Eupelmus (Eupelmus) cushmani (Crawford)

Gibson, Gary A. P., 2011, 2951, Zootaxa 2951, pp. 1-97 : 33-43

5. Eupelmus (Eupelmus) cushmani (Crawford)

Figs 9, 28, 37, 42, 50, 59, 63, 75, 76; Map 7

Cerambycobius cushmani Crawford, 1908: 158 . Syntypes, female (BMNH and USNM, examined). Type data: USA, Texas, Victoria ; reared from Anthonomus grandis Boh.

Cerambycobius townsendi Crawford, 1912: 166−167 . Holotype, female (USNM, examined). Type data: Peru: Department of Piura; reared from? Anthonomus vestitus . Synonymy by Gahan, 1951: 172.

Eupelmus cyaniceps amicus Girault, 1916b: 244 . Syntypes, female (USNM, examined). Type data: USA, New Mexico, Las Cruces ; reared from Bruchus amicus Horn. N. syn.

Eupelmus cushmani ; Cotton & Good, 1937: 42, 61.

Eupelmus amicus ; Burks, 1979: 881.

Description. FEMALE. ( Fig. 37) Length about 2.1−5.2 mm. Head sometimes mostly dark brown, usually in smaller specimens, but more commonly ( Fig. 28) with frontovertex dark mesally between scrobal depression and posterior ocelli except for variably extensive and distinct longitudinal greenish region along inner orbit (inner margin of region sometimes more yellowish, coppery or violaceous) extending mesally to at most about level of inner margin of posterior ocellus, though rarely mesal region green and laterally purple along inner orbit; parascrobal region, interantennal region and clypeal region also often variably extensively dark, and vertex posteriorly or along upper orbit more purple; maxillary and labial palpi dark brown. Antenna dark brown with slight metallic luster on scape and pedicel or often scape with at least slight yellowish-orange tinge apically to entirely yellowish-orange ( Fig. 28). Mesosoma similar in color to head with tegula brown or dark with slight metallic luster, otherwise mostly dark brown with slight metallic green luster under some angles of light, usually in smaller specimens, to quite distinctly metallic green except lateral lobe at least dorsolongitudinally, scutellum variably extensively, and often convex or concave part of medial mesoscutal lobe variably dark, brownish to reddish-violaceous. Forewing hyaline; venation yellowish-brown; setae uniformly brownish or sometimes lighter in basal cell and on submarginal vein. Front leg with trochanter yellowish to dark brown; femur sometimes entirely dark brown in smaller specimens, more commonly with trochantellus and extreme apex yellowish; tibia often completely yellow or almost completely yellowish except for dorsal and ventral brown bands or, in smaller specimens, more extensively brown except knee and apex more widely yellowish; tarsus yellowish except for apical brown tarsomere to more or less uniformly yellowish-brown. Middle leg with mesotibial apical pegs and mesotarsal pegs dark, otherwise often yellowish to yellowish-orange at least beyond coxa and sometimes including coxa or, particularly in smaller specimens, femur and tibia in part more distinctly brownish but knee, tibia apically, and at least basal tarsomere lighter yellowish to white. Hind leg with trochanter yellowish to brown; femur commonly brown basally and yellowishorange apically to a greater or lesser extent, though rarely entirely yellowish-orange to entirely brown; tibia usually yellowish-orange to variably extensively brown (except often ventral margin) except for knee and apically more whitish, though sometimes entirely brown except for about apical 0.2; tarsus with at least basitarsus white and apical tarsomere brown, but tarsomeres 2−4 variably white to dark brown. Gaster entirely dark brown or at most with metallic green luster anteriorly on basal tergum and sometimes laterally on terga, and with brown hairlike setae similar in color to cuticle; dark inner plate of ovipositor sometimes projecting distinctly beyond apex, but at least sheaths with short dark basal region abruptly delineated from longer medial whitish region, the whitish region variably abruptly and distinctly delineated from yellowish-brown to dark brown apical region, but at least apical darker region subequal in length or longer than lighter colored medial region.

Head with frons variably distinctly and coarsely sculptured, sometimes almost meshlike coriaceous ( Fig. 28), though usually at least some cells slightly sunken so as to be delineated by rounded ridges or somewhat imbricate along inner orbits, to more extensively and conspicuously reticulate or transversely imbricate-wrinkled to reticulate-strigose along inner orbits, and merged into parascrobal region through slight undulation; vertex variably transversely alutaceous-reticulate in smaller specimens and rounded into occiput to quite strongly transversely strigose-carinate in larger specimens and continued above posterior ocelli in similar plane as frons and quite narrowly and abruptly angled relative to occiput ( Fig. 37); scrobal depression reticulate-rugulose to transversely reticulatestrigose; IOD = 0.36−0.44× head width; OOL: POL: LOL: MPOD = 0.5−0.9: 2.3−2.9: 1.7−1.8: 1.0. Antenna with combined length of pedicel + flagellum = 1.06−1.3× head width; scape about 4.2−5.4× as long as wide, in outer view ventral margin often almost straight and angulate but sometimes slightly sinuate with very slender, inconspicuous flange over about apical half; pedicel in lateral view about 1.9−2.1× as long as wide; fl1 slightly transverse to quadrate; fl2 about 1.8−2.5× as long as wide and about 3.0−5.0× as long as fl1; subsequent funiculars increasing in width to quadrate or slightly longer than wide fl8; clava about 1.9−3.0× as long as wide, 0.8−1.2× combined length of apical three funiculars, and 0.25−0.45× length of funicle. Mesoscutum almost uniformly meshlike reticulate except lateral lobe more minutely coriaceous mediolongitudinally and mesoscutal medial lobe usually more transversely alutaceous-reticulate in smaller to reticulate-imbricate anteriorly in larger specimens. Scutellar-axillar complex with axillae obliquely alutaceous-reticulate to more coriaceous-imbricate, but at least scutellum mostly meshlike coriaceous to coriaceous- or somewhat reticulate-imbricate laterally. Prepectus sometimes superficially bare or only very inconspicuously setose, but if with 2 or 3 distinct lines of white, hairlike setae mediolongitudinally then setal apices not extending to margins (Fig. 42). Acropleuron finely meshlike reticulate anterior and posterior of medial microsculptured region, the cells sometimes somewhat larger posteriorly than anteriorly but with flat surfaces defined by slightly raised ridges. Forewing with linea calva variable, often a distinctly isolated bare band (cf. Fig. 45) but sometimes continuous or almost continuous through vanal area (cf. Fig. 48); costal cell ventrally setose along length, with 2 or 3 lines of setae medially, and dorsally either bare or much more commonly setose apically along leading margin for distance usually little greater than length of parastigma (cf. Fig. 45) or at most half length of cell; cc: mv: pmv: stv = 4.1−5.2: 4.1−5.3: 1.0−1.5: 1.0. Mesotibia with apical row of 3−8 pegs; mesotarsus ventrally with pegs on basal four tarsomeres, basitarsus with 10−24 pegs arranged distally in double row on either side, second tarsomere with 3−7, third tarsomere with 1−4, and apical tarsomere with 1 or very rarely 2 pegs on one side. Propodeum with U-shaped plical depression extending to foramen; callus with white, comparatively sparse hairlike setae not obscuring cuticle or sculpture (Fig. 50). Gaster with inner plate of ovipositor variable, not projecting beyond apex if ovipositor sheaths relatively short but usually extending beyond apex quite distinctly if sheaths long; ovipositor sheaths about 0.8−1.33× length of metatibia and about 0.97−1.7× length of marginal vein.

MALE (Fig. 59). Length about 1.5−3.3 mm. Head dark metallic green to bluish-green or smaller specimens more brown with variably distinct metallic luster; maxillary and labial palpi dark brown or apex of apical palpomeres at most yellowish-brown. Antenna dark brown except often scape and pedicel with metallic luster similar to head capsule, and outer surface of scape sometimes yellowish ventrally along scapular scrobe. Mesosoma similar in color to head, the tegula at least dark brown and often also variably distinctly metallic. Front leg with trochanter, trochantellus, and femur except sometimes narrowly apically, brown to dark except with at least slight metallic luster similar to mesosoma; tibia almost completely dark except for extreme base and apex to almost completely yellowish except usually for dorsolongitudinal dark band; tarsus yellow to brownish-yellow or with apical 1 or 2 tarsomeres more distinctly brown. Middle leg mostly dark except knee and apex of tibia variably distinctly lighter, and basal 1−4 tarsomeres white. Hind leg with similar color pattern as middle leg. Forewing hyaline. Gaster with basal tergum sometimes metallic green to bluish-green basally, but remainder brown.

