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Nov 18, 2005 - Head and mesosoma with large umbilicate punctures (Figs. 2, 4, 6); ... Color: Head and mesosoma metallic purple to blue and green, fading to ...
Zootaxa 1082: 45–55 (2005) www.mapress.com/zootaxa/

ISSN 1175-5326 (print edition)

Copyright © 2005 Magnolia Press

ISSN 1175-5334 (online edition)

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Nomenclatural changes in Neotropical Eulophidae, Eupelmidae and Torymidae (Hymenoptera: Chalcidoidea) relating to parasitoids of Cecidoses eremita (Lepidoptera: Cecidosidae) R.A. BURKS1, G.A.P. GIBSON2 & J. LA SALLE3 1. University of California, Riverside, CA 92521, USA; [email protected] 2. Biodiversity Theme, Agriculture and Agri-Food Canada, 960 Carling Avenue, K. W. Neatby Bldg., Ottawa, Ontario, Canada, K1A 0C6 3. CSIRO Entomology, GPO Box 1700, Canberra, ACT 2601, Australia

Abstract Two monospecific genera described by Brèthes (1927) in Eulophidae are newly synonymized under genera in Torymidae and Eupelmidae; respectively, Parasympiesis Brèthes is recognized as a junior synonym of Torymus Dalman n. syn. and Tropimius Brèthes as a junior synonym of Brasema Cameron n. syn. Torymus cecidicolus (Brèthes) n. comb. and Brasema willei (Brèthes) n. comb. are rediagnosed, and Torymus alegrensis (Brèthes) n. syn. is recognized as a junior synonym of T. cecidicolus. All three species were described as parasitoids of the gall-forming lepidopteran, Cecidoses eremita Curtis (Cecidosidae). Brasema willei is one of 11 species of Brasema described from South America, including B. acauda (Ashmead) n. comb., B. aprilis (Ashmead) n. comb., B. basicuprea (Walker) n. comb., B. chapadae (Ashmead) n. comb., B. corumbae (Ashmead) n. comb., B. peruviana (Crawford) n. comb., and B. proxima (Ashmead) n. comb. from Eupelmus Dalman, and B. cyanea (Ashmead) n. comb. from Zaischnopsis Ashmead. Key words. Chalcidoidea, Cecidoses, Eulophidae, Neotropical, parasitoid

Introduction Brèthes (1927) described Parasympiesis cecidicola and Tropimius willei as new genera and species of Eulophidae, as hyperparasitoid and parasitoid, respectively, of Cecidoses eremita Curtis, 1835 (Lepidoptera: Cecidosidae), a Neotropical gall-inducer on Schinus (Anacardiaceae) that produces fruit-like stem galls (Parra 1998). Both taxa were regarded as near Ophelimus Haliday, although reasons for this assessment were not given. Gauthier et al. (2000) listed the two genera as unplaced genera of Eulophinae and the taxa have been considered dubiously placed in Eulophidae because the description of P. cecidicola Accepted by M. Buffintong: 29 Sept. 2005; published: 18 Nov. 2005

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stated that it had 13 antennal segments, a state not present in any known eulophid, and both descriptions included other features seldom found in eulophids. Brèthes (1927) did not state the sex on which he based the descriptions of P. cecidicola or T. willei. Upon examination, it was discovered that the syntype series of P. cecidicola consisted of males, that they belonged to the genus Torymus Dalman (Torymidae), and that they were conspecific with females of another species described by Brèthes (1927) as a parasitoid of C. eremita. Another author (GAPG) had previously discovered that the single known specimen of T. willei was also a male and that it belonged to Brasema Cameron (Eupelmidae). In this article, we rediagnose and transfer the species to their proper families and genera and discuss the species within the context of other members of the respective genera.

