Calosota aestivalis Curtis

Gary A. P., Gibson, 2010, Calosota Curtis (Hymenoptera, Chalcidoidea, Eupelmidae) - review of the New World and European fauna including revision of species from the West Indies and Central and North America, ZooKeys 55, pp. 1-75 : 13-19

Calosota aestivalis Curtis Figs 213, 143346556072

Calosota aestivalis Curtis 1836: folio 596. Type data: England: Southgate. Lectotype ♀ (MVMA; not examined), designated by Graham 1969: 90-91.

Calosoter vernalis Walker 1837: 359. Type data: England: near London; Ireland: Holywood. Syntypes, ♀ and ♂ (BMNH, type no. 5.1621; not examined). Synonymy by Graham 1969: 90-91. Homonym of Calosota vernalis Curtis (1836).

Calosota fumipennis Bolívar y Pieltain 1923: 65-67. Type data: Spain: Villaviciosa de Odón. Holotype ♀ (MNCN; examined), by original designation. Synonymy by Askew and Nieves-Aldrey 2006: 89.

Calosota septentrionalis Hedqvist 1956: 96-97. Type data: Sweden: Fundort: Dalekarlien, Älvdalen 1955. Holotype ♀ (K. Hedqvist personal collection; examined). syn. n.

Calosota kentra Burks 1973: 30-31. Holotype ♀ (USNM, type no. 72482; examined), by original designation. syn. n.

Calosota montana Burks 1973: 31. Holotype ♀ (USNM, type no. 72483; examined), by original designation. syn. n.

Description.

FEMALE (Figs 13, 33). Length about 3-6.5 mm. Color. Head (Fig. 2) variably dark greenish-blue to purple, including spot below anterior ocellus, but with more or less complete coppery or dark transverse band on vertex between inner orbits (Fig. 14) (sometimes reduced to large spot behind ocellar triangle and adjacent to each upper inner orbit) and with M-like coppery or dark region on upper face (region very rarely narrowly divided medially below anterior ocellus), with lateral arm of region usually extending dorsally to or toward posterior ocellus (sometimes filling ocellar triangle and rarely contiguous with transverse band on vertex) and ventrally abutting inner orbit. Maxillary and labial palpi dark. Antenna dark brown except scape sometimes with metallic luster similar to lower face. Tegula dark. Mesoscutum (Figs 13, 14) variably dark greenish-blue to purple similar to head except almost always with at least obscurely differentiated dark, coppery or greenish paramedial longitudinal bands (Fig. 13) (anteriorly each band sometimes subdivided into band occupying region between parapsidal line and notaulus, and narrower band extending posteriorly from anteroadmedian line, and posteriorly paramedial bands sometimes broadly contiguous); scutellar-axillar complex or at least scutellum mostly same color as mesoscutal paramedial bands, the axillae more commonly and often margins of scutellum similar in color to remaining mesoscutum. Acropleuron (Fig. 55) dark with slight coppery luster to variably greenish-blue or purple similar to most of head and mesoscutum. Legs (Fig. 33) mostly brown with knees, apices of tibiae, and at least basal tarsomeres of meso- and metatarsi yellowish, but usually tarsi more extensively yellowish and meso- and metatibiae sometimes also mostly or entirely yellowish-brown to yellowish. Fore wing hyaline or disc variably extensively and conspicuously infuscate; setae uniformly brown. Gaster (Fig. 33) usually dark brown with slight reddish-coppery luster under some angles of light or laterally partly blue to purple.

