Systematics of the parasitic wasp genus Oxyscelio Kieffer (Hymenoptera, Platygastridae s.l.), Part I: Indo-Malayan and Palearctic fauna

Abstract The Indo-Malayan and Palearctic species of Oxyscelio (Hymenoptera: Platygastridae s.l.) are revised. A total of 90 species are recognized as valid, 19 of which are redescribed - Oxyscelio acutiventris (Kieffer), Oxyscelio brevinervis (Kieffer), Oxyscelio carinatus (Kieffer), Oxyscelio ceylonensis (Dodd), Oxyscelio consobrinus (Kieffer), Oxyscelio crassicornis (Kieffer), Oxyscelio cupularis (Kieffer), Oxyscelio dorsalis (Kieffer), Oxyscelio excavatus (Kieffer), Oxyscelio flavipennis (Kieffer), Oxyscelio florus Kononova, Oxyscelio foveatus Kieffer, Oxyscelio kiefferi Dodd, Oxyscelio magnus (Kieffer), Oxyscelio marginalis (Kieffer), Oxyscelio naraws Kozlov & Lê, Oxyscelio perpensus Kononova, Oxyscelio rugosus (Kieffer) and Oxyscelio spinosiceps (Kieffer), and 71 which are described as new - Oxyscelio aclavae Burks, sp. n., Oxyscelio amrichae Burks, sp. n., Oxyscelio anguli Burks, sp. n., Oxyscelio angustifrons Burks, sp. n., Oxyscelio angustinubbin Burks, sp. n., Oxyscelio arcus Burks, sp. n., Oxyscelio arvi Burks, sp. n., Oxyscelio asperi Burks, sp. n., Oxyscelio aureamediocritas Burks, sp. n., Oxyscelio bipunctuum Burks, sp. n., Oxyscelio brevidentis Burks, sp. n., Oxyscelio caesitas Burks, sp. n., Oxyscelio capilli Burks, sp. n., Oxyscelio capitis Burks, sp. n., Oxyscelio cavinetrion Burks, sp. n., Oxyscelio chimaerae Burks, sp. n., Oxyscelio codae Burks, sp. n., Oxyscelio convergens Burks, sp. n., Oxyscelio cordis Burks, sp. n., Oxyscelio crateris Burks, sp. n., Oxyscelio crebritas Burks, sp. n., Oxyscelio crustum Burks, sp. n., Oxyscelio cuculli Burks, sp. n., Oxyscelio cyrtomesos Burks, sp. n., Oxyscelio dasymesos Burks, sp. n., Oxyscelio dasynoton Burks, sp. n., Oxyscelio dermatoglyphes Burks, sp. n., Oxyscelio doumao Burks, sp. n., Oxyscelio fistulae Burks, sp. n., Oxyscelio flabellae Burks, sp. n., Oxyscelio flaviventris Burks, sp. n., Oxyscelio fodiens Burks, sp. n., Oxyscelio fossarum Burks, sp. n., Oxyscelio fossularum Burks, sp. n., Oxyscelio genae Burks, sp. n., Oxyscelio granorum Burks, sp. n., Oxyscelio granuli Burks, sp. n., Oxyscelio greenacus Burks, sp. n., Oxyscelio halmaherae Burks, sp. n., Oxyscelio intermedietas Burks, sp. n., Oxyscelio jaune Burks, sp. n., Oxyscelio jugi Burks, sp. n., Oxyscelio kramatos Burks, sp. n., Oxyscelio labis Burks, sp. n., Oxyscelio lacunae Burks, sp. n., Oxyscelio latinubbin Burks, sp. n., Oxyscelio latitudinis Burks, sp. n., Oxyscelio limae Burks, sp. n., Oxyscelio longiventris Burks, sp. n., Oxyscelio mesiodentis Burks, sp. n., Oxyscelio mollitia Burks, sp. n., Oxyscelio nasolabii Burks, sp. n., Oxyscelio nodorum Burks, sp. n., Oxyscelio noduli Burks, sp. n., Oxyscelio nubbin Burks, sp. n., Oxyscelio obsidiani Burks, sp. n., Oxyscelio ogive Burks, sp. n., Oxyscelio operimenti Burks, sp. n., Oxyscelio peludo Burks, sp. n., Oxyscelio planocarinae Burks, sp. n., Oxyscelio praecipitis Burks, sp. n., Oxyscelio reflectens Burks, sp. n., Oxyscelio regionis Burks, sp. n., Oxyscelio sinuum Burks, sp. n., Oxyscelio spinae Burks, sp. n., Oxyscelio striarum Burks, sp. n., Oxyscelio tecti Burks, sp. n., Oxyscelio unguis Burks, sp. n., Oxyscelio vadorum Burks, sp. n., Oxyscelio vittae Burks, sp. n. and Oxyscelio zeuctomesos. Neotypes are designated for nine species, including the type species O. foveatus Kieffer, Oxyscelio brevinervis (Kieffer), Oxyscelio bifurcatus (Kieffer), Oxyscelio frontalis (Kieffer), Oxyscelio crassicornis (Kieffer), Oxyscelio cupularis (Kieffer), Oxyscelio foveatus Kieffer, Oxyscelio kiefferi Dodd, Oxyscelio magnus (Kieffer) and Oxyscelio marginalis (Kieffer). Oxyscelio bifurcatus (Kieffer) syn. n. and Oxyscelio frontalis (Kieffer) syn. n. are synonymized under Oxyscelio consobrinus (Kieffer). The fauna is divided into 13 species groups, with six species unplaced to a group. A phylogenetic analysis employing 73 morphological characters did not find most of these groups to be monophyletic, but they are retained to aid in specimen identification. Potential biogeographical patterns are discussed, including regional variation in surface sculpture and a morphological link between Sri Lankan and Australian species.