Head with frons reticulate; scrobal depression reticulate at least though scrobes and sometimes over about dorsal half of interantennal region above level of toruli; vertex rounded into occiput, reticulate to transversely reticulate-strigose but without evident transverse carina. Head with IOD about 0.5× head width: OOL: POL: LOL: MPOD = 0.5−0.7: 2.5−3.1: 1.4−1.6: 1.0; lower face with uniformly distributed white setae, the setae usually longer and more distinctly curved laterally toward malar sulcus but sometimes with only one distinctly longer seta opposite long seta below malar sulcus. Antenna (Fig. 63) with scape ovoid, about 2.2−2.5× as long as maximum width, the outer surface ventrally with variably crowded, more or less pinprick micropunctures apically ( Fig. 75), or if sculpture more extensive along length of scrobe then shallow and more or less elongate-strigose or rugulose rather than distinct pits ( Fig. 76); length of pedicel + flagellum about 1.4−1.6× head width; pedicel subglobular, only about 1.2−1.4× as long as wide, ventrally with line of 4 or 5 distally curved setae; flagellum conspicuously, densely setose with projecting, curved setae so as to appear robust-filiform, the flagellomeres of equal width or including setae slightly tapered to narrower clava; fl1 very strongly transverse, disc-like, and at least superficially bare, at most with single line of setae along apical margin; fl2 and fl8 both oblong, about 1.1−1.8× and 1.3−1.5× as long as wide, respectively; clava lanceolate with micropilose sensory region occupying apical two claval subsegments (usually collapsed in air-dried specimens), about 2.2−2.8× as long as wide and about 0.6−0.7× as long as apical three funiculars. Mesoscutum variably deeply meshlike reticulate; axillae and scutellum usually more shallowly reticulate-imbricate to meshlike coriaceous in small specimens. Propodeum finely coriaceous to almost smooth and shiny, and callus similarly sculptured with setae originating from tiny bumps. Forewing with cc: mv: pmv: stv = 4.0−4.5: 3.0−3.9: 1.0−1.2: 1.0; costal cell dorsally with line of dark setae extending over only about apical onequarter to one-third and ventrally with dark setae continuously along length, mesally with 1 or 2 lines.

Regional material examined (715 ♀, 290♂). CANADA. ONTARIO: Chatham, 26.VII.54, K.G. Davey, host plant red clover (1♀) . Mainfleet Bog, 8 km S. Welland, 14-21.IX.87, A. Stirling (1♀). Kent Co., Rondeau Prov. Park, Southpoint Trail near East parking lot, 7.X.2003, S. Marshall (1♀) .

