New reared rogadine wasp species from New Guinea 85 Colastomion Baker ( Braconidae , Rogadinae ) : nine new species from Papua New Guinea reared from Crambidae

Nine new species of Colastomion Baker are described, illustrated and keyed based on series of specimens reared from caterpillars of crambid moths from lowland Papua New Guinea plus one additional field collected specimen, viz. C. cheesmanae Quicke sp. n., C. crambidiphagus Quicke sp. n., C. gregarius Quicke sp. n., C. maclayi Quicke sp. n., C. madangensis Quicke sp. n., C. masalaii Quicke sp. n., C. parotiphagus Quicke sp. n., C. pukpuk Quicke sp. n. and C. wanang Quicke sp. n. Most species are morphologically easily distinguished but DNA barcoding additionally reveals a pair of exceedingly similar species (C. pukpuk sp. n. and C. maclayi sp. n.) that might otherwise have gone unrecognised. The new species each appear to be relatively specialised on their host species and all parasitize only caterpillars of Lepidoptera: Crambidae: Spilomelinae. JHR 28: 85–121 (2012) doi: 10.3897/JHR.28.3484 www.pensoft.net/journals/jhr Copyright Donald L.J. Quicke et al. This is an open access article distributed under the terms of the Creative Commons Attribution License 3.0 (CC-BY), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. ReseARCh ARtiCle Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012) 86


introduction
Colastomion Baker (1917) is a relatively uncommon genus of rogadine parasitic wasp that occurs throughout the southern East Palaearctic, S. E. Asia and Africa.It belongs to a group of genera which additionally includes the tropical Old World genera Cystomastacoides van Achterberg, Macrostomion Szépligeti, Megarhogas Szépligeti and Myocron van Achterberg (1991) all of which share a more or less deep and ventrally strongly curved hypopyium with a strongly down-curved ovipositor (also present in various other SE Asian genera).Until recently nothing was known about its biology though Macrostomion has been reared from various Sphingidae caterpillars within which it is gregarious with 20-50 individuals typically emerging from a single mummified host larva (Shaw 2002, Maeto andArakaki 2005).
Here we describe nine new species of Colastomion, eight of which are based on reared specimens from caterpillars feeding on various trees and shrubs in lowland Papua New Guinea.These were obtained during an extensive caterpillar collecting and rearing programme which has additionally yielded a number of other interesting rogadine parasitoids including the recently described, highly distinctive genus Vojtechirogas Quicke & van Achterberg (Quicke et al. 2012).
The host records for some of the species include a degree of fuzziness because identifications are based on caterpillar morphology.The actual parasitized caterpillars of course never produce an adult moth, and so there is always some degree of uncertainty with host records acquired by rearing.Because in the course of the New Guinea rearing programme there were several hundred thousand caterpillars reared, we are able to quantify the inevitable uncertainty.For any reared specimen we present only host records which are at least 95% sure, i.e. at least 95% of the tens or hundreds of moth adults reared from the same combination of caterpillar morphotype and host plant as the parasitoid agree with the reported host identification.Two rearing records for C. masalaii sp.n. and C. parotiphagus sp.n. were confirmed by sequencing host remnants (Hrcek et al. 2011).
The systematics of several of the moth taxa involved here, all of which belong to the crambid subfamily Spilomelinae, is also far from complete and there is no meaningful phylogeny as yet.Thus some genera may well be poly-or paraphyletic assemblages, and in these cases we have sometimes been able to assign hosts to species groups, that in the future may shift into other generic combinations.terminology Terminology follows van Achterberg (1979van Achterberg ( , 1988)).Measurements of the height and horizontal length of the eye are approximations because the very bulbous face and very large eyes make it difficult to measure consistently; in our attempts to do so, the head was orientated so that the face (defined as running from the anterior edge of the toruli to the dorsal margin of the clypeus) was horizontal or perpendicular to the measurement axis.
Abbreviations: NHM (The Natural History Museum, London); USNM (United States National Museum, Washington D.C.).Note, some paratypes will be repatriated to PNG when analysis is complete.