Introduction
The genus Oxyscelio Kiefer comprises relatively robust platygastoid wasps that occur across equatorial and east Africa, the south-eastern part of the Palearctic, the Indo-Malayian and Australasian regions. They are relatively easily identified by the fore wing submarginal vein being distant from wing margin, very short marginal vein, virtually absent postmarginal vein, posteriorly rounded vertex, and distinct metascutellum. In addition, many species can be recognised by the pronounced frontal depression on the head which is often rimmed by a carina.
The genus was first erected for a single species from Indonesia, Oxyscelio foveatus, by Kieffer (1907), which he later treated as a subgenus of Chromoteleia Ashmead (Kieffer 1910a) but later raised again to generic rank (Kieffer 1926). Following its original description the status of Oxyscelio and the genera that surround it was particularly confusing. This is clearly evident in the fact that, other than the type species, all taxa described prior to 1930 that are currently accommodated in Oxyscelio were described under other generic names. This situation was resolved by Dodd (1931) who recognized a range of species from the Oriental and Australian regions as being congeneric based on several key characters, the form of the fore wing venation (outlined above) and the structure of the metanotal plate. In so doing he treated Dicroteleia Kieffer, Camptoteleia Kieffer and Xenoteleia Kieffer as junior synomyms of Oxyscelio, and transferred 32 species to that genus including all Australian taxa that he had preciously described under Sceliomorpha Ashmead (sensu Kieffer 1926). This work by Dodd (1931) and his other studies on various scelionid genera around that time provided significant taxonomic stability and are testament to his thoughtful and perceptive approach to discriminating genera and species.
Since the descriptive work of Kieffer and Dodd prior to1920, only three additional species of Oxyscelio have been added to the world fauna (Kozlov and Lê 2000;Kononova 2007). Masner (1976) provided a diagnosis of the genus and key to separate putative related genera, and Johnson (1992) catalogued the world's species.
The current study is the first of three papers that aim to fully revise the world species of Oyscelio, focusing on describing the large number of new taxa. This first paper deals with the Indo-Malayan and Palearctic species; the second one will treat the Australasian taxa, and the third one the African species. This work has arisen from our Platygastroidea Planetary Biodiversity Inventory (see below) which aims to revise all species on a worldwide basis for a number of important platygastroid genera.
The contributions of the individual authors are as follows; R.A. Burks: character definition, species concept development; key development, imaging, capture of specimen data, manuscript preparation, phylogenetic analysis and illustration; L. Masner: specimen acquisition, and generic overview; N.F. Johnson: generic concept development, software and database development and manuscript preparation; A.D. Austin: initial species concept development, manuscript preparation, and taxonomic overview.
This revision is a product of the Platygastroidea Planetary Biodiversity Inventory, funded by the U.S. National Science Foundation (N.F. Johnson, Ohio State University; Andy Austin, University of Adelaide; Principal Investigators). An objective of this project is to use biodiversity informatics resources to accelerate taxonomic work, making real-time collaboration possible. Data associated with specimens examined in this study can be accessed at hol.osu.edu and entering the unique specimen identifier (e.g. OSUC 247918) in the search form. Life science identifiers (LSIDs) can be resolved at http://lsid.tdwg.org (i.e. urn:lsid:zoobank.org:act: 99E3E72E-DA88-4740-9ECB-2D03BCD1DACE).
Terminology. Morphological terminology follows Mikó et al. (2007) except where noted. Antennal terminology follows Bin (1981). Anteclypeus and postclypeus are used sensu Dangerfield et al. (2001). Dorsal epomial carina and vertical epomial carina were mentioned by Masner and Johnson (2007) and are here used as illustrated by Talamas et al. (2011). Ovipositor terminology is used as described by Austin and Field (1997). "Middle genal carina" is the largest carina subparallel to the eye but between the genal carina and the carina immediately encircling the eye; it has proven to be recognizable as homologous (when present) in Oxyscelio. T1 midlobe refers to the raised antero-medial area of T1 that is flanked by depressed lateral areas. This is usually flat and only weakly elevated in Oxyscelio, and therefore is not strictly the same as a T1 horn, but a T1 midlobe can be expressed as a T1 horn.
Surface sculpture terminology referring to repeated sculptural elements follows Eady (1968) when possible, with a novel set of designations and distinctions to increase specificity and descriptive value. Diminutive terms such as "foveolate" and "rugulose" were avoided because of a nearly total lack of criteria for separating them from nondiminutive alternatives. "Major" surface sculpture is here treated as repeated sculptural patterns that interact with seta placement. It does not include non-repeated elements or those which are repeated only once due to bilateral symmetry. Umbilicate-foveate sculpture refers to rounded crater-like sculptural elements, each surrounding a setiferous pit, with each fovea being much larger than its setiferous pit and spatially separated from that pit. Umbilicate-punctate sculpture indicates that no sculptural element accompanies the setiferous pit. Rugose sculpture refers to branching or wrinkling elevations that flank setiferous pits but do not fully surround them. Rugose sculpture can coexist with umbilicate sculpture in the same area of the sclerite, in which case the rugae occur on spaces between umbilicate sculptural elements. Note that "rugose" refers to a distribution of sculptural elements, and therefore can be "irregular" or "regular" even though rugae (the elements themselves) are by definition at least slightly irregular. Where both umbilicate-foveate and umbilicate-punctate sculpture are reported for the same sclerite, this should be interpreted as variable sculpture where some setiferous pits are surrounded by foveae while others are not. Under this scheme, "major" surface sculpture cannot occur in any part of the sclerite that lacks setae.
"Microsculpture" refers to repeated tiny sculptural elements that do not interact with seta placement. Microsculpture can occur on "major" sculptural elements, such as on rugae and on all surfaces of foveae. Punctate microsculpture refers to tiny round pits that do not bear setae. Granulate microsculpture refers to sculpture that is similar to that of leather or skin, with areas enclosed by tiny grooves (= sunken septa). Microsculpture can occur in areas that lack setae.
Sculptural terms for repeated sculpture that are not included in the above categories are 1) "carinae" which refers to elevations that are sharp and not branched or wrinkled, 2) "striae" which refers to repeated elevations that are not sharp and do not branch or exhibit wrinkling. These sculptural elements do not interact with setiferous pit placement, but umbilicate sculpture can occur between them. While alternative logic may suggest that rugose sculpture is better classed within this category, this choice was avoided because rugose sculptural patterns did apparently interact with umbilicate sculptural patterns. For the occipital carina, "crenulate" means that short carinae radiate from the occipital carina. For carinae in general: the carina may be described using the phrase "as a ruga" if it is expressed as a wrinkled and/or irregularly meandering elevation.
Illustrations and data citations. Photographs were taken using one of the following systems: 1) Visionary Digital BK+ Imaging System, November 2010 model, with either a K2 Long Distance Microscope or a 65 mm varifocal lens; 2) Synoptics, Ltd. system using a Leica Z16 APO microscope and a JVC KY-F75U 3-CCD camera; or 3) GT EntoVision Mobile Imaging System. Source photos were stacked using Zerene Stacker version 1.04 or Auto-Montage Pro version 5.01.0005, and enhanced using Adobe Photoshop CS5 or CS6.

