A review of New World Eurytenes s. str. (Hymenoptera, Braconidae, Opiinae)

Th e New World species of Eurytenes Foerster sensu stricto (Hymenoptera: Braconidae, Opiinae) are revised, and a key to these species is presented. Four new species are described: Eurytenes (Eurytenes) dichromus sp. n. from Texas, E. (E.) microsomus sp. n. from Texas, E. (E.) pachycephalus sp. n. from Mexico, and E. (E.) ormenus sp. n. from Mexico. Eurytenes abnormis (Wesmael) is redescribed for comparison, and its host records are reviewed.

In addition to the type species, three other species are currently included in Eurytenes s. str.: E. orientalis Fischer, 1966, E. cratospilum Chen & Weng, 2005, and E. basinervis Wu & Chen, 2006.Eurytenes abnormis is Holarctic (Fischer 1972(Fischer , 1977)), E. orientalis is from the Philippines (Fischer 1966), and the other two species are from China (Chen and Weng 2005;Wu and Chen 2006).Th e purpose of this paper is to expand the known distribution of Eurytenes s. str.by describing new species from central Texas and central Mexico.We also discuss morphological features useful for discriminating both New and Old World species.

Materials and methods
Nearly all of the material used in this revision is from the Texas A&M University Insect Collection (TAMU).Material for comparison was obtained from or examined at the following institutions: American Entomological Institute (AEI), Gainesville, FL, USA; Canadian National Collection, Ottawa, Canada; Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium; Naturhistorisches Museum, Vienna, Austria (NHMW); U. S. National Museum of Natural History, Washington, D. C. (USNM).Th e newly described species were compared with several Old World specimens, including E. abnormis from eight European localities, the holotype of E. orientalis, two undetermined specimens from Taiwan, and one undetermined specimen from the Kurile Islands.
Descriptive terminology largely follows Sharkey and Wharton (1997), with modifi cations as in Wharton (2006).For clarity, the fi rst metasomal tergite (T1) is referred to as the petiole with the following median tergites referred to as T2, T3, etc. Th e mesoscutum consists of an anterior declivity and the more readily visible, relatively fl at portion posteriorly referred to in the descriptions as the mesoscutal disc.Measurements were taken using a reticle on a Zeiss Stemi, DRC microscope and converted to ratios or millimeters.Flagellomere width was measured at the narrowest point.Clypeus was measured as in Fig. 1 and the width compared to the distance from the inner margins of anterior tentorial pits to the eye.Face width was measured as the shortest distance between eyes and compared to face height from epistomal sulcus at top of clypeus to the lower margin of the antennal socket.Angle of precoxal sulcus was estimated with head-mesosoma and mesosoma-petiole attachments aligned horizontally.Hind tibia was measured from attachment point of femur to attachment point of tarsus vs maximum width at apex.Petiole (=T1) length was measured laterally from the base of the dorsope to attachment point of T2 (Fig. 2).Total ovipositor length, measured in lateral view, was estimated using specimens whose hypopygium was extended, thus exposing the majority of the ovipositor.Mesosomal length was measured as in Fig. 3. Measurements are presented as ranges followed by the mean (m).Th e term butterscotch is used to describe the tawny, yellowish-brown color of many of the body parts.
In the material examined sections, label data are presented in a uniform format for specimens other than the holotypes of new species.For holotypes of newly described species, data are recorded exactly as given on specimen labels, with square brackets for additional data not on the labels.
Images were acquired digitally using Syncroscopy's Auto-Montage Pro 5.01.0005 (Copyright Synoptics Ltd.) and PictureFrame (TM) Application 2.3 in combination with a ProgRes 3008 digital camera mounted on a Leica MZ APO dissecting microscope.All images were further processed using Adobe Photoshop® CS5.Images are stored in mx, a web-based content management system that facilitates data management and dissemination for taxonomic and phylogenetic works (e. g.Yoder et al. 2006).Th e mx project is open source, with code and further documentation available at http://sourceforge.net/projects/mx-database/.Mesosoma.Pronotum dorsally with narrow, transverse, crenulate sulcus extending continuously along lateral pronotum to ventral corner; weak to deep median pit interrupting sulcus dorsally.Mesoscutum in profi le with anterior declivity very slightly concave, nearly vertical; notaulus present on anterior portion of mesoscutal disc, angled laterally at anterior end, laterally-directed portion carinate along anterior margin; notaulus continuous with weakly to distinctly crenulate impression bordering lateral mesoscutal margin, the impression extending posteriorly at least to level of tegula; midpit of mesoscutum well-developed, discrete.Scutellar sulcus (Fig. 31) narrow, though not exceptionally so, 3-4 times wider than mid-length, crenulate, with numerous closely-spaced ridges.Mesopleuron smooth, shiny, posterior margin not crenulate; precoxal sulcus distinctly impressed, crenulate.Propodeum with large median areola, variously obscured by sculpture.
