Systematics of Old World Odontacolus Kieffer s.l. (Hymenoptera, Platygastridae s.l.): parasitoids of spider eggs

Abstract The genera Odontacolus Kieffer and Cyphacolus Priesner are among the most distinctive platygastroid wasps because of their laterally compressed metasomal horn; however, their generic status has remained unclear. We present a morphological phylogenetic analysis comprising all 38 Old World and four Neotropical Odontacolus species and 13 Cyphacolus species, which demonstrates that the latter is monophyletic but nested within a somewhat poorly resolved Odontacolus. Based on these results Cyphacolus syn. n. is placed as a junior synonym of Odontacolus which is here redefined. The taxonomy of Old World Odontacolus s.str. is revised; the previously known species Odontacolus longiceps Kieffer (Seychelles), Odontacolus markadicus Veenakumari (India), Odontacolus spinosus (Dodd) (Australia) and Odontacolus hackeri (Dodd) (Australia) are re-described, and 32 new species are described: Odontacolus africanus Valerio & Austin sp. n. (Congo, Guinea, Kenya, Madagascar, Mozambique, South Africa, Uganda, Zimbabwe), Odontacolus aldrovandii Valerio & Austin sp. n. (Nepal), Odontacolus anningae Valerio & Austin sp. n. (Cameroon), Odontacolus australiensis Valerio & Austin sp. n. (Australia), Odontacolus baeri Valerio & Austin sp. n. (Australia), Odontacolus berryae Valerio & Austin sp. n. (Australia, New Zealand, Norfolk Island), Odontacolus bosei Valerio & Austin sp. n. (India, Malaysia, Sri Lanka), Odontacolus cardaleae Valerio & Austin sp. n. (Australia), Odontacolus darwini Valerio & Austin sp. n. (Thailand), Odontacolus dayi Valerio & Austin sp. n. (Indonesia), Odontacolus gallowayi Valerio & Austin sp. n. (Australia), Odontacolus gentingensis Valerio & Austin sp. n. (Malaysia), Odontacolus guineensis Valerio & Austin sp. n. (Guinea), Odontacolus harveyi Valerio & Austin sp. n. (Australia), Odontacolus heratyi Valerio & Austin sp. n. (Fiji), Odontacolus heydoni Valerio & Austin sp. n. (Malaysia, Thailand), Odontacolus irwini Valerio & Austin sp. n. (Fiji), Odontacolus jacksonae Valerio & Austin sp. n. (Cameroon, Guinea, Madagascar), Odontacolus kiau Valerio & Austin sp. n. (Papua New Guinea), Odontacolus lamarcki Valerio & Austin sp. n. (Thailand), Odontacolus madagascarensis Valerio & Austin sp. n. (Madagascar), Odontacolus mayri Valerio & Austin sp. n. (Indonesia, Thailand), Odontacolus mot Valerio & Austin sp. n. (India), Odontacolus noyesi Valerio & Austin sp. n. (India, Indonesia), Odontacolus pintoi Valerio & Austin sp. n. (Australia, New Zealand, Norfolk Island), Odontacolus schlingeri Valerio & Austin sp. n. (Fiji), Odontacolus sharkeyi Valerio & Austin sp. n. (Thailand), Odontacolus veroae Valerio & Austin sp. n. (Fiji), Odontacolus wallacei Valerio & Austin sp. n. (Australia, Indonesia, Malawi, Papua New Guinea), Odontacolus whitfieldi Valerio & Austin sp. n. (China, India, Indonesia, Sulawesi, Malaysia, Thailand, Vietnam), Odontacolus zborowskii Valerio & Austin sp. n. (Australia), and Odontacolus zimi Valerio & Austin sp. n. (Madagascar). In addition, all species of Cyphacolus are here transferred to Odontacolus: Odontacolus asheri (Valerio, Masner & Austin) comb. n. (Sri Lanka), Odontacolus axfordi (Valerio, Masner & Austin) comb. n. (Australia), Odontacolus bhowaliensis (Mani & Mukerjee) comb. n. (India), Odontacolus bouceki (Austin & Iqbal) comb. n. (Australia), Odontacolus copelandi (Valerio, Masner & Austin) comb. n. (Kenya, Nigeria, Zimbabwe, Thailand), Odontacolus diazae (Valerio, Masner & Austin) comb. n. (Kenya), Odontacolus harteni (Valerio, Masner & Austin) comb. n. (Yemen, Ivory Coast, Paskistan), Odontacolus jenningsi (Valerio, Masner & Austin) comb. n. (Australia), Odontacolus leblanci (Valerio, Masner & Austin) comb. n. (Guinea), Odontacolus lucianae (Valerio, Masner & Austin) comb. n. (Ivory Coast, Madagascar, South Africa, Swaziland, Zimbabwe), Odontacolus normani (Valerio, Masner & Austin) comb. n. (India, United Arab Emirates), Odontacolus sallyae (Valerio, Masner & Austin) comb. n. (Australia), Odontacolus tessae (Valerio, Masner & Austin) comb. n. (Australia), Odontacolus tullyae (Valerio, Masner & Austin) comb. n. (Australia), Odontacolus veniprivus (Priesner) comb. n. (Egypt), and Odontacolus watshami (Valerio, Masner & Austin) comb. n. (Africa, Madagascar). Two species of Odontacolus are transferred to the genus Idris Förster: Idris longispinosus (Girault) comb. n. and Idris amoenus (Kononova) comb. n., and Odontacolus doddi Austin syn. n. is placed as a junior synonym of Odontacolus spinosus (Dodd). Odontacolus markadicus, previously only known from India, is here recorded from Brunei, Malaysia, Sri Lanka, Thailand and Vietnam. The relationships, distribution and biology of Odontacolus are discussed, and a key is provided to identify all species.