USA. 4♀, 4♂ (MEMS). 25.VIII.20, 4, 5, 6, 9, 13, 14, 15, 16, 18, 19.X.20 (11♀, 2♂ MEMS). 218 01 (1♀ USNM). 5893 01 (2♀ USNM). 3593g Hopk U.S. (3♀ USNM). 4028 o, 17.IX.1886 (1♂ USNM). P1906.0, [?], par. Anthonomus grandis (1♀ USNM). 19, 30.VIII.1893, 10.XI.1893, par. Bruchus obsoletus (4♀ USNM). Hunter No. — 1334, 17.IX.07, par. Tyloderma foveolatum (1♀ USNM); 1410, 28.VI.07, 21.VIII.07, par. Bruchus prosopis (1♀, 1♂ USNM); 1450 (2♀ USNM); 1454, 29, 30.IX.07, 1, 9, 10.X.07, par. Bruchus (11♀ USNM); 1455, 30.IX.07, 1, 10.X.07, par. Bruchus (3♀, 4♂ USNM); 1456 [?] (1♀ USNM). Calvert, June 1909, [?] July 7 (1♀ USNM). ALABAMA: Etowah Co., Attalla, 16.VIII.65, T.L. Chesnut, em. 9.II.65 from fallen cotton squares and bolls (1♀ USNM). Talladega Co., Coleta, H.H. Smith (1♀ USNM). ARIZONA: 2552, C.F. Baker (2♀ USNM). Huachuca Mountains, 6000', 14.VI.28, A.A. Nicol (1♀ CASC). S. Catalina Mountains, 21.XI.13, J.J. Thornber, par. Anthon [omus] gr. [andis] thurb.[eriae] (1♀ USNM). From USDA colony, southern Arizona, 1968, R.E. Fye (1♀ MEMS). Cochise Co., Cochise Stronghold , 14.X.73, C.D. Johnson, em. by 13.XI.73, reared seeds Mimosa biuncifera (1♂ TAMU). Douglas, W.W. Jones (1♀ USNM). 6 mi. NW Huachuca City, 6.X.72, em. by 18.XII.72, C.D. Johnson, reared seeds Acacia constricta (1♀ USNM). San Simion Road, 4 mi. NNW Portal, 31º58'N 109º9'W, 4600', 25.V.88, em. VI.88, H.A. Hespenheide, ex Acacia constricta Benth. seed (3♀, 1♂; 1♀ with adult Stator pruininus (Horn) glued to point below it). Gila Co., Roosevelt , 18.XI.40, R.L. Furniss, Cercidium microphyllum (2♀ USNM). Maricopa Co., Lone Mountain Road, 25.VII.83, M. Hetz, reared seeds Cercidium microphyllum (1♀ USNM, CNC Photo 2010-45). Pima Co., Quijotoa , 1 mi. N., 8.X.77, em. 17.VII.77, C.D. Johnson, reared seeds Acacia constricta (1♂ TAMU). South of Sells, 8.X.77, C.D. Johnson, em. 17.X.77, reared seeds Nissolia schottii (1♂ TAMU). Tucson — Kunze, Baker coll. (1♀ USNM); 1, 4.VIII.21, parasite of mesquite weevil (4♀ UAIC); 3.VIII.31, G.D. Butler, swept from cotton (1♀ UAIC). Pinal Co., E. of Florence, fall 1976, J. Holcomb, seeds in crop of Gambol’s [?], Stator pruininus (1♀ UAIC). McCleary’s Ranch, Stone Cabin Canyon, Santa Rita Mountains , 4.XII.13, ex Anthonomus (1♀ USNM). Santa Cruz Co., Pena Blanca Lake , 3 mi. W., 13.VI.72, em. by 5.VII.72, C.D. Johnson, reared seeds Calliandra eriophylla (1♂ TAMU). Ranch, 24.VI-14.VII.2009, E.E. Grissell (1♀). Ruby, 3 mi. NW, ca. 4200', 15.X.76, C.D. Johnson, em. by 10.XII.76, reared seeds Mimosa biuncifera (1♂ TAMU). Santa Rita Mountains, Amado , 8.III.67, R.E. Fye, em. from Thurberia weevils in Gossypium thurberi before 14.III.67 (1♀ MEMS). Y avapai Co., Bumble Bee Road, 25.VII.83, M. Hetz, reared seeds Cercidium microphyllum (2♀, 1♂ TAMU). Bull Pen Road, 2.IX.83, M. Hetz, reared seeds Acacia constricta (1♀ TAMU). Camp Verde, ca. 5 mi. S., 30.VI.69, C.W. Forister, em. 3.VII.69, reared seeds Condalia lycioides (1♂ UCDC). W. Clear Creek Campground, 7.5 mi. ESE Camp Verde, 3500', 8.IX.83, 10.X.83, M. Hetz, reared seeds Acacia constricta (2♀ TAMU). ARKANSAS: Benton Co., Bentonville, 23, 24.VI.18, 2, 4.VII.18, 15, 19, 29, 30.VI.19, 2.VII.19, VII.20, VII.23, D. Isley, par. Mineola (19♀, 5♂ USNM). Siloam Springs, 22.VIII.08, S.W. Foster (1♀ USNM). Clark Co., Gurdon, 26.IX.10, Hunter No. 1326, bred cotton square (1♂ USNM). Johnson Co., 8 mi. N. Clarksville, T10N, R23W, Sec. 2, 6.VII.97, R.L. Brown, sweeping Coreopsis tinctoria (1♀ MEMS). Miller Co., Texar- kana, 11.VIII.09, C.E. Hood, par. A. grandis (1♀ USNM). Ouachita Co., Camden, 9.IX.08, C.E. Hood, par. A. grandis (1♀ USNM). CALIFORNIA: Camp Angeles, Hopk. US 20816-b, c, Amorpha californica (1♀, 2♂ USNM). Riverside Co., Gavilan Hills, NW of crossing Gavilan & Idaleona Roads, 19.III.98, D.C. Hawks & J.M. Heraty (1♂ UCRC). Menifee Valley, 1.V.81, J.D. Pinto (1♂). Menifee Valley, hills on W. end, 1800', 33º39'N 117º13'W — 27.VIII-1.IX.80 (3♂), 8-15.V.82 (1♂), J.D. Pinto; 1.X-1.XI.80, J.B. Woolley, J.D. Pinto & J. LaSalle (1♂ TAMU). Rubidoux Jurupa Rancho Co. Park, 27.VI.86, J. LaSalle (1♂). San Bernardino Co., 6 mi. E. Yermo, dry bed Mojave River, 18, 22.IX.52, H.B. Leech & J.W. Green, reared from seed pods of Prosopis pubescens Benth. infested by Algarobius prosopis (Leconte) (1♀, 1♂ CASC). San Diego Co., Bonsall, 1 mi. W., 8.VIII.79 — G. Gordh (1♂ UCRC); C. Melton (1♂ UCRC). Santee, Carlton Oaks Golf Course, 22.I.81, G.T. Scriven (1♀). Sentenac Canyon, 22.IV.81 — J. Woolley (1♂); J.T. Huber (2♂ UCRC). Ventura Co., Oxnard, 26.XII.34, Dunning, bred from Mylabris pruininus (2♀, 1♂ CASC). DELAWARE: New Castle Co., Wilmington, 18.VI.71, P.P. Babiy (1♀ ZSMC). DISTRICT OF COLUMBIA: Chain Bridge , 18.VIII.20, J.C. Bridwell, ex Acanthoscelides perforatus (3♀ USNM). Washington — 12.VII.08, 31.VII.07, par. Tyloderma foveolatum (4♀ USNM); 15.IX.07, on Baptisia sp. (3♀ USNM); 16, 17.IX.07, J.A. Hylsop, bred from Baptisia tinctoria (1♀, 1♂ USNM). FLORIDA: Alachua Co. , 15.I.30, R.R. Mathews, Florida Fruit Fly Trap Survey (1♀ USNM). Gainesville, 13.V.75, Rhyanconia frustrana , Pinus taeda (1♀ FSCA). Baker Co., Glen St. Mary, 24.X.2007, E. Zoll & S. Fullerton (1♀ UCFC). Jefferson Co., Monticello — 22.VI.16, A.I. Fabis, bred 5, 7, 12.VII.16 from A. [crobasis] hebescella Hulst (2♀, 2♂ USNM); 27, 30.IV (3♀ FSCA), 1.VI.28 (4♂ FSCA), 30.VI.28 (1♂ FSCA), F.W. Walker; 3.VII.27, F.W. Walker, ex Acrobasis caryae Grote , secondary parasite (6♀ FSCA). Monroe Co., Big Pine Key, 6.III.92, D.M. Lott, D.H. Habbeck & J. Gillmore, Rhyncosia [!] minima seeds (1♀ FSCA). Orange Co., Orlando, 28.VII.199 [!] (1♀ UCFC), 5.VII.96 (1♂ UCFC) S.M. Fullerton. Wekiwa Springs State Park, 19.II.2001, 21.III.2001, P. Russell & S. Fullerton (2♀ UCFC). Pinellas Co., Dunedin, L.J. Bottimer — 15.II.30 (1♂ AEIC); 15, 24.II.30, 11.III.30, 18.IV.30, ex Acanthoscelides floridae in Amorpha seeds (3♀, 6♂ CNC; 4♀, 6♂ USNM). Seminole Co., Econ Wildlife Area, 6.V.2000, T.Smith, P. Russell & S.M. Fullerton (3♀ UCFC). Suwannee Co., Branford, 10.III.56, H.F. Howden, host Acanthoscelides floridae Horn (2♀). GEORGIA: Chattahoochee Co., Fort Benning, 8, 14.VI.95, B.T. Sullivan (2♀ USNM). Chattooga Co., Summerville, 16.VIII.65, T.L. Chestnut, em. 31.VIII.65 from fallen cotton squares and bolls (1♀ MEMS). Clarke Co., Athens, 9.VIII.91, J. Pickering, ex Ceratomia catalpae (1♀). Mitchell Co., Dewitt, 6.III.13, J.B. Gill, bred from E. [narmonia] caryana (1♀ USNM). Peach Co., Byron, 27.II.86, A.A. Amis, ex gall Dryocosmus kuriphilus (1♀ USNM). Fort Valley 15.III.37. G.F. Moznette, ex Laspeyresia caryana in pecan shucks (1♀, 1♂ USNM). Putnam Co., Oconee National Forest, 18.VII.95, B.T. Sullivan (1♀). Spalding Co., 1.VII.37, T.L. Bissell, bred from Bruchus pisorum (2♀ USNM). Thomas Co., Thomasville, G.D. Smith, par. A. grandis (1♀, 1♂ USNM). Tift Co., Tifton (1♀ USNM). ILLINOIS: Greene Co., Carrollton, 5.VII.96, H.E. Anderson (1♀ UCRC). Iroquois & Ford Co., Loda and Prospect Cemetaries Prairie, VI.2001, J. Tooker, reared from Silphium laciniatum and S. terebinthinaceum (1♂ UCDC). Moultrie Co., XI.31, reared IV.32, Bigger, reared from Desmoris fulvus (1♀ USNM). INDIANA: Tippecanoe Co., Lafayette, X.38, Montgomery, issued from seed on red bud (1♀ USNM). KANSAS: Doniphan Co., Wathena, 27.VII.39, S.C. Schell, ironweed (4♀ USNM, 1♂). Riley Co., Manhattan, S.C. Schell — X.37, em. 28.III.38 from seed of Hibiscus (1♀ USNM); VIII.37, em. 28.III.38 from mummified apples (1♀ USNM); VIII.37, from pod Cercis canadensis (1♀ USNM); X.37, em. 20.IV.38 (2♀ USNM). KENTUCKY: Mason Co., 30.VII.50, N. Taylor (1♂ USNM). LOUISIANA: Avoyelles Parish, Mansura,3.II.10, C.E. Hood, in Spanish moss (1♀ USNM). Caddo Parish, Forbing , 18.IX.07, par. Spermophagus robiniae (1♀ USNM). Shreveport, 1907 (1♀ MEMS). Cameron Parish, Jhnsn’s Byu [Johnson Bayou], 26.VIII.06. par. Anth. albopilosus (1♀ USNM). DeSoto Parish, Mansfield, 23, 24.VIII.06, par. A. grandis (18♀, 3♂ USNM). East Carroll Parish, [Lake] Providence, 15, 25.VIII.14, 1.IX.14 (2♀, 1♂ MEMS). Lincoln Parish, Ruston, 9.IX.08, W.D. Pierce, bred 10.IX.08, A. grandis (1♀ USNM). Madison Parish, Tallulah — 3.IX.14, 28.VIII.14 (3♀ MEMS); 8, 10, 12, 13, 22, 