Structure/setation. Head in dorsal view about 1.8−2.1 × as wide as long, with IOD about 0.33−0.45 × head width, LOL at least slightly greater than and sometimes up to about 2 × OOL and slightly less than to slightly greater than MPOD, and POL about 1.2−1.7 × MPOD; in frontal view about 1.2−1.3 × as wide as high, with dorsal margin of torulus about at level of lower orbits; malar space about 0.5−0.7 × eye height. Head (Fig. 2) with frontovertex and upper parascrobal region meshlike reticulate to about level of dorsal limit of interantennal region, medially the reticulations tapered ventrally between dorsal limits of smooth and shiny scrobes; lower parascrobal region and interantennal region much shallower meshlike reticulate to coriaceous-reticulate; clypeal region microcoriaceous to granular and paraclypeal region obliquely reticulate-alutaceous. Head sometimes with whitish setae except for bare scrobal depression but more commonly with brownish setae on frontovertex and more conspicuous white setae on parascrobal region, interantennal region and lower face. Antenna (Fig. 33) with scape about 4.6−5.3 × as long as wide; pedicel about 2.5−3.3 × as long as wide; flagellum variably distinctly clavate (funiculars all about same width and clava usually only slightly wider than funicle except if compressed) with length of flagellum + pedicel about 1.3−1.75 × head width; combined length of fu1 + fu2 about 1−1.7 × as long as pedicel; fu1 about 1.3−2.3 × as long as wide; subsequent funiculars all longer than wide with fu2 about 2−3.1 × and fu8 at least slightly and sometimes up to about 1.5 × as long as wide; clava often collapsed, but about as long as apical 2.5−3.5 funiculars. Mesoscutum (Figs 13, 14, 72) more or less uniformly meshlike reticulate, with inconspicuous white setae; notaulus extending from spiracle as curved furrow on inclined anterior surface, its posterior limit contiguous dorsally with posterior limit of anteroadmedian line; parapsidal line usually quite a distinct region of microsculpture posterior to spiracle. Axillae elongate-triangular, separated by about 3−4 × own width (Fig. 72). Scutellum flat to low convex and at least slightly (up to about 1.2x) longer than wide; similarly reticulate as mesoscutum (Fig. 72); with inconspicuous white setae. Mesopleuron (Fig. 55) with exposed, bare lower mesepimeron; acropleuron variably deeply and distinctly meshlike reticulate anterior to oblique microsculptured region and longitudinally coriaceous-alutaceous posteriorly. Fore wing (Fig. 60) with cc: mv: stv: pmv about 40−60: 23−31: 10: 14−16, and perpendicular distance between apex of stigmal vein and anterior margin of wing usually about 0.7−0.8x, only very rarely up to about 9.5x, length of stigmal vein; basal cell entirely setose; cubital area bare except sometimes anteriorly near mediocubital fold, and up to about apical half closed by setae along posterior margin; disc setose except usually for short region of mediocubital fold just beyond basal fold or with variably broad and distinct, often lunate bare region along basal fold, the bare region sometimes continuous with cubital area. Metacoxa setose along dorsal, ventral and usually basal margins, but sometimes up to about basal third of outer surface setose. Propodeum with callus setose to posterior margin; bare anteriorly between spiracle and foramen. Gaster (Figs 13, 33) about 1.8−2.2 × as long as mesosoma; sparsely setose dorsally and more densely setose laterally with white to brownish hairlike setae; penultimate tergum with posterior margin extending to or slightly beyond level of syntergum; syntergum about 1.5−2.6 × as long as transcercal width, variably distinctly compressed depending on length, and about 0.7−1.3 × as long as penultimate tergum.

MALE (Fig. 46). Similar to female except as follows. Length about 2.5−3.8 mm. Color. Legs (Fig. 46) always extensively dark with knees and tarsi often yellowish but tibiae usually only slightly lighter apically; all males examined with M-like region on upper face and with lateral arms extending ventrally to inner orbits (Fig. 2); mesonotum more commonly (particularly smaller individuals) without distinct paramedial bands; fore wing disc often, but at most only very slightly infuscate.