Wings.Slightly more than twice as long as wide.Stigma long, very narrow, nearly parallel-sided basally, widening distally; thickest part of stigma twice maximum width of proximal half.Radial cross-vein (r) thickened, weakly to strongly bowed anteriorly, arising from extreme base of stigma, nearly in line with 3RSa; RS+M weakly to distinctly sinuate; second submarginal cell nearly parallel-sided, not or only very weakly converging distally; m-cu nearly always postfurcal, entering second submarginal cell; 2CUa distinct, shorter than 2cu-a.Hind wing with both RS and M distinct nearly to wing margin, usually nebulous: very weakly pigmented; m-cu varying from indistinct in smaller individuals to present in larger individuals as a weakly pigmented impression extending more than half way to wing margin.
Hosts.Agromyzidae.See comments under E. abnormis, the only species with host records.
Comments.Fischer (1972) provides the most recent detailed description of Eurytenes s. str.(in German); Wharton (1988Wharton ( , 2006)), Fischer (1998), and Wu and Chen (2006) provide diagnoses for Eurytenes s. str.and s. l.Wu and Chen (2006) were the fi rst to use morphological features other than color for discriminating between species of Eurytenes s. str.Diagnosis.Eurytenes abnormis is most readily recognized by the pale coloration of the petiole and metasoma and is further distinguished from the four North American species described below by the narrower, more ventrally concave clypeus (Fig. 10).Th e petiole is narrower than in E. dichromus, sp.n. and E. microsomus, sp.n. and is thus more similar in shape to the darker Mexican species described below.
Mesosoma.Posterior-ventral margin of lateral pronotum crenulate for most of length.Precoxal sulcus parallel-sided, narrowly crenulate along most of length, usually weakly impressed anteriorly, often extending very close to anterior margin of mesopleuron; precoxal sulcus inclined at a 35 degree angle.Notaulus distinctly impressed over anterior third of mesoscutal disc, crenulate over anterior 0.2-0.3;with cluster of short setae at rugulose base of anterior declivity; with widely spaced line of 3-5 longer setae extending posteriorly towards but not usually reaching cluster of scattered setae around midpit.Propodeum with median carina present anteriorly, bifurcating near middle to form fi vesided median areola over posterior 0.6, surface rugose laterally and posterior-medially, partly obscuring areola, but posterior-lateral fi elds largely smooth, as in Fig. 11.Wings.Fore wing r-m tubular and pigmented only at extreme anterior end, otherwise unpigmented, with lateral boundaries often only weakly indicated; (RS+M) b absent, m-cu entering extreme base of second submarginal cell; 3M very weakly pigmented basally in available material, spectral over most of length.Hind wing m-cu varying from indistinct in smaller individuals to present as a spectral impression extending more than half way to wing margin in larger individuals.
Color.Head and mesosoma dark reddish-brown to black.Scape, pedicel and fi rst fl agellomere yellow, antenna quickly darkening distally to dark brown; palps and tegula pale yellow; mandible and petiole tawny (darker yellow).Metasoma posteriad petiole usually bright yellow, sometimes with faint slight butterscotch banding to lighter brown banding.Hind femur and tibia darkening distally, femur transitioning from yellow to dark yellow or yellow-brown, tibia mostly infuscated, tarsi infuscated; legs otherwise yellow.Ovipositor sheath dark brown to black; ovipositor light brown throughout.Wings hyaline.