29.VIII.32 (9♀ MEMS); par. A. grandis (1♀ USNM); 3.IX.09, 20.IX.13, bred cotton boll weevil (1♀, 1♂ USNM); 19.IX.09, 20.XII.09, bred cotton square (2♀ USNM); 24.IX.09, bred square (1♀ USNM); 10.VIII.10, R.A. Cushman, par. A. grandis (2♀ USNM). Rapides Parish, Alexandria, 2, 8, 16, 20, 22.VIII.07, par. A. grandis (6♀, 3♂ USNM). Glenmora, 22, 23.VIII.07, par. A. grandis (5♀, 2♂ USNM). Sabine Parish, Many, 23, 24.VIII.06, par. A. grandis (22♀, 6♂). near Noble, III.60, M.W. Sanderson, ex Caryobruchus in Sabel (1♀ USNM). MARYLAND: Calvert Co., Chesapeake Beach, 13.VI.85, L. Masner (1♀). Charles Co., Newburg, 17.VII.73, 3.VII.74, J. Lasomb (1♀, 1♂ MEMS). Harford Co., Riverside, iss. XII.26, seed pot of Oenothera (1♂ USNM). Montgomery Co., Ashton, 4 mi. SW, 39º06'36"N 77º01'30"W, 10.IV.2004, G. Gibson, on dead Pru- nus infested with scolytids/cerambycids (3♂). Plummers Island, III.61, Spangler, Acanthoscelides alboscutellatus in Ludwigia (3♀, 6♂ USNM). Prince George’s Co., Beltsville, III.61, Rhyanconia frustrana material (1♀ USNM). Berwyn, 27.IX.1897, bred from B. obtectus in beans (5♀ USNM). College Park, 22.VII.18, from L. molesta (1♀ USNM). Patuxent Wildlife Preserve, 24.VI.86, D. Wahl (2♂). Wicomico Co., Salisbury, 2, 8, 11.IX.29, H.S. McConnell, Laspeyresia molesta (secondary) (1♀, 2♂ USNM). MICHIGAN: Washtenaw Co., Ann Arbor — 21- 23.V.60, H. & M. Townes (1♀ AEIC); 12-21.VII.82, R. Wharton (1♀ TAMU). MISSISSIPPI: Adams Co., Natchez — 20, 21.VIII.33, Station (1♀, 1♂ MEMS); 30.VIII.33, 4, 5, 6.IX.33, Junkins (3♀, 1♂ MEMS). Sibley, 5.VII.20, A. Flemming (1♀ MEMS). Bolivar Co., Shaw , 1.IX.14 (1♂ MEMS). Carroll Co., Slaton Field — 9.I.65, em. fallen cotton squares and bolls (1♀ MEMS); 10.IX.65, 16.VIII.66,W.H. Cross, cultivated cotton (1♀ MEMS); 10.IX.66, experimental cotton plot, em. 21, 23, 25.I.67, H.C. Mitchell (1♀, 3♂ MEMS). Claiborne Co., Port Gibson, 29.VIII.14, 2.IX.14 (2♀, 1♂). Harrison Co., Gulfport, 28.III.57, J.F. Coyne, Rhyanconia frustrana (1♀ USNM). Hinds Co., Utica, VIII (1♀ USNM). Jackson Co., Pascagoula, 29.V.26, R.P. Colmer, reared from casebearer ex larva, 19.VI.26 (1♀ MEMS). Jones Co., Laurel , 21.VII.12, Ellesville, W.D. Pierce coll. (1♀ USNM). Lauderdale Co., Arundel [Spring?], 20.VIII.12, W.D. Pierce, par. A. grandis (1♀ USNM). Oktibbeha Co., 8.III.68, D.D. Hardee, in alfalfa (2♀ MEMS). A & M College, J.M. Langston, reared from L771, pecan — 16.VI.30 (1♀ MEMS); 28.VI.30, reared from Olethreutidae (1♀ MEMS); 20.VIII.30 (1♀ USNM). Bradley, 4 mi. WNW, 2.III.74, W.L. McGovern, em. 25.III.74 from boll weevil infested cotton (1♀, 1♂ MEMS). Dorman Lk, 16.IX.81, R.L. Brown, em.8.X.81. Helianthus angustifolius (1♀ MEMS). Maben, 4 mi. E., 4.VI.71, J.E. Leggett (1♀ MEMS). near Oktoc, 22, 27.II.64, W.H. Cross (2♀ MEMS). Pontotoc Co., Ecru, 20.XI.80, R.L. Brown, ex stems of Oenothera biennis infested with Mompha eloisella , em. 6.I.81 (1♀ MEMS). Sharkey Co., Rolling Fork, 27.VIII.14 (1♂ MEMS). Stone Co., Wiggins, 23, 25, 29.V.29, 1, 3, 8.VI.29, J.P. Kislanko, reared from A. juglandis (5♀, 1♂, MEMS; 1♀, 2♂ USNM). Washington Co., Delta Experimental Forest near Stoneville, 33º29'N 90º54'W, 18.IV-9.V.88, N.N. Schiff (1♀ UCDC). Stoneville — VI.78, W.H. Cross, lab. colony (14♀, 13♂ MEMS); 3 mi. W., 18.I.68, W.H. Cross, em. dry sqs + bolls from standing stalks cotton, 3.II.68 (1♀ MEMS); 3 mi. W., 28.IX.67, W.L. McGovern, em. Croton lindheimeri , dead in box, 2.II.67 (1♀ MEMS). Yalobusha Co., Water Valley, 27.II.25, C.G. Wallace (1♀ MEMS). MISSOURI: Boone Co., Columbia , 19, 25, 27.VIII.77, W.S. Craig, ex seed red bud (1♀, 2♂ UMRM). Henry Co., Chapel View Prairie, 29.VI.99, M. Gates (1♂ UCRC). Phelps Co., St. James, 14.VII.53, P.J. Spangler, ex Chlamys gibbosa cocoons (1♀ USNM). NEW JERSEY: Burlington Co., Beverly, 1934, Oriental Fruit Moth Par. Invest., Grapholita molesta (1♀ USNM). Bridgeboro, 1945, Oriental Fruit Moth Par. Invest., ex G. molesta (1♀ USNM). Browns Mills, 24.VII.35, L.J. Bottimer (1♀ USNM). Moorestown, 1931, Oriental Fruit Moth Par. Invest., ex Macrocentrus cocoon (1♀ USNM). Camden Co., Haddon Heights, VIII.31 (1♀ AEIC; 8♀, 9♂ CNC; 6♀, 3♂ USNM; 1♀ WFBM), 8.VIII.31 (5♀ USNM; 1♂ WFBM), L.J. Bottimer, ex Bruchus brachialis , in Vicia villosa . Cumberland Co., Cedarville, 2.VIII.76, K.W. Thorpe (1♀ USNM). NEW MEXICO: 2♀ USNM. Dona Ana Co., Las Cruces, from Bruchus amicus Horn (2♀, 2♂ syntypes E. cyaniceps amicus ). Hidalgo Co., 9.3 mi. W. Animas, 26-30.VII.92, 4400', G. Gibson (1♂). NEW YORK: Nassau Co., Farmingdale, 31.VII.38, H. & M. Townes (1♀ USNM). Tompkins Co., Ithaca, F.H. Chittenden (1♀ USNM). NORTH CAROLINA: Beaufort Co., Washington, J.P. Weyerhaeuser seed orchard, coll. as pupae 4.V.76, G.F. Fedde, in dying Pinus taeda L. twigs associated with weevil galleries (1♀ USNM). Burke Co., Pineola bridge at Linville Falls P.O., reared 24.X.80, A.T. Drooz, ex Diprion similis (6♀ NCSU). Cabarrus Co., 8.X.55, T. Daggy (1♀ NCSU). Craven Co., 2.VII.17, R.W. Leiby, bred material exp. No. 39-57 (1♀ NCSU). Cumberland Co., Fort Bragg, 16.V.67, J.D. Birchim (1♂ CASC). Edgecombe Co. — reared 314 (1♀ USNM); 26.VI.40, A. Barnes, pecan leaves (1♀ USNM). Guilford Co., Greensboro, J.S. Pinckney, reared from M. [ompha] eloisella (1♀ USNM). Harnett Co., Lillington, 13.VI.41, S.C. Schell (2♀ USNM). Iredell Co., Statesville, 12.VII.48, C.C. Hill, bred from B. brachialis (5♀, 1♂ USNM; ♂ CNC Photo 2010-25). Johnston Co., Clayton, Central Crops Research Station, 12.VIII.58, H.H. Neunzig (1♀ NCSU). Martin Co., S. of Jamesville, 19.XI.74, L.K. Lawrence, reared from pupa Rhyacionia frustrana (1♀ NCSU). Mecklenburg Co., Beverly Woods, P.P. Babiy — 2.VI.64, on Dacus car . (1♂ ZSMC), 12.VI.64 (1♀ ZSMC). Montgomery Co., Uwharrie National Forest, junction State Road 1179 & Government Road 597, 25.VI.93, R.L. Blinn, taken on Pinus virginiana (1♀ NCSU). Moore Co., Eagle Springs, 28.III.50, M.H. Farrier, shaken from peach (1♀ NCSU). Person Co., 18.VIII.84 — H.D. Blocker (1♀ NCSU); J.F. Cornell, Solidago (1♀ NCSU). Pitt Co., Grifton, 2.VII.75, R.J. Galloway (1♀ NCSU). Polk Co., Tryon, W.F. Fiske (3♀ USNM). Rowan Co., Salisbury, J.S. Pinckney, reared from B. brachialis (7♀, 3♂ USNM; ♂ CNCI LB-specm 2010-004). Scotland Co., 34.98013ºN 79.55366ºW, 138 m, 17.X.2010, G. Gibson (1♀). Swain Co., Smokemont, 6.VI.77, N.C.D.A. (1♀ NCSU). Wake Co., 27.XI.62, P. Kamnerdratana (1♀ NCSU). Raleigh — 27.VII.40, S.C. Schell (1♀ USNM); VI.41, S.C. Schell, par. Pine moth (1♀ USNM); II.59, H.H. Neuhzig (1♀ NCSU); IX.63-IV.64, J.F. Cornell, alfalfa field sticky trap (1♀ NCSU). OHIO: Athens Co., Athens, 4.XII.96, A.C. 2738 (14♀ USNM). Butler Co., Oxford, F.J. Rickey, em. 16.I.95, from Cercis canadensis seedpod (1♀ UCDC). Eire Co., Sandusky, H.G. Ingerson, 22.VIII.17, par. grape berry moth (1♀ USNM). Ottawa Co., Oak Harbor, 20.VII.32, ex Macrocentrus cocoon, Oriental Fruit Moth Par. Invest. (3♀ USNM). Wayne Co., Wooster, 8.II.97, A.C. 2738 (5♀ USNM). OKLAHOMA: Latimer Co., Red Oak, X.90, IX.94, K. Stephan (2♀). OREGON: Benton Co.,Corvallis, em. 18.I.83 (1♀). PENNSYLVANIA: Adams Co., Arendtsville, VI-VII.40, J.H. Beacher, Coleophora malivorella (1♀, 9♂ USNM). Berks Co., em. 17.VIII.38, S.C. Schell, par. Apion rostrum (4♀ USNM). 