Structure/setation. Antenna with scape more robust, only about 3.4−3.9 × as long as wide; flagellum of smaller individuals sometimes more distinctly clavate, the funicle evenly widened toward clava; fu1 variably distinctly widened distally and sometimes only about as long as wide, but usually quite obviously (up to about 1.5x) longer than wide; fu2 about 1−1.9 × as long as wide or length of fu1; combined length of fu1 + fu2 about 0.65 × (smallest specimens) to about 1.25 × length of pedicel; and fu8 quadrate to slightly longer than wide. Fore wing venation similar to female with cc: mv: stv: pmv about 38−52: 22−31: 10: 13−14, and perpendicular distance between apex of stigmal vein and anterior margin of wing about 0.8−0.9 × length of stigmal vein; basal cell and disc completely setose or sometimes disc with arcuate bare band along basal fold, but bare region only rarely continuous with cubital area. Propodeal callus sometimes setose to posterior margin, and then rarely with one or more setae anteriorly between spiracle and foramen, but more often setose only to level about equal with posterior margin of spiracle, with 1 or 2 setae often behind spiracle but bare anteriorly between spiracle and foramen.

Biology.

Burks (1973) stated that the type specimens of Calosota montana were reared from an unidentified gall on Pinus contorta (lodgepole pine). The female from British Columbia labeled as reared from Dioryctia sp. ( Lepidoptera : Pyralidae ) is a Forest Insect Survey specimen that very likely represents an incorrect host association; however, the California host record of Anthaxia sp. ( Coleoptera : Buprestidae ) reared from Quercus (oak) is more likely correct even though coniferous trees such as Pinus (pine) and Pseudotsuga (Douglas-fir) are more commonly indicated as tree associates in North America. Noyes (2003) listed several host species in Europe in Anobiidae , Cerambycidae , Cleridae , and Curculionidae including Scolytinae ( Coleoptera ) emerging from Betulaceae , Fabaceae , Fagaceae , Pinaceae and Tamaricaceae . The cited records of Trichodes leucopsideus (Olivier) ( Cleridae ) and associated Megachile sp. ( Hymenoptera : Apidae ) are incorrect because of previous historical misidentification of Calosota vernalis as Calosota aestivalis (see below and under Calosota vernalis ). All documented host records indicate Calosota aestivalis is a primary parasitoid ( Noyes 2003).

