The genus Vipio Latreille (Hymenoptera, Braconidae) in the Neotropical Region

Abstract The genus Vipio Latreille is revised for the Neotropical region (south of Nicaragua). All species are fully illustrated. Thirteen species are recognised of which five (V. boliviensis, V. carinatus, V. godoyi, V. hansoni, and V. lavignei) are described as new, all with descriptions attributable to Inayatullah, Shaw & Quicke. All previously described Neotropical species are redescribed. A key is included for the identification of the Vipio species known from the Americas south of Nicaragua, and all species are illustrated.


Introduction
The braconid genus Vipio Latreille is most diverse in the Holarctic Region but has a significant representation in the Neotropical Region. However, little is known of these tropical species, only a very few of which are described (Brullé 1846, Ashmead 1900, Szépligeti 1906, Bréthes 1909, 1913, nor are there any host records for the species from this part of the world. Because of a recent upsurge in Hymenoptera studies in South America, the publication of keys to the genera of New World Braconidae , and a growing number of biodiversity studies, there is a need to provide identification keys to the Neotropical fauna. Here we present a revision of the nine Vipio species now known to occur in South America and southern Central America (south of Nicaragua), which includes descriptions of four new species, and redescriptions of the five previously described ones whose original descriptions do not mention many important characters. An illustrated key to the species is also provided. The Nearctic species were revised (Inayatullah et al., 1997) and since then only one additional New World species, V. porteri Inayatullah et al., 2015, has been described. Species from northern Central America, Mexico, and the Caribbean are diverse and comprise a complex assemblage that will be treated in a subsequent paper.

Terminology and collections
Terminology follows van Achterberg (1979,1988) except for the relative heights of eye (EH) and malar space (MS) follow Inayatullah (1992) and Inayatullah et al. (2012), and wing venation nomenclature which follows ; see also fig. 2.2 in Quicke (2015) for comparison of wing venation naming systems. Sculpture terminology follows Harris (1979). The following abbreviations are used to save space: EH eye height; FH face height measured between anterior margin of antennal socket and anterior tentorial pit; FW face width; HH head height as least distance between base of mandible and lateral ocellus (inclusive), in lateral view; HW head width; HL head length: ITD inter-tentorial distance; LMC labiomaxillary complex; LRC length of radial cell measured between apex of pterostigma and 3RSb; MS malar space; PL pterostigma length; PW maximum width of pterostigma; T tergite; TOD tentorio-ocular distance.
Description. Female. Length of body 5.4 mm; of fore wing 5.6 mm; of ovipositor (part exserted beyond apex of abdomen) 6.4 mm.
Mesosoma. Length of mesosoma 1.43 × height. Pronotum carinate antero-laterally. Notauli smooth. Propodeum rugulose medially, with a blunt and short median longitudinal carina not reaching posterior end, and with a pair of short, longitudinal, anteriorly diverging submedial carinae not reaching middle of propodeum; remainder of mesosoma laterally smooth and shiny.
Wings. Fore wing: length of fore wing 1.03 × body length; PL/LRC 0.75; PW/ PL 0.19; length of vein 1M 0.71 × length of (RS+M)a; length of vein 3RSb 0.82 × combined length of r-rs and 3RSa; vein 3RSa reaching wing margin 0.55 × distance between apex of pterostigma and wing tip. Hind wing: with a glabrous area distal to cu-a; apex of C+SC+R with one basal hamule Legs. Claw without pointed basal lobe. Metasoma. First metasomal tergite 1.20 × longer than wide, raised median area of T I oval, rugose, with a median longitudinal carina posteriorly, surrounding area with short, transverse carinae, dorso-lateral carina present; T II 1.5 × wider than long medially, longitudinally striate, basal areas rugulose, oblique furrow strongly impressed; T III 1.45 × wider than long medially, anteriorly with longitudinal striations running postero-laterally, and with transverse striations posteriorly; T IV entirely with transverse striation but not reaching lateral margin which is smooth and shiny; T V-VII smooth and shiny; hypopygium short, ending at apex of metasoma; ovipositor sheath with sparse setae; ovipositor 1.17 × body length. Colour. Yellowish red except head, antenna, palpi, prosternum, T IV posteriorly, and T V-VII, metasomal laterotergites and ovipositor sheath black. Wings smoky.