Host Records.Fischer (1959) listed 12 species of Agromyzidae, one Anthomyiidae, and one microlepidopteran as hosts but with no records of host plants.In this publication Fischer also noted that the anthomyiid and especially the microlepidopteran host (Coleophora nigricella Rondani) need verifi cation.Fischer (1964) added four more dipterans to the list and later (Fischer 1969a, b) provided additional records, including host plant information for nearly all of the known hosts.Nomenclatural updates for agromyzids and plant hosts from Fischer (1972) and Yu et al. (2005) are incorporated in the list of confi rmed hosts given below, with additional updates from Ellis (2007).Host plants for these 23 agromyzid hosts are split unevenly between monocots (8 fl y species) and dicots (15 fl y species).Four of the host fl y species were reared from Asteraceae and four from Poaceae, whereas Lamiaceae, Ranunculaceae, and Cyperaceae each harbored three host fl y species.
Th e agromyzid host records found in Fischer (1964Fischer ( , 1969a, b) , b) have a relatively high degree of confi dence because these records pertain to rearings by Buhr, Groschke, and Nowakowski, respectively.Fischer identifi ed the Eurytenes reared from these hosts (specimens in NHMW) and the hosts and host plants correspond well with information in Ellis (2007).Earlier literature, and several compilations based on the earlier primary sources, are problematic, however, because of the potential for misidentifi cation of the wasp and/or host fl y, as well as the absence of voucher specimens.We follow Fischer (1959) and treat the published host records for Pegomya bicolor (Wiedemann) (Diptera: Anthomyiidae) and Amauromyza verbasci (Bouché) dating to Bouché (1834), Ratzeburg (1848), andRondani (1872) as almost certainly erroneous and likely based on misidentifi cation of the other opiines that routinely attack these hosts or possibly on misidentifi cation of the host.Records of non-dipteran hosts are clearly erroneous since members of the Opiinae are all parasitoids of cyclorrhaphous Diptera.
Host: Plant.Agromyza albitarsis Meigen: host plant for E. abnormis has not been recorded previously but since this fl y is known to attack several trees in the family Salicaceae, the record may need to be verifi ed; Agromyza woerzi Groschke: Knautia arvensis (L.) Coult, Caprifoliaceae; Amauromyza labiatarum (Hendel): Galeopsis tetrahit L., Lamiaceae; Amauromyza lamii (Kaltenbach): Lamiastrum galeobdolon (L.Distribution.Previously recorded from throughout most of Europe (specifi cally Austria, Belgium, Bulgaria, Croatia, England, Finland, Germany, Hungary, Ireland, Italy, Lithuania, Poland, western Russia as far as the Urals, and Ukraine).Also recorded from eastern Palaearctic (Korea and Sakhalin Island), central to eastern Canada (Ontario, Saskatchewan) and USA (Minnesota, Missouri, North Dakota, South Carolina).Specifi c references to individual records can be found in Yu et al. (2005) for the most part; the record from the Urals is from Tobias and Jakimavicius (1986).Fischer (1970) recorded E. abnormis from Montana; however, the specimen on which it is based was collected in Missouri (label information only indicated the state as Mo.).Th e records from Sakhalin (Tobias 1998) and Korea (Papp 1985) may need to be verifi ed in light of other species described from that general region.Th e specimens we have examined from Taiwan and the Kuril Islands diff er in wing venation, body coloration, and sculpture from typical E. abnormis.Papp (1985) also noted the darker coloration of the petiole of his Korean specimen.