Materials and methods Structural terminology follows that of Gibson (1997) except: "interocular distance" is the distance between the eyes across the face, either at the dorsal or ventral extreme of the eyes; "interspiracular distance" is the greatest distance between the propodeal spiracles measured from dorsal view; LOL is the distance from the median ocellus to a lateral ocellus; POL is the distance between the lateral ocelli; OOL is the shortest distance from a lateral ocellus to the eye. Sculptural terminology follows that of Gibson (1989, 2003) except: "alveolate" describes sculpture divided into pentagonal to heptagonal cells by raised, sloping septa as figured by Harris (1979); "imbricate" describes rows of shinglelike sculpture divided into cells by sunken septa that partially overlap the next (more anterior, posterior, central, or peripheral) row of cells; and "umbilicate puncture" is a setal socket and its surrounding concavity whose diameter is relatively large, nearly or greater in apparent diameter than sculptural meshes on the same sclerite. The redescription of T. cecidicolus is by the senior author; relative measurements were taken using all available specimens. Color photographs were taken using AutoMontage software (version 4.04.0128 BETA, Synoptics, Ltd., UK 1997...2003) using a 3-CCD digital videocamera attached to either a stereoscope or compound microscope. The redescription of B. willei is by GAPG; relative measurements of the unique males were taken with a Nikon SMU-Z stereoscope fitted with a 10mm. ocular grid having 100 divisions.

Torymus Dalman Parasympiesis Brèthes, 1927: 328. Type species: Parasympiesis cecidicola Brèthes, by monotypy. n. syn. For additional synonymy, see Grissell (1995) and Graham & Gijswijt (1998).

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Torymus cecidicolus (Brèthes) n. comb. (Figs. 1–6)

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Parasympiesis cecidicola Brèthes, 1927: 328. Syntomaspis (?) Alegrensis Brèthes, 1927: 333. n. syn. Callimome (?Syntomaspis) alegrensis (Brèthes); Bondar, 1930: 112. Callimome alegrensis (Brèthes); Costa Lima, 1936: 378. Torymus alegrensis (Brèthes); Costa Lima, 1962: 280. Torymus alegrensis (Brèthes); DeSantis, 1980: 218. Torymus alegrensis (Brèthes); Grissell, 1995: 274.

Type material (Deutsches Entomologisches Institut, Eberswalde). Syntypes (8 %%): "No. 2" / "No. 2 Überparasit von 1." / "Porto Allegre" / "Syntypus" / "Wille 1924.25" / "type!" / "Parasympiesis cecidicola Brèthes, Brèthes det. 1926" / "Torymus cecidicolus (Brèthes), det. R.A. Burks 2003" (4%%); "2. Überparasit von 1" / "Porto Allegre" / "Syntypus" / "Wille 1924.25" / "type!" / "Parasympiesis cecidicola Brèthes, Brèthes det. 1926" / "8 ST cecidicola" / "Torymus cecidicolus (Brèthes), det. R.A. Burks 2003" (4%%). Condition of types. The syntype series consists of eight males, four each glued to two cards on separate pins. All are glued more or less obliquely venter downward by the head and/or the side of the body, with the face fully visible in only one, in which it is dorsally collapsed. Three have at least one complete antenna. One consists only of the mesosoma, complete except for the absence of one foreleg beyond the first trochanter and one entire midleg. The female type of Syntomaspis (?) Alegrensis (Fig. 3) is glued by the right legs onto a card. It is intact except for the antennal flagellum (of which only the anellus of the right antenna remains), the cercal setae (even though the cerci are intact), the tip of the left hind wing, and the extreme tips of the ovipositor sheaths (but the stylet tips remain). The gaster appears distorted, as discussed below. Diagnosis. Head and mesosoma with large umbilicate punctures (Figs. 2, 4, 6); clypeal margin truncate or slightly emarginate; first funicular segment subequal in length with pedicel in male; anellus transverse; frenal groove distinct, frenum bare; propodeum without median carina or plicae; propodeal spiracle less than its own diameter from metanotum; stigmal vein sessile (Fig. 5); metafemur without denticles; ovipositor sheaths longer than rest of body (Fig. 3). Description. Male (Figs. 1–2). Total body length = 1.70–1.89 mm (9.2–10.2 in relative units used below). All ranges represent measurements from all syntypes that could be measured accurately. Color: Head and mesosoma metallic purple to blue and green, fading to brownish (more so in smaller specimens), except tegula and humeral plate pale to dark brown, and with stout white setae. Antenna with scape dull brownish-yellow, becoming slightly more brownish apically; pedicel and flagellum dark brown, longitudinal sensilla nearly white. Mandible brownish-white basally and dark brown apically; palps brown. Legs with coxae, femora, and apical tarsomeres dark brown with same metallic luster as body (weaker on