Remarks. This species appears to be closely related to V. paraguayensis Szépligeti, because of the presence of a median longitudinal carina on propodeum and similar sculpture of T1 and T2. Vipio boliviensis can be distinguished by the carinate pronotum (smooth and shiny in paraguayensis), absence of pointed basal lobe on the claw (present in paraguayensis), transverse striations on T III and T IV (longitudinal in paraguayensis), and short hypopygium (long in paraguayensis).
Etymology. Named after the country of Bolivia, where the holotype was collected. We have retained this name despite the recent discovery of a specimen from Argentina because of its use in Inayatullah's MS thesis (1992). Diagnosis. Ovipositor shorter than fore wing, propodeum with an anteriorly blunt but complete mid-longitudinal carina, claw with pointed basal lobe, ovipositor approximately half length of fore wing, body except head, predominantly yellow.
Legs. Claw with pointed basal lobe. Metasoma. First metasomal tergite 1.2 × longer than wide, raised median area oval, slightly rugulose, with a blunt and irregular dorso-lateral carinae which are more pronounced anteriorly, surrounding area with short transverse carinae, dorso-lateral carina present; T II-IV longitudinally striate (Fig. 6C); T II 1.8 × wider than medially long, basal areas smooth and shiny, oblique furrow impressed; T III 1.7 × wider than medially long; T V-VII smooth and shiny; hypopygium ending at same level as tergites; ovipositor 0.47-0.5 × body length.
Colour. Predominantly yellow to orange-yellow, head and antenna black, except maxillary palp, face laterally, and basal half of mandible blackish red, legs blackish red except fore tibia and tarsi yellow (Fig. 5A). Wings brown with dark brown venation, pterostigma entirely dark brown (Fig. 6A). 1.6-1.9 × TOD; MS 0.28-0.30 × EH. Face yellowish white with a black spot above clypeus; carinae on propodeum more pronounced than in female.
Etymology. Named for the presence of distinctive carinae on the propodeum which are diagnostic.
Diagnosis. This species can be distinguished from other Neotropical species with very long ovipositors (> 2.0 × body length) by having a yellow-red head, densely striate metasoma and a pointed basal lobe to the claw.
Colour. Yellow to reddish yellow, except tip of mandible, labial palp, basal two segments of maxillary palp, labio-maxillary complex, antenna basally, fore trochanter, middle and hind legs and ovipositor sheath black. slightly smoky to dark brown, pterostigma black, yellow basally.
Male. Unknown. Remarks. Based on the long body, ovipositor length, similar propodeal and metasomal sculpture, this species is closely related to V. melanocephalus Brullé. The longer MS (0.42-0.46 × EH) and yellow head in fiebrigi will separate it from melanocephalus, in which the MS/EH ratio is 0.39-0.41 and the head is black.
Description. Holotype ♀ length of body 7.1 mm; fore wing 7.1 mm and of ovipositor 3.8 mm.
Wings. Length of fore wing: 1.0 × body length; PL/LRC 0.8; PW/PL 0.25; length of vein 3RSb 0.88 × combined length of r-rs and 3RSa; length of vein 1M 0.7 × length of (RS+M)a; vein 3RSa reaching anterior wing margin 0.71 × distance between apex of pterostigma and wing tip. Hind wing: uniformly setose; apex of vein C+SC+R with two basal hamules.
Legs. Claw with pointed basal lobe. Metasoma. First tergite 1.1 × longer than posteriorly wide; raised median area oval, rugulose, with a median longitudinal ridge posteriorly, surrounding area smooth and shiny; dorso-lateral carina laminate, area below smooth and shiny, carina absent above spiracle; T II 1.75 × wider than medially long, longitudinally striate, basal areas smooth and shiny, oblique furrows impressed, striate; T III 1.9 × wider than medially long, longitudinally striate except apex smooth, anterolateral area smooth; all metasomal spiracles large, those of T III 0.57 × the diameter of median ocellus; T IV with short longitudinal striae at base and posterior to anterolateral area, remainder of ter- Figure 10. Montaged light micrographs of Vipio godoyi sp. nov. A Propodeum, oblique dorsal view B metasoma, lateral view C metasomal tergites II-III, dorso-lateral view D metasomal tergite I, dorso-lateral view E metasomal tergites III-IV, postero-dorsal view F male paratype, lateral habitus G holotype labels H labels. gum, smooth and shiny; T V-VII smooth and shiny, mostly retracted; hypopygium barely extending beyond apex of metasoma (Fig. 10B); ovipositor 0.54 × body length.
