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Two new species, Heterospilus belokobylskiji Kula, sp. n. and Heterospilus vincenti Kula, sp. n., from the Nearctic Region are described and differentiated from all other New World species of Doryctinae that exhibit brachyptery or aptery. They are the first brachypterous species of Heterospilus Haliday known in the New World and increase the total number of brachypterous species in the genus to four worldwide.
Apterous, aptery, brachyptery, parasitoid, taxonomy
Of the 1, 335 species of Doryctinae listed as valid in
Heterospilus
Haliday, the richest doryctine genus in the New World considering the
number of undescribed species (P. Marsh in litt.), is one genus for
which brachypterous and apterous species are not known in the New World.
However, Heterospilus brachyptera (Jakimavicius), with the female brachypterous and known only from the holotype, and Heterospilus hemipterus have been reported from the Palearctic Region (
The author discovered two new brachypterous species of Doryctinae in the Nearctic Region through a study testing pan trap color preference for selected Hymenoptera. The two species fit Heterospilus sensu
Specimens were collected using blue, red, and white 12 ounce Solo™ (Urbana, Illinois) party bowls placed in an ~100 m wide power line right-of-way ~two miles east of Prince Frederick, Maryland. The clearing runs roughly north-south and is bordered to the east and west by eastern deciduous forest. The flora within the clearing was not surveyed. Topographically, it contains upland areas primarily with herbaceous plants and lowland areas primarily with woody plants. All traps were placed in upland areas. The bowls were filled with a solution of water and Liqui-Nox (Alconox, Inc., White Plains, New York) detergent; the latter served as a surfactant for the water. Contents of the bowls were collected every other day, and the bowls were refilled with water-detergent solution at that time.
Specimens were dehydrated using hexamethyldisilazane (HMDS) as in
Terminology for morphological features and setation largely follows
Measurements were taken with an ocular micrometer as in
Abbreviations used in diagnoses and descriptions are as in
Habitus images were obtained using a Visionary Digital imaging system. The system consists of an Infinity Optics K2 long distance microscope affixed to a Canon EOS 40D digital SLR camera. A Dynalite M2000er power pack and Microptics ML1000 light box provided illumination. Image capture software is Visionary Digital’s proprietary application with images saved as TIF with the RAW conversion occurring in Adobe Photoshop Lightroom 1.4. Image stacks were montaged with Helicon Focus 4.2.1. Final images were prepared using Adobe Illustrator CS4 and are deposited in Morphbank (image ID numbers 581765, 581772, 581777, and 581782).
Results and discussionHeterospilus belokobylskiji Kula, sp. n. and Heterospilus vincenti Kula, sp. n. can be differentiated from other brachypterous or apterous doryctines in the New World (excluding ypsistocerines) using form of the wings (Table 1). Additionally, the scutellar disc is flat in Heterospilus belokobylskiji and Heterospilus vincenti; it is convex in Aptenobracon formicoides and conical in Ecphylopsis costaricensis. A tubercle is present at the base of the hind coxa in Heterospilus belokobylskiji and Heterospilus vincenti; it is round at the base in Ecphylopsis costaricensis, Ecphylus caudatus, Ecphylus lepturgi, Ecphylus pacificus, and Ecphylus schwarzii. The propodeal bridge is absentin Heterospilus belokobylskiji and Heterospilus vincenti; the metasoma articulates with the mesosoma directly above the metacoxae. The propodeal bridge is present between the metacoxae and petiolein Oroceguera andersoni; the metasoma articulates with the mesosoma high above the metacoxae resulting in a large gap between those features (cf. cenocoeliine braconids). The femora are not enlarged in Heterospilus belokobylskiji and Heterospilus vincenti; all femora are enlarged in Pambolidea yuma, Psenobolus ficarius, Psenobolus parapygmaeus, and Psenobolus triangularis.
Species of Doryctinae in the New World, excluding ypsistocerines, that exhibit brachyptery or aptery.