Comments.Fischer (1972) treated Opius paradoxus Ratzeburg, 1848 as a nomen nudum, while Dalla Torre (1898) listed it with a query as a synonym under E. abnormis, undoubtedly following Marshall (1891).Ratzeburg (1848), in his treatment of Opius, separated abnormis from all other species on the basis of the wing venation features that we now use to defi ne Eurytenes s. str.Ratzeburg (1848) initially states that only a single species, abnormis, belongs to the section of Opius with the radius arising from the base of the stigma.In the following sentence, however, Ratzeburg introduces the name paradoxus, indicating that it also should be placed here.Th ough this can be interpreted to mean that Ratzeburg was treating paradoxus as a synonym of abnormis, nevertheless he also mentioned body coloration (dark) and clypeal characters (lack of opening between clypeus and mandibles) that diff er from typical abnormis.Ratzeburg referred to Bouché throughout when discussing paradoxus and abnormis, and at the end of his treatment gives information on a more typical pale specimen of abnormis reared by Bouché.Whether intentional or otherwise, it would appear that Ratzeburg (1848) did provide a valid description of paradoxus with two characters that could be used to diff erentiate it from abnormis.However, his text could just as easily be interpreted to mean that paradoxus is invalid since it was fi rst proposed as a synonym of abnormis.We prefer the latter interpretation.Wu and Chen (2006) were the fi rst to use morphological features other than color for discriminating between species of Eurytenes s. str.Th ey used propodeal sculpture and the extent of the precoxal sulcus to diff erentiate their newly described E. basinervis from E. orientalis.Previously, Fischer (1966Fischer ( , 1998) ) used only color diff erences to distinguish between E. orientalis and E. abnormis.We have noted diff erences among species in the shape of the clypeus, but the appearance of the ventral margin of the clypeus changes with angle of view, and the diff erences are subtle.All species appear to have a concave ventral margin if the ventral part of the head is strongly rotated anteriorly.When placed in the same plane of view, however, the clypeus of the New World species described here is more truncate ventrally than that of E. abnormis.
In addition to specimens listed in the material examined section above, two specimens from Poland and one from Germany (NHMW) were also briefl y examined; data for these specimens were previously recorded by Fischer (1969a, b).In our summary of references above, we have not included several papers that provide only distribution information.Th ese can be found in Yu et al. (2005).Wharton.Additional specimens, not paratypes (TAMU): 2 ♀, Florida, Alachua Co., Hague Dairy, 29 47.311'N, 82 24.880'W, 28.iii.2007, J. Sivinski;1 ♀, same data except 29 47.328'N, 82 24.969'W, 29.iii.2007;1 ♀, Texas, Anderson Co., 10 mi SW Elkhart, 5-6.vi.1976, H. R. Burke.Diagnosis.Th is species is most readily recognized by its broader, bicolored petiole and bicolored, ventrally truncate clypeus.Based on the shape and color pattern of the petiole, as well as the shape of the clypeus, E. dichromus is most similar to E. microsomus, sp.n.Th e propodeum is nearly always more heavily sculptured in E. dichromus than in E. microsomus and usually slightly more rugulose posterior-laterally than in E. abnormis.
Mesosoma.Posterior-ventral margin of lateral pronotum crenulate for most of length.Precoxal sulcus narrowly crenulate anteriorly, sculptured area broadening posteriorly, usually weakly impressed anteriorly, often extending very close to anterior margin of mesopleuron; precoxal sulcus approximately 45 degrees, inclined more vertically than in E. abnormis.Notaulus distinctly impressed over anterior third of mesoscutal disc, crenulate over anterior 0.2-0.3;with dense cluster of short setae at rugulose base of anterior declivity extending ventrally to cover most of anterior declivity; with widely spaced line of 3-5 longer setae extending posteriorly towards but not usually reaching cluster of scattered setae around midpit as in Fig. 30.Propodeum with median carina present anteriorly, bifurcating near basal 0.3 to form fi ve-sided median areola over posterior 0.6-0.7,surface extensively rugose laterally and posterior-medially (Figs 32,33), partly obscuring areola, posterior-lateral fi elds often completely rugose.
Wings.Fore wing r-m pigmented basally, less commonly over anterior 0.5, otherwise unpigmented, largely tubular, with lateral boundaries usually distinct for most of length; m-cu usually postfurcal, entering base of second submarginal cell, less commonly interstitial; 3M distinctly pigmented, nearly tubular in basal third, gradually weakening and becoming depigmented distally.Hind wing m-cu usually poorly developed, varying from very weakly to distinctly impressed.
Color.Head and mesosoma black, with small red-brown spot adjacent eye dorsalmedially near ocelli, face at base of antennae also usually red-brown.Scape and pedicel yellow, fl agellomeres dark brown; mandible butterscotch with distal tip infuscated; clypeus infuscated, dark brown dorsally, butterscotch ventrally; palps and tegula yellow.Petiole dark brown dorsally, posterior fi fth and ventral-lateral region usually yellow.T2+3 butterscotch medially; T2 and T3 each with a medium brown lateral splotch; T4 and successive tergites each with dark brown transverse banding anteriorly fading to butterscotch posteriorly.Hind tibia pale yellow to whitish over about basal 0.15, remainder infuscated to medium brown, tarsus completely medium brown, legs otherwise yellow to nearly white, with femur and trochantellus often (though not in holotype) darker yellow than coxa and trochanter.Ovipositor sheath dark brown; ovipositor light brown.Wings hyaline.