Colour. Reddish yellow, except head, including mouthparts and antenna, legs and ovipositor sheath black. Wings black.
Biology. Unknown. Distribution and seasonality. Costa Rica, Honduras, Nicaragua and Panama. Recorded flying from February through August in Costa Rica, November in Honduras and Panama, and October in Nicaragua. May occur sympatrically with V. hansoni sp. nov. (in one case, specimens of both species were taken from the same Malaise trap sample).
Comments. Vipio godoyi is apparently closely related to V. hansoni sp. nov. based on similar body colour, stout antennae, smooth and shiny propodeum, lamelliform dorsolateral carinae of T I, deeply impressed oblique furrows, oval and posteriorly narrowed raised median area of T I, short hypopygium, and short ovipositors in both species. Females of V. godoyi sp. nov. can be separated from those of V. hansoni sp. nov. by the presence of a pointed basal lobe on claw (absent in hansoni), and the setosity of ovipositor sheath (Fig. 10B). Males of V. godoyi have visibly larger and broader spiracles on metasomal T I-III as compared with those of other species. The diameter of spiracle on T III in males of V. godoyi is 0.6-1.0 × the diameter of median ocellus (0.35 × in hansoni).
Etymology. Vipio godoyi is named after Ms. Carolina Godoy, currently of the Instituto Nacional de Biodiversidad (INBio), who assisted with collection of the holotype specimen.  Diagnosis. Vipio hansoni sp. nov. can be recognised by the combination of the predominantly reddish yellow colour, claw with rounded basal lobe, and the presence of two anterior carinae on raised median area of first metasomal tergite. Description. Holotype ♀ length of body 5.5 mm, of fore wing 5.5 mm and of ovipositor 2.9 mm.
Wings. Fore wing: length of fore wing 1.0 × body length. PL/LRC 0.87; PW/PL 0.18; length of vein 3RSb 0.88 × combined length of r-rs and 3RSa; length of vein 1M 0.61 × length of (RS+M)a; vein 3RSa reaching anterior wing margin 0.67 × distance between apex of pterostigma and wing tip. Hind wings: uniformly setose; apex of vein C+SC+R with one basal hamule.
Legs. Claw with small, rounded basal lobe. Metasoma. First tergite 1.1 × longer than posteriorly wide, raised median area oval, rugulose, anteriorly with two carinae joining posteriorly and becoming a single median longitudinal carina reaching apex of disc; surrounding area with short transverse carinae; dorso-lateral carina present, area below rugulose; T II 1.7 × wider than long, longitudinally striate, basal areas smooth and shiny, oblique furrow impressed, striate; T III 1.15 × wider than medially long, longitudinally striate, baso-lateral areas smooth and shiny for the most part; T IV longitudinally striate, T V-VII smooth and shiny; hypopygium ending at the apex of metasoma; ovipositor 0.53 × body length.
Colour. Reddish yellow except head, legs, propleuron, and ovipositor sheath black. Wings dark brown.
Host. Unknown. Distribution and seasonality. So far recorded only from Limon, Alajuela, and Heredia Provinces in Costa Rica. Specimens were collected in March, April, and September.
Remarks. This species is closely related to V. godoyi the explanation given under godoyi distinguishes both species. This species also is similar to V. lavignei sp. nov., but the comments given under lavignei separate these species.
Etymology. Vipio hansoni is named after Professor Paul Hanson, of the Universidad de Costa Rica, who collected the holotype specimen. Diagnosis. Vipio lavignei can be recognised by the black head and pronotum, rugo-punctate face, and rectangular, dorso-laterally carinate raised median area of first metasomal tergite.
Wings. Fore wing: length of fore wing 0.96 × body length; PW/PL 0.28; PL/LRC 0.71; length of vein 3RSb 1.1 × combined length of r-rs and 3RSa; length of vein 1M 0.82 × length of (RS+M)a; vein 3RSa reaching anterior wing margin 0.74 × distance between apex of pterostigma and wing tip. Hind wing: uniformly setose; apex of vein C+SC+R with one basal hamule.