introduction
The genera Odontacolus Kieffer and Cyphacolus Priesner are easily recognized from other platygastroid wasps by the obvious lateral compression of the T1 horn in the female. Previously considered to be relatively rare based on material available in collections, recent intensive collecting using Malaise and yellow-pan traps has revealed that some species of Odontacolus are moderately common. The likely phylogenetic affinities of the genera based on morphology are somewhat equivocal (Austin and Iqbal 2005;Valerio et al. 2010); however, a preliminary molecular analysis (Carey et al. 2006) has confirmed that Odontacolus belongs to the Baeini and is related to Baeus Haliday and Idris Förster s.l., the latter representing a huge assemblage, possibly numbering in excess of a thousand species. Host data are scant, but available records confirm that, like all members of the Baeini, both genera are endoparasitoids of spider eggs (e.g. Masner 1976;Austin 1984Austin , 1985Galloway and Austin 1984;Stevens and Austin 2007;Valerio et al. 2010).
The taxonomic status of both genera has largely been stable since their description, in part because they have received little attention. Odontacolus was originally described by Kieffer (1910a) for O. longiceps from the Seychelle Islands with only a few species described since then. Cyphacolus was described by Priener (1951) for C. veniprivus from Egypt, but recently Valerio et al. (2010) have completely revised the genus and recognized 16 species distributed from Africa, the Middle East, India, south-east Asia to Australia. Cyphacolus + Odontacolus clearly form a monophyletic group within the Baeini, based on the unique shape of the T1 horn, the presence of large blunt spines on the propodeum, and the subpedunculate metasoma. As discussed by Austin and Iqbal (2005), the shape of the horn is apparently linked to the functional mechanics of the ovipositor system, as the ovipositor is retracted within the metasoma and is curled around in the curved head of the horn so that it forms an elongate, U-shape (Austin 1983;Valerio et al. 2010) (Fig. 1C). However, although Cyphacolus is putatively monophyletic based on the fore wing being spoon-shaped and contoured to the convex surface of metasoma (subelytriform), with a dark infuscate patch at the fore wing margin, and the distal venation being absent, Odontacolus does not possess any obvious synapomorphies, and thus may be paraphyletic with respect to the former genus.
The current study builds on our recent taxonomic treatment of Cyphacolus (Valerio et al. 2010) by undertaking a phylogenetic analysis to examine the relationships between the two genera and their component species using a larger morphological dataset, and by completing a taxonomic revision of Odontacolus s.l. for the Old World, describing a large number of new species, and documenting their biology and distribution.
The contributions of the individual authors are as follows; A.A. Valerio: character definition, character development, character coding, imaging, species concept development; key development, capture of specimen data, phylogenetic analysis and manu- lecting data for specimens are available at Hymenoptera On-Line (http://hol.osu.ed), the Global Biodiversity Information Facility (http://www.gbif.org) and in the Darwin Core Archive 27 file available as a supplement to this paper.
Appendix I list terms associated with identifiers in the Hymenoptera Anatomy Ontology (http://glossary.hymao.org). Identifiers in the format HAO_XXXXXXX represent concepts in the HAO in August 2012 and are provided to enable readers to confirm their understanding of the concepts being referenced. The identifier can also be used as a URI (universal resource identifier) by appending the identifier to 'http://purl.obolibrary.org/obo/' (e.g. http://purl.obolibrary.org/obo/HAO_0000124), which resolves to the HAO's community-based resource that includes additional images, notes, and other metadata. The external hyperlinks are explicitly cited in the endnotes so that users of the printed version of the paper have access to the same resources. As possible the external information conforms to standards developed and maintained through the organization Biodiversity Information Standards (Taxonomic Database Working Group). All new species have been registered with Zoobank (http://www.zoobank.org), and other taxonomic names, where appropriate, have been retrospectively registered.
Illustrations and data citation. Images were taken with a JVC 3 CCD camera (model KY-575U) attached to a Leica Z16 APO with a Planapo 1.0× objective alone or in combination with a 2× magnifying lens. Specimens were illuminated with a 4 channel LED dome light from Advanced Illumination. Figures were produced using Auto-Montage Pro versions 5.10 and post-processed with Adobe Photoshop CS5. All images are archived at Morphbank 28 and in Specimage 29 (the image database at The Ohio State University).
Phylogenetic analysis. For the phylogenetic analysis Idris s. str. (Idris_sp. 1, OSUC 233340, and Idris_sp. 2, OSUC 228297), Idris (Ceratobaeus) (Idris_sp. 3, OSUC 190640), and I. floris (Kononova & Fursov) were used as outgroups; I. floris was used to root the tree. The ingroup comprised 1) all 36 species of Old World Odontacolus s.str. treated herein, 2) 13 of the 16 species of Cyphacolus treated by Valerio et al. (2011), and 3) six Neotropical species: Odontacolus flavissimus Megyaszai and O. szaboi Megyaszai and four undescribed species: Od_sp. 4 (OSUC 246525), Od_sp. 5 (OSUC 246524), Od_sp. 6 (OSUC 253001) and Od_sp. 7 (OSUC 233343), A total of 41 morphological characters were scored for the 55 ingroup and four outgroup taxa (see Appendices II and III). Only character 9 (shape of the medial area of the vertex) was uninformative under the parsimony criterion. Characters 37 (ovipositor shape) and 41 (number of antennal segments) where coded directly from speci-mens for some species, and from the literature for others (e.g. Masner 1976;Galloway and Austin 1984;Valerio et al. 2010). The analysis was performed using the software TNT -Tree analysis using New Technology ver. 1.1 (Goloboff et al. 2008) under the command 'xmult'and all characters were treated as unordered. In addition to the Maximum Parsimony analysis an implied weighting analysis was performed (under options K=3, 5 and 7) to explore possible structure of the data matrix with the resulting tree(s). The trees obtained under TNT were edited using Adobe Illustrator CS5.

Phylogeny
Maximum parsimony analysis resulted in 10,000 trees of 348 steps (CI= 0.18, RI= 0.58). The strict consensus tree is largely unresolved except for a monophyletic group containing all 13 Cyphacolus species (tree not shown). The dataset clearly is very homoplasious, and so to explore possible relationships further, we compared the 50% majority rule consensus (MR) tree ( Fig. 2) with that obtained from the implied weighting (IW, with K=3) analysis (4 trees, score=20.52) (Fig. 3). In both trees Odontacolus+Cyphacolus are monophyletic with respect to the outgroup taxa. In the MR tree three Odontacolus species from Fiji (O. veroae, O. schlingeri, O. irwini) form a grade at the base of the tree; there is a single large clade (A) with relative low support (65%) which contains Cyphacolus and 19 Odontacolus species, while the remaining taxa form a polytomy. In the IW tree the taxa in clade B largely correspond to clade A in the MR tree but with some exceptions, and Clade C encloses Cyphacolus plus O. bouceki, O. gallowayi, O. harveyi and O. zborowskii. The six Neotropical species occur in different places between the two trees but always with some taxa occurring among the Old World Odontacolus (i.e. O. sp. 7). In the MR tree four species (O. flavissimus, O. szaboi, O. sp. 4, O. sp. 5) are found in clade A but as a paraphyletic grade, while in the IW tree these same species are found througthout clade B, and O. sp. 6 and O. sp. 7 are found basally within Clade B next to O. sp.4. Further, in the IW tree the Fijian species, which are among very few taxa that lack any traces of notauli, form a monophyletic group within clade B with O. kiau and O. baeri. This placement of these species would seem to make more sense biogeographically, given they are endemic to an isolated oceanic island, rather than three of them being at the base of the Odontacolus+Cyphacolus clade (as in the MR tree).
In both trees the same group of Australian species is sister to Cyphacolus, though they differ in their specific relationships. In the MR Cyphacolus seems to be the sister clade to the species O. australiensis, O. berryae, O. harveyi, O. zborowskii and O. gallowayi, while Cyphacolus and O. australiensis, O. berryae are place at the base of the Cyphacolus + Odontacolus clade. Interestingly, these five species have a short stigmal vein (r-rs), the head is moderately short and strongly narrowed towards mouth, and they have somewhat setose eyes. These characters are also shared between other Odontacolus and Cyphacolus species. One of the main characters previously used to differentiate Cyphacolus from Odontacolus was its convex distal fore wing surface and conspicuous constriction at the base of the wing. However, it is now evident that this wing shape is also found in some Odontacolus species (e.g. O. bosei, O. veroae) although not as pronounced.  Additionally, the topology of the IW trees for K=5 and K=7 were similar to that for K=3, with Cyphacolus always being nested within Odontacolus, however the relationships among the most basal taxa for the clade Odontacolus+Cyphacolus clade varied among the trees. Also, the number of Odontacolus species placed as sister species to the Cyphacolus clade changes.
Mesosoma. Mesoscutum wider than long, usually flat or virtually so, sometimes dorsally convex. Notauli usually present as distinct grooves reaching no more than about half way to anterior margin of mesoscutum, sometimes virtually absent or hidden by coarse longitudinal sculpture. Mesoscutellum either flat or transverse, with posterior margin usually straight medially, or dorsally convex and semicircular or oval in shape. Propodeum with pair of broad, elongate spines which are blunt or truncate apically and flank the T1 horn.
Wings. Macropterous, never brachypterous. Fore wing narrow basally, broad in apical half, sometimes fore margin sinuate and surface of apical half convex (spoonshaped) and molded to convex dorsal surface of metasoma (i.e. subelytriform). Fore wing venation with tubular submarginal (Sc+R) and stigmal veins (r-rs), marginal vien (C+R) very short, postmarginal vein (R1) very short or absent; in O. veniprivus species group venation lacking except for incomplete submarginal vein, with pronounced infuscate patch at position of marginal (C+R) and stigmal veins (r-rs). Fore wing color varying from hyaline to having dark infuscate bands.
Metasoma in dorsal view subpedunculate. T1 square or longitudinally slightly longer than wide (rarely more transverse), with parallel or slightly curved lateral margins. Metasoma widest in posterior half; in lateral view dorsal surface slightly to strongly convex. T1 with large, laterally compressed hornlike process (i.e. ellipsoidal in cross-section) which reaches to level of posterior mesocutellum or higher. T3 the largest tergite, slightly longer than T2, sometimes subequal in length with T2. Ovipositor at least 1.5× length of metasoma, with shaft curled back on itself within rounded head of the T1 horn. Gonoplacs elongate, approximately 0.75× length of metasoma.
Male. Antenna short, 11-segmented, often appearing to be 9-or 10-segmented as distal antenomeres are closely joined, distal antenna becoming progressively broader so as to be subclavate. Metasomal horn absent, but anterior part of T1 inflected upwards.
Diagnosis. Odontacolus s.l. can be distinguished from all other genera of Platygastroidea by three unique characters: T1 of the female has a laterally compressed T1 horn (elliptical in cross-section); the ovipositor is retracted within the metasoma and curled around within the curved head of the T1 horn so that it forms an elongate, U-shape (compared to Idris species where it is straight or slightly curved, see Figs 1A-B); the propodeum in both sexes has a pair of spinelike flanges that (in the female) flank the T1 horn. In addition, all species are macropterous, never brachypterous, and the metasoma is subpedunculate in shape. Distribution and regional diversity. Odontacolus is clearly more widespread than previously thought and, in the Old World, is found throughout Africa south of the Sahara, the Middle East (O. veniprivus group only; see Valerio et al. 2010), India, Sri Lanka, Nepal, south-east Asia (with one species extending north into China), and Australasia extending east in the Pacific to Fiji (Fig. 4, link to distribution map 30 ). Australasia is the most species rich region (19 spp., four of which are endemic to Fiji), followed by India/south-east Asia (14 spp.) and Africa/Madagascar (11 spp.), the remainder of species have a broader distributions across regions.
Biology. Like other members of the Baeini, Odontacolus species are endoparasitoids of spider eggs. However, of the 52 species here recorded from the Old World only five species have been reared. Veenakumari and Mohanraj (2011) record O. markadicus from eggs of an unknown salticid spider in southern India; O. pintoi has been reared from Clubiona cycladata Simon from under tree bark in South Australia (recorded as Odontacolus sp. in Austin (1984Austin ( , 1985 (specimens in WINC); O. berryae has been reared from eggs of the salticid Trite planiceps Simon (specimens in LUNZ); O. whitfieldi has been reared from an unknown spider from China (specimens in UCRC), while O. harteni has also been reared from an unknown spider from Pakistan (specimens in BMNH) (Valerio et al. 2010). Although hosts are only from two spider families, Clubionidae and Salticidae, it is likely that Odontacolus species attack a much greater range of host spiders given the number of species and diversity of habitats from which they have been collected (e.g. rainforest, tropic dryforest, eucalypt forest, coastal dunes, mangroves, grasslands, etc.; see Appendix IV).
Although nothing is known about the ovipositional behavior of Odontacolus species, we predict that females employ their long ovipositor to parasitize host eggs from outside the egg sac, by pushing the ovipositor through the silk wall, in a similar way as has been described for Idris (Ceratobaeus) species (Austin 1984(Austin , 1985 that also have a T1 horn, albeit of a different shape (Figs 1B, C). Although documented for only a few species, the stategy of Baeus species (Stevens and Austin 2007), Idris s.str. and other baeines that lack a T1 horn, is to burrow through the silk wall of the egg sac and oviposit into eggs while in direct contact with them.
Species excluded from Odontacolus. The previously described species Odontacolus longispinosus Girault (Fig. 1) and O. amoenus Kononova are here transferred to the genus Idris Förster and should therefore be treated as new combinations. We were unable to borrow the holotype of O. amoenus but one of us (NFJ) during a recent visit to the UASK was able to examine a female paratype and confirm that the species does indeed belong to Idris. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: largely smooth ventrally, sparsely granulate ventrally. Surface of torular triangle: slightly bulging. Development of central keel on frons: present, elongate (equal to or greater than1/3× height of frons), but not reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: granulose throughout. Sculpture of malar space: granulose throughout; with fan-like striae, striae not extending into antennal scrobe. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: large. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: granulate. Sculpture of occipital carina: largely simple, at most with sparse weak crenulae medially. Distance from occipital carina to orbital carina: at least 2× width of occipital carina; slightly greater than width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weak, small granulae. Sculpture of gena: granulose.