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smaller structures), except femora pale brown at apices; trochanters, protibia, and basal 4 tarsomeres of all legs pale brownish-white; mesotibia and all tibial spurs varying from pale to dark brown. Forewing venation brownish-white. Gaster brownish, only slightly paler at base than at apex. Sculpture: Head and body imbricate with regularly distributed small umbilicate punctures separated by much more than their own diameter, the punctures less distinct in smaller specimens; scrobal depression alveolate in sharp contrast with surrounding areas of face, with scrobe very finely carinate laterally; vertex finely and shallowly rugulose; prepectus with a fovea along its dorsal two-thirds, set off ventrally by a tiny carina; metacoxa alveolate, without a dorsal carina. Head (Fig. 2) height: width: length = 2.3–2.9: 2.7–3.2: 1.5–1.6 Clypeal margin truncate to very slightly convex. Ventral margin of torulus 1.0–1.4 distance to mouth and about three torular diameters above lower eye margin, distance from dorsal margin of torulus to vertex = 1.1–1.5; eye height = 1.5–1.9, eye length (measured dorsally) = 1.0–1.1; dorsal interocular distance = 1.7–2.1, ventral interocular distance = 2.0–2.3; POL: OOL = 0.8–1.0: 0.2–0.3; temple = 0.3; malar space = 0.6–0.8. Flagellum (Fig. 2) with 1 basal anelliform segment and 7 funicular segments; funicular segments cylindrical, each slightly longer than broad, and parallel-sided except the first slightly broadened apically, all with 2 partially overlapping rows of longitudinal sensilla and with setae appressed except for a few tiny setae near apex of each segment; first funicular segment slightly longer to slightly shorter than pedicel; second funicular segment the longest, with succeeding segments gradually shorter but hardly narrower; club not broader than funicle, shorter than the preceding two segments combined; length: width of scape = 1.0–1.1: 0.3–0.4, of pedicel = 0.3–0.4: 0.3. Mesosoma (Fig. 1) length = 4.2–5.0. Pronotum gradually sloping, reaching top of mesoscutum but not forming a collar, maximum height = 1.6–2.1; lateral panel without spatulate setae. Mesoscutum length: width = 1.7–2.5: 2.4–3.0; notauli complete, more distinctly so in smaller specimens. Scutellum length: width = 1.8–2: 1.5–1.8, with a posterior foveolate rim and with a slight medial posterior concavity or truncation, not overhanging propodeum; frenum bare, separated from scutellum by a fine groove curved posteriad mesally (as in Fig. 6); axillae separated from scutellum by a simple groove. Dorsellum small but slightly convex. Dorsal length of prepectus = 0.7–1.0, appearing slightly longer to slightly shorter than tegula. Propodeum length: width = 0.6 –0.9: 2.2–2.6; without median carina, plicae or longitudinal foveae; callus with 9–12 setae in 2–3 rows; spiracle about its own lesser diameter from metanotum; interspiracular distance = 1.4 –1.8. Length: width of profemur = 1.9–2.3: 0.5–0.7, of protibia = 1.8–2.2: 0.3–0.4, of mesofemur = 2.0–2.5: 0.4–0.6, of mesotibia = 2.2–2.6: 0.3, of metafemur = 2.4–3.0: 0.6–0.9, of metatibia 2.6–3.0: 0.4–0.5; length of mesotibial spur = 0.4, longest metatibial spur 0.40.5x basal metatarsomere length; metacoxa without dorso-basal setae; metafemur without denticles. Forewing length: width = 8.6–9.6: 3.9; cc: mv: pm = 3.4–3.8: 2.3–2.6:

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0.2–0.3; costal cell with a single row of ventral setae, apparently interrupted in a more or less long central gap (difficult to assess because of debris), without dorsal setae; basal fold bare; basal cell with at most a single dorsal seta and no ventral setae; speculum bare; admarginal area bare ventrally; stigma oblong (as in Fig. 5), not petiolate, with distance from stigmal apex to postmarginal vein = 0.20.3, and from uncus apex to postmarginal vein = 0.1; setal tracks arising from stigmal apex indistinct. Hind wing length: width = 6.1–6.5: 2–2.2. Gaster At least slightly shorter than mesosoma. Female (Figs. 36). Total body length 6.55 mm (35.4 in relative units used below), including ovipositor and distorted gaster. Color like male except: pedicel, pronotum, propleuron, prosternum, tegula, humeral plate, all legs, and most of gaster yellow; gaster with 2 separated dorsal brownish blotches in posterior half, and a sublateral blotch on either side and just anteriad of the first dorsal blotch; extreme gastral apex, cerci, and ovipositor sheaths dark brown. Sculpture like male except: face and mesoscutum more densely supplied with umbilicate punctures, giving them a nearly rugulose appearance; vertex finely rugulose with umbilicate punctures; short strip posterior to malar sulcus without punctures; antennal scrobe punctate, with oval to teardrop-shaped meshes; area below occipital carina regularly imbricate and with punctures not interrupting the sculpture. Head (Fig. 4) height: width: length = 3.7: 4.3: 2.2. Clypeal margin shallowly emarginate. Distance from top of torulus to vertex = 1.6, from bottom of torulus to mouth = 1.7; eye height = 2.7, length = 1.8; temple = 0.3; malar space = 0.9; POL: OOL = 0.6: 0.3; dorsal interocular distance = 2.3, ventral interocular distance 3.0. Occipital carina a slightly exaggerated arch. Anellus transverse; scape length: width = 1.5: 0.5, reaching median ocellus; pedicel = 0.6: 0.3. Mesosoma (Fig. 6) length = 6.8. Pronotum maximum height = 2.0, length: width = 2.6: 2.8, not forming a collar, lateral panel without scrobe; mesoscutum length: width = 3.5: 4.3; scutellum = 2.6: 2.5; dorsellum rectangular, slightly convex; propodeum = 0.8: 3.9. Dorsal margin of prepectus = 1.2, slightly shorter than tegula. Notauli complete. Scutellum narrowly rounded anteriorly but reaching mesoscutum, not overhanging propodeum, with a posterior foveolate rim; axillae separated by a simple groove; frenum bare, frenal groove distinct and curved posteriad mesally. Propodeum without median carina, plicae, or longitudinal foveae but anterior margin with a row of tiny longitudinal carinae; callus with 17–20 setae in a patch extending to mesally to spiracle level; spiracle less than its own lesser diameter from metanotum; interspiracular distance = 2.2. Profemur length: width = 3.1: 0.9; protibia = 3.0: 0.5; mesofemur = 3.3: 0.7; mesotibia = 3.9: 0.5; mesotibial spur = 0.7; metacoxa = 3.6: 1.7; metafemur = 0.4: 1.1; metatibia = 4.5: 0.6; longest metatibial spur no greater than metatibial width, less than half basal metatarsomere length. Forewing = 14.1: 5.5; cc: mv: pm = 5.4: 4.2: 1.0. Costal cell with two rows of ventral setae in apical half, separated from a basal (anterior) row of setae in the basal third

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PLATE 1. Figs. 1–6. Torymus cecidicolus. Fig. 1. Habitus of syntype male. Fig. 2. Face and antenna of syntype male. Figs. 36. Female. Fig. 3. Habitus (wings omitted from photo). Fig. 4. Face. Fig. 5. Stigmal and postmarginal veins. Fig. 6. Mesosomal dorsum. Fig. 7. Brasema willei, habitus of type male. 50

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by a long central gap, without dorsal setae, length; stigma short (Fig. 5), not petiolate, distance from stigmal apex to postmarginal vein = 0.3, from uncus apex to postmarginal vein = 0.1, three relatively distinct setal tracks radiating from stigmal apex; speculum bare; admarginal area mostly bare dorsally, but with two irregular rows of admarginal setae ventrally; basal fold with 01 seta; basal cell bare. Hind wing = 8.5: 3.0. All other characters as in male. Gaster strongly arched dorsally (Fig. 3), but this is likely due to distortion of the specimen at death. From gastral base to tip of hypopygium = 5.7. Large membranous area visible between 6th an 7th gastral tergites. Ovipositor sheaths = 22.0 (entire length if straight), with strong brown bristles that are slightly longer than width of sheath. Biology. In addition to Brèthe’s report, this species has also been reared from cecidosid galls on Inga sp. (Fabaceae) (Costa Lima 1936), and Eugenia sp. (Myrtaceae) (DeSantis 1980). Discussion. Both Parasympiesis cecidicola and Syntomaspis alegrensis were reared from the same host material, with the former being based on males and the latter on a female. Our synonymy of T. alegrensis (Brèthes) with T. cecidicolus (Brèthes) n. comb., n. syn. is based on the similarity between the type material comprising the two taxa, as well as the two being reared from the same host material. Athough the Neotropical fauna of Torymus is incompletely known, no other torymids are known to be associated with Cecidosidae and the features given in the diagnosis, especially the form and sculpture of the face, mesoscutum, and scutellum, indicate T. cecidicolus is different from all other species of the genus. The Neotropical torymids described by Ashmead (1904) under the current generic synonym Syntomaspis Förster are similar to T. cecidicolus, but are distinguished from it by much shorter ovipositor sheaths, partially setose frenum, and/or lack of umbilicate punctures. In recent keys to species of Torymus, males of T. cecidicolus key to T. druparum Boheman and females unsatisfactorily to T. ramicola Rushka in Graham & Gisjwijt's (1998) key to European Torymus, unsatisfactorily to T. osborni (Huber) or T. warreni (Cockerell) in Grissell's (1976) key to western Nearctic species, and to T. varians (Walker) in Nikol'skaya & Zerova's (1978) key to torymids of the European part of the former Soviet Union (if the gaster were not distorted). However, T. cecidicolus differs from all of these species in having imbricate to rugulose mesonotal sculpture as opposed to alveolate or aciculate sculpture, and in the unusual coloration of the female. Further, it differs from T. druparum and T. varians in having a much more transverse anellus, from T. ramicola in that the scape is 3x as long as broad in females, from T. osborni in lacking a median propodeal carina, and from T. warreni in lacking metafemoral serrations.