Biology.
Although there are a large number of species known from Asia and Australia there are apparently no available rearing records from hosts identified beyond ordinal level (see Kononova and Fursov 2007, for photograph of egg of orthopteran host of Oxyscelio perpensus Kononova). However, given the large size of most species, the diversity of habitats in which they have been collected, and the structure of the ovipositor, we presume that all Oxyscelio species parasitise orthopteran eggs of some type.
Phylogenetic relationships. Masner (1976) postulated that Oxyscelio is related to Bracalba Dodd, Chromoteleia Ashmead and Baryconus Foerster, and he provided a key to separate them. Based on the structure of the ovipositor system it is unlikely that Baryconus is related to this group of genera given it has a Ceratobaeus-type system, where the other genera all have a Scelio type ovipositor system (Austin and Fields 1997). The molecular phylogenetic study of Murphy et al. (2007) included three of these genera, Oxyscelio, Chromoteleia and Baryconus, and none showed a sister-group relationship to each other, although the support on the branches that linked these genera were far from robust. Two of us (NFJ and ADA) are currently coordinating a significantly expanded molecular analysis of the Platygastroidea involving additional sequence data and a trebling of taxa, and this should help resolved the relationships among these and other genera.
The species level phylogeny generated as a part of this study ( Fig. 1) does not always uphold monophyly of the species groups described below. The authors do not see this as a major problem, as the species groups are informal groupings that are not necessarily meant to be strictly monophyletic. They are meant to be useful for species diagnostics, but can be seen as potentially valid alternatives to the included phylogeny.

Species Groups of Oxyscelio
For the Indo-Malayan and Palearctic fauna of Oxyscelio we recognize 13 species groups. These groups are discussed below to indicate intuitively our perception of the structure within the genus and to serve as an aid in specimen identification.

Oxyscelio carinatus Species Group
Characteristics: Frontal depression flat or nearly so. Hyperoccipital carina complete as a strong ruga, continuous with the anteriormost genal carina, laterally not con- Figure 1. Strict consensus phylogram of four most parsimonious trees for Indo-Malayan and Palearctic species of Oxyscelio using TNT New Technology with Implied Weighting. Search with set initial level = 95, best score = 16.98. Bootstrap support values above 50% indicated above branches, found using TNT new technology search (set initial level = 95). nected with occipital carina. Occipital carina complete or incomplete, but without strong lateral corners. Metascutellum with a pair of subapical dorsal setae, concave dorsally, slightly emarginate apically with rounded posterolateral lobes. T7 in males with acuminate posterolateral corners.
Comments: The carinatus-group is very similar to the cuculli-group, but differs in that the hyperoccipital carina is defined by a ruga and in having a deeper frontal depression. The O. mesiodentis-complex within the cuculli-group has a much more densely setose and differently shaped metascutellum than in the carinatus-group. The general trend towards sculptural reduction in Philippine species (a more pronounced reduction occurs in species from the Maluku Islands of Indonesia) leads one to consider the possibility that the carinatus-group could be weakly sculptured species of the cuculli-group. However, it does not seem proper to lump these groups without additional data supporting this hypothesis. Another possibility exists, that the carinatusgroup could be closely related to the dasymesos-group, as both groups contain species with a setose metascutellum and nearly flat frontal depression. Includes: O. carinatus, O. praecipitis, O. spinosiceps, O. vittae.

Oxyscelio crateris Species Group
Characteristics: Hyperoccipital carina complete, continuous with the anteriormost genal carina, laterally connected with occipital carina by a distinct longitudinal carina or elevation; area between hyperoccipital and occipital carinae slightly sunken and crater-like. Metascutellum about as long as broad, concave dorsally and with little or no median sculpture, rounded apically. T7 in males without posterolateral spines.