Regional material examined

(Map 2). CANADA. British Columbia: Anahim Lake to Redstone, 1000-1500 m., 17.VII.88, S&J Peck (1♂ CNC). Beaverdell, 7.VII.61, FIS (Forest Insect Survey), 60-8056-01, ex. Dioryctria sp. (1♀ CNC). Brookmere, 21.VII.33, K. Graham (1♀ CNC, CNC Photo 2009-47). Kaslo, 24.VI.03, R.P. Currie (1♂ USNM). O.K. Centre, 2.V.30, 17461 Lot1, A.C. Thrupp (1♀ CNC, CNC Photo 2009-44). Manitoba: Onah, 10.VII.84, R.M. White, Tamarack (1♀ CNC). Quebec: Gatineau Pk, Ridge Rd, 27. V– 10.VI.6[?] (1♀ CNC). Pontiac, Eardley, 5.VI.91, S. Laplante (1♀ CNC, CNC Photo 2009-16). USA. California: Argus Mts, V.1891 (1♂ paratype of Calosota longiventris , USNM). Santa Cruz Mts, antenna on slide (1♂ USNM). Kern Co., Glennville, em. III, IV.67, J.W. Tilden (1♀ CNC Photo 2009-20, 1♂ EMEC). Marin Co., Lagunitas, 14.VII.28, E.H. Nast (1♀ EMEC). Mendocino Co., NCCRP, 3 mi. E Branscomb, 1400', 21-23.V.82, C. Besette (1♀ EMEC, CNC Photo 2009-19). Montery Co., Santa Lucia Mts, Junipero Serra Pk, on peak ca. 5800', 27, 28 (CNC Photo 2009-17).VIII, 4.IX.56, 4.IV, 16.V.57, H.B. Leech, ex. dead branches Pinus coulteri (1♀, 4♂ CASC). Napa Co., 2 mi. NNE Angwin, N side Howell Mt., 1300', 17, 20.V.76, R.R. Leech, ex. log Pseudotsuga menziessii (2♂ CASC); Butts Cyn Rd, 0.5 mi. S Snell Valley Rd, 11, 15 (CNC Photo 2009-18), 17, 23, 29.IV.78, R.B. Leech, ex. dead branches Pinus sabiniana (1♀, 4♂ CASC). San Bernardino Co., Burns Piñon Ridge Reserve, 1260 m., 34°08'57N; 116°27'11W, 21-23.V.05, K. Will et al. (2♀ RLZC), 23.V.05, R.L. Zuparko, Chilopsis linearis arcuata (2♀ RLZC). San Luis Obispo Co., 6 mi. SE Poso, R16E, T315, sects. 4-5, 1800', 2. IV– 4.V.89 (1♂ CNC), 1500', 23. IV– 4.V.89 (1♀, CNC), W.E. Wahl. Shasta Co., 2 mi. W Shingletown, 2750', 15-20.VII. 85, R. Miller, damp open meadow with Rush ( Juncus ) and wild flower along stream in pine forest (1♂ FSCA). Siskiyou Co., Yreka, 4820 a’ Hopk. U.S., reared, Quercus , probably ex. Anthaxia (1♀ USNM). Tulare Co., Tulare, 20.III.75, emerged 10.IV.75 (1♂ CDFA). Ventura Co., 25.IV.05, 2771 Hopk. U.S., bred, Pinus (1♀ USNM). Idaho: Craters of the Moon Nat. Mon., summer '64, reared from Pinus flexilis (1♀ USNM). Latah Co., Moscow, Paradise Ridge, 3000', 9.V.31, P. Rice (1♂ USNM). Montana: Granite Co., Rock Creek, 10 (allotype), 11 (holotype).II.69, from unidentified gall on Pinus contorta , by J. G. Bringuel (♀ holotype, ♂ allotype of Calosota montana ). New Hampshire: Carroll Co., Albany, 2.VII.58, W.J. Morse (♀ holotype of Calosota kentra ). North Carolina: Northhampton Co., 7 km. S Jackson, 23. IX– 15.XI.87, bald cypress swamp, BRC Hym. team (1♂ CNC). Oklahoma: Latimer Co., Red Oak, IX.90, K. Stephan (1♀ CNC). Virginia: Fairfax Co., near Annandale, 1-17.IV.90, D.R. Smith (1♂ CNC).

Distribution.

Noyes (2003) listed several countries in the Palaearctic region; I saw specimens from Corsica (CNC SEM 2009-48, ZSMC), Cyprus (NMPC), England (BMNH), France (BMNH, CNC), Jordan (CNC), Morocco (CNC), Slovakia (CNC), Spain (CNC) and Sweden (CNC). Its extensive, apparently transcontinental distribution (Map 2) and morphological variability in North America (see further below) suggest that it has been present in the region for a long time as a naturally occurring Holarctic species. Its presence in North America for a long time may also be supported by head sculpture pattern, which is shared with four other similar species in North America ( Calosota bicolorata , Calosota elongata , Calosota longivena and Calosota longiventris ), but not with any other species in at least Western Europe. This suggests that Calosota aestivalis may be more closely related to the four North American species than it is to any species from Western Europe. This hypothesis needs be tested through phylogenetic or molecular analyses.

Remarks.

Burks (1973: 31) inadvertently published the binomen Cecidostiba montana as the name of his new species preceding its description. However, this certainly was an unintentional printing error based on the title of the paper, proper generic placement of the species in the abstract and key, and other typographical errors in the type locality and type sections of the preceding description of Calosota kentra .

Recognition.