Key to the females of Old World species of Odontacolus Kieffer
Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), welldefined foveae. Sculpture of pronotal lateral area: largely smooth, dorsal margin with dense, weak punctulae. Netrion: present, smooth, linear. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: anterior half coriaceous, otherwise densely granulose. Sculpture of mesoscutellum: with weak, fine, granulate sculpture. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: densely, finely rugulose. Shape of propodeal anterior spines: elongate, narrow, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: weakly coriaceous. Sculpture of upper 1/4 of mesopleuron: with dense, well-defined longitudinally costate sculpture reaching just half of its width. Metapleuron sculpture: largely smooth except lower half with longitudinal carinae.
Wings. Male. Body length: 1.36-1.60 mm (n=3). Body color: head, mesoscutum, edges of T1-T2, T4-T6 dark brown, reminder of terga light honey yellow as remainder of mesosoma. Sculpture of antennal scrobe: upper half with transverse carinae mixed with granulae, remainder of scrobe smooth. Shape and size of anterior ocellus: large, oblong in shape. Vertex posterior area sculpture: densely granulose. Occipital carina dorsal area: cristate, conspicuously present. Netrion: well-defined, suboval. Sculpture of mesepisternum: absent (smooth). Sculpture of pronotal lateral areas: with thin, longitudinal carinae. Length of fore wing stigmal vein: conspicuously elongate. Angle of stigmal vein in relation to anterior margin of fore wing: at an angle of approximately 45° with respect to anterior margin of wing. Sculpture of T2: mesal area with weal longitudinal carinae, remainder of tergum with longitudinal carinae mixed with coriaceous sculpture.
Diagnosis. Odontacolus africanus is very similar to O. anningae, but the latter species can be identified by the body being completely yellow and the mesal area of T2 with a conspicuous smooth area, and the remainder of the tergum with thin, somewhat sparse, well-defined, longitudinal costae throughout their length, and without coriaceous sculpture in the background.
Etymology. This species is named in reference to its broad distribution across southern and east Africa.
Comments. The holotype has the posterior right leg detached from the body and glued to the triangle; the right side wings are detached and in a gelatin capsule; the left antenna is missing. Most of the paratypes are in perfect condition.
In some female specimens the color pattern differs from that described above, the metasoma is a slightly lighter tone of dark brown compared to the head or the mesosoma (i.e. OSUC 238763). Additionally, the mesal area of T2 normally is covered by fine, dense, longitudinal costae (as in the holotype) but some specimens exhibit a smooth area with weak coriaceous sculpture in the background (in the absence of longitudinal costae), or a narrow smooth area with the lateral areas of the tergite with weak, longitudinal costae mixed with weak coriaceous sculpture in the background. Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: largely granulose, ventral 1/4 with broad costae. Netrion: present, smooth, well developed, sub-obovate. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: weakly rugulose mixed with weak granulae. Sculpture of mesoscutellum: granulose. Mesoscutellar profile: elevated, anterior margin higher than posterior. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: sparsely transversely carinate. Shape of propodeal anterior spine: elongate, narrow, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: weakly coriaceous. Sculpture of upper 1/4 of mesopleuron: densely longitudinally costate across entire width. Metapleural sculpture: largely smooth except lower half with longitudinal carinae.
Male. Unknown. Diagnosis. This is the only species of Odontacolus known from Nepal so far. It has a small but evident lagrimal as in O. mot; however, the lagrimal is not as large as in O. gentingensis. Odontacolus aldrovandii can be distinguished from O. mot by its smaller body size and the honey yellow coloration of the metasoma. In O. mot the metasoma in mainly dark brown except for the light orange T1.
Etymology.This species is named to honor the Renaissance Italian naturalist Ulisse Aldrovandi. The epithet is used as noun in the genitive case.
Comments. The holotype is in good condition except that the right hind wing and right antennal clava are missing. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: largely smooth ventrally, sparsely granulate ventrally. Surface of torular triangle: slightly bulging. Development of central keel on frons: present, elongate (equal to or greater than1/3× height of frons), but not reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: with sparse, transverse costae mixed with weak, dense granulae. Sculpture of malar space: coriaceous throughout, without fan-like striae. Furrow at lateral portion of antennal scrobe: absent. Shape of medial area of vertexmedial area of vertex: flat to weakly convex. Size of lateral ocelli: small. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: granulate. Sculpture of occipital carina: largely simple, at most with sparse weak crenulae medially. Distance from occipital carina to orbital carina: slightly greater than width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weak, small granulae. Sculpture of gena: granulose.
Wings. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head moderately short and strongly narrowed towards mouth, head appearing short and broad. Sculpture of antennal scrobe: smooth throughout. Surface of torular triangle: flat. Development of central keel on frons: present, elongate, reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: with weak rugulose-aciculate sculpture. Sculpture of malar space: with fan-like striae, striae extending into antennal scrobe. Furrow at lateral portion of antennal scrobe: absent. Shape of medial area of vertex: flat to weakly convex. Size of lateral ocelli: minute. Distance between lateral ocellus and occipital carina: greater than 1.5× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: with rugulose sculpture mixed with weak granulae. Sculpture of occipital carina: strongly crenulate throughout. Distance from occipital carina to orbital carina: at least 2× width of occipital carina; slightly greater than width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weak, small granulae. Sculpture of gena: with weak rugulose sculpture and granulate background sculpture.
Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: transversely costate. Netrion: absent, obscured by longitudinal sculpture of lateral pronotum. Notaulus: present, with crenulae that extend completely through depth of furrow. Length of notaulus: approximately 2/5× length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: finely granulose. Sculpture of mesoscutellum: granulose. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Male. Body length: 1. 39 mm (n=1). Body color: Antenna yellow as legs, metasoma light brown, head dark brown as mesosoma, fore wing with light infuscate color, no dark bands present. Sculpture of antennal scrobe: mesal area smooth, remainder with weakly coriaceous sculpture mixed with weak rugulose sculpture except upper 1/6 more rugulose than coriaceous. Shape and size of anterior ocellus: minute, very round. Vertex posterior area sculpture: with dense, small granulate sculpture. Occipital carina dorsal area: well-defined, conspicuously present. Netrion: practically absent by presence of longitudinal carinae on pronotal lateral areas. Sculpture of mesepisternum: mostly weakly coriaceous. Sculpture of pronotal lateral areas: with dense, fine, straight, transverse carinae. Length of fore wing stigmal vein: short. Angle of stigmal vein in relation to anterior margin of fore wing: at an angle of almost 90˚. Sculpture of T2: medially smooth, sublateral areas with semi-curved longitudinal carinae.
Diagnosis. This is one of two species within a group of taxa in which the netrion is obscured and its marginal crenulae are not well-defined, the mesoscutum is completely sculptured, the mesoscutellum is never with smooth areas, and has short, dense setation. Odontacolus australiensis can be separated from O. gallowayi by the deep impressed notauli that exhibit few sparse crenulae throughout their length in combina-tion with its brown body color; O. gallowayi has a shallow notauli with dense crenulae throughout their length, and its body color is yellow.
Etymology. This species is named after the collection locality of the species, Australia. The epithet is used as an adjective.
Link to distribution map. Comments. The holotype is in perfect condition; all paratypes are in good condition except for specimen OSUC 239143 which is missing the metasoma. Description. Female. Body length: 1.44 mm (n=1). Antenna color: completely yellow. Body color: mainly yellow, T1 horn dark brown, anteromesal mesoscutum honey yellow. Coxae color: whitish yellow. Leg color (excluding coxae): whitish yellow. Fore wing color: slightly infuscate throughout.