Species | Female wing form | Male wing form |
---|---|---|
Aptenobracon formicoides | apterous | apterous |
Ecphylopsis costaricensis | scalelike pads | scalelike pads |
Ecphylus caudatus | apterous, macropterous | apterous, macropterous |
Ecphylus lepturgi | apterous, macropterous | apterous |
Ecphylus pacificus | unknown | apterous |
Ecphylus schwarzii | apterous | apterous |
Heterospilus belokobylskiji Kula, sp. n. | brachypterous | brachypterous |
Heterospilus vincenti Kula, sp. n. | brachypterous | brachypterous |
Oroceguera andersoni | apterous | unknown |
Pambolidea yuma | apterous, macropterous | apterous |
Psenobolus ficarius | macropterous | brachypterous |
Psenobolus parapygmaeus | macropterous | brachypterous |
Psenobolus triangularis | macropterous | brachypterous |
urn:lsid:zoobank.org:act:09A27F4E-4564-4FA1-95B2-ED283F49D341
Figs 1–2female. U.S.A., “MARYLAND:Calvert Co. [;] 2 mi E Prince Frederick [;] 38°33'3.83"N, 76°33'3.09"W [;] 14.v.-16.v.2007 SEL Hym Unit [;] pan trap, transect6 treatment B" (USNM).
1 ♂ same data as holotype except 38°32'57.95"N, 76°33'1.43"W, transect8 treatment W (USNM).
The vertex is smooth except a pair of small strigulate areas posterolaterad the lateral ocelli in Heterospilus belokobylskiji; the vertex is entirely strigate to strigate-coriaceous in Heterospilus hemipterus, and it is entirely coriaceous in Heterospilus vincenti. The face is smooth in Heterospilus belokobylskiji; the face is at least partially strigate in Heterospilus hemipterus, and it is smooth mesally and coriaceous laterally in Heterospilus vincenti. The frons is partially strigulate in Heterospilus belokobylskiji; the frons is entirely coriaceous in Heterospilus vincenti. The mesopleuron (excluding subalar groove, precoxal sulcus, and posterior mesopleural furrow) is weakly coriaceous with some areas nearly smooth in Heterospilus belokobylskiji; the mesopleuron is at least partially strigate in Heterospilus hemipterus. The hind wing stigma of the male is located slightly basad the middle of the wing in Heterospilus belokobylskiji; the stigma is located at the wing apex in Heterospilus vincenti. Transverse grooves are absent on T3 in Heterospilus belokobylskiji; a crenulate transverse groove is present on T3 in Heterospilus hemipterus. The head (excluding mouthparts and antenna) is brown in Heterospilus belokobylskiji; the head is yellow in Heterospilus vincenti.
Female (Fig. 1).
Body length. 2.28 mm.
Head. HL 0.81× HW, HW 1.09× TW, FW 1.92× FH, EL 1.00× EH, MSH 0.80× EH, F1L 0.85× F2L, PMPL 0.45× F1L; antenna broken at eighth flagellomere; mandible with two teeth, tooth closest to labiomaxillary complex shorter than other tooth, setiferous; malar space smooth, setiferous, malar suture absent; clypeus with roughly apical 1/2 setiferous and basal 1/2 glabrous; face smooth, glabrous mesally and setiferous laterally; frons partially strigulate and partially smooth, sculpture strongest in depressions dorsad antennal sockets, glabrous except one to two setae along margin of eye; vertex mostly smooth but with pair of small strigulate areas posterolaterad lateral ocelli, setiferous; ocelli present but small (cf. Heterospilus striatus Muesebeck & Walkley); gena smooth, setiferous; occiput smooth, glabrous except pair of setae ventrally on both sides of head.