Comments:
The crateris-group contains a few species with a crater-like area, between the occipital and hyperoccipital carinae, that is fully outlined by carinae. This area also has distinctive sculpture that is different from that of surrounding areas. Some members of the latitudinis-group may have a similarly weakly concave or partially outlined crater-like area as well, but these species have a very different, broad and strongly sculptured metascutellum.

Oxyscelio crebritas Species Group
Characteristics: Hyperoccipital carina absent or weakly indicated by rugae, laterally not connected with occipital carina. Occipital carina complete or incomplete, but without strong lateral corners. Frons without oblique flange; frontal depression without transverse carinae or grooves in ventral half. Metascutellum medially concave and smooth or with transverse carinae. T7 in males usually with sharp posterolateral corners, rarely with short spines or without spines.
plete, but without strong lateral corners. Metasomal depression setose. T7 in males with acuminate posterolateral corners. Comments: A setose metasomal depression does not occur in any other Asian species of Oxyscelio. The dasymesos-group is otherwise difficult to compare with other Oxyscelio species groups, but it bears some general resemblance to the carinatus-group and crebritas-group.

Oxyscelio florus Species Group
Characteristics: Hyperoccipital carina absent or indicated by poorly defined rugae, laterally not connected with occipital carina. Occipital carina complete or incomplete, but without strong lateral corners. Frons without oblique flange. Metascutellum with longitudinal rugae and without any strong transverse carinae. T2 without longitudinal depressions or strong curved striae.

Comments:
The florus-group contains species that are similar to the crebritasgroup in having a dark radicle and uniformly curved occipital carina, but differ in having a rugose metascutellum and a generally longer metasoma. The latitudinis-group is also similar to this group based on most of the above-mentioned features, but differs in having an occipital carina with strong lateral corners. Includes:O. arvi, O. dermatoglyphes, O. florus, O. jaune, O. regionis.

Oxyscelio fossarum Species Group
Characteristics: Hyperoccipital carina incomplete or indicated by weak rugae. Occipital carina with sharp protruding lateral corners. T2 with long sublateral depressions.

Comments:
The fossarum-group is similar to the foveatus-group, latitudinis-group, and striarum-group, but is distinguished by the T2 depressions that occur in females (and in males of some species). These groups differ in metascutellar form as well, with the fossarum-group having a generally narrower metascutellum. The defining feature of this group can be difficult to discern, but is best verified by finding the strong medial borders of the depressions. Includes:O. aclavae, O. acutiventris, O. cyrtomesos, O. fistulae, O. fodiens, O. fossarum, O. fossularum, O. rugosus, O. zeuctomesos.

Oxyscelio foveatus Species Group
Characteristics: Hyperoccipital carina incomplete or indicated by weak rugae. Occipital carina with strong lateral corners. Ventral frons with oblique flange. Metascutellum tiny and concave, or broad and convex, or elongate with a smooth channel. T2 without longitudinal depressions or strong curved striae.
Comments: The foveatus-group likely is a non-monophyletic group containing species with an oblique facial flange and an occipital carina with protruding lateral corners, but with none of the defining features of some other species groups. Species with longitudinal T2 depressions, but which would otherwise agree with this group, have been placed in the fossarum-group. Other species with an oblique facial flange occur in the carinatus-group, crateris-group, and cuculli-group, but differ strongly from these species. Includes:O. angustinubbin, O. cupularis, O. foveatus, O. greenacus, O. latinubbin, O. nasolabii, O. operimenti.
Comments: The latitudinis-group is essentially negatively defined among Oxyscelio that have strong lateral corners of the occipital carina. The best distinctive feature of this group is the broad, rugose metascutellum of most species, but a few have a narrower metascutellum that more closely approaches that of the fossarum-group. Most members of this group have a metallic green luster, but this is lost in some specimens. Except where noted in species descriptions, color seems to be a highly unreliable character for identification of Oxyscelio. Includes:O. dorsalis, O. latitudinis, O. naraws, O. peludo, O. perpensus.

Comments:
The limae-group contains species from India and Sri Lanka, all with a strongly elevated and anteriorly steep mesoscutum. These species strongly resemble the crebritas-group, but differ in having very short fore wing venation with no sign of a postmarginal vein. Some Australian species, including O. montanus (Dodd) strongly resemble this group, but differ in having a short metascutellum with dorsally protruding posterolateral corners. Tiny but sharp and slightly protruding posterolateral corners of T4 or T5 in females of Oxyscelio limae and Oxyscelio anguli indicate that the limae-group may be the closest relative of an otherwise Australian clade containing O. montanus and Oxyscelio mirellus (Dodd). Includes:O. anguli, O. flaviventris, O. limae.

Oxyscelio noduli Species Group:
Characteristics: Hyperoccipital carina absent or weakly indicated by rugae, laterally not connected with occipital carina. Occipital carina complete, but without strong lateral corners. Frons without oblique flange; frontal depression crossed by many carinae. Metascutellum medially concave and smooth.

Comments:
The noduli-group contains some species that are resemble the latitudinis-group in metasomal length and frontal depression sculpture, but which have a small and medially smooth metascutellum and an occipital carina without strong lateral corners. The latter features are similar to those in the crebritas-group, and therefore these species may be phylogenetically intermediate between that group and the latitudinis-group. Alternatively, they may be reduced apomorphic members of the latitudinis-group.

Oxyscelio ogive Species Group
Characteristics: Hyperoccipital carina incomplete or indicated by rugae. Occipital carina complete medially, but with sharp lateral corners and concave medial sections that meet at a median peak. Comments: Members of the ogive-group superficially resemble the crebritas-group, but differs in the sinuate occipital carina with sharp lateral corners. It differs from the latitudinis group in having a sharp and rounded submedian carina. Includes:O. cavinetrion, O. flabelli, O. labis, O. ogive, O. sinuum.