Methodology
Phylogenetic inference and bar-coding discrimination of species were based upon maximum likelihood (ML) analysis of approximately 657 base pairs of the 5' end of the mitochondrial cytochrome oxidase I gene, using the programme RAxML (Stamatakis 2006) with the GTRGAMMA model and using a sequence from the related genus Myocron van Achterberg as outgroup.DNA sequencing was performed at the Biodiversity Institute of Ontario, University of Guelph, using their standard methods (Hrcek et al. 2011).
Most specimens were imaged using Cell^D® imaging facility.C. gregarious sp.n. and C. cheesmanae sp.n. were imaged using a Canon EOS 7D camera, Adobe Lightroom software and edited using Adobe Photoshop CS4.

Results
DNA barcodes were generated from all but one of the species described below, the exception being a nearly 80 year old specimen found in the NHM collection.Fig. 1 shows a ML tree from these, with eight clearly separated molecular species, and intraspecific variation was limited to two individuals of C. masalaii sp.n. (vouchers USNM ENT 00503254 and 00643295) differing from the remainder at a single base position.Paratypes.4 males and 3 females, 1 unknown (missing metasoma), rest of reared series from the same parasitized host, mounted on total of 3 cards and on same pin as holotype.(USNM).
Head.Antenna with 31-32 flagellomeres.Terminal flagellomere acuminate.Median flagellomeres approximately 2 × longer than wide.3 rd segment of maxillary palp of female weakly swollen, approximately 4.5 longer than maximally wide.5 th segment approximately same length as 6 th .Base of mandible well removed from eye, closest point approximately 1.3 distance from middle of anterior tentorial pit and eye.Distance between anterior tentorial pits approximately 2.7 × shortest distance between pit and eye.Width of face: width of head across eyes: height of eye = 1.0 : 2.25 : 1.3.Face rather shiny, with fine transverse striation laterally above level of clypeus.Shortest distance between posterior ocelli: transverse diameter of posterior ocellus: shortest distance between posterior ocellus and eye = 1.0: 1.2 : 1.4.Occipital carina broadly obliterated or extremely weak mediodorsally Mesosoma.Notauli narrow, moderately impressed, crenulated.Mesoscutum longitudinally striate-rugulose at confluence of notauli.Mesopleuron largely smooth and shiny.Precoxal sulcus deeply impressed and strongly crenulate.Propodeum with very short midlongitudinal carina anteriorly that splits giving rise to pair of weak carinae bordering a widening midlongitudinal depression with moderately well developed transverse striation.Propodeum with distinct mediolateral projections.
Hind wing.Vein M+CU 0.9 × length of 1-M.Vein 2-SC+R thickened and distinctly inclivous.Vein SR moderately strongly curved on basal half, almost reaching wing margin approximately half distance to wing tip, then running more or less parallel to wing margin.
Head.Antenna with 35-37 flagellomeres.Terminal flagellomere pointed, not or hardly acuminate.Median flagellomeres 1.8 × longer laterally than wide.3 rd segment of maxillary palp of female not swollen, approximately 7 × longer than maximally wide.5 th segment approximately 0.7 length of 6 th .Base of mandible well separated from eye, closest point approximately 1.2 distance from middle of anterior tentorial pit and eye.Distance between anterior tentorial pits approximately 3.0 × shortest distance between pit and eye.Width of face: width of head across eyes: height of eye = 1.0 : 2.3 : 1.3.Face with distinct transverse striation laterally.Frons with distinct pattern of elongate pits forming chevrons on either side of midlongitudinal sulcus.Shortest distance between posterior ocelli: transverse diameter of posterior ocellus: shortest distance between posterior ocellus and eye = 1.0: 1.6: 2.0.Occipital carina complete but irregular and weak mediodorsally Mesosoma.Notauli more or less impressed along whole length, and crenulated, converging medioposteriorly and mesoscutum with some irregular rugosity at their confluence.Mesopleuron smooth and shiny.Precoxal sulcus strongly impressed, rather narrow, foveate.Propodeum with short midlongitudinal carina anteriorly that splits giving rise to a widening midlongitudinal depression with moderately well developed transverse striation.Propodeum with distinct mediolateral projections.
Coloration.Body largely pale brown yellow; scapus and pedicellus yellow, flagellum largely black becoming more rufous distally; legs entirely pale brown yellow.Wing membrane hyaline, venation largely brown, pterostigma dark brown.
Etymology.Named after the local forest spirits called 'masalai' in Melanesian Pidgin.
Hind wing.Vein M+CU 0.6 × length of 1-M.Vein 2-SC+R thickened, distinctly inclivous.Vein SR weakly curved on basal half, getting closest to wing margin approximately 0.6 distance to wing tip, then running more or less parallel to wing margin.
Coloration.Body largely brown yellow, metasoma somewhat more reddish; antennae largely cream white, darkening to blackish near tip; fore legs entirely, mid-and hind legs from apex of coxa white.Wing membrane hyaline, venation largely brown, pterostigma largely brown, paler at base.
Etymology.Named in reference to the province where specimens were reared.