Mesosoma. ML 2.83× MW, ML 1.97× MH, MW 0.70× MH, SSL 0.50× SSW; pronotal collar without transverse carina, anterior portion rugulose and posterior portion smooth mesally and crenulate laterally, anterior and posterior portions both glabrous except setiferous along anterior margins, pronope absent, lateral portion of pronotum (including pronotal groove) rugose, roughly setiferous along margins and glabrous mesally; notauli complete and meeting posteromesally, bearing a few weak crenulae; mesoscutal midpit absent; mesoscutum (excluding lateral margin and notauli) coriaceous, setiferous along margins and notauli; scutellar sulcus with median longitudinal carina and pair of crenulae adjacent to carina; scutellar disc weakly coriaceous, glabrous except five setae along lateral and posterior margins; propodeum strongly carinate, setiferous, carinae forming hastate areola mesally, sculpture within areola areolate-rugose, mesolaterally with transverse carina dividing propodeum into roughly basal and apical halves, dorsal lateral carinae dividing basal 1/2 into median and lateral areas, median area with weak indication of coriaceous sculpture and lateral area areolate-rugulose, apical 1/2 areolate-rugose; subalar groove crenulate; precoxal sulcus present in roughly anterior 1/2 of mesopleuron and crenulate, posterior 1/2 of mesopleuron without impression but with rugosities extending to mesocoxa; posterior mesopleural furrow crenulate; mesopleuron (excluding subalar groove, precoxal sulcus, and posterior mesopleural furrow) weakly coriaceous with some areas nearly smooth, setiferous except glabrous area dorsomesally roughly between subalar groove and posterior mesopleural furrow to level of episternal scrobe; metapleuron areolate-rugose, setiferous; metacoxa with anteroventral basal tubercle.
Forewing. Brachypterous, extending to posterior margin of T2 (including fringe); hyaline; stigma present anterodistally, posterior margin difficult to differentiate from R1 vein; with following veins complete and tubular: C+SC+R, M+CU, 1-1A, 1RS, 1M, and 1CU; (RS+M) and m-cu veins complete but nebulous resulting in distinct 1st discal cell; one wing with 3RS vein minute but tubular and clearly differentiated from stigma and R1 vein.
Hind wing. Brachypterous, extending to posterior margin of T2 (including fringe); hyaline; basal and subbasal cells enclosed by tubular veins, veins enclosing cells differ in width and degree of sclerotization; R1 vein tubular; M+CU vein shorter than 1M vein.
Metasoma. T1L 1.15× T1W; subcylindrical; ovipositor with minute teeth ventrally, EOL about 2.23× T2+T3L; ovipositor sheaths setiferous, setae increasing in density anteriorly to posteriorly; T1 costate, dorsal carinae extending posteriorly about 3/4 length of tergum, setiferous, dorsope present; T2 costate, setiferous; transverse groove between T2+T3 weakly impressed, smooth; T3–T7 smooth, setae forming single transverse row in middle or posterior 1/2 of tergum; T8 smooth, setae in no apparent pattern.
Color. Head (excluding mouthparts and antenna) brown, mouthparts whitish yellow except mandible yellow with teeth brown, scape and pedicel yellow, flagellum yellow proximally transitioning to brown distally; mesosoma orangish brown except pronotum and propleuron yellowish brown; wing venation tan; legs yellow; T1–T2 entirely yellowish brown, T3–T5 mostly brown with posterior edge slightly darker but all with some irregular yellow coloration, T6 yellow anteromesally but otherwise yellowish brown, T7 yellowish brown, T8 yellow.
Lateral habitus images of Heterospilus belokobylskiji, scale bars = 1.00 mm. 1 Female 2 Male.
(Fig. 2). As in female except:
Body length. 2.04 mm.
Head. HL 0.78× HW, HW 1.06× TW, EL 0.93× EH, MSH 0.73× EH, F1L 0.93× F2L, PMPL 0.38× F1L; antenna with 17 flagellomeres; frons glabrous except a few setae along margin of eye.
Mesosoma. ML 2.03× MH, MW 0.72× MH; pronotal collar with anterior portion coriaceous and posterior portion rugulose, scutellar sulcus with median longitudinal carina and pair of shorter longitudinal carinae adjacent to median carina; propodeum with basal and dorsal lateral carinae distinct, basal median area rugulose, remainder of propodeum areolate-rugose, areola (if present) obscured by surrounding sculpture.
Forewing. Extending nearly to end of T3 (including fringe).
Hind wing. Extending nearly to end of T3 (including fringe); stigma slightly basad middle of wing, subelliptical; basal and subbasal cells enclosed by tubular veins except delimited distally by stigma, basal cell delimited ventrally by M+CU vein, 1M vein absent; R1 vein tubular.
Metasoma. T1–T2 costate-rugose; T2 sculpture extending into transverse groove between T2+T3; T3 smooth except band of carinulae anteromesally.
Color. Mesosoma brownish yellow; T1–T2 entirely brownish yellow, T3 brown with posterior edge slightly darker except yellow anteromesally, T4 brown with posterior edge slightly darker, T5–T6 yellow with posterior edge brown, T7–T8 yellow.