Odontacolus baeri
Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: weakly coriaceous throughout. Surface of torular triangle: slightly bulging. Development of central keel on frons: present, short (less than1/3 of frons height). Sculpture on upper frons below anterior ocellus: granulose throughout. Sculpture of malar space: coriaceous throughout, without fanlike striae. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: small. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: granulate. Sculpture of occipital carina: largely simple, at most with sparse weak crenulae medially. Distance from occipital carina to orbital carina: at least 2× width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weak, small granulae. Sculpture of gena: granulose.
Wings. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head moderately short and strongly narrowed towards mouth, head appearing short and broad. Sculpture of antennal scrobe: smooth throughout. Surface of torular triangle: slightly bulging. Development of central keel on frons: present, elongate (equal to or greater than1/3× height of frons), but not reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: with weak rugulose-aciculate sculpture. Sculpture of malar space: with fan-like striae, striae extending into antennal scrobe. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: small. Distance between lateral ocellus and occipital carina: greater than 1.5× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: with abundant areolate sculpture mixed with dense, fine granulae. Sculpture of occipital carina: strongly crenulate throughout. Distance from occipital carina to orbital carina: at least 2× width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weakly rugulo aciculate sculpture. Sculpture of gena: with weak rugulose sculpture and granulate background sculpture.
Male. Unknown. Diagnosis. This species may be distinguished from all others with an obscured netrion by the combination of the very wide notauli that have a few, broad and well-defined crenulae; the mesoscutum broadly smooth in its posterior half; and the mesoscutellum mesally with a smooth patch and its conspicuously long but somewhat sparse setae.
Etymology. This species is named after our entomological colleague Dr Jo Berry from New Zealand. The epithet is a noun in the genitive case.
Link to distribution map. 42 Biology. Recorded as an egg parasitoid of Trite planiceps Simon (Araneae: Salticidae) in New Zealand. Comments. The holotype is in perfect condition as are the remainder of the paratypes except for specimen OSUC 238564 which is covered with dust. Description. Female. Body length: 1.11-1.56 mm (n=20). Antenna color: clava and A2 light honey yellow, remainder of antenna yellow. Body color: completely dark brown. Coxae color: yellow. Leg color (excluding coxae): yellow. Fore wing color: slightly infuscate throughout.

Odontacolus bosei
Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: largely smooth ventrally, dorsally with sinuate, transverse ridges. Surface of torular triangle: slightly bulging. Development of central keel on frons: present, elongate, reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: with sparse, transverse costae mixed with weak, dense granulae. Sculpture of malar space: with weak rugulose sculpture mixed with granulate sculpture. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: normal. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: with rugulose sculpture mixed with weak granulae. Sculpture of occipital carina: weakly crenulate throughout. Distance from occipital carina to orbital carina: at least 2× width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weakly rugulo aciculate sculpture. Sculpture of gena: with weak rugulose sculpture and granulate background sculpture.
Male. Unknown. Diagnosis. This species is very similar to O. markadicus, but can be distinguished by the completely dark brown body and the conspicuously crenulate lateral occipital carina. In O. markadicus the body color is yellow and the lateral occipital carina is mainly smooth.
Etymology. This species is name after the amazing Bengali painter Nandalal Bose, recipient of the "Padma Vibhushan". The epithet is a noun in the genitive case. Comments. The holotype is in perfect condition as are most of the paratypes except for specimen OSUC 238789 which has the left fore wing glued to the point, and OSUC 238790 which has the metascutellum detached from the body. Description. Female. Body length: 1.65 mm (n=1). Antenna color: completely yellow. Body color: mostly yellow, propodeum, propodeal anterior spines, T1 horn dark brown. Coxae color: yellow. Leg color (excluding coxae): yellow. Fore wing color: completely hyaline.

