and outer surface coriaceous to coriaceous-strigose apically but lacking distinct setiferous punctures (Fig. 360); pedicel subglobular or at most about 1.4x as long as apical width; flagellum with setae much shorter than width of respective segment; funicle with fu 1 about 2.0–3.4x as long as wide and about 1.4–2.3x as long as pedicel, and subsequent funicular segments all oblong. Otherwise similar to female except as follows. Head in anterior view subequal in height and width; in lateral view (Fig. 348) with malar space about 0.5–0.6x eye height and about 0.7–0.8x eye width; inclined lateral surface of scrobal depression and parascrobal region (Fig. 349) as well as gena (Fig. 348) sometimes more or less reticulate-rugose, with crowded, multisided punctures separated by linear ridges. Fore wing with at least 3 setae in line on mediocubital fold or basal cell extensively setose. Petiole (Fig. 356) 1.8–2.5x as long as medial width.

Material examined. Nearctic and Neotropical (1830 specimens in AMNH, BMNH, BPBM, CASC, CISC, CNC, CUAC, CUIC, FSCA, IRCW, LACM, MLPA, NCSU, TAMU, UATV, UCRC, USNM, WFBM). Complete collection records are not provided; those with host data: Nearctic. CANADA: Alberta, Leduc, 18.IX.80, R. Medved, M. domestica (UCRC, USNM) . Ontario, Ottawa vicinity, various localities and dates 2000 and 2001, G. Gibson & L. Bartels, M. domestica and S. calcitrans (see Gibson and Floate 2004). Yorks Corners Road, S Kenmore, G. Gibson & L. Bartels,? Physiphora demandata — Donevelyn farm, 45º 11.758'N 75º 23.547'W, 30.VIII.01; Melenhorst farm, 45º 12.996'N 75º 24.452'W, 16, 30.VIII.01. USA: Arizona, Tucson, 4.VI.80, R. Medved, M. domestica (UCRC) . California, Imperial Co., IX.23, N.J. Osborne, H. irritans (BPBM) . Los Angeles Co., 24.VII.51, S. calcitrans (LACM) . Colorado, R. Medved, M. domestica — Boulder, 19.IX.80 ( UCRC, USNM); Lafayette, 19, 20.IX.80 ( UCRC, USNM); Parker, 23.IX.80 ( UCRC). Florida, Alachua Co. — Gainesville, VII.74, 13.III.75, 6.V.75, 15, 19.VI.75, 31.VII.75, 14.IX.75, R.L. Escher, H. irritans (FSCA) ; High Springs, 19.XII.74, M. domestica (FSCA) . Bradford Co., Starke, 14.XI.74, em. 18.XI.74, R.D. Kramer, M. domestica (FSCA) . Columbia Co., 6.8 mi. NE Fort White, 27.IV.83, 6.V.83, 16, 24.VI.83, 1, 7, 15.VII.83, 27.VIII.83, J.T. Vaughan, S. calcitrans (FSCA) . Union Co., Lake Butler, V.73, C. Morgan, M. domestica (FSCA, LACM) . Illinois, Bloomington, IX.09, M. domestica . Minnesota, St. Paul, 21.VIII.80, A.M. Paprocki, M. domestica (UCRC) . Mississippi, Starkville, 12.VII.72, K.J. Watts, H. irritans (USNM) . Missouri, Boone Co., Columbia, 12.X.81, D.E. Figg, Anthomyiidae (UNML) . Nebraska, Lincoln, VII.58, C.M. Jones, S. calcitrans (USNM) . New Jersey, Woodfern, 5.VII.67, D. Shibles, M. domestica (USNM) . New York, Cayuga Co., L. Smith, M. domestica (CUIC) — LI, 5.IX.84; Waterman 4, 9.VIII.84. Cornell University, Exp. 841, M. domestica (CUIC) . Sullivan Co., 24.IX.87, C. Henderson, M. domestica (CUIC) . North Carolina, Craven Co., Spring Hope, 29.VI.77, T.D. Edwards, M. domestica (CUIC) . Wake Co., 15, 23.VIII.83, M. domestica (NCSU) . Wilkes Co., N. Wilkesboro, 17.VIII.77, T.D. Edwards, M. domestica (CUIC) . Ohio, Columbus — 25.XI.16, Muscina assimilis (USNM) ; 22.II.17,? L. serricata (USNM) ; 26.II.17, M. domestica (USNM) . Pennsylvania, Wannamakers, 19.VIII.66, J. Drea, Ravinia querula (USNM) . South Carolina, Clemson, 17.VII.68, E.F. Legner, M. domestica (UCRC) . Oconee Co., South Union, 12VI.73, J.R. Ables, M. domestica (CUAC) . Texas, Cuero, V.40, L.F. Hitchcock, H. irritans (USNM) . Dallas, 8.XII.12, S. calcitrans (USNM) . Laredo, 26.III.29, R.A. Roberts, M. domestica (USNM) . Virginia, Blacksburg, 25.VIII.67, R.P. Burton, Orthellia caesarion (USNM) . Wisconsin, Neillsville, 7.IX.65, E.F. Legner, Trichopria (?) sp., original ( UCRC).