Unknown.
This species is named in honor of Dr. Sergey A. Belokobylskij for his contributions to braconid systematics and for providing information on brachypterous and apterous doryctines critical to completion of this article.
urn:lsid:zoobank.org:act:0C6454A6-4090-4032-8C26-EF5A3A587CCE
Figs 3–4female. U.S.A., “MARYLAND:Calvert Co. [;] 2 mi E Prince Frederick [;] 38°33'4.19"N, 76°33'3.96"W [;] 30.v.-1.vi.2007 SEL Hym Unit [;] pan trap, transect5 treatment B" (USNM).
1 ♂ same data as holotype except 38°33'15.72"N, 76°33'8.73"W, 14.v.-16.v.2007, transect4 treatment R (USNM).
Heterospilus vincenti can be differentiated from Heterospilus belokobylskiji using the diagnosis for Heterospilus belokobylskiji. The vertex is coriaceous in Heterospilus vincenti; the vertex is strigate to strigate-coriaceous in Heterospilus hemipterus. The face is smooth mesally and coriaceous laterally in Heterospilus vincenti; the face is at least partially strigate in Heterospilus hemipterus. The frons is coriaceous in Heterospilus vincenti; the frons is strigate Heterospilus hemipterus. Transverse grooves are absent on T3 in Heterospilus vincenti; a crenulate transverse groove is present on T3 in Heterospilus hemipterus. The head (excluding mouthparts and antenna) is yellow in Heterospilus vincenti; the head is dark reddish brown except yellowish brown along eye and ventrally in Heterospilus hemipterus.
Female (Fig. 3).
Body length. 2.36 mm.
Head. HL 0.77× HW, HW 1.09× TW, FW 1.83× FH, EL 1.00× EH, MSH 1.00× EH, F1L 0.93× F2L, PMPL 0.31× F1L; antenna with 17 flagellomeres; mandible with two teeth, tooth closest to labiomaxillary complex shorter than other tooth, setiferous; malar space coriaceous, setiferous, malar suture absent; clypeus with roughly apical 1/2 setiferous and basal 1/2 glabrous; face smooth mesally and coriaceous laterally, glabrous mesally and setiferous laterally; frons coriaceous, glabrous except a few setae along margin of eye; vertex coriaceous, setiferous; ocelli present but small (cf. Heterospilus striatus); gena coriaceous, setiferous; occiput smooth, glabrous except pair of setae ventrally on both sides of head.
Mesosoma. ML 3.58× MW, ML 2.27× MH, MW 0.63× MH, SSL 0.57× SSW; pronotal collar with transverse carina, anterior portion smooth and posterior portion rugose, anterior and posterior portions both glabrous except setiferous along anterior margins, pronope absent, lateral portion of pronotum (including pronotal groove) rugose except small coriaceous area dorsally, roughly setiferous along ventral and posterior margins but otherwise glabrous; notauli complete to transscutal articulation and separated by carina posteromesally, more strongly impressed anteriorly than posteriorly, bearing a few weak crenulae; mesoscutal midpit absent; mesoscutum (excluding lateral margin and notauli) coriaceous, setiferous along margins and notauli; scutellar sulcus with median longitudinal carina; scutellar disc coriaceous, glabrous except pair of setae laterally; propodeum strongly sculptured, setiferous, with basal and dorsal lateral carinae distinct, basal median area coriaceous, basal lateral area rugose, remainder of propodeum areolate-rugose, areola absent; subalar groove crenulate; precoxal sulcus complete to mesocoxa, more strongly impressed anteriorly than posteriorly, transitioning from crenulate anteriorly to rugose posteriorly; posterior mesopleural furrow crenulate; mesopleuron (excluding subalar groove, precoxal sulcus, and posterior mesopleural furrow) coriaceous, setiferous with setae largely confined to margins, subalar groove, and precoxal sulcus; metapleuron areolate-rugose, setiferous; metacoxa with anteroventral basal tubercle.
Forewing. Brachypterous, extending to end of mesosoma (including fringe); hyaline; stigma absent; venation limited to tubular vein along anterior margin complete to wing apex and vein along posterior margin transitioning from nebulous proximally to tubular distally and bending anteriorly near wing apex to intersect vein along anterior margin.