Odontacolus cardaleae
Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: with weak, sinuate, transverse ridges throughout. Surface of torular triangle: slightly bulging. Development of central keel on frons: present, elongate (equal to or greater than1/3× height of frons), but not reaching anterior ocellus. Sculpture of vertex: with abundant areolate sculpture mixed with dense, fine granulae sculpture. Sculpture on upper frons below anterior ocellus: covered by sinuate, transverse, fine costae. Sculpture of malar space: granulose throughout, without fan-like striae. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: normal. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of occipital carina: largely simple, at most with sparse weak crenulae medially. Distance from occipital carina to orbital carina: at least 2× width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weakly rugulo aciculate sculpture. Sculpture of gena: granulose.
Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: largely smooth, dorsal margin with dense, weak punctulae. Netrion: present, smooth, linear. Notaulus: present, with crenulae that extend completely through depth of furrow. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: finely granulose. Sculpture of mesoscutellum: with weak, fine, granulate sculpture. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: densely, finely rugulose. Shape of propodeal anterior spine: elongate, narrow, apex rounded. Sculpture of propodeum between anterior spines: rugulose throughout. Sculpture of ventral half of mesepisternum: weakly coriaceous. Sculpture of upper 1/4 of mesopleuron: densely longitudinally costate across entire width. Metapleural sculpture: mainly with weak coriaceous sculpture, lower 1/3 without longitudinal costae. Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: coriaceous. Netrion: present, smooth, well developed, not looking -linear. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: densely, finely granulate, posterior 1/4 more coarsely coriaceous. Sculpture of mesoscutellum: with weak, fine, granulate sculpture. Mesoscutellar profile: elevated, anterior margin higher than posterior. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: sparsely transversely carinate. Shape of propodeal anterior spine: short, broad, apex subtriangular. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: smooth or nearly so. Sculpture of upper 1/4 of mesopleuron: densely longitudinally costate across half width. Metapleural sculpture: sparsely longitudinally costate.
Male. Unknown. Diagnosis. This species is easily identified based on its unique color pattern; there is no other known species that has the head and metasoma black in combination with an orange mesosoma and yellow legs.
Etymology. This species is named after the famed British biologist Charles Robert Darwin. The epithet is a noun in the genitive case.
Link Comments. The holotype is in perfect condition. Paratype OSUC 250602 is smaller in size than other specimens, and also has the mesal area of T2 smooth, the anterior mesal area of T3 mainly smooth with the exception of some weak, sparse longitudinal costae contrasting with the rest of specimens in which the sculpturing of T3 is as follows: anterior third is weakly costate sublaterally, weakly coriaceous mesally but otherwise granulose. Specimen OSUC 339589 has the metasoma lighter in color than other specimens. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: largely smooth ventrally, sparsely granulate ventrally. Surface of torular triangle: flat. Development of central keel on frons: present, elongate (equal to or greater than 1/3× height of frons), but not reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: granulose throughout. Sculpture of malar space: with sparse, short fan-like striae, striae not extending into scrobal area, mixed with weak coriaceous sculpture. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: slightly depressed. Size of lateral ocelli: large. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: granulate. Sculpture of occipital carina: largely simple, at most with sparse weak crenulae medially. Distance from occipital carina to orbital carina: at least 2× width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weak, small granulae. Sculpture of gena: granulose.

Odontacolus dayi
Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: coriaceous. Netrion: present, smooth, well developed, sub-obovate. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: weakly rugulose mixed with weak granulae. Sculpture of mesoscutellum: granulose. Mesoscutellar profile: elevated, anterior margin higher than posterior. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: coarsely rugulose. Shape of propodeal anterior spine: elongate, narrow, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: smooth or nearly so. Sculpture of upper 1/4 of mesopleuron: densely longitudinally costate across half width. Metapleural sculpture: largely smooth except lower half with longitudinal carinae.
Wings. Male. Unknown. Diagnosis. This is the only known species that has a longitudinally depressed vertex. In the Neotropics there is one undescribed species (in CNCI) with a very conspicuous longitudinal depression on the vertex, but the general sculpture of the body is very different from O. dayi.
Etymology. This species is named after the hymenopterist M. C. Day, now retired from the Natural History Museum, London. The epithet is a noun in the genitive case.
Comments. The holotype is in perfect condition. Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: largely covered by longitudinal costae except upper 1/5 foveate. Netrion: absent, obscured by longitudinal sculpture of lateral pronotum. Notaulus: present, with low crenulae that do not extend through depth of furrow. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: finely granulose. Sculpture of mesoscutellum: with weak, fine, granulate sculpture. Mesoscutellar profile: elevated, anterior margin higher than posterior. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: sparsely transversely carinate. Shape of propodeal anterior spine: short, broad, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: smooth or nearly so. Male. Unknown.

Odontacolus gallowayi
Diagnosis. This is one of two species within a group of taxa that has an obscured netrion, notaular crenulae poorly defined, and the mesoscutum completely sculptured (without smooth areas) and entirely covered by short, abundant setae. Odontacolus australiensis can be separated from O. gallowayi by the deep impressed notauli that exhibit few sparse crenulae throughout their length in combination with its brown body color; O. gallowayi has a shallow notauli with dense crenulae throughout its length, and its body color is yellow.
Comments. The holotype is in perfect condition; the paratype has the right hind wing and metasoma detached but glued to a card point. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: smooth throughout. Surface of torular triangle: flat. Development of central keel on frons: present, elongate (equal to or greater than1/3× height of frons), but not reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: granulose throughout. Sculpture of malar space: with fan-like striae, striae extending into antennal scrobe. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: large. Distance between lateral ocellus and occipital carina: less than0.5× maximum ocellar diameter. Lagrimal: conspicuously present (approximately 0.4× of malar sulcus length). Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: granulate. Sculpture of occipital carina: largely simple, at most with sparse weak crenulae medially. Distance from occipital carina to orbital carina: at least 2× width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weakly rugulo aciculate sculpture. Sculpture of gena: granulose.

Odontacolus gentingensis
Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), welldefined foveae. Sculpture of pronotal lateral area: smooth or nearly so. Netrion: present, smooth, linear. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: anterior half coriaceous, otherwise densely granulose. Sculpture of mesoscutellum: granulose. Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: weakly punctate, smooth posteroventrally. Netrion: present, smooth, linear. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: weakly rugulose mixed with weak granulae. Sculpture of mesoscutellum: granulose. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: depressed. Lateral propodeal area: coarsely rugulose. Shape of propodeal anterior spine: elongate, narrow, apex sharply acute. Sculpture of propodeum between anterior spines: longitudinally costate. Sculpture of ventral half of mesepisternum: longitudinally costate on weakly coriaceous background. Sculpture of upper 1/4 of mesopleuron: sparsely longitudinally costate across entire width. Metapleural sculpture: mainly with weak coriaceous sculpture, lower 1/3 without longitudinal costae.
Wings. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head moderately short and strongly narrowed towards mouth, head appearing short and broad. Sculpture of antennal scrobe: weakly rugulose throughout. Surface of torular triangle: flat. Development of central keel on frons: present, short (less than1/3 of frons height). Sculpture on upper frons below anterior ocellus: coriaceous throughout. Sculpture of malar space: weakly rugulose throughout, without fan-like striae. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: normal. Lagrimal: absent or minute. Distance from occipital carina to orbital carina: slightly greater than width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of gena: granulose.
Mesosoma. Sculpture of pronotal lateral area: upper 1/3 granulose, lower 1/3 with transverse foveae, otherwise smooth. Netrion: present, smooth, linear. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: weakly rugulose mixed with weak granulae. Sculpture of mesoscutellum: weakly rugulose mixed with granulate sculpture. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: sparsely transversely carinate. Shape of propodeal anterior spine: short, broad, apex rounded. Sculpture of ventral half of mesepisternum: smooth or nearly so. Sculpture of upper 1/4 of mesopleuron: densely longitudinally costate across half width. Metapleural sculpture: smooth. Male. Unknown. Diagnosis. This species can be separated from all other species with an obscured netrion by the subtriangular area below the eyes (seen in anterior view) that narrows towards the mandibles, unsculptured notauli, and non-ornamented occipital carina.
Etymology. This species is named after our colleague and arachnologist Dr Mark Harvey from the Western Australian Museum, who also manages to collect numerous parasitic Hymenoptera. The epithet is a noun in the genitive case.
Link to distribution map. Comments. The holotype is in good condition except for the right wings which are missing. The paratypes are in good condition except OSUC 237919 which has the metasoma detached and glued to the wings.
In some female specimens the color of the mesoscutum varies from completely yellow to almost completely brown. Additionally, the color of the lateral portions of the T1 horn may vary from completely yellow to almost completely light brown. Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: coriaceous. Netrion: present, smooth, well developed, sub-obovate. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: finely granulose. Sculpture of mesoscutellum: with weak, fine, granulate sculpture. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: sparsely transversely carinate. Shape of propodeal anterior spine: elongate, narrow, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: weakly coriaceous. Male. Unknown. Diagnosis. Odontacolus heydoni belongs to a group of species that has the occipital carina separated from the orbital carina, a well-defined netrion, a central keel on the frons, and well-defined notauli. Along with O. cardaleae it is the only species in the group that has a few, broad crenulae across the notauli and the antennal scrobes with transverse sinuate ridges. Odontacolus heydoni can be separated from O. cardaleae by its large ocelli (Fig. 40) and the short distance between the lateral ocellus and occipital carina (approximately 0.5× ocellar diameter); O. cardaleae has small ocelli and the distance between lateral ocellus and occipital carina is approximately 1.5× the ocellar diameter.