Oxyscelio striarum Species Group
Characteristics: Hyperoccipital carina incomplete or indicated by rugae. Occipital carina with sharp lateral corners. Metascutellum rugose. T2 (at least) in females with strong curved longitudinal striae submedially that flank a triangular area without striae.

Comments:
The striarum-group is similar to the latitudinis-group, but differs in the strong curved striae of T2 and T3 in females. Males may be difficult to recognize, because those of O. caesitas reveal that they do not possess these strong striae. They do have slightly more distinct sublateral striae of S2 and S3, but these striae are straight and do not distinctly differ from those of other species groups.

Oxyscelio angustifrons
Etymology. Compound noun meaning "narrow nubbin." Refers to the smaller than usual, smooth oblique flange between the antennal foramen and the eye.
Link to distribution map. Metapleuron not crossed by carinae above lower metapleural area, instead with rough irregular sculpture. Female: A4, A5 longer than broad. T1 midlobe with 5 longitudinal carinae. T6 longer than broad. Male: A11 longer than broad. A5 tyloid expanded, sinuate or teardrop-shaped. T7 with short, sharp and protruding posterolateral corners. Oxyscelio asperi is very similar to O. crebritas, but is entirely dark brown in color, lacks straight carinae on the metapleuron, and has at most one straight carina crossing the mesofemoral depression. It is also distinctive within the crebritas-group in having relatively long A4 and A5.
Comments. The strong similarity between O. brevidentis and O. mesiodentis may indicate that they are really one species attacking a wide variety of hosts, but it seems best to verify this possibility before combining these two species. ( Camptoteleia brevinervis Kieffer, 1916: 171, 175 (original description, keyed);Kieffer 1926: 380, 384 (description, keyed Comments. The type material of Camptoteleia brevinervis Kieffer, collected from Mindanao (Butuan) in the Philippines, could not be found after an extensive search of collections known to house Kieffer type material. The neotype of Camptoteleia brevinervis is presently designated to clarify the taxonomic status of the species. It was selected because of its collection locality, its short stigmal vein relative to the postmarginal vein, and for its long flagellomeres. Diagnosis. Both sexes: Mesoscutellum without granulate areas. Metascutellum nearly square, rugose. Female: T1 with a strong anterior horn. T2 and T3 with long, approximated curved striate that for much of their length are not separated by setal pits. Fore wings long enough to reach middle or nearly to apex of T4. Male: A11 longer than broad. T1 midlobe with 6 or more longitudinal carinae. T7 with sharp, protruding posterolateral corners.
Comments. Oxyscelio caesitas is the only known species of Oxyscelio with any metallic blue luster. It is also the only member of the striarum-group in which males are definitively known. These male specimens do not exhibit the distinctive curved T2 and T3 striae found in females. Male. Body length 3.5-4.6 mm (n=20). A5 tyloid: carina-like, not expanded. A11: longer than broad; as long as broad. Median tooth of frontal depression: absent. Median lobe of T1: with 3 longitudinal carinae. Metasomal apex: with acuminate lateral corners.

Oxyscelio capilli
Diagnosis. Both sexes: Middle genal carina subparallel with eye margin. Hyperoccipital carina indicated by rugae. Mesoscutellum without granulate sculpture. Metascutellum concave dorsally, smooth aside from some transverse carinae. Female: A5 broader than long. T1 midlobe with 5 longitudinal carinae. T6 rounded apically. Mesopleuron, along ventral margin of femoral depression, with many fine setae arising from tiny pits, some of these setae arising from the femoral depression itself. Male: A11 longer than broad. T1 midlobe with 3 longitudinal carinae. T7 with short, sharp and protruding posterolateral corners that are very widely separated. Oxyscelio capilli is very similar to O. crebritas, and males of these two species (plus some others) are very difficult to separate due to variation in what constitute reliable diagnostic features for other species. Females of O. capilli can be recognized by the extensive setation along the ventral edge of the femoral depression, which also occurs in O. reflectens and some other species of Oxyscelio.
Etymology. Latin noun, genitive case, meaning "hair." Refers to the unusually extensive setation of the mesopleuron ventrally.
Comments. There are two distinct size fractions of specimens included in this species, but these variants exhibit no other apparent differences. T1 midlobe: obscured by other raised sculpture. T1: with long anterior bulge, reaching metascutellum. T2: with straight longitudinal striae or rugae. T6: longer than broad; as long as broad. Apical flange of T6: exposed apically. Metasomal apex: rounded. Major sculpture of T6: umbilicate-punctate; longitudinally striate or rugose. Microsculpture of T6: absent.
Diagnosis. Both sexes: Middle genal carina angled towards genal carina dorsally. Metascutellum flat but with one or more transverse carinae. Female: A4, A5 broader than long. T1 midlobe with well-developed anterior horn. Male: A11 broader than long. A5 tyloid expanded, sinuate or teardrop-shaped. T7 with short, sharp and protruding posterolateral corners. Oxyscelio capitis is very similar to O. reflectens, but is smaller-bodied, with a relatively larger head, a flat metascutellum, a T1 horn in females, and acuminate posterolateral corners on T7 in males.
Etymology. Latin noun, genitive case, meaning "head." Emphasizes the large head of this species.
Link to distribution map. Diagnosis. Both sexes: Occipital carina complete as a distinct carina, but medial portions concave and meeting at a peak. Mesoscutellum with a few flattened longitudinal carinae. Netrion concave anteriorly. Metascutellum tiny, dorsally concave. Female: A4, A5 broader than long. Fore wings long enough to reach middle or apex of T5. T1 midlobe without anterior horn. Male: A11 longer than broad. Fore wings long enough to reach middle of T5. T7 with short, sharp and protruding posterolateral corners. Oxyscelio cavinetrion is very similar to O. flavipennis, but has a shorter metasoma which lacks the anterior T1 horn in females, and a differently shaped metascutellum.
Etymology. Compound noun intended to mean "concave netrion." Link to distribution map.  Comments. The lateral propodeal carinae exhibit strong variation in O. ceylonensis, being narrowly separated and subparallel in some specimens (especially those from Nepal) and strongly divergent in others. This variation did not prove consistent enough to serve as a convincing feature for species separation. Description. Female. Body length 3.6-3.9 mm (n=20).
Interantennal process: not elongate. Median longitudinal elevation in frontal depression: absent. Diagnosis. Both sexes: Middle genal carina subparallel with eye margin. Hyperoccipital carina indicated by rugae. Mesoscutellum without granulate sculpture. Metascutellum concave dorsally, smooth aside from some transverse carinae. Female: A5 broader than long. T1 midlobe with a small anterior horn obscuring the longitudinal carinae. T6 rounded apically but longer than broad. Male: A11 longer than broad. T1 midlobe with 3 longitudinal carinae. T7 with short, sharp and protruding posterolateral corners that are curved and not widely separated. Oxyscelio codae is distinguished from most members of the crebritas-group in having a long T6 in females. Males of O. codae are especially difficult to distinguish from those of O. capilli, another species from Sulawesi. They differ in that O. codae has a more elongate, tapering metasoma in which T7 is rounded apically, with acuminate apical projections angled slightly towards one another. In O. capilli, the metasoma is usually shorter and broader, with T7 more truncate apically, and with the inner margins of the acuminate apical projections being at right angles and not angled towards one another.