Hind wing. Brachypterous, extending to end of mesosoma (including fringe); hyaline; basal cell distinct but open, SC+R vein spectral distally; subbasal cell enclosed by tubular veins; SC+R vein and 1M vein converge distally to form thickening roughly width of two veins.
Metasoma. T1L 1.14× T1W; subcylindrical; ovipositor with minute teeth ventrally, EOL about 3.28× T2+T3L; ovipositor sheaths setiferous, setae increasing in density anteriorly to posteriorly; T1 costate-rugose, dorsal carinae blending with ground sculpture posteriorly, setiferous, dorsope present; T2 carinate-rugulose, setiferous; transverse groove between T2+T3 absent, T2 and T3 indicated by inconspicuous break in sculpture; T3 carinulate in roughly anterior 1/2, smooth in roughly posterior 1/2, setiferous with most setae forming transverse row in middle of tergum; T4–T8 smooth, setae forming single transverse row in middle or posterior 1/2 of tergum.
Color. Head (excluding mouthparts and antenna) yellow, mouthparts whitish yellow except mandible yellow with teeth brown, scape and pedicel yellow, flagellum yellow proximally transitioning to brown distally; mesosoma yellow with pronotum and propleuron slightly lighter; wing venation and legs yellow; T1–T2 entirely yellow, T3–T4 yellow with posterior edge brownish yellow, T5–T8 entirely yellow.
Lateral habitus images of Heterospilus vincenti, scale bars = 1.00 mm. 3 Female 4 Male.
(Fig. 4). As in female except:
Body length. 2.38 mm.
Head. HL 0.80× HW, HW 1.11× TW, FW 1.92× FH, EL 0.94× EH, MSH 0.94× EH, F1L 0.88× F2L, PMPL 0.29× F1L; antenna with 20 flagellomeres; gena weakly coriaceous, sculpture barely discernable in some areas; occiput glabrous except a few setae ventrally on both sides of head.
Mesosoma. ML 3.48× MW, ML 2.29× MH, MW 0.66× MH; pronotal collar without transverse carina, anterior portion coriaceous-rugulose and posterior portion rugulose; notauli weakly impressed but complete and meeting posteromesally, bearing a few rugosities; scutellar sulcus with pair of crenulae; scutellar disc bearing three setae; propodeum largely obscured by hind wings but strongly sculptured, basal carina distinct, outer-most dorsal lateral carina indistinct, inner-most dorsal lateral carina (if present) obscured by hind wings, visible portion of basal median area coriaceous, remaining visible portions areolate-rugose, areola (if present) obscured by hind wings.
Forewing: Additional tubular vein located above vein along posterior margin, additional vein arising at base of wing and terminating into vein along posterior margin roughly at its midpoint.
Hind wing: Base of wing membranous with minute veins along anterior and posterior margins; apex of wing with stigmalike swelling bearing flap of wing membrane at distal end of swelling.
Metasoma: T1L 1.26× T1W; T1 carinate-rugose; T2 carinulate-rugose; transverse groove between T2+T3 weakly impressed, T2 and T3 also indicated by inconspicuous break in sculpture; T3 carinulate-rugose in roughly anterior 1/2, smooth in roughly posterior 1/2; T4 with a few crenulae anteriorly but otherwise smooth.
Color: T3 roughly anterior 1/3 yellow and posterior 2/3 brown, T4 yellow anteriorly and brown posteriorly (partially retracted under T3).
Unknown.
This species is named for the author’s son, Vincent Marion Kula.
I thank Erin Kolski (private contractor; Bowie, Maryland) for sorting pan trap samples and dehydrating, mounting, and labeling specimens. I also thank Matthew Kweskin (formerly Systematic Entomology Laboratory [SEL]) for mounting and labeling specimens. Sergey A. Belokobylskij (Zoological Institute of Russian Academy of Sciences, St. Petersburg) graciously provided information, including unpublished data, for brachypterous and apterous doryctines. I am grateful to Taina Litwak (SEL) for capturing and digitally enhancing the lateral habitus images. Michael G. Pogue (SEL), Thomas J. Henry (SEL), and Paul M. Marsh (Kansas State University) kindly reviewed a presubmission version of the manuscript.