Colastomion crambidiphagus
Head.Antenna with 45 flagellomeres.Terminal flagellomere strongly acuminate.Median flagellomeres approximately 2.5 × longer than wide.3 rd segment of maxillary palp of female swollen, approximately 4.0 longer than maximally wide.5 th segment reduced, approximately 0.5 length of 6 th .Base of mandible separated from eye by approximately same distance as from middle of anterior tentorial pit and eye.Distance between anterior tentorial pits approximately 4.0 × shortest distance between pit and eye.Width of face: width of head across eyes: height of eye = 1.0 : 2.5 : 1.6.Face weakly transversely to obliquely striate dorsolaterally.Shortest distance between posterior ocelli: transverse diameter of posterior ocellus: shortest distance between posterior ocellus and eye = 1.0: 2.0 : 1.2.Occipital carina narrowly obliterated medially Mesosoma.Notauli moderately deep, narrow and crenulated, uniting shortly before posterior of mesoscutum.Mesoscutum with distinct narrow midlongitudinal groove posteriorly.Propodeum with midlongitudinal carina at least on anterior 0.5, posteriorly either complete with strong transverse rugae lateral to it, or replaced by transverse rugosity.
Hind wing.Vein M+CU 1.1 × length of 1-M.Vein 1-SC+R slightly thickened, almost transverse.Vein SR strongly curved on basal half, almost reaching wing margin approximately half distance to wing tip, then running more or less parallel to wing margin.
Head.Antenna with 50 flagellomeres.Terminal flagellomere acuminate.Median flagellomeres 2 × longer than wide.3 rd segment of maxillary palp of female distinctly swollen, approximately 3.5 × longer than maximally wide.5 th segment reduced and approximately half length of 6 th .Base of mandible very close to eye, closest point approximately 0.2 distance from middle of anterior tentorial pit and eye.Distance between anterior tentorial pits approximately 5 × shortest distance between pit and eye.Width of face: width of head across eyes: height of eye = 1.0 : 2.5 : 1.6.Face transversely striate except for smooth triangular area above clypeus extending nearly to antennal sockets.Frons largely smooth but with well developed ridge running parallel to eye margin and reaching stemmaticum.Shortest distance between posterior ocelli: transverse diameter of posterior ocellus: shortest distance between posterior ocellus and eye = 1.5: 2.5:1.0.Occipital carina complete, well developed dorsally.
Mesosoma.Notauli deep, complete, finely punctate anteriorly becoming obliquely crenulated posteriorly, uniting to form groove before posterior margin of mesoscutum.Mesoscutum with an elongate groove between notauli anterior to the point of their fusion.Propodeum with complete midlongitudinal carina.Mesopleuron largely smooth.Precoxal sulcus narrow and deep on posterior half of mesopleuron, finely crenulated.Propodeum with complete midlongitudinal carina.
Hind wing.Vein M+CU 1.75 × length of 1-M.Vein 2-SC+R weakly thickened, distinctly reclivous.Vein SR strongly curved on basal half, almost reaching wing margin approximately half distance to wing tip, then distinctly diverging again towards wing tip.
Coloration.Head and body ochreous yellow, stemmaticum black, hypopygium usually brown.Scapus and pedicellus bright yellow, contrasting with flagellum which is usually largely black but with variable paler brown-yellow to yellow zone from approximately middle to near the apex.Wing membrane yellowish on basal half, pale grey distally; pterostigma bicolorous, dark brown distally but with large yellow zone basally and posterobasally.
Etymology.Named after the known host genus.Head.Antenna with 56-57 flagellomeres.Terminal flagellomere acuminate.Median flagellomeres approximately 2 × longer than wide.3 rd segment of maxillary palp of female weakly swollen, approximately 6 longer than maximally wide.5 th segment approximately 0.8 length of 6 th .Base of mandible very close to eye, closest point approximately same as distance from middle of anterior tentorial pit and eye.Distance between anterior tentorial pits approximately 10 × shortest distance between pit and eye.Width of face: width of head across eyes: height of eye = 1.0 : 3.3 : 2.0.Face with coarse transverse striae ventrolaterally, becoming finer and reaching near to midline dorsally.Frons with rather well developed ridge paralleling margin of eye; generally with weak sublongitudinal sculpture Shortest distance between posterior ocelli: transverse diameter of posterior ocellus: shortest distance between posterior ocellus and eye = 1.0: 3.5: 1.7.Occiput with deep crescent-shaped depressions behind posterior ocelli; with only weak midlongitudinal groove.Occipital carina complete and strongly lamelliform.
Hind wing.Vein M+CU 2.5 × length of 1-M.Vein 2-SC+R very thick, short transverse.Vein SR gently curving basally and gradually getting closer to anterior margin towards wing tip.
Etymology.Named after collection locality as a noun in apposition.Biology.Host unknown for holotype.