10.IX.38, S.C. Schell, em. III.39, par. moth on Smilax (1♀ USNM). Chester Co., West Chester, 30.VI.25, F.F. Smith, Malacasoma americana Fab. (2♀ USNM). York Co., Hanover, 15.VI.30, Bridwell & Barber (1♀ USNM). SOUTH CAROLINA: Florence Co., Florence — 1924, E.W. Dunnam (11♀, 3♂ MEMS); 22.VII.26, A.R. Ground (1♀ MEMS); 2.VIII.29, C.F. Rainwater (3♀ MEMS). Greenville Co., Cleveland, 16.IX.75, 12.V.76, G.F. Townes (2♀ AEIC). Greenville, 7.VI.52, 5.IX.52, G. & L. Townes (2♀ AEIC). Jasper Co., Tillman, 10 mi. NW, 26.IV.87, H. Goulet (1♀). Marlboro Co., Bennettsville, cotton boll (2♀ USNM). Clio, cotton boll (3♂ USNM). Pickens Co., Clemson — III.75, R. Fox, Rhyanconia sp. on pine (1♀ UCRC); IV.93, R73-91, ex R. frustrana (1♂ UCRC). Clemson Collage, G. Ainslie, Webster no. 1831 (2♀ USNM). Richland Co., Columbia , 18.VIII.51, L. & G. Townes (1♀ AEIC). TENNESSEE: Blount Co., GSMNP [Great Smoky Mountains National Park], Cades Cove, Mill Creek, N35.34.60/W83.50.8, Steck & Sutton — 1.X.2003, 1792' (1♀ UCFC), 1.X.2004, 1720' (1♀ UCFC). TEXAS: Kings Ranch, 4, 6.XI.78, R. Medved, ex Acacia seed (2♀, 1♂ UCRC). Anderson Co., Palestine, 10.VIII.06, par. A. grandis (21♀ USNM). Bee Co., Beeville, 12.VII.06, bred cotton square, par. Bracon mellitor (1♀ USNM). Bexar Co., San Antonio, 13.XII.84, Bezark, Kitayama & Orsack, ex legume pods (1♀, 2♂ UCDC). Bosque Co., Roswell, 28.X.06, par. A. grandis (1♀ USNM). Brazos Co., College Station — 1923, S.W. Bilsing, par. Acrobasis caryivorella (1♀ USNM); 2.VII.75, B. Cutter, emerged from nutlet with A. nuxvorella (1♀ TAMU); 15.VIII.15 (1♀ TAMU). Brewster Co., 17 mi. S. Alpine, 8.VIII.82, J.C. Schaffner (1♀ TAMU). Big Bend National Park — Basin, 23.V.83, R.S. Anderson (1♀, 1♂); 2.7 km NE Castolon, 760 m, 14.VIII.82, G. Gibson (1♀); Cottonwood campsite, 2300', 13-14.VII.82, G. Gibson (3♀), 13.VII.82, R.S. Anderson (1♀); Glenn Spring Road, 0.5 mi. in, 3000', 23.VI.82, G. Gibson (1♀); Govt, Springs, Grapevine Hills Road, 1.IX.71, E.E. Grissell & R.F. Denno (4♀ UCDC, CNC Photo 2010-24); 12.5 mi. SE Panther Junction, 2500', 10- 18.VII.82, G. Gibson (1♀, 2♂); Rosillos Mountains, Buttrill Spring, 4-5.X.91, R. Wharton (1♀); Rosillos Mountains, Nine Mile Draw, 10 mi. W. 385, 29º35'N 103º16'W, 12.IX.93, E.E. Grissell & R.F. Denno (1♂ USNM). Brown Co., Brownwood, 25.VI.18, 4.VII.18, A.I. Fabis, par. Acrobasis sp. (1♀ USNM). Callahan Co., Pecan Bayou, 2.VII.18, 22.VII.18, A.I. Fabis, par. Acrobasis sp. (1♀ USNM). Cameron Co., Brownsville — 16.I.42, on Malvastrum coromandelianum (1♀ USNM). 15.VI.68, Board & Hefernik (1♀ TAMU); 11.VI.71, W.E. Clark (1♀ TAMU); 5.IX.06, par. A. grandis (1♂ USNM); 21.I.16, R.A. Vickery, reared from Mantis eggs (1♀ USNM); 18.X.09, par. Bruchus exigua bred from Amorpha fructiosa pods (1♀ USNM); IV.21, J.C. Bridwell, bred from Bruchus nr limbatus from Texas ebony (12♀, 15♂ USNM); 2.VII.27, Bridwell, ex Bruchus sallei in huisache (1♀, 1♂ USNM). Sabal Palm Grove, 20-22.X.89, R. Anderson & E. Riley (1♀). Chambers Co., N. shore Lake Charlotte, 10.IX.16, C. Heinrich, in Cephelanthus occidentalis (1♀ USNM). Collin Co., Plano, 12.XII.08, E.S. Tucker, bred from mistletoe I.09 (1♀ USNM). Colorado Co., 14.VIII.07, par. A. grandis (1♀ USNM). Dallas Co., Dallas — 29.I.03, 12.IX.06, 27.VII.07, par. A. grandis (2♀, 1♂ USNM); 12.IX.07, par. Bruchus exiguua (14♀, 26♂ USNM; ♂ CNCI LB-specm 2010-008). DeWitt Co., Cuero — 12, 24.VIII.07, par. A. grandis (2♀, 2♂ USNM); 12.VIII.07, par. Bracon mellitor (1♂ USNM). Fort Bend Co., Richmond, 15.VIII.07, par. A. grandis (2♂ USNM, CNCI LBspecm 2010-002). Goliad Co., Goliad, 3.IX.08, par. A. grandis (1♀ USNM). Grayson Co., Denison, 27.VIII.07, par. Bracon mellitor (1♂ USNM). Rock Creek, 5 mi. NW Gordonville, 22.VII.72, E.E. Grissell, on Ambrosia psilostachya DC (1♂ TAMU). Harrison Co., Marshall, 22.VIII.06, 10.VIII.07, par. A. grandis (8♀, 1♂ USNM). Hidalgo Co., Bentsen-Rio Grande [Valley] State Park, River hiking trail, 19.VI.86, J.B. Woolley (1♀). McAllen, 1921, J.C. Bridwell, bred from Megacerus piger (1♀, 1♂ USNM). Mercedes Hoblitzelle Farm, 21.V.83, C. Melton (1♂). Santa Ana National Wildlife Refuge, 150', 3.VII.82, G. Gibson (1♀). Hood Co., Granbury, 7, 17.VIII.07, par. A. grandis (2♂ USNM). Kaufman Co., Terrell, 26.VIII.07, par. A. grandis (2♀, 1♂ USNM). Kleberg Co., Kingsville, V.23, M.M. High, ex beans huisache (3♀ USNM). Lavaca Co., Hallettsville, 9, 30, 31.VIII.06, 13, 17, 18.VIII.07, par. A. grandis (9♀, 3♂ USNM). McLean Co., Waco, 25.VII.06, 28, 29.VIII.06, 20.IX.06, 2.VIII.07, par. A. grandis (13♀, 4♂ USNM), 16.IX.65, T.L. Chestnut (1♀ MEMS). Medina Co., D’Hanis , 11.VII.65, C.W. O’Brian, ex pupa Merobruchus julianus (1♀ TAMU). Navarro Co., Corsicana — 21.VIII.08, par. A. grandis (1♀ USNM); III.04, C.M. Walker, par. A. grandis (1♀ USNM). Nacogdoches Co., 12 km SW Nacogdoches, SFA [Stephen F. Austin] Experimental Forest, Angelina National Forest, 47º31'N 94º47'W, 30.IV-16.V.2005, N. Schiif & C. Rudolph (1♀ UCDC). Nueces Co., 24.II.30 (1♀ TAMU). Presidio Co., Big Bend Ranch — SNA, 6mi. SE La Sauceda, 19.V.90, Wharton & Judd (1♀); 29º29'40"N 104º06'00"W, Agua Adentio Sprg, 3560', 20.VI.90, G. Zolnerowich (1♀). Big Bend Ranch State Natural Area, E.E. Grissell & R.F. Denno — Duck Pond 5 mi. W. Sauceda Ranch, 29º29'N 104º1'W, 4-5.IX.93, on Juglans sp (1♂ USNM); 1.7 mi. NE McGurks Tanks, 29º29'N 103º48'W, 4.IX.93, on Rhus virens (1♂ USNM). Presidio, 7.III.45, 14.VII.45, J.N. Russell, from tunnel in twig of Koeberlinia spinosa (2♀ USNM). Robertson Co., Calvert — 28.VIII.06, 30.IX.06, 21.VII.07, par. A. grandis (2♀, 1♂ USNM); 28.VIII.06, par. (accident) Catolaccus incertus (1♀ USNM). Easterly, 11 mi. N., Camp Cooley Ranch, 5.XII.80. J.C. Cuda, host: Trichobaris bridwelli Barber , Texas A&M voucher specimen #23 (1♂ TAMU). Rusk Co., Overton, 23.VIII.06, par. A. grandis (1♀ USNM). Starr Co., 30.VII.28, R.H. Beamer (1♀ USNM). Tarrant Co., Arlington — 6, 17, 25, 28.VIII.07, 2.IX.07, par. A. grandis (12♀, 8♂ USNM); 6, 28.VIII.07, par. Bracon mellitor (1♀, 2♂ USNM); 6.VIII.07, par. Catolaccus incertus (1♀ USNM); 6.VIII.08, F.C. Bishopp, par. A. grandis (1♀ USNM). Travis Co., vicinity Long Hollow Creek, 30º27'43"N 97º52'19"W, 23.VI.93, Alexander, Quinn, Riley, Wharton et al., on Juniperus ashei (1♀ TAMU). Trinity Co., Trinity, 9, 30.VIII.06, par. A. grandis (4♀, 2♂ USNM). Victoria Co., Victoria — 14.3, E.A. Schwarz (1♀ USNM); 1.IX.06, 10.VIII.07, par. A. grandis (4♀ syntypes E. cushmani : 1 BMNH, 3 USNM); 25.VII.03, 1.IX.06, 5, 10, 11.VIII.07, par. A. grandis (17♀, 7♂ USNM); 22.IV.07, 24.VI.07, 10, 22.VII.07, 10.VIII.07 par. Bruchus prosopis (14♀, 2♂); 13.IX.07, par. Bracon mellitor (1♀, 1♂ USNM); 12.V.07, bred Melia azed . berry (1♀, 1♂ USNM); 10.VIII.07, par. Anthonomus quadrigibbus (1♂ USNM); 13.III.08, J.D. Mitchell (3♀ USNM); 13.III.08, J.D. Mitchell, bred Iva ciliata stem (1♂ USNM); 14.III.08, J.D. Mitchell, bred Xanthium stem (1♀ USNM). Williamson Co., Taylor, 22.X.33, J.E. Gillaspy (1♀ TAMU). Wood Co., Mineola , 20.VIII.07, par. A. grandis (1♀ USNM). Zapata Co., Falcon State Park, 250', 2.VII.82, G. Gibson (1♀). UTAH: Washington Co., Pintura, 11.VIII.29, R.H. Beamer (1♀ USNM). VIRGINIA: 29.VII.63, D.M. Harmon, Pissodes strobi (1♀ USNM). Arlington Co., Rosslyn, No. 1175.2 (1♀ USNM). Augusta Co., Deerfield , 6, 15, 16.VII.63, H.M. Kulman, ex Pissodes strobi on white pine (3♀). Clarke Co., Blandy Experimental Farm, 2 mi. S. Boyce, 15- 30.IV.90, D.R. Smith (1♂). Cumberland Co., Cumberland State Park, 26.VI.64, O. Peck (1♀). Essex Co., 1 mi. SE Dunnsville, D.R. Smith — 37º52'N 76º48'W, 23.IV-6.V.96 (1♀ USNM); 27.III-11.IV.91, 12-24.IV.91, 30.IV- 13.V.91, 14-24.V.91, 6-14.VI.91, 17.IX-10.X.91, 12-29.IV.92, 17-29.IV.92, 30.IV-13.V.91 (CNC Photo 2010-64), 13-29.V.92, 12-24.VI.92, 30.V-11.VI.92, 17.VII-2.VIII.96 (15♀, 9♂). Fairfax Co., near Annandale, D.R. Smith — 21-27.VIII.88, 9-15.VIII.89 (2♀); 6.VIII.83,19-25.VI.88 (2♀ USNM). Falls Church, reared 14.VIII.15, Heinrich, on Trifolium (1♂ USNM).Vienna, L.J. Bottimer — XII.26 (1♂), 9.XII.26 (1♀); 5.I.27 (1♀ USNM). Louisa Co. , 4 mi. S. Cuckoo, J. Kloke & D.R. Smith — 6.VI.86, 16-31.VII.87, 21.VIII-2.IX.87, 21.IX-13.X.87, 1-12.VII.88, 12- 23.VIII.88 (7♀); 25.V.85, 7, 14, 21.VII.85, 12-25.IV.88 (3♀, 2♂ USNM). Montgomery Co., 22.IX.64, 2.IV.65, K.R. Lewis, host R. frustrana (1♀, 1♂ TAMU). Prince William Co., Manassas, 15.X.86, J. Cronin, hyperparasitoid Thyridopteryx ephemeraeformis (1♀ USNM). Pulaski Co., 16.VII.64, 16, 21, 22.IX.64, K.R. Lewis, host R. frustrana (4♀, 3♂ TAMU). WEST VIRGINIA: A.D. Hopkins (1♂ USNM). Hardy Co., 3 mi. NE Mathias, 38º55'N 78º49'W, D.R. Smith — 22.VIII-13.IX.2002 (1♀, CNC Photo 2010-46); 2400', 14.VIII-4.IX.95 (1♀). Jefferson Co., VI-VII.31, E. Gould, ex Coleophora malivorella (11♀, 3♂ USNM). Ritchie Co., Cairo, reared 28.V.13, A.D. Hopkins, Hopk. U.S. 9423b5, Hicoria (1♂ USNM). WISCONSIN: Fond du Lac Co., T18N, R19E, S23, 19- 26.VII.17, Gypsy Moth M.T. (1♀ IRCW). Grant Co., T 6N, R6W, S17, 26.VII-9.VIII.76, 19-26.VII.76, Gypsy Moth M.T. (3♀ IRCW). Green Co., T4N, R6E, S5, 29.VI.76, Gypsy moth proj. #760030A, em. 26.VII.76, Juglans nigra (1♀ IRCW).