Oxyscelio consobrinus
Comments. The weak sculpture of O. consobrinus, which can resemble melted plastic, is a distinctive trait common to many Philippine species. Recently collected specimens of this species have indicated that this was not an artefact of any unusual collecting or preservation methods.
The type material of Camptoteleia bifurcata Kieffer, collected from Mindanao (Butuan) in the Philippines, could not be found after an extensive search of collections known to house Kieffer type material. The neotype of Camptoteleia bifurcata is presently designated to clarify the taxonomic status of the species. It was selected because of its collection locality, and because it resembles Kieffer's (1916) description in having a shiny mesosoma. In assigning a neotype for Camptoteleia bifurcata, we presumed that Kieffer (1916) was mistaken in his description of the fore wing venation. The "forked submarginal vein" seems to refer to the strongly tilted venation in which only the postmarginal vein closely approaches the anterior wing margin (the marginal vein is distant from the wing margin). This state is variable in many species of Oxyscelio, and therefore likely only indicates that a postmarginal vein is present. Females of O. consobrinus can have a variably emarginate metascutellum, with extreme cases seeming bifurcate. This, and the description of the thorax as shiny, leads us to conclude that the lost type series of C. bifurcata represented specimens of O. consobrinus corresponding to the above criteria, which proved to fit within intraspecifc variation.

Oxyscelio cyrtomesos
Diagnosis. Both sexes: Mesoscutellum laterally granulate. Metascutellum long and tongue-shaped. Propodeum forming a nearly complete arch over the base of T1, but with a narrow break along middle of the arch. Female: A4, A5 broader than long. T1 midlobe with 6-7 longitudinal carinae. T2 with sublateral depressions. T6 strongly tapering to a narrow point. Male: A11 slightly broader than long. T1 midlobe with 5 longitudinal carinae. T7 with sharp, protruding posterolateral corners. Oxyscelio cyrtomesos is very similar to O. zeuctomesos, but differs in development of the propodeal arch.
Etymology. Compound noun based on Greek, meaning "convex middle." Refers to the way that the propodeum arches medially over the base of T1.
Link to distribution map.  228701-228702, 228704, 228732, 228739, 240920, 247815-247816, 248922 (OSUC);OSUC 228699, 247837, 247933, 247957, 257039, 257041, 257064, 257066, 257068, 257071, 257079 (ROME) Comments. Oxyscelio dermatoglyphes is part of a species complex occurring also in Japan and Korea. This complex can be characterized by the elongate body, dark antennal radicle, weak occipital carina without protruding lateral corners, subrectangular flat metascutellum (reduced in males), and very strong T1 horn in females. ( Comments. Oxyscelio fistulae is unusual in having an elongate interantennal process. This state is also found in a species from New Guinea and New Britain, but the process is differently shaped in that species. longer than broad, tapering to a rounded apex. Oxyscelio florus differs from O. mollitia, a similar Japanese species, in sculpture, metasomal length, and in having a much stronger T1 horn in females. Especially, the mesofemoral depression lacks a row of foveae along its anterior limit. Oxyscelio florus is very similar to the Taiwanese species O. dermatoglyphes as well, especially in having extra carinae parallel to the submedian carina; these species differ in that O. dermatoglyphes has no median carina on the mesoscutellum, only a very weak and indistinct one on the mesoscutum, and has a much shorter metasoma in females (fore wing long enough to reach T6 or apex of T5).
Etymology. Latin noun, genitive case, meaning "that which is between." Refers to its similarity to several other species in the cuculli-group.
Comments. Oxyscelio intermedietas seems to be a small-bodied relative of O. granorum. It otherwise strongly resembles several other species nearer to its own size, including O. cuculli. It differs from these species in several subtle ways, including surface sculpture, shallowness of the frontal depression, and the very narrow anterior separation of the lateral propodeal carinae. While the possibility remains that O. intermedietas is just a small form of O. granorum, these features proved convincing enough to allow recognition of it as a separate species.  T1 midlobe: obscured by other raised sculpture. T1: with long anterior bulge, reaching metascutellum. T2: with straight longitudinal striae or rugae. T6: longer than broad. Apical flange of T6: exposed apically. Metasomal apex: rounded. Major sculpture of T6: umbilicate-punctate. Microsculpture of T6: granulate.