Legs. Claw without pointed basal lobe; Metasoma. First metasomal tergite 1.14 × longer than wide; raised median area rectangular with dorso-lateral carina; slightly rugulose, surrounding area with widely spaced transverse carinae, dorso-lateral carina laminate, area below smooth and shiny; T II 1.8 × wider than long, sparsely longitudinally striate; basal areas smooth and shiny; OF wide, deep, and striate; posterior of tergum smooth; second metasomal suture wide, striate; T III 1.9 × wider than medially long, smooth and shiny posteriorly, baso-lateral areas smooth and shiny for the most part with surrounding area strongly striate; T IV smooth and shiny, except for crenulate transverse basal groove; remainder of metasoma smooth and shiny; hypopygium short, ending at apex of metasoma; ovipositor 0.37 × body length.
Distribution and seasonality. Argentina and Peru. Two specimens from Argentina were collected in March and May and two from Peru in October.
Etymology. Named after UW Professor Emeritus Robert J. Lavigne, in honour of his diverse contributions to entomological research and his role in promoting the insect systematics program at the University of Wyoming.
Remarks. This species is apparently closely related to V. hansoni sp. nov. because of its strongly sclerotised metasoma, and wide, deep 2 nd metasomal suture. V. lavignei can be separated by the rugo-punctate face (smooth and shiny in hansoni sp. nov.), the black mesonotum (yellow in hansoni), and the rectangular T I raised median area (oval in hansoni). Diagnosis. Vipio melanocephalus can be recognised by the combination of its size (body length > 1cm), ovipositor length (≥ 2 × body length), largely black head and claw with large, acutely pointed basal lobe.
Colour. Predominantly orange; head largely black, face sometimes orange; frons sometimes yellow laterally; antenna rufous or black; labial and maxillary palp reddish black; legs black or reddish black, except fore legs beyond trochanter and middle tarsi orange.
Remarks. This species is similar to V. fiebrigi Bréthes, but can be readily separated by the characters discussed under V. fiebrigi. Szépligeti, 1906 Figures 17-19 Vipio paraguayensis Szépligeti, 1906: 157;Shenefelt, 1978Shenefelt, : 1857. Diagnosis. May be distinguished from other Neotropical Vipio species by the combination of long ovipositor (1.5-1.9 × body length), presence of an acutely pointed basal lobe to claw and a short mid-anterior, rather wide, carina on the propodeum.
Legs. Claw with pointed basal lobe. Metasoma. T I 1.34-1.38 × longer than wide, raised median area oval, anterior smooth area narrowing posteriorly, becoming a median longitudinal carina with short transverse carinae posteriorly; carinate at lateral margin; surrounding area with short transverse striae; dorso-lateral carina present, area below crenulate; T II 1.35-1.50 × wider than long, baso-lateral areas smooth and triangular; baso-medial area becoming a median longitudinal carina posteriorly and reaching a small raised smooth area at the apex of tergum; remainder of the tergum longitudinally striate, oblique furrows impressed, striate; T III 1.3-1.7 × wider than medially long longitudinally striate, basolateral areas distinct; T IV longitudinally striate with small baso-lateral area; T V-VII smooth and shiny; hypopygium extending 0.4-0.7 mm beyond apex of metasoma; ovipositor 1.1-1.4 × body length.
Remarks. Vipio paraguayensis can be easily recognised by the combination of the presence of a pointed basal lobe on the claw, the presence of a median longitudinal carina on the propodeum, the densely striate T II-IV, and the long ovipositor. Based on the presence of the median longitudinal carina on propodeum, this species may be closely related to V. boliviensis sp. nov. However, the presence of a pointed basal lobe on the claw, longitudinal striations on the metasoma, and longer hypopygium in paraguayensis separate it from boliviensis sp. nov. (in which the basal lobe of the claw is rounded, T III and IV are transversely striated, and the hypopygium is short). Males of this species can be confused with males of V. belfragei because of the expanded third and fourth maxillary segments, but the clavate antenna in paraguayensis (as opposed to a filiform antenna in belfragei) readily separate these two species. Another useful character is the presence of a median longitudinal carina on the propodeum in this species, as opposed to several short carinae posteriorly in belfragei. Figure 20 Vipio porteri Inayatullah, Sabahatullah & Ain Tahira, 2015: 132-133, fig. 4 xi.1967;2 ♀, same data, except 18.xi-4.xii.1967;1 ♀, same data, except 27.v-14.viii.1968;1 ♀, same data, except 24.ix-17.x.1968;1 ♀, same data, except xii.1967;4 ♀, 1 ♂, same data, except 24.ix-17.x.1968 (ESUW); 1 ♀, Las Cejas, 11 km. W, 12.iv-5.v.1968
Colour. Head black, except a yellowish red and/or yellowish stripe surrounding the eye and basal half mandible reddish yellow to yellow; antenna, maxillary and labial palpi, prosternum, propleuron, legs, and metasoma black. Wings brownish black, pterostigma black.