Distribution. Widely distributed in southern USA and eastern North America (Map 7). In addition, E. cushmani has been recorded from throughout Central and South America as well as the Caribbean islands ( Noyes 2010). It was also introduced intentionally into Hawaii from Guatemala as a biocontrol agent of the pepper weevil, Anthonomus eugenii Cano ( Coleoptera : Curculionidae ) ( Swezey et al. 1939, Bess and Haramoto 1959, Duan et al. 1996).

Biology. Noyes (2010) listed E. cushmani as a primary parasitoid or, through Hymenoptera and Diptera , as a hyperparasitoid of 34 species in 14 families of 5 insect orders. However, the lists of hosts given for E. cushmani and E. cyaniceps are both partly inaccurate because Noyes (2010) included E. cyaniceps amicus as a synonym of E. cyaniceps and therefore hosts recorded in the original literature under E. cyaniceps amicus or E. amicus were listed for E. cyaniceps rather than E. cushmani . Most of the host records in Noyes (2010) were validated based on reference to such catalogs as Peck (1963) and Burks (1979), whose information came either from the original literature or label data of specimens in the USNM. Both of these authors listed E. amicus and its hosts separate from E. cyaniceps . I have seen specimens of E. cushmani labelled as from the following arthropod hosts and plant associates. Specimens indicated by an exclamation mark (!) are confirmed hosts for E. cushmani that were listed as hosts for E. cyaniceps in Noyes (2010).