Male and Host.Unknown.Distribution.Known only from central Texas.
Etymology.Th e name dichromus is derived from Greek: di, two; chromus, color.Th e name refers to the color of the clypeus.

Eurytenes microsomus
Diagnosis.Th is species is nearly identical to E. dichromus but E. dichromus is 1.25 × larger.Th e body is less heavily sculptured than in E. dichromus, there are fewer fl agellomeres, and T2+3 tends to be paler in coloration.
Mesosoma.Posterior-ventral margin of lateral pronotum weakly crenulate, nearly smooth for most of length.Precoxal sulcus parallel-sided, narrowly crenulate, short, weakly impressed anteriorly, not extending close to anterior margin of mesopleuron; precoxal sulcus at 45 degree angle, inclined more vertically than in E. abnormis.Notaulus distinctly impressed over anterior third of mesoscutal disc, crenulate over anterior 0.2-0.3;with moderately dense cluster of short setae at rugulose base of anterior declivity extending ventrally to cover much of anterior declivity; with widely spaced line of 3-4 longer setae extending posteriorly towards but not reaching cluster of scattered setae around midpit (Fig. 3).Propodeum with median carina extending over anterior 0.3 before bifurcating to form fi ve-sided areola over posterior 0.7.Surface smooth to weakly rugose laterally and posteriorly, carinae forming areola not obscured by sculpture, entirely visible, areola varying from smooth to weakly rugose (Figs 34,35).
Wings.Fore wing r-m at most pigmented at extreme base, largely tubular (with lateral boundaries distinct) over anterior half; m-cu distinctly postfurcal; 3M distinctly pigmented in basal third, gradually weakening and becoming depigmented distally.Hind wing m-cu indistinct.
Color.Head and mesosoma dark reddish-brown as in E. abnormis, but with pale spot adjacent eye similar to though weaker than the spot in E. dichromus.Scape and pedical butterscotch, fl agellomeres medium brown; clypeus butterscotch with slight infuscation dorsally.Palps, mandible, tegula, petiole, and ovipositor as in E. dichromus.Metasoma posteriad petiole patterned as in E. dichromus but T2+3 paler, whitish medially and T2 more lightly infuscate laterally.Legs about as in E. dichromus, with hind legs often a little paler.Ovipositor sheath dark red-brown.Wings hyaline.
Etymology.Th e name microsomus is derived from Greek: micro, small; somus, body.Th e name refers to the smaller size of this species compared to other species of Eurytenes.
Comments.Eurytenes microsomus and E. dichromus both occur in Austin, the westernmost locality for either species.Diagnosis.Th is species is most readily recognized by the dark brown hind femur.All other species from the New World have relatively pale (whitish to dark yellow) hind femora.Th e petiole is completely dark, as in E. pachycephalus, sp.n., but the latter is a much larger species with a distinctly broader gena.
Mesosoma.Posterior-ventral margin of lateral pronotum distinctly impressed, varying from crenulate to nearly smooth for most of length.Precoxal sulcus weakly impressed, not extending close to anterior margin of mesopleuron; precoxal sulcus approximately 30 degrees, inclined slightly less vertically than E. abnormis.Notaulus narrow, weakly impressed, crenulate over anterior 0.3 of mesoscutal disc; with relatively sparse cluster of short setae at fi nely rugulose base of anterior declivity and 1-2 widely spaced longer setae extending posteriorly.Propodeum with median carina present anteriorly, bifurcating near anterior 0.2 to form fi ve-sided areola over posterior 0.8; surface densely punctate-rugose to coarsely granular laterally and posteriorly, obscuring carinae, weakly sculptured to nearly smooth anteriorly on either side of short median carina.
Wings.Fore wing r-m very weakly pigmented at extreme base; somewhat tubular (with lateral boundaries distinct) over anterior 0.3-0.5;m-cu distinctly postfurcal; 3M distinctly pigmented in basal third, gradually weakening and becoming depigmented distally.Hind wing m-cu indistinct.