Colastomion cheesmanae
Coloration.Head, hind trochanter and trochantellus and metasomal tergite 1 posteriorly, 2-4 entirely, anterior half of tergite 5, and extreme base of tergite 6 black, hind tibia dark brown, mesosoma and rest of legs orange red, metasomal tergites 5 posteriorly and 6 largely cream-white.Antenna largely black with a distinct brown-yellow zone occupying flagellomeres 22-34.Wing membrane largely pale brown, venation largely (except more basal veins, dark brown; pterostigma dark brown. Biology.Unknown.Etymology.Named in honour of Miss L. E. Cheesman who collected much interesting material in New Guinea in expeditions in the 1930s (see Kimmins 1962).
Notes.Very similar to C. wanang sp.n. except lacks the thickened and strongly curved fore wing vein M+CU.Morphology.Length of body 10.0 mm, of fore wing 8.5-9.5 mm and of antenna 9.5-11.2mm.

Colastomion pukpuk
Head.Antenna with 56 flagellomeres.Terminal flagellomere strongly acuminate.Median flagellomeres 2 × longer laterally than wide.3 rd segment of maxillary palp of female moderately swollen, approximately 4 longer than maximally wide.5 th segment approximately 0.9 length of 6 th .Base of mandible close to eye, closest point approximately 0.75 distance from middle of anterior tentorial pit and eye.Distance between anterior tentorial pits approximately 6 × shortest distance between pit and eye.Width of face: width of head across eyes: height of eye = 1.0 : 3.1 : 2.0.Face with a smooth triangular area above clypeus bordered by well developed transverse striation that more or less meets medially at level of antennal sockets.Shortest distance between posterior ocelli: transverse diameter of posterior ocellus: shortest distance between posterior ocellus and eye = 1.0: 2.0: 1.2.Occipital carina complete, rather wavy when viewed laterally.
Etymology.Named after Nicholai Miklucho-Maklaj, a Russian explorer of New Guinea who lived around the Madang area for some time.