Map 7. Regional distribution of E. cushmani .

Arthropod hosts — COLEOPTERA . Apionidae :! Apion rostrum Say ( Pierce 1908b) . Bruchidae :! Acanthoscelides alboscutellatus (Horn) ( Bissell 1940, Ott 1991),! Acanthoscelides floridae (Horn) ( Brett 1946) , * Acanthoscelides obsoletus (Say) ,! Acanthoscelides obtectus (Say) ( Chittenden 1898) , * Acanthoscelides perforatus (Horn) , * Algarobius prosopis (Leconte) , * Acanthoscelides submuticus (Sharp) (= Bruchus exiguus Horn ) ( Cushman 1911),! Bruchus brachialis Fahraeus ( Bridwell 1933, Pinckney 1937),! Bruchus pisorum (L.) ( Bissell 1938), * Caryobruchus sp. , * Megacerus ? impiger (Horn) (= M. piger ?), Merobruchus julianus (Horn) ( Forister and Johnson 1971) ,! Mimosestes amicus (Horn) ( Girault 1916b) , Mimosestes sallaei (Sharp) ( Pierce 1908b, Cushman 1911),! Amblymerus robiniae (Fabricius) ( Pierce 1908b, Cushman 1911), * Stator nr limbatus (Horn), * Stator pruininus (Horn) . Chrysomelidae : * Neochlamisus bebbianae (Brown) (= Chlamys gibbosa (Fabricius)) . Curculionidae :! Anthonomus albopilosus Dietz ( Pierce 1907, 1908a),! Anthonomus grandis Boheman ( Crawford 1907, Hunter and Hooker 1907, Mitchell and Pierce 1911, Pierce et al. 1912, Fenton and Dunnam 1929, Chestnut and Cross 1971, Cross and Chestnut 1971, Mitchell et al. 1974), * Anthonomus grandis thurberiae Pierce , * Anthonomus quadrigibbus Say , * Pissodes strobi (Peck) ,! Smicronyx (= Desmoris ) fulvus LeConte ( Bigger 1933) ,! Tyloderma foveolatum (Say) ( Pierce 1908a) , * Trichobaris bridwelli Barber. HYMENOPTERA . Braconidae :! Bracon mellitor Say ( Pierce 1908a, McGovern et al. 1974),! Macrocentrus sp. ( McConnell 1934) . Cynipidae : * Dryocosmus kuriphilus Yasumatsu. Diprionidae :! Diprion similis (Hartig) ( Finlayson 1962) . Pteromalidae :! Lyrcus incertus (Ashmead) ( Pierce 1908a) . LEPIDOPTERA . Coleophoridae :! Coleophora malivorella Riley ( Gould and Geissler 1940, Beacher 1947). Lasiocampidae : * Malacasoma americana (Fabricius) . Momphidae :! Mompha eloisella (Clemens) (Allan 1962) . Pyralidae : Acrobasis (= Mineola ) sp., * Acrobasis betulella Hulst (= A. hebescella Hulst ),! Acrobasis caryae Grote ( Nickels et al. 1950) ,! Acrobasis caryivorella Ragonot ( Nickels 1951) , * Acrobasis juglandis (LeBaron) ,! Acrobasis nuxvorella Neuzig ( Gunasena and Harris 1988) . Psychidae :! Thyridopteryx ephemeraeformis (Haworth) ( Baerg 1928) . Sphingidae : * Ceratomia catalpae (Boisduval) . Tortricidae :! Cydia caryana (Fitch) ( Reyes-Villanueva 1987) ,! Grapholita molesta (Busck) ( McConnell 1932, Brunson and Allen 1948),! Rhyanconia frustrana (Scudder) ( Schaffner 1959) . MANTOIDEA. Mantidae : * Mantis eggs.

In addition to the above confirmed host records, the following hosts listed by Noyes (2010) for E. cyaniceps likely also refer to E. cushmani based on the original identification of the parasitoid as E. cyaniceps amicus or E. amicus — COLEOPTERA . Curculionidae : Chalcodermus aeneus Boheman ( Bissell 1940) , Chalcodermus inaequicollis Horn ( Bissell 1940) . Scolytidae : Scolytus multistriatus (Marsham) ( Hoffmann 1942) . HYMENOPTERA : Bethylidae : Goniosus punctaticeps (Kieffer) ( Nickels et al. 1950). Braconidae : Agathis acrobasidis (Cushman) ( Nickels et al. 1950) , Bracon variabilis (Provancher) ( Nickels et al. 1950) , Macrocentrus ancylivorus Rohwer (Brunson 1948) , Macrocentrus instabilis Muesebeck ( Nickels et al. 1950) . Ichneumonidae : Calliephialtes grapholithae (Cresson) ( Nickels et al. 1950) , Glypta sp. ( Driggers 1929) . LEPIDOPTERA . Pyralidae : Dioryctria disclusa Heinrich ( Farrier and Tauber 1953) , Pyrausta (= Ulophora ) grotei Munroe ( Bissell 1940). Tortricidae : Epiblema stenuana (Walker) ( Haden 1935) .

Based on examined specimens it is also likely that at least two hosts recorded for E. cushmani actually belong to E. cyaniceps — COLEOPTERA . Curculionidae : Microlarinus lypriformis (Wollaston) and Trichobaris texana LeConte (see under E. cyaniceps ).

Host records suggest that E. cushmani is usually a primary parasitoid or, through at least Braconidae , Ichneumonidae and rarely Pteromalidae , a hyperparasitoid of various Coleoptera and Lepidoptera . It appears to commonly be a hyperparasitoid and very possibly is an obligate hyperparasitoid of at least the cotton boll weevil, Anthonomus grandis , based on McGovern et al. (1974). Although McGovern et al. (1974) identified the hyperparasitoid as E. cyaniceps , examination of voucher specimens shows that this was a misidentification of E. cushmani , as appears to be all other literature recording E. cyaniceps from the cotton boll weevil. Similarly, all literature recording E. cyaniceps from various Bruchidae appear to be misidentifications of E. cushmani . Anomalous regional host records include Mantis eggs and Dryocosmus kuriphilus .

I can not confirm the extralimital host records for Diptera listed by Noyes (2010), though E. cushmani was stated to have quickly spread to other coleopteran hosts on all the major Hawaiian islands after being introduced ( Stein 1983) as well as to Procecidochares utilis Stone and P. alani Steyskal (Tephritidae) ( Duan et al. 1996). Both of these gall making species were introduced into Hawaii, the former species as a weed biocontrol agent. Eupelmus cushmani has also been reported from South America as a hyperparasitoid through Paratheresia claripalpis (Wulp) (Tachinidae) ( De Santis 1979) and Sarcodexia lambens (Wiedemann) (= S. sternodontis Townsend ) ( Parker et al. 1953).

Plant associates — Asteraceae : Helianthus angustifolius L. (swamp sunflower), Iva annua L. (marshelder), Vernonia altissima Nuttal (ironweed), Xanthium L. (cocklebur),? Silphium laciniatum L./ S. terebinthinaceum L. [based on reared male]. Euphorbiaceae : Crotoncapitatus var. lindheimeri (Engelmann & Gray) (hogwort). Fabaceae : Acacia constricta Bentham (whitehorn acacia), Amorpha californica Nuttall (California false indigo), Amorpha fructiosa L. (desert false indigo), Baptisia tinctoria (L.) (yellow false indigo), Calliandra eriophylla Bentham (fairy duster), Cercis canadensis L. (eastern redbud),? Mimosa biuncifera Bentham [based on reared males] (catclaw mimosa),? Niasolia schottii (Torrey) [based on reared male] (Schott’s yellowhead), Parkinsonia microphylla (Torrey) (yellow palo verde), Pithecolobium flexicaule (BenthAM) (Texas ebony), Prosopis pubescens Bentham (screwbean mesquite), Rhynchosia minima (L.) seeds (least rhychosia), Trifolium pratense L. (red clover), Vachellia farnesiana (L.) Fabaceae (huisache), Vicia villosa Roth (hairy vetch). Juglandaceae : Hicoria sp. , Juglans nigra L. (eastern black walnut). Koeberiniaceae: Koeberlinia spinosa Zuccarini (crown of thorns). Malvaceae : Gossypium thurberi Todaro (Thurber’s cotton). Meliaceae : Melia azedarach L. (Persian lilac). Fabaceae . Onagraceae : Ludwigia L. (primrose-willow), Oenothera biennis L. (common evening primrose). Pinaceae : Pinus strobus L. (white pine), Pinus taeda L. (loblolly pine). Rhamnaceae :? Zisiphus obtusifolia (Hooker ex Torrey & A. Gray) (= Condalia lycioides ) [based on reared male] (lotebush). Rubiaceae : Cephelanthus occidentalis L. (buttonbush). Santalales :? Phoradendron leucarpum Reveal (mistletoe). Smilacaceae : Smilax L.

Remarks. Through crossing experiments Pierce and Cushman (1915) showed that E. cushmani and E. cyaniceps are reproductively isolated. Furthermore, E. pervius is supported as a distinct species from both E. cushmani and E. cyaniceps by host differences and a difference in male flagellar structure/setation (cf. Figs 62–64, 66). However, males of E. cushmani and E. cyaniceps are very similar except for what appears to be a slight difference in sculpture pattern of the scape. Typical E. cyaniceps males have an elongate region of quite distinct micropunctures ventrally along the entire length of the scapular scrobe ( Fig. 77), whereas males of E. cushmani have this region more extensively smooth and shiny or at least with much shallower sculpture that appears more as tiny pin pricks and/or somewhat strigose ( Figs 75, 76) rather than being pitted by distinct punctures. However, there is no distinct division between the two extremes and it is very possible my identification is unreliable for some males not definitely associated with females through rearing. For example, females collected from Menifee Valley (California, Riverside Co.) are identified as E. cyaniceps , whereas males from the same locality and similar dates are tentatively identified as E. cushmani .