Color.Head, thorax, and petiole dark red-brown.Scape and pedicel yellow, fi rst four fl agellomeres light brown, quickly darkening distally to dark brown; palps, mandible, clypeus, and tegula yellow.Metasoma posteriad petiole medium brown.Legs yellow except hind femur medium to dark brown medially with apical and basal 0.1-0.15pale, tibia and tarsus almost completely medium brown, tibia variously pale brown dorsally.Ovipositor sheath dark brown, ovipositor light brown.Wings hyaline.
Male and Host.Unknown.Distribution.South central Mexico.
Etymology.Th e name ormenus is derived from Greek: ormenus, petiolated.Th e name refers to the elongate petiole of the species.Comments.Th is is a small-bodied species similar in size to E. microsomus but with a more heavily sculptured propodeum and darker hind femur.Eurytenes ormenus is characterized by the long, narrow petiole, similar in form to the petiole of E. pachycephalus sp.n. and E. abnormis and unlike the broader petiole of E. microsomus and E. dichromus.As in E. pachycephalus sp.n., and unlike the other three species, the petiole is uniformly very dark in coloration.Th e anterior tentorial pits of E. ormenus are slightly larger in this species than in the others treated here.Diagnosis.Th is species is most readily recognized by its broad clypeus and infl ated gena.It is a much larger species than E. ormenus, which was also collected at high elevation sites in central Mexico.Although both E. pachycephalus and E. ormenus have a uniformly dark petiole, the hind femur is dark in E. ormenus and more lightly colored in E. pachycephalus.
Mesosoma.Posterior-ventral margin of lateral pronotum strigose for most of length, the sculpture extending towards middle of sclerite.Precoxal sulcus extending very close to anterior margin of mesopleuron; deeply crenulate anteriorly, sculptured area broadening posteriorly; precoxal sulcus approximately 45 degrees, inclined slightly more vertically than E. abnormis.Notaulus distinctly impressed and crenulate over anterior 0.3-0.4 of mesoscutal disc; with cluster of short setae at rugulose base of anterior declivity extending ventrally to some extent onto anterior declivity at each side; longer setae absent posteriorly.Median carina extending over anterior 0.2 before bifurcating to form fi ve-sided areola; surface of areola and lateral margin of propodeum rugose, posterior-lateral fi elds and region anteriorad areola smooth or nearly so.
Wings.Fore wing r-m very weakly pigmented at extreme base, largely spectral (with lateral boundaries indistinct); m-cu distinctly postfurcal; 3M distinctly pigmented and largely tubular in basal third, gradually weakening distally.Hind wing m-cu extending nearly half way to wing margin as a very weakly pigmented and impressed curved line.
Legs.Hind tibia 8.3 × longer than maximum width.Metasoma.Petiole 2.1 × longer than apical width.Female ovipositor sheath barely visible due to postmortem changes in position; visible portion densely setose.
Color.Head, mesosoma, and petiole black.Scape and pedicel yellow, fl agellomeres dark brown; mandible butterscotch with distal tip infuscated; clypeus dark brown dorsally, ventral half butterscotch; palps and tegula butterscotch.Metasoma with T2 and T3 brown, middle tergites with brown and yellow transverse banding, apical tergites yellow.Legs yellow except hind tibia butterscotch to weakly infuscate, tarsus entirely medium brown.Wings largely hyaline, though appearing very slightly darker than other species treated here.

Taxonomy Genus Eurytenes Foerster s. str.
Eurytenes Foerster 1862: 259.Type species Opius abnormis Wesmael 1835 by original designation and monotypy.Description.Head.Antenna fi liform, longer than body.Frons, vertex, and temple smooth, shiny; frons bare, vertex and upper temple nearly so.Labrum exposed.Clypeus weakly to distinctly protruding in profi le.Malar sulcus deeply impressed.Mandible gradually to somewhat more abruptly widening from apex to base, carinate ventrally over most of basal half, never with distinct basal tooth as in Opius s. str.Maxillary palp longer than head, reaching mid coxa.Occipital carina present laterally, extending dorsal-medially at least to level of inner eye margin, broadly absent mid-dorsally; widely separated from hypostomal carina at base of mandible.