Brasema Cameron Brasema Cameron, 1884: 124. Type species: Brasema brevispina Cameron, by monotypy.

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Cerambycobius Ashmead, 1896: 17. Type species: Eupelmus cleri Ashmead, by monotypy and original designation. Synonymy by Gibson, 1995: 154, 156. Ooderelloides Girault, 1913: 89. Type species: Ooderelloides nigripurpurea Girault, by original designation. Synonymy by Bouèek, 1988: 556. Tropimius Brèthes, 1927: 329. Type species: Tropimius Willei Brèthes, by monotypy. n. syn. Anickia Kalina, 1984: 11–12. Type species: Anickia rara Kalina, by monotypy and original designation. Synonymy by Gibson, 1995: 154, 157.

Brasema willei (Brèthes) n. comb. (Figs. 7–11) Tropimius Willei Brèthes, 1927: 329.

Type material (Deutsches Entomologisches Institut, Eberswalde). Syntype (%): "Porto Allegre" / "Wille 1924.25" / "5. Parasite on Cecidoses eremita" / "No. 5" / "Tropimius willei Brèthes, Brèthes det. 1926" / "TYPUS" / "Syntype" / "type !" / "Eupelmidae Bk. 1957" / "Dtsch. Entomol. Institut Berlin" / "Brasema willei (Brèthes), det. G.A.P. Gibson 2003".

PLATE 2. Figs. 8–11. Brasema willei, type male. Fig. 8. Head, frontolateral (stb = strigose band). Fig. 9. Head, dorsal. Fig. 10. Head and mesosoma, dorsal. Fig. 11. Mesosoma, d 52