The concept of the names Calosota aestivalis and Calosota vernalis were incorrectly reversed in the literature beginning with Walker (1837) until corrected by Graham (1969). Although I did not examine the lectotype of Calosota aestivalis , my concept of this name is based on the keys of Graham (1969) and Askew and Nieves-Aldrey (2006) plus authoritatively identified specimens in the collections listed above. I examined the holotype female of Calosota septentrionalis in 1984 and at that time considered that it probably was a synonym of Calosota aestivalis , noting that the vertex was reticulate in both. Hedqvist (1956) compared his new species to Calosota aestivalis (as Calosota vernalis) and differentiated the latter species from Calosota septentrionalis and Calosota fumipennis based on the fore wing being either yellowish or more or less infuscate, respectively. Graham (1969) noted that European females of Calosota aestivalis often have the fore wings partly infuscate, which is the form that Bolívar y Pieltain (1923) described as Calosota fumipennis . Hedqvist (1956) differentiated Calosota septentrionalis from Calosota fumipennis on color differences as well as equally variable differences in the ratios of the marginal, postmarginal and stigmal veins, but stated specifically that Calosota septentrionalis has paramedial longitudinal greenish-bronze bands on the mesoscutum. Based on this color pattern, head sculpture pattern, and other features Hedqvist (1956) used in his key to differentiate species, I hereby synonymize Calosota septentrionalis under Calosota aestivalis syn. n., as was done previously for Calosota fumipennis by Askew and Nieves-Aldrey (2006).

Askew and Nieves-Aldrey (2006) suggested that another European name, Calosota ariasi Bolívar y Pieltain (1929), whose unique female holotype had been lost from its mount in MNCN, might also be a synonym of Calosota acron . They refrained from formalizing the synonymy because of some significant discrepancies between the original description of Calosota ariasi and specimens of Calosota acron . Based on its original description, Calosota ariasi more likely is a synonym of Calosota aestivalis than Calosota acron . The description of Calosota ariasi states that the head had a transverse coppery band behind the ocelli as well as a coppery band extending from each posterior ocellus to surround the anterior ocellus ventrally and widen on the upper face to about its mid height. This color pattern describes very accurately the color pattern of Calosota aestivalis (Fig. 2) but not Calosota acron (Fig. 1), which has the frontovertex mostly dark. The described bluish-black femora and brownish-black tibiae except for the knees and apex of the tibiae and very slightly infuscate fore wings also more strongly suggest Calosota aestivalis (Fig. 33) than Calosota acron . Askew and Nieves-Aldrey (2006) keyed Calosota ariasi along with Calosota vernalis , primarily because the original description states that the distance between the eyes was the same as the width of an eye. Description of head sculpture for Calosota ariasi (weakly shagreened dorsally and face strongly shagreened, almost scale-like) might also be interpreted as descriptive of the head sculpture of Calosota vernalis (see under this species), but the acropleuron is described as being very shagreened over its basal two-fifths, especially dorsally. This is characteristic of Calosota aestivalis (Fig. 55) but not Calosota vernalis (Fig. 54), which suggests description of the head sculpture actually referred to a somewhat coarser reticulate sculpture below than above the anterior ocellus that is also typical of Calosota aestivalis (Fig. 2). As noted by Askew and Nieves-Aldrey (2006), described shape and width of its axillae, which Bolívar y Pieltain (1929) used as key features to differentiate Calosota ariasi and Calosota aestivalis (as Calosota vernalis + Calosota fumipennis) from other Spanish Calosota , also contradicts the possibility of Calosota ariasi being synonymous with Calosota vernalis . The features used to distinguish Calosota ariasi from Calosota aestivalis by Bolívar y Pieltain (1929) (anellus only about one-third longer than wide versus twice as long as wide and syntergum twice as long versus only about as long as wide) certainly are within the range of variation noted for North American specimens I identify as Calosota aestivalis . It seems likely that Calosota ariasi is a synonym of Calosota aestivalis , but I hesitate to formalize the synonymy prior to a more comprehensive revision of Spanish and European Calosota .