Discussion
The species described here comprise three groups based both on morphology and DNA (Fig. 1).Colastomion gregarious sp.n., C. madangensis sp.n. and C. masalaii sp.n. are all small, at least facultatively gregarious species with an antefurcal fore wing vein cu-a (Fig. 15) and an inverted 'V'-shaped carina on the propodeum (Fig. 7); C. crambidiphagus sp.n. and C. parotiphagus sp.n. are medium-sized, predominantly yellow species; C. cheesmanae sp.n., C. maclayi sp.n., C. pukpuk sp.n. and C. wanang sp.n. are all large, black headed species with largely black metasomas with whitish tips.The amount of morphological variation observed between these three groups of species is considerable, and might in the past have been deemed sufficient by some taxonomists to warrant division into different genera or subgenera, and the relatively long internal branches on the molecular phylogeny support their high level of differentiation.Addition molecular studies on the whole subfamily will be needed to determine whether, despite the vertical hind wing vein 2-SC+R, these taxa form a natural group to the exclusion of other members of the complex.
As with many rogadine wasps, Colastomion had no published host records prior to the current caterpillar rearing campaign in Papua New Guinea, despite it being widely distributed and occurring also in tropical Africa and S. E. Asia.In common with the closely related genus Macrostomion Baker, several of the new species described here are gregarious.This biology is unusual among the Rogadinae.A few species of the cosmopolitan genus Aleiodes Wesmael are gregarious (specifically A. pallescens on various Notodontidae, A. stigmator (Say) on Noctuidae, and A. leptocarina Fortier on an unidentified hairy host caterpillar).Another aspect of the gregariousness exhibited by some Colastomion is that the brood size is rather small, and at least one of the new species, C. masalaii sp.n., is facultatively gregarious.Thus they may make interesting study organisms for investigating the evolutionary transition from solitary to gregarious life histories (Mayhew and van Alphen 1999, Mayhew and Glaizot 2001, Guinee et al. 2005).
It is interesting that all the reared Colastomion species are parasitoids of Crambidae, though these collectively feeding on a wide range of host plant families.Few tropical genera of Rogadinae have multiple host records, but of those that do, several appear to be relatively restricted in the range of host families attacked suggesting at least some degree of co-evolutionary tuning between host defence mechanisms and the parasitoid's ability to overcome them.However, before this can be investigated much further, a far larger body of host-parasitoid association data will be required.

Figure 1 .
Figure 1.Maximum likelihood tree (using GTR+G parameter model) of the reared and barcoded species of Colastomion (final Gamma -based likelihood -2578.386754).
Collection data, sex, host and host plant associations of paratypes of Colastomion parotiphagus sp.n.

Quicke, sp. n.
Antenna largely black with paler brown zone approximately occupying flagellomeres 22-39.Wing membrane largely pale brown, venation largely (except more basal veins, dark brown; pterostigma dark brown.Etymology.Pukpuk in Melanesian Pidgin means crocodile, a favourite of the indigenous people living around Sepik river.Biology.Unknown.Antenna with 57 flagellomeres.Terminal flagellomere distinctly acuminate.Median flagellomeres 1.8 × longer laterally than wide.3 rd segment of maxillary palp of female very weakly swollen medially swollen, approximately ?7 × longer than maximally wide.5 th segment approximately 0.65 length of 6 th .Base of mandible very close to eye, closest point approximately 0.8 distance from middle of anterior tentorial pit and eye.Distance between anterior tentorial pits approximately 5.0 × shortest distance between pit and eye.Width of face: width of head across eyes: height of eye = 1.0 : 2.7