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Condition of type. Specimen (Fig. 7) glued by venter to card rectangle, with mouth in glue and mandibles not visible; left antenna beyond scape, right mesotarsus, apical 2 tarsomeres of left mesotarsus, and apex of left forewing missing; eyes, forelegs and gaster collapsed; both scapes collapsed and bent, with right antenna covered in glue and attached to card beyond Fl4, and broken between Fl6 and Fl7; right hind wing glued to top of gaster and left hind wing to bottom of forewing by oily film, which covers wings and parts of the body (Figs. 8, 9). Description. Body 1.4 mm in length. Head and mesosoma variably distinctly metallic green (Fig. 7) to brownish under different angles of light; antenna dark brown; tegula yellowish-brown; legs beyond coxae (mesocoxae concealed) bright yellow except metatibia, and less distinctly mesotibia, with about medial two-thirds light brown. Head (Figs. 8, 9) alutaceous; lower face, interantennal region and parascrobal region with translucent-white spatulate setae, the setae well separated and not obscuring cuticle; frontovertex with scattered, dark brown, unmodified setae on vertex and, based on setal pores, originally with single row of setae adjacent to inner orbit (Fig. 8); scrobal depression arch-shaped, separated from inner orbit by less than one ocellar diameter at about mid-height and dorsally from median ocellus by about one ocellar diameter, with lateral margin rounded and with sulcus extending between dorsal angle and median ocellus (Figs. 8, 9); eye microsetose. Head height: width: length = 51: 55: 33; POL: OOL: LOL = 11: 2: 9; eye height: width = 32: 27; minimum distance between eyes = 22. Torulus with dorsal margin slightly below level of lower margin of eyes (Fig. 8). Antenna clavate; scape ovoid, mesal (convex) surface alutaceous and evenly setose except for longitudinally strigose, bare band ventrally (Fig. 8, stb); pedicel subequal to combined length of basal 4 flagellar segments; flagellum with at least basal 3 segments transverse (see discussion), with short, straight setae; clava collapsed, but about 1.5x as long as wide with broadly rounded apical margin. Mesosoma (Figs. 10, 11) with pronotum vertical medially; mesonotum, dorsellum and propodeum finely alutaceous; mesonotum uniformly covered by brown setae; propodeum with median carina, with plical region bare except for single seta medially about midway between spiracle and median carina, and with callus at most sparsely setose laterally. Forewing hyaline, without speculum; cc: mv: st: pm = 47: 32: 16: 23. Discussion. Males of Brasema can be identified to genus using the key to genera of E upelminae based on males given by Gibson (1995). Although there are about 50 described world species, most were described and currently are classified in Eupelmus Dalman. Only 11 species of Brasema have been described from South America, including B. peruviana (Crawford 1912) n. comb. from Peru and the following 10 species described from Brazil: B. acauda (Ashmead 1904) n. comb., B. aprilis (Ashmead 1904) n. comb., B. basicuprea (Walker 1852) n. comb., B. chapadae (Ashmead 1904) n. comb., B. corumbae (Ashmead 1904) n. comb., B. cyanea (Ashmead 1904) n. comb., B. longicauda Gibson 1995, B. proxima (Ashmead 1904) n. comb., B. schizomorpha Gibson 1995, and B. willei (Brèthes)

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n. comb. All of these species were described originally in Eupelmus except for B. cyanea, described in Zaischnopsis Ashmead, and B. willei. All the names except for B. willei were also based on females. Extreme sexual dimorphism characterizes Eupelminae and species are typically characterized using female morphology within this subfamily of Eupelmidae (Gibson 1995). For these reasons it will not be possible to place B. willei until the sexes are associated for species of Brasema from Brazil and these are revised. The description provided above, particularly the measurements, may not be entirely accurate because of the condition of the type. For example, Brèthes (1927, fig. 2) illustrated the flagellum with the basal four segments quadrate to slightly longer than wide, whereas at least the basal three segments appear distinctly transverse in the one remaining antenna. The single measurement Brèthes included in the original description, of length of the specimen, is accurate and it is possible that his drawing is also accurate, the flagellar segments now appearing transverse because of shrinkage. The antennal drawing by Brèthes is similar to the antennae of some other specimens seen from South America that are otherwise very similar to the type of B. willei. Brasema willei is the only eupelmid yet reared from C. eremita and further rearings from this host could serve not only to associate the sexes but also to provide fresh males for comparison with the type specimen and a more accurate description.

Acknowlegments We thank Andreas Taeger (DEI) for loaning the Brèthes types, Ms. Jennifer Read (AAFC) for the SEM plates, Matthew Buffington (UCR) for assistance in digital imaging, and John Heraty (UCR) for reviewing an initial version of this manuscript.

References Ashmead, W.H. (1896) On the genera of the Eupelminae. Proceedings of the Entomological Society of Washington, 4, 4–20. Ashmead, W.H. (1904) Classification of the chalcid flies, or the superfamily Chalcidoidea, with descriptions of new species in the Carnegie Museum, collected in South America by Herbert H. Smith. Memoirs of the Carnegie Museum, 1: ixi, 225–551, pls. xxi–xxxix. Bondar, G. (1930) Contribuçao para o conhecimento dos Hymenopteros phytophagos Chalcidoideos. Boletim do Museu Nacional de Rio de Janeiro, 6(2), 113–117. Bouèek, Z. (1988) Australasian Chalcidoidea (Hymenoptera). A Biosystematic Revision of Genera of Fourteen Families, with a Reclassification of Species. Wallingford, England:C.A.B International, 832 pp. Brèthes, J. (1927) Hyménoptères Sud-Américains du Deutsches Entomologisches Institut: Terebrantia. Entomologische Mitteilungen, 16, 319–335. Cameron, P. (1884) Hymenoptera (Families Tenthredinidae--Chrysididae). Biologia CentraliAmericana. Insecta, 1, 1–1487 + 20 pls.

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