When Burks (1973: 31) described Calosota montana he stated that it "greatly resembles the European species vernalis Curtis", but it is apparent he was still misinterpreting the name Calosota vernalis in the sense of Calosota aestivalis based on other features listed as shared between " Calosota vernalis " and Calosota montana . The presence or absence of longitudinal banding on the mesoscutum (cf. Figs 13, 14), shading on the fore wing, and a bare region on the fore wing disc adjacent to the basal fold were all used by Burks (1973) to differentiate Calosota montana (western USA: Montana) from Calosota kentra (eastern USA: New Hampshire). Based on observation of more material, females with infuscate wings normally have the disc more or less uniformly setose to the basal cell whereas females with hyaline wings typically have a more definite bare band adjacent to the basal fold, but all the features used by Burks (1973) to differentiate Calosota montana and Calosota kentra appear to be present in different combinations, including structure of the stigma and uncus. Burks (1973, fig. 2) partly characterized the holotype of Calosota montana as having an enlarged stigma with a long, slender uncus. The holotypes of Calosota montana and Calosota kentra are both similar to European females of Calosota aestivalis in having the stigmal vein comparatively long and at an acute angle to the postmarginal vein such that the perpendicular distance between its apex and the anterior margin of the wing is only about 0.8 × its length (Fig. 60). Both holotypes also have an M-like region differentiated on the upper face (cf. Fig. 2) as well as at least an obscurely differentiated transverse region on the vertex (Fig. 14), and the marginal vein is only about 2.4 × as long as the stigmal vein (cf. Fig. 60). I consider the morphological differences between the type material of Calosota kentra and Calosota montana to represent intraspecific variation in one quite variable and widely distributed species in North America and therefore synonymize both Calosota kentra and Calosota montana under Calosota aestivalis syn. n. However, I also describe a new species, Calosota longivena , based on females that are morphologically very similar to some females I include in Calosota aestivalis except for relative length of the marginal vein and, usually, a somewhat differently structured stigmal vein. Furthermore, the single female from Oklahoma (Map 2) that I include in Calosota aestivalis is intermediate in some features between those I identify as Calosota aestivalis and Calosota longivena . In addition to having distinctly infuscate fore wings and an obvious bare band adjacent to the basal cell, the Oklahoma female is unusual in having a comparatively short and straight stigmal vein (perpendicular length from its apex to anterior margin of wing almost equal to its length) and an unusually long costal cell and marginal vein as compared to other females I identify as Calosota aestivalis . The Oklahoma female has the marginal vein about 3.2 × and the costal cell about 6 × as long as the stigmal vein compared to up to about 2.8 × and up to about 4.8x, respectively, for other Calosota aestivalis females. Females of Calosota longivena have a shorter and more obtusely angled stigmal vein such that the marginal vein is at least about 3.6 × the length of the stigmal vein and stigmal vein length is subequal to the distance from its apex to the anterior margin of the wing. Although the paramedial bands on the mesoscutum of the Oklahoma female are also somewhat obscurely differentiated it has quite a distinct M-like coppery region on the upper face. The combination of features of the Oklahoma female suggests the possibility of introgression or a morphological cline in what I identify as Calosota aestivalis and Calosota longivena . The southeastern Oklahoma origin of the aberrant female, between the known distribution of Calosota longivena (Florida to southwestern Texas) and what appears to be a more northern transcontinental and further eastern (California) distribution for Calosota aestivalis (Map 2), might also support a hypothesis of introgression or morphological cline. However, if females I describe as Calosota longivena represent only a highly modified southern form of Calosota aestivalis in North America then Calosota aestivalis is far more variable in North America than in Europe. Collection of males with females in the known range of Calosota longivena could help clarify morphological limits and species status, as could molecular analysis of specimens from throughout North America and Europe. Males I identify as Calosota aestivalis are most similar to those of Calosota longiventris (see further under latter species and Calosota bicolorata ).