Distribution and seasonality.
Ranging from northern Mexico southwards to Ecuador (with records from Costa Rica, El Salvador, Ecuador, Guatemala, Honduras, Mexico and Panama). Specimens from Costa Rica were collected from January through March, June in El Salvador, May in Guatemala, July in Honduras, August through December in Panama, and from June through September in Mexico.
Remarks. Vipio quadrirugulosus appears closely related to the Nearctic V. rugator because of presence of a raised median area on the face, a strongly sclerotised and densely longitudinally striate metasoma, short ovipositor, and the presence of a pointed basal lobe on the claw. However, the black metasoma and the presence of carinae on the raised median area in V. quadrirugulosus will readily separate this species from V. rugator in which the metasoma is yellow or reddish yellow and the raised median area of the first tergite is areolate and rugose and lacks such a carina.
Colour. Largely red; head, including antenna and palpi, black except basal half of mandible reddish yellow and a yellow or yellowish red stripe surrounding the eye; pronotum reddish black with pronotal furrow red; prosternum, propleuron, basal 0.8 of mesopleuron, middle and lateral lobe of mesonotum laterally, scutellum apically, legs, and ovipositor sheath black. Wings brownish-black.
Male (N = 12). As in female, except length of body 4.6-6.5 mm, fore wing as long as body length; antennae 1.0-1. Remarks. Vipio strigator can be recognised by the combination of reddish black markings on the mesosoma, the short hypopygium, and the short ovipositor. This species is similar to the Nearctic V. rugator because of the presence of a raised area on face, short hypopygium, and short ovipositor in both species. However, the red coloration with reddish black markings on the mesosoma and a carinate propodeum in strigator will readily separate it from rugator (in which the mesosoma lacks black markings and propodeum lacks such carinae).
Colour. Face yellow, except a black, raised median area; palpi, tip of mandible, vertex, temple, occiput, and antenna black; metasoma reddish yellow to black. Wings smoky.
Remarks. This species resembles the Nearctic species V. rugator because of the presence of a raised area on the face, strong dorso-lateral carina of T I, widely ovate and densely striate metasoma, and short hypopygium. The relatively longer ovipositor (ovipositor length/body length 0.5) and reddish black mesosoma separate thoracica from rugator, in which the ovipositor is shorter (ovipositor length/body length 0.29-0.35) and the metasoma is yellow or reddish yellow.

Conclusions
The genus Vipio in the Neotropics is generally uncommon and most of the material available for examination is rather old. It can be noted that at the time of writing, not one Neotropical Vipio barcode sequence is listed on The Barcode of Life Data System (BOLD) (www. barcodinglife.org) (Ratnasingham and Hebert 2007) despite extensive Malaise trapping in Costa Rica, French Guiana, and Honduras, and examination of various other Neotropical samples. Currently, the BOLD database contains only 13 specimens of Vipio, one from a European Malaise trap and the remainder from an extremely extensive sampling of North America (Young et al. 2012, Steinke et al. 2017, Global Malaise Trap Program 2019. New and freshly collected material would be desirable for carrying out molecular investigation including DNA barcoding to test the current morphological taxonomic hypotheses. School, Chulalongkorn University, for the award of a Senior Postdoctoral Fellowship to DLJQ, Office of Higher Education Commission (BDCPG2-160007) and RSPG to BAB. Finally, special thanks to Sadaf for patience, understanding, and support. Any opinions, findings, and conclusions expressed are those of the authors and do not necessarily reflect the views of the National Science Foundation.