I am equally uncertain of the morphological limits and differentiation of all E. cushmani and E. cyaniceps females. Although ‘typical’ females of E. cushmani and E. cyaniceps are morphologically quite distinct, all structural, color and sculptural features by which they are recognized appear to intergrade. I primarily use setal pattern of the prepectus in combination with relative length of the ovipositor sheaths to differentiate females. The prepectus of females that I include in E. cushmani is often virtually bare or only quite obscurely setose, but at most there are only 2 or 3 rows of comparatively inconspicuous setae mediolongitudinally such that the apices of the setae do not attain the margins and there is quite a wide bare band ventrally (Fig. 42). Most females, or at least larger females of E. cyaniceps , have the prepectus much more conspicuously and extensively setose (Fig. 44), usually with at least four or five rows of white setae and with the ventral-most row of setae near the ventral margin so that there is only a narrow bare band ventrally and/or the apices of the setae generally attain the ventral and/or dorsal margins of the prepectus. Females of E. cyaniceps usually also have the propodeal callus much more densely and conspicuously setose (cf. Fig. 51) than do E. cushmani females (Fig. 50). Because the setae are less numerous they are quite easy to count for E. cushmani females, but not so for typical E. cyaniceps females with a more densely setose callus. However, there is no distinct division between the two states and propodeal setation, although often diagnostic, is an even less reliable identifier of E. cushmani females than is prepectal setation.

It is sometimes difficult to assess the setal pattern of the prepectus in museum specimens because the setae are abraded, but regardless I believe this feature is insufficient to differentiate all females of E. cushmani and E. cyaniceps , at least in eastern North America . Almost all females that I include in E. cushmani based on quite a sparsely setose prepectus have ovipositor sheaths that are at least as long as the marginal vein ( Fig. 37), whereas in eastern North America females that I identify as E. cyaniceps based on an obviously, extensively setose prepectus have ovipositor sheaths that are at least slightly shorter than (usually less than 0.9×) the length of the marginal vein ( Fig. 39). Figure 38 is of a female from Texas that is about 3.2 mm in length, which has short ovipositor sheaths only about 0.88× the length of the marginal vein, but a comparatively inconspicuously setose prepectus with only about 3 rows of seta similar to some E. cushmani females. In other respects it differs little from other eastern North American females of E. cyaniceps having a more conspicuously setose prepectus, including having a uniformly brown scape, the frons somewhat meshlike coriaceous-imbricate (also almost entirely green except narrowly bluish-purple along inner orbits), the vertex broadly rounded into the occiput (see further below), the protibia extensively brown, and the metatibia mostly brown except for up to about the apical quarter white (cf. Figs 38, 39). I consider that smaller females of E. cyaniceps can sometimes have the prepectus less extensively and conspicuously setose than is typical for larger specimens and include in E. cyaniceps females from eastern North America with ovipositor sheaths less than 0.9× the length of the marginal vein in combination with a comparatively inconspicuously, sparsely setose prepectus.

Large females of E. cushmani tend to have longer ovipositor sheaths and the longest sheaths are usually correlated with the inner plate of the ovipositor projecting conspicuously beyond the apex of the gaster ( Fig. 9). For example, the inner plate of the ovipositor projects obviously beyond the apex of the gaster in the syntypic series of E. amicus whose ovipositor sheaths are about 1.53−1.6× the length of the marginal vein, but not in the syntypes of E. cushmani whose ovipositor sheaths are only about 1.08−1.1× the length of the marginal vein. Although females of both are quite large, about 4.3 mm in length, ovipositor structure apparently is not entirely correlated with body size because the female from Camp Angeles (California) associated with Amorpha california Nuttal (Fabaceae) is only 3 mm in length, yet the ovipositor sheaths are about 1.3× the length of the marginal vein and the inner plate of the ovipositor extends conspicuously beyond the apex of the gaster such that the apparent sheath length is about 1.6× the length of marginal vein.

In addition to typically different prepectal and propodeal setal patterns and relatively longer ovipositor sheaths, various other features help to distinguish females of E. cushmani from those of E. cyaniceps , though any single feature appears to form a continuum or overlaps in the two species. Females of E. cushmani are often distinguished by an entirely or almost entirely yellowish-orange scape ( Fig. 28), whereas all except a single female of E. cyaniceps I examined (see under E. cyaniceps ) have a uniformly and comparatively dark brown to somewhat orangey-brown scape that has at least a slight metallic green luster under some angles of light. However, scape color is highly variable. Females of E. cushmani from more southern localities commonly have at least a partly yellowish scape, though smaller females almost always have a dark scape similar to E. cyaniceps females and scape color does not seem to be completely correlated with either body size or geographical locality. For example, type specimens of E. townsendi and E. cushmani from Peru and Texas, respectively, have a yellowish-orange scape whereas the syntypes of E. amicus from New Mexico have quite a dark, only slightly orangey-brown scape. Larger females of E. cushmani usually also have the frontovertex dark mesally in distinct contrast with bright green or often variably distinctly multicolored metallic bands ( Fig. 28) along the inner orbits, though the frons can be more uniformly colored without or with less conspicuous lateral bands, particularly in smaller females. Larger females typically also have the frons coriaceous to coriaceous-imbricate mesally and variously extensively reticulate-imbricate to roughened or transversely wrinkled with scattered setiferous punctures laterally toward the inner orbits ( Fig. 28), though smaller females often have the frons more uniformly meshlike coriaceous. Larger, typical females of E. cushmani usually also have the vertex transversely reticulate-strigose to quite strongly strigose-carinate and in lateral view the vertex continued beyond the ocelli in a similar plane as the frons so that the highest point of the head is above the posterior ocelli or at least the vertex is quite abruptly and narrowly angled relative to the occiput ( Fig. 37). Smaller females have the vertex variably sculptured but at least broadly rounded into the occiput such that in lateral view the posterior ocelli are dorsal on the head. Typical females of E. cyaniceps have a more uniformly colored frontovertex, often without or at least with less abruptly and distinctly delineated bluish to purple regions along the inner orbits ( Fig. 29). Some females have a similar frontovertex color pattern to typical E. cushmani females, but unlike many such females the lateral metallic region is a single color. The frons is often also more extensively and distinctly reticulate in E. cyaniceps females ( Fig. 29), though sometimes more or less uniformly meshlike coriaceous to slightly coriaceous-reticulate in smaller specimens similar to smaller specimens of E. cushmani . The vertex is always broadly rounded into the occiput in E. cyaniceps females ( Fig. 39) and usually is more transversely reticulate-imbricate than distinctly strigose, though again this is similar to some smaller E. cushmani females.

Correct identification of E. cushmani from western North America is also complicated by some females, particularly those reared from Acacia seeds, having an elongate linea calva that is open or virtually so to the vanal area similar to females of E. pervius (cf. Fig. 45). However, these E. cushmani females have typical sparse setal patterns of the prepectus (Fig. 42) and propodeal callus (Fig. 50) compared to the much more extensively setose prepectus (cf. Fig. 44) and propodeal callus (Fig. 51) of E. pervius females, in addition to often having a yellowish-orange scape. It is likely such E. cushmani specimens are parasitoids of Bruchidae in the Acacia seeds whereas E. pervius are parasitoids of Prodoxus moths in Yucca . The similarity in forewing setal pattern either indicates introgression or that what I presently interpret as just three species, E. cushmani , E. cyaniceps and E. pervius , represents a species complex of more than three morphologically similar species. Female body size varies considerably within each putative species. I currently interpret this variation as at least partly correlated with different females being reared from different sized hosts and/or being a primary or a hyperparasitoid. However, observed variation could reflect unrecognized sibling species associated with different hosts. The collection of more material that is reliably associated with hosts through rearing and particularly the collection of morphologically diverse specimens from throughout North America that are suitable for molecular analysis are needed to test the species concepts proposed here to better assess diversity.

In addition to the similarities noted above, some females of E. cushmani with a dark scape and long ovipositor sheaths might be mistaken for E. utahensis females, but differ by several setal features as discussed under the latter species. Because of scape color and usually quite long ovipositor sheaths, females of E. cushmani are also similar to E. cerris Förster in Europe, but E. cerris females have the frons entirely finely coriaceous and a line of setae over most of the length of the costal cell dorsally, more similar to E. annulatus females, as well as the propodeal callus more densely setose.