Odontacolus heratyi
Etymology. This species is named after Dr Steve Heydon at the Bohart Museum, who collected this magnificent species. The epithet is a noun in the genitive case. Comments. The holotype is in perfect condition as are the paratypes except for OSUC 238768 which has the right antenna missing. Some specimens have the body color completely yellow and without darker areas on the metasomal terga. Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), weakly defined foveae. Sculpture of pronotal lateral area: largely smooth, dorsal margin with dense, weak punctulae. Netrion: present, smooth, well-developed, not looking linear. Notaulus: absent. Length of notaulus: not applicable, notauli absent. Width of notaulus: not applicable, notauli absent. Sculpture of mesoscutum: finely granulose. Sculpture of mesoscutellum: with weak, fine, granulate sculpture. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: longitudinally costate. Shape of propodeal anterior spine: short, broad, apex subtriangular. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: smooth or nearly so. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: largely smooth ventrally, sparsely granulate ventrally. Surface of torular triangle: slightly bulging. Development of central keel on frons: present, elongate (equal to or greater than1/3× height of frons), but not reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: with weak rugulose-aciculate sculpture. Sculpture of malar space: with sparse, short fan-like striae, striae not extending into scrobal area, mixed with weak coriaceous sculpture. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: normal. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: granulate. Sculpture of occipital carina: weakly crenulate throughout. Distance from occipital carina to orbital carina: contiguous or nearly so, subequal to width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weakly rugulo aciculate sculpture. Sculpture of gena: granulose.

Odontacolus irwini
Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: dorsally punctate, otherwise smooth. Netrion: present, smooth, linear. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: finely granulose. Sculpture of mesoscutellum: granulose. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: coarsely rugulose. Shape of propodeal anterior spine: elongate, narrow, apex sharply acute. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: weakly, finely coriaceous. Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: coriaceous. Netrion: present, smooth, well developed, sub-obovate. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: weakly rugulose mixed with weak granulae. Sculpture of mesoscutellum: granulose. Mesoscutellar profile: elevated, anterior margin higher than posterior. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: densely, finely rugulose. Shape of propodeal anterior spine: short, broad, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: smooth or nearly so. Sculpture of upper 1/4 of mesopleuron: densely longitudinally costate across entire width. Metapleural sculpture: mainly with weak coriaceous sculpture, lower 1/3 without longitudinal costae.
Wings. These species belong to a group that have short, smooth notauli, a well-defined netrion, a central keel on the frons, and the sculpture of the frons always has transverse costae. Additionally, O. kiau can be separated from O. mayri by the dark brown coxae and the slightly bulging torular triangle observed on the former; in O. mayri the coxae are yellow and the torular triangle is flat.
Etymology. This species is named after the East New Britain word 'kiau' (in Kuanua language) which means 'egg', and refers to the stage of the host parasitized by Odontacolus. The name is used as a noun in apposition.
Link Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), weakly defined foveae. Sculpture of pronotal lateral area: upper 1/3 granulose, lower 1/3 with transverse foveae, otherwise smooth. Netrion: present, smooth, well developed, subobovate. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: weakly rugulose mixed with weak granulae. Sculpture of mesoscutellum: granulose. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: longitudinally costate. Shape of propodeal anterior spine: elongate, narrow, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: smooth or nearly so. Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal lateral area: largely smooth, dorsal margin with dense, weak punctulae. Netrion: present, smooth, linear. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: coriaceous. Sculpture of mesoscutellum: weakly coriaceous. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: longitudinally costate. Shape of propodeal anterior spine: short, broad, apex subtriangular. Sculpture of propodeum between anterior spines: smooth or largely smooth; with strong, oblique ridges laterally. Sculpture of ventral half of mesepisternum: smooth or nearly so. Male. Unknown. Diagnosis. This species can be easily separated from all other Odontacolus with smooth, short notauli and a well-developed netrion by the combination of the completely dark brown body, the nearly smooth posterior portion of the mesoscutum (between the notaulus and tegula), and the nearly smooth posterior 1/3 and mesal region of the mesoscutellum.
Comments. The holotype is in good condition except that the right hind wing and right antenna are detached from body and glued to the point. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: smooth throughout. Surface of torular triangle: flat. Development of central keel on frons: present, elongate (equal to or greater than1/3× height of frons), but not reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: largely smooth, with sparse coriaceous sculpture close to compound eyes. Sculpture of malar space: largely smooth, coriaceous sculpture present near compound eyes (somewhat weak). Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: small. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: coriaceous. Sculpture of occipital carina: largely simple, at most with sparse weak crenulae medially. Distance from occipital carina to orbital carina: slightly greater than width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weakly rugulo aciculate sculpture. Sculpture of gena: coriaceous dorsally, otherwise smooth.

Odontacolus madagascarensis
Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: largely smooth, dorsal margin with dense, weak punctulae. Netrion: present, smooth, linear. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: coriaceous. Sculpture of mesoscutellum: weakly coriaceous. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: densely, finely rugulose. Shape of propodeal anterior spine: short, broad, apex subtriangular. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: smooth or nearly so. Sculpture of upper 1/4 of mesopleuron: densely longitudinally costate across half width. Metapleural sculpture: smooth.
Wings. Male. Unknown. Diagnosis. This species can be distinguished from all other Odontacolus species by the combination of the completely smooth gena, the well-defined netrion, the presence of notauli, and the confused, dense rugulose sculpture of the lateral propodeum.
Etymology. This species is named after the island from which the species was collected, Madagascar. The epithet is used as an adjective.
Wings. Comments. We were unable to obtain any of the type series, and so our interpretation of O. markadicus is based on comparision of the additional material (listed above) with the images and description in Veenakumari & Mohanraj (2011) as well as images kindly sent to us by our colleague Dr Rajmohana K.
When taking all of the available material into account some variation is evident in the females of this species: the antennal clava can be completely yellow or the distal half of the clava slightly darker, and the vertex and mesoscutum vary from completely yellow to slightly honey yellow in color. Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: upper 1/3 granulose, lower 1/3 with transverse foveae, otherwise smooth. Netrion: present, smooth, well-developed, not looking linear. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: weakly granulose. Sculpture of mesoscutellum: with weak, fine, granulate sculpture. Mesoscutellar profile: elevated, anterior margin higher than posterior. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: densely, finely rugulose. Shape of propodeal anterior spine: elongate, narrow, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: smooth or nearly so. Sculpture of upper 1/4 of mesopleuron: densely longitudinally costate across half width. Metapleural sculpture: mainly smooth, lower third sparsely longitudinally carinate.
Male. Unknown. Diagnosis. This species belongs to a group that has short, smooth notauli, a welldefined netrion, central keel present, and sculptured antennal scrobe always with transverse costae in the background. Within this group Odontacolus mayri can be separated from O. kiau and O. whitfieldi by its completely yellow body and the weakly granulose sculpture of the mesoscutum.
Etymology. This species is named after the German biologist Ernst Walter Mayr. The epithet is a noun in the genitive case.
Link to distribution map. Comments. The holotype is in perfect condition. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: largely smooth ventrally, dorsally with sinuate, transverse ridges. Surface of torular triangle: slightly bulging. Development of central keel on frons: present, elongate, reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: covered by sinuate, transverse, fine costae. Sculpture of malar space: granulose throughout. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: normal. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: present, well-defined but small (length less than 0.3 of malar sulcus length). Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: granulate. Sculpture of occipital carina: largely simple, at most with sparse weak crenulae medially. Distance from occipital carina to orbital carina: at least 2× width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weak, small granulae. Sculpture of gena: granulose.
Male. Unknown. Diagnosis. This species is very similar to O. markadicus; however, O. mot differs from it by the larger body size (1.92 mm vs. 1.74 mm or less), the broader transverse sinuate carinae on the upper frons (in O. markadicus the sculpturing is completely absent or if present then the carinae are thin and straighter than in O. mot), and by the denser and less evenly distributed setae on the metasoma. Additionally, T2 in O. mot is more quadrate, while it is more elongate and less broad posteriorly in O. markadicus.
Etymology. The name of this species is an arbitrary combination of letters and is used as a noun in apposition.
Comments. The holotype is in perfect condition. The differences between O. mot and O. markadicus could be associated with the difference in body size. However, since O. mot also differs in metasomal setation and sculpture, we have chosen to treat them as separate species until additional material of O. mot is available. Head. Size of compound eye: reduced, approximately less than or equal to 1/3× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: coriaceous throughout. Surface of torular triangle: flat. Development of central keel on frons: present, elongate (equal to or greater than1/3× height of frons), but not reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: with sparse, transverse costae mixed with weak, dense granulae. Sculpture of malar space: coriaceous throughout, without fan-like striae. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: normal. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: granulate. Sculpture of occipital carina: largely simple, at most with sparse weak crenulae medially. Distance from occipital carina to orbital carina: at least 2× width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weak, small granulae. Sculpture of gena: granulose.