Oxyscelio jaune
Male. Unknown. Diagnosis. Female: Upper frons without additional carinae near the strong submedian carina. Hyperoccipital carina indicated by rugae. Mesoscutellum without granulate sculpture. Mesofemoral depression crossed by 3 carinae below speculum. Metascutellum subrectangular, with scattered weak rugae. T1 midlobe with long anterior bulge. T2 without sublateral depressions or curved striae. T6 longer than broad, tapering to a rounded apex. Oxyscelio jaune is similar to O. longiventris and O. regionis, in that they have a dark antennal radicle, long body, and very strong T1 horn in females. The color of the holotype is distinctive but may not be constant within the species, meaning that the small number of carinae (3)  Radicle color: darker than scape. Scape color: Yellowish. A4: longer than broad; as long as broad. A5: broader than long. Antennal club: formed, segments compact.
Diagnosis. Both sexes: Middle genal carina subparallel with eye margin. Hyperoccipital carina indicated by rugae. Mesoscutellum with granulate sculpture laterally but not medially. Metascutellum concave dorsally, smooth aside from some transverse carinae. Female: A5 broader than long. T1 midlobe with 5 longitudinal carinae. T6 acuminate apically. Male: A11 longer than broad. T1 midlobe with 4 longitudinal carinae. T7 with short, sharp and protruding posterolateral corners. Oxyscelio jugi can be distinguished from similar species by its strong mesoscutal sculpture and laterally granulate mesoscutellum. In males, the medial pair of longitudinal carinae of the T1 midlobe are usually curved to closely approach one another anteriorly.
Etymology. Latin noun, genitive case, meaning "ridge." Refers to the extra longitudinal carinae found on the mesoscutum.
Comments. Only a few females of O. jugi are known, and these appear to be variable in sculpture and metasomal length. This variation is attributed to differences in body size. Comments. The type material of Camptoteleia flavipennis Kieffer, collected from Laguna, Los Baños, in the Philippines, could not be found after an extensive search of collections known to house Kieffer type material. The neotype of Camptoteleia flavipennis is presently designated to clarify the taxonomic status of the species. It was selected because it was collected in the Philippines and resembles Kieffer's (1913b) description in having a short metasoma and smooth frontal depression. We presume that Kieffer was mistaken when he mentioned that T7 lacked armature. Radicle color: darker than scape. Scape color: Yellowish. A4: longer than broad. A5: longer than broad; as long as broad. Antennal club: formed, segments compact.

Oxyscelio kramatos
Interantennal process: not elongate. Median longitudinal elevation in frontal depression: absent.  Diagnosis. Both sexes: Frons without elevation between antennal foramen and eye. Hyperoccipital carina present, continuous with vague anterior genal carina. Gena with some strong sculpture, at least one strong ruga along middle. Mesoscutellum strongly granulate. Metascutellum subrectangular, weakly emarginate. Metasomal depression elongate, without extensive sculpture; lateral propodeal carinae narrowly separated anteriorly. Female: T1 midlobe with 5 longitudinal carinae. T6 rounded apically. Male: T1 midlobe with 4 longitudinal carinae. T7 with posterolateral tubercles. Oxyscelio kramatos shares several characters with O. vadorum, but has stronger genal sculpture and little or no postmarginal vein. It also strongly resembles the craterisgroup in having a vaguely crater-like occiput, but this is not as laterally well-defined as in that group.
Etymology. Noun based on Greek, meaning "mixture." Meant to indicate the morphological similarity to both O. vadorum and the crateris-group.
Interantennal process: not elongate. Median longitudinal elevation in frontal depression: absent. Male. Unknown. Diagnosis. Female: A4, A5 broader than long. Frontal depression crossed by many carinae. Submedian carina undefined. Hyperoccipital carina indicated by rugae; occipital carina without distinct lateral corners. Mesoscutellum without granulate sculpture. Metascutellum tiny, narrow. T1 midlobe with 5 longitudinal carinae. Fore wings long enough to reach apex of T5. T6 broader than long. Oxyscelio lacunae is an unusual species that resembles O. crebritas in some ways, but has an indistinct submedian carina, tiny metascutellum, and irregular metapleural sculpture. The overall body color is dark, with the scape, wings, and coxae brownish.
Male. Unknown. Diagnosis. Female: Antennal club formed. A4, A5 longer than broad. Face with broad oblique expanded flange between antennal foramen and eye. Metascutellum longer than broad, with central smooth channel. Oxyscelio latinubbin is similar to O. aclavae, in having a long metascutellum with a median channel and a propodeum that forms a short arch above the anterior part of T1. It differs in having a swollen, compact antennal club.
Etymology. Compound noun intended to mean "broad nubbin." Refers to the unusual broad oblique flange between the antennal foramen and eye.