Neotropical. BRAZIL: Goias, Santa Isabel, Ilha do Bananal, Rio Araguaia, 15-20.VII.57, M. domestica (CASC) . CHILE: Santiago, 1958, prob. M. domestica (USNM) . MEXICO: Morelos, Cuernavaca, S end, 4500̓, 7.IV.59, H.E. Evans, M. domestica (CUIC) . PUERTO RICO: Añasco, 25.VIII.80, B. Hawkins, M. domestica (UCRC) . Cayey, 26.VIII.80, B. Hawkins, M. domestica (UCRC) . Hormigueros, 28.II.50, H.K. Plank, M. domestica (USNM) . Juana Diaz, 25.VIII.80, B. Hawkins, M. domestica (UCRC) . Mani, 22.VIII.80, B. Hawkins, M. domestica (UCRC) . Mayaguez, 27.III.50, W. Gaud, M. domestica (USNM) . Naguabo, 10.VII.36, H.L. Dozier, Sarcophagula occidua (USNM) . Parguera, 23.VIII.80, B. Hawkins, M. domestica (UCRC) .

Distribution. A cosmopolitan species that Noyes (2003) records from all six biogeographic regions. Within the New World (Fig. 365) I saw specimens from North America ( Canada, USA, Mexico), Central America ( Costa Rica, Guatemala), West Indies ( Puerto Rico, Santa Lucia) and South America ( Argentina, Brazil, Chile, Peru).

Biology. Noyes (2003) lists S. nigroaenea as a parasitoid of about 50 species in 8 families of Diptera and, probably as a hyperparasitoid, from Diatraea saccharalis (Pyralidae) . Almost all of the host records I saw from the New World were from the three principal filth-fly pests of livestock, the house fly, stable fly and horn fly, but I also saw specimens labelled as reared from? Lucilia serricata (Meigen) , Muscina levida (Harris) (= M. assimilis ) and Neomyia cornicina (Fabricius) (= Orthellia caesarion ) ( Muscidae ), plus Ravinia querula (Walker) and Sarcophagula occidua (Fabricius) (Sarcophagidae) , and? Trichopria sp. ( Hymenoptera : Diapriidae ). The Missouri Anthomyiidae record is based on one male voucher specimen from Figg et al. (1983) that was incorrectly identified as S. nigra . I did not see voucher specimens from Figg et al. (1983) for Gymnodia arcuata (Stein) (Muscidae) or Ravinia spp. , which were the two listed host records for S. nigroaenea . The tentative record of Physiphora demandata (Fabricius) by Gibson and Floate (2004) was based on comparison of pupal remains correlated with flies emerged from similar puparia. The tentative Trichopria record suggests that S. nigroaenea can sometimes develop as a hyperparasitoid of flies through hymenopterous primary parasitoids.

Recognition. I include S. nigroaenea as one of six species in the nigra species group as discussed under S. nigra . It is most likely to be mistaken for S. chontalensis because both species share an anteriorly margined pronotal collar (see under latter species), but it might also be mistaken for S. masneri or S. nigroides if the sometimes only obscurely developed collar margin is missed (see under respective species for differential features). Among the six nigra -group species, only females of S. nigroaenea have the outer surface of the scape distinctly punctate-setiferous (Fig. 362) and the inner surface smooth and shiny over at least its basal half (Fig. 361). It is also the only species of the group that is sexually dimorphic in fore wing setation, the mediocubital fold being bare in females but males having a line of at least a few setae. Also characteristic of S. nigroaenea is its usually obviously subdivided, lunate episternal scrobe (Fig. 354), and the median smooth and shiny band that extends from the anterior convex region of the mesoscutal median lobe through the internotaular region (Fig. 353).

Spalangia nigroaenea is also very similar to the European species S. slovaca Bouček. My concept of S. slovaca is based on a NMPC female paratype and specimens from Kazakhstan and Russia that were compared with other nigra -group species in Taylor et al. (2006). Males of S. slovaca lack any setae on the mediocubital fold, which differentiates them from S. nigroaenea males, whereas a different sculpture pattern of the scape differentiates females of the two species. Female S. slovaca have the outer surface of the scape rugulose-strigose (Fig. 364; Taylor et al. 2006, figs 9, 15), lacking the distinct setiferous punctures that characterize S. nigroaenea females (Fig. 362; Taylor et al. 2006, figs 11, 13). Furthermore, the inner surface is at least finely (Fig. 363; Taylor et al. 2006, figs 10, 16) and sometimes distinctly longitudinally striate rather than smooth and shiny medially as in S. nigroaenea (Fig. 361; Taylor et al. 2006, figs 12, 14).

Fig. 365. Distribution of Spalangia nigroaenea Curtis.