Odontacolus noyesi
Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: coriaceous. Netrion: present, smooth, well developed, sub-obovate. Notaulus: present, with crenulae that extend completely through depth of furrow. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: finely granulose. Sculpture of mesoscutellum: with weak, fine, granulate sculpture. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: densely, finely rugulose. Shape of propodeal anterior spine: short, broad, apex subtriangular. Sculpture of propodeum between anterior spines: smooth or largely smooth; longitudinally costate. Sculpture of ventral half of mesepisternum: weakly coriaceous. Sculpture of upper 1/4 of mesopleuron: densely longitudinally costate across entire width. Metapleural sculpture: mainly with weak coriaceous sculpture, lower 1/3 without longitudinal costae.
Male. Unknown. Diagnosis. This is the only species of Odontacolus that has the compound eyes unusually small, approximately 1/3 of the height of the head.
Etymology. This species is named after our friend and colleague, the chalcid specialist and great insect collector Dr John Noyes from the Natural History Museum, London. The epithet is a noun in the genitive case. Comments. The holotype has the left legs separate and glued to the point. Otherwise it is in good condition. The paratype in perfect condition: it has the metasoma completely honey yellow and the remainder of the body, including the legs, yellow.
Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: smooth throughout. Surface of torular triangle: slightly bulging. Development of central keel on frons: present, elongate (equal to or greater than1/3× height of frons), but not reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: granulose throughout. Sculpture of malar space: largely smooth, with sparse, weak, fan-like striae. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: minute. Distance between lateral ocellus and occipital carina: greater than 1.5× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: granulate. Sculpture of occipital carina: largely simple, at most with sparse weak crenulae medially. Distance from occipital carina to orbital carina: at least 2× width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weak, small granulae. Sculpture of gena: coriaceous dorsally, otherwise smooth.
Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: transversely costate. Netrion: present, smooth, linear. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: coriaceous. Sculpture of mesoscutellum: smooth or nearly so. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: sparsely transversely carinate. Shape of propodeal anterior spine: short, broad, apex subtriangular. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: smooth or nearly so. Sculpture of upper 1/4 of mesopleuron: densely longitudinally costate across entire width. Metapleural sculpture: smooth.
Male. Body length: 1.27-1.30 mm (n= 3). Body color: antenna yellow as legs and T1, remainder of metasoma dark brown as dorsal mesosoma as head, remainder of body dark honey yellow. Sculpture of antennal scrobe: with weakly sinuate, fine, weak dorsoventral carinae, except area below anterior ocellus with granulate sculpture. Shape and size of anterior ocellus: minute, very round. Vertex posterior area sculpture: dense granulate sculpture. Occipital carina dorsal area: cristate, conspicuously present. Netrion: well-defined, suboval, broad longitudinal carinae present before netrion. Sculpture of mesepisternum: absent (smooth). Sculpture of pronotal lateral areas: with few, sinuate, transverse carinae. Length of fore wing stigmal vein: conspicuously elongate. Angle of stigmal vein in relation to anterior margin of fore wing: at an angle of approximately 45°. Sculpture of T2: mesal area smooth, remainder of tergum with weak longitudinal carinae.

Diagnosis.
Odontacolus pintoi can be distinguished from all other species that have an obscured netrion in combination with the elongate and broad ventral portion of the head (below the eyes in anterior view) (Fig. 243) by the mostly smooth gena (shiny, sometimes with large coriaceous sculpture present but restricted to upper 1/3), and the smooth antennal scrobe (except for the presence of a central keel).
Etymology. This species is named after the trichogrammatid specialist Dr John Pinto, formerly of the University of California, Riverside. The epithet is a noun in the genitive case.
Link to distribution map. 86 Biology. Specimens from South Australia have been reared from eggs of Clubiona cycladata Simon (Araneae: Clubionidae) under the bark of eucalypt trees (recorded as Odontacolus sp. in Austin 1984& Austin 1985. Comments. The holotype is in perfect condition; the paratypes are for the most part in very good condition except for OSUC 239063 and OSUC 239074 which have the heads detached from body and glued to the point, and OSUC 239046 which is missing the metasoma. Some females have a different color pattern in that the whole body is dark brown with the anterior edge of T1 (except the horn), T2 and the legs yellow.
Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: weakly granulose throughout. Surface of torular triangle: flat. Development of central keel on frons: present, short (less than1/3 of frons height). Sculpture on upper frons below anterior ocellus: granulose throughout. Sculpture of malar space: granulose throughout, without fan-like striae. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: large. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: granulate. Sculpture of occipital carina: weakly crenulate throughout. Distance from occipital carina to orbital carina: slightly greater than width of occipital carina. Shape of occipital carina: weakly sinuate medially. Sculpture of occiput: with weak, small granulae. Sculpture of gena: with weak rugulose sculpture and granulate background sculpture.
Mesosoma. Dorsal mesosoma in lateral view: conspicuously flattened. Sculpture of pronotal lateral area: largely smooth, dorsal margin with dense, weak punctulae. Netrion: present, smooth, linear. Notaulus: absent. Length of notaulus: not applicable, notauli absent. Width of notaulus: not applicable, notauli absent. Sculpture of mesoscutum: weakly rugulose, carinae thin in shape. Sculpture of mesoscutellum: rugulose. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: longitudinally costate. Shape of propodeal anterior spine: short, broad, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: smooth or nearly so. Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), welldefined foveae. Sculpture of pronotal lateral area: upper 1/3 granulose, lower 1/3 with transverse foveae, otherwise smooth. Netrion: present, smooth, linear. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: weakly rugulose mixed with weak granulae. Sculpture of mesoscutellum: granulose. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: coarsely rugulose. Shape of propodeal anterior spine: elongate, narrow, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: smooth or nearly so. Male. Unknown. Diagnosis. This is the only known species to have a furrow at the lateral areas of the antennal scrobes in combination with a completely dark brown body.
Etymology. This species is named after our friend and colleague, hymenopterist Dr Mike Sharkey from the University of Kentucky. The epithet is a noun in the genitive case.

Odontacolus spinosus
Comments. The holotype of O. spinosus has the head and most of the wings detached from the body; the head and first pair of legs are glued to the point; the remainder of the legs has the distal tarsomeres missing. The holotype of O. doddi is slide mounted and the specimen was partly destroyed in the process.
The following color variations occur in this species: the body ranges from yellow with the tips of the propodeal spines and T1 horn dark brown to mostly dark brown with the posterior area of the mesoscutum and mesoscutellum light yellow. The occipital carina of the specimens OSUC 239180 and OSUC 239173 is present but very fine which renders it difficult to see; in all other specimens it is well developed. The sculpture also varies between the anterior propodeal spines, from being completely smooth to completely punctate in some specimens with darker color (i.e. OSUC 239165, 265167, 239170-239172). The sculpture of the mesal portion of T2 varies from in the development of the costae. In any case there is always coriaceous background sculpture. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head elongate and broad at mouth, head appearing elongate and somewhat thin. Sculpture of antennal scrobe: weakly granulose throughout. Surface of torular triangle: flat. Development of central keel on frons: present, elongate (equal to or greater than1/3× height of frons), but not reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: granulose throughout. Sculpture of malar space: granulose throughout, without fan-like striae. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: large. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: granulate. Sculpture of occipital carina: largely simple, at most with sparse weak crenulae medially. Distance from occipital carina to orbital carina: contiguous or nearly so, subequal to width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: uncertain, with weakly rugulo aciculate sculpture. Sculpture of gena: granulose.
Wings. Etymology. This species is named after Veronica Valerio, sister of the first author. The epithet is a noun in the genitive case. Comments. The holotype has all but one leg missing, but otherwise is in perfect condition; the paratypes are in good condition. The color of the metasoma varies from completely dark brown (matching the color of the head and mesosoma) to being conspicuously lighter than the head and mesosoma. The color of the antenna may vary from yellow to brown. Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), welldefined foveae. Sculpture of pronotal lateral area: largely smooth, dorsal margin with dense, weak punctulae. Netrion: present, smooth, well developed, sub-obovate. No-taulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: finely granulose. Sculpture of mesoscutellum: with weak, fine, granulate sculpture. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: sparsely transversely carinate. Shape of propodeal anterior spine: elongate, narrow, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: weakly coriaceous. Sculpture of upper 1/4 of mesopleuron: with sparse, broad, smooth costate sculpture reaching just half of its width. Metapleural sculpture: mainly with weak coriaceous sculpture, lower 1/3 with sparse longitudinal carinae.