Oxyscelio mesiodentis
Comments. The vertical elevation between the antennal foramen and eye in O. mesiodentis and similar species bears a granulate patch. Some males from Borneo and Laos were intermediate in these features, but could not be associated with females. These were left unassigned to species until corresponding females can be found. While O. mesiodentis and O. brevidentis may represent variant forms of the same species, the differences in metatarsomere length indicate that they are kept separate pending new information. Diagnosis. Both sexes: Upper frons without additional carinae dorsal to submedian carina. Hyperoccipital carina indicated by rugae. Mesoscutellum without granulate sculpture. Mesofemoral depression crossed by more than 3 carinae below speculum. Mesopleuron along anteroventral edge of femoral depression with rows of foveae. Female: Metascutellum subrectangular, with scattered rugae. T1 midlobe with weak anterior bulge. T2 without sublateral depressions or curved striae. Fore wing long enough to reach middle of T5. T6 longer than broad, tapering to a rounded apex. Male: Flagellomeres longer than broad. T1 midlobe with 5 longitudinal carinae. T7 with sharp, protruding posterolateral corners. Oxyscelio mollitia is similar to O. florus, another Palearctic species. It differs in having one or more rows of foveae along the anterior limit of the femoral depression, a transversely carinate metascutellum, a usually shorter anterior horn on the T1 midlobe, and shorter metasoma in females (fore wing long enough to reach middle of T5).

Oxyscelio mollitia
Etymology. Latin noun in apposition, meaning "flexibility." Refers to the tendency of various body parts to buckle when specimens of this species are dried.
Comments. Oxyscelio naraws exhibits a wide range of variation in the lengths of A4, A5, the metascutellum, and the metasoma in females. This may indicate that it represents a suite of very similar species, but no convincing consistent features were found to support any separation. Male. Unknown. Diagnosis. Female: A4, A5 not longer than broad. Radicle darker than scape. Face with long expanded flange between antennal foramen and eye. Mesoscutum and mesoscutellum without median carina. Metascutellum broad but smooth medially. T1 midlobe elevated but with 3 strong carinae. T2 without sublateral depressions or curved striae.T6 longer than broad.

Oxyscelio nasolabii
Etymology. Latin noun, genitive case, intended to mean "nasolabial fold." Refers to the similarity of the ridge between the antennal foramen and eye to the nasolabial fold of the human face.
Link to distribution map. Comments. Oxyscelio nasolabii is unusual within its genus in having a medially smooth metascutellum but lacking median carinae of the mesoscutum and mesoscutellum. The coxae of the holotype are darker than other parts of the legs, but this character is variable in Asian species. culpture of mesoscutellum laterally: absent; granulate. Mesoscutellar apex: convex or straight. Setae along anterior limit of femoral depression: arising from rows of foveae. Number of carinae crossing speculum above femoral depression: 3. Number of carinae crossing femoral depression: more than 5. Mesepimeral sulcus pits: more than 5. Metascutellum dorsally: flat. Metascutellar sculpture dorsally: with scattered rugae. Median carina of metascutellum: absent or branched. Metascutellar setae: with many dorsal setae. Metascutellar apex: convex or straight. Metapleuron above ventral metapleural area: crossed by carinae. Metasomal depression setae: absent. Lateral propodeal carinae anteromedially: strongly diverging. Anterior areoles of metasomal depression: absent. Anterior longitudinal carinae in metasomal depression: absent. Lateral propodeal areas: separated medially. Postmarginal vein: present. Fore wing apex: reaching middle of T5. T1 midlobe: obscured by other raised sculpture. T1: with small rounded anterior bulge, not reaching metascutellum. T2: with straight longitudinal striae or rugae. T6: longer than broad. Apical flange of T6: not exposed apically. Metasomal apex: tapering to a sharp point. Major sculpture of T6: umbilicate-punctate; longitudinally striate or rugose. Microsculpture of T6: granulate.
Diagnosis. Both sexes: A4 longer than broad, A5 about as long as broad. Frontal depression crossed by many carinae. Mesoscutellum strongly granulate. Metascutellum dorsally setose. Female: Metascutellum fingernail-shaped, rugose. T1 with strong anterior horn. Fore wings long enough to reach middle of T5. T6 strongly narrowing towards nearly acuminate apex. Male: T1 midlobe with 5 longitudinal carinae. T7 with sharp, protruding posterolateral corners. The metascutellar setae of O. peludo are easily overlooked in dorsal view, but are more apparent from an oblique or lateral view. They are present in males as well, making males of O. peludo much more easily recognizable than those of most other species. Among species with a bare metascutellum, O. naraws very strongly resembles O. peludo.
Link to distribution map.  Kononova: Kononova and Kozlov 2008: 190, 192 (description, keyed). Comments. The mesoscutal median carina is less visible in our figure (Fig. 374) than in that of Kononova and Fursov (2007: Fig. 9.1), but this is because the carina is relatively weak and rounded, becoming less visible under diffused lighting. Dicroteleia rugosa Kieffer, 1908: 92 (original description);Kieffer 1926: 387, 388 (description, keyed). Chromoteleia (Oxyscelio)  Comments. Oxyscelio rugosus possesses the characters of a few different distinctive species groups, and therefore is difficult to place based on the strength of the single existing specimen. Because of the T2 sublateral depression medially bordered by a strong carina, O. rugosus is provisionally placed in the fossarum-group. Psilanteris spinosiceps Kieffer, 1916: 177, 178 (original description, keyed). Camptoteleia spinosiceps (Kieffer): Kieffer 1926: 379, 386 (generic transfer, description, keyed Comments. Oxyscelio vadorum superficially resembles many weakly sculptured species outside the cuculli-group. However, it has the elongate, sculptured metasomal depression found in O. convergens and similar species. Even though only a single, damaged specimen is known, this species is described because of its many unusual character states, including the chiefly granulate gena.