Odontacolus wallacei
Wings. Comments. The holotype generally is in good condition: the left front and hind legs are glued to the point, and the right hind leg is covered in glue. The metasomal color pattern varies between specimens from almost completely yellow (i.e. OSUC 238004) to being mostly honey yellow with whitish areas on T3 and dark brown T4-T6 (i.e. OSUC 238001). In some specimens (i.e. OSUC 239020) the upper area of the T1 horn is dark brown with the remainder of the metasoma yellow, and the mesoscutum (which is normally light yellow) being a honey yellow color mesally. Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: coriaceous. Netrion: present, smooth, well developed, sub-obovate. Notaulus: present, simple. Length of notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: weakly rugulose mixed with weak granulae. Sculpture of mesoscutellum: granulose. Mesoscutellar profile: elevated, anterior margin higher than posterior. Mesoscutellar shape: flat, not depressed. propodeal area: densely, finely rugulose. Shape of propodeal anterior spine: elongate, narrow, apex rounded. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: smooth or nearly so. Sculpture of upper 1/4 of mesopleuron: with sparse, broad, smooth costate sculpture reaching just half of its width. Metapleural sculpture: largely smooth except lower half with longitudinal carinae.

Odontacolus whitfieldi
Wings. Male. Body length: 1.29-1.37 mm (n=5). Body color: antenna yellow as legs, head, mesoscutum, mesoscutellum and metasoma dark brown, remainder of body light honey-yellow. Sculpture of antennal scrobe: lower mesal area smooth, remainder with weak coriaceous sculpture mixed with somewhat sinuate carinae (especially at upper 1/5), except area below anterior ocellus with granulose sculpture. Shape and size of anterior ocellus: small, round. Vertex posterior area sculpture: with dense, small granulae. Occipital carina dorsal area: cristate, conspicuously present. Netrion: well-defined, suboval. Sculpture of mesepisternum: with few, thin, transverse carinae. Sculpture of pronotal lateral areas: with few, thin transverse carinae, otherwise mainly smooth. Length of fore wing stigmal vein: conspicuously elongate. Angle of stigmal vein in relation to anterior margin of fore wing: at an angle of approximately 45°. Sculpture of T2: mostly smooth, lateral areas with few sparse longitudinal carinae.
Diagnosis. Odontacolus whitfieldi is very similar to O. kiau but the former has a complete S2 anterior carina; O. kiau lacks this carina. These species belong to a group that have short and smooth notauli, a well-defined netrion, a central keel present on the frons and sculptured antennal scrobes always with transverse costae; within this group O. mayri can be separated from O. whitfieldi and O. kiau by its completely yellow body and the weakly granulose sculpture of the mesoscutum.
Etymology. This species is named after our friend and colleague, the microgastrine systematist Dr Jim Whitfield at the University of Illinois. The epithet is a noun in the genitive case.
Comments. Holotype specimen is in perfect condition. Most of paratype specimens are in good condition. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head moderately short and strongly narrowed towards mouth, head appearing short and broad. Sculpture of antennal scrobe: with weak, sinuate, transverse ridges throughout. Surface of torular triangle: slightly bulging. Development of central keel on frons: present, short (less than1/3 of frons height); present, elongate, reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: covered by sinuate, transverse, fine costae. Sculpture of malar space: with fan-like striae, striae not extending into antennal scrobe. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: small. Distance between lateral ocellus and occipital carina: greater than 1.5× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: with sinuate, transverse, fine ridges. Sculpture of occipital carina: largely simple, at most with sparse weak crenulae medially. Distance from occipital carina to orbital carina: at least 2× width of occipital carina. Shape of occipital carina: weakly sinuate medially. Sculpture of occiput: with dense, fine transverse costae across its width. Sculpture of gena: with sinuate dorsoventral costae.

Odontacolus zborowskii
Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal cervical area: with small (at most as large as crenulae on anterior edge of mesoscutum), well-defined foveae. Sculpture of pronotal lateral area: transversely costate. Netrion: absent, obscured by longitudinal sculpture of lateral pronotum. Notaulus: present, with low crenulae that do not extend through depth of furrow. Length of notaulus: ap-Comments. The holotype is in perfect condition as are the paratypes. In some specimens (e.g. OSUC 237956) the area between the propodeal anterior spines has some dark honey yellow color instead of the typical yellow. Head. Size of compound eye: approximately 1/2× height of head. Head shape in lateral view: lower head moderately short and strongly narrowed towards mouth, head appearing short and broad. Sculpture of antennal scrobe: smooth throughout. Surface of torular triangle: slightly bulging. Development of central keel on frons: present, elongate (equal to or greater than1/3× height of frons), but not reaching anterior ocellus. Sculpture on upper frons below anterior ocellus: coriaceous throughout. Sculpture of malar space: with fan-like striae, striae not extending into antennal scrobe. Furrow at lateral portion of antennal scrobe: absent. Mesal surface of vertex: flat to weakly convex. Size of lateral ocelli: minute. Distance between lateral ocellus and occipital carina: 0.5-1.2× maximum ocellar diameter. Lagrimal: absent or minute. Length of OOL: less than or equal to 1/3× width of ocellus. Sculpture of vertex: coriaceous. Sculpture of occipital carina: weakly crenulate throughout. Distance from occipital carina to orbital carina: contiguous or nearly so, subequal to width of occipital carina. Shape of occipital carina: simply arcuate medially. Sculpture of occiput: with weakly rugulo aciculate sculpture. Sculpture of gena: coriaceous dorsally, otherwise weakly rugulose.

Odontacolus zimi
Mesosoma. Dorsal mesosoma in lateral view: convex. Sculpture of pronotal lateral area: dorsal 2/3 punctate, otherwise smooth. Netrion: present, smooth, linear. Notaulus: present, simple. Length of `notaulus: approximately less than or equal to 1/3 of length of mesoscutum. Width of notaulus: narrow (notaulus width less than or equal to half the width of tegula). Sculpture of mesoscutum: anterior half coriaceous, otherwise densely granulose. Sculpture of mesoscutellum: granulose. Mesoscutellar profile: mainly flat, anterior and posterior edge at same height or nearly so. Mesoscutellar shape: flat, not depressed. Lateral propodeal area: sparsely transversely carinate. Shape of propodeal anterior spine: short, broad, apex subtriangular. Sculpture of propodeum between anterior spines: smooth or largely smooth. Sculpture of ventral half of mesepisternum: longitudinally costate. Sculpture of upper 1/4 of mesopleuron: densely longitudinally costate across entire width. Metapleural sculpture: smooth. Male. Unknown. Diagnosis. This species can be separated from all the species which have the occipital carina almost touching the orbital carina by the absence of the S2 anterior carina, the posterior medial area of the mesoscutellum not depressed, and the complete absence of the lagrimal. The closest species is O. jacksonae, but the body of this species is completely dark brown in contrast with the honey yellow metasoma and dark brown head and mesosoma of O. zimi.
Etymology. This species is named after the anime character 'Invader Zim', in reference to the invasion of the spider egg sacs that occurs when Odontacolus oviposit. The epithet is a noun in the genitive case.
Comments. The holotype is in perfect condition.