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New reared rogadine wasp species from New Guinea 85
Colastomion Baker (Braconidae, Rogadinae): nine new
species from Papua New Guinea reared from Crambidae
Donald L.J. Quicke
1,2,†
, M. Alex Smith
3,‡
, Scott E. Miller
4,§
, Jan Hrcek
5,|
,
Buntika Butcher
6,¶
1 Department of Biological Sciences, Imperial College London Silwood Park Campus, Ascot, Berkshire SL5
7PY, UK 2 Department of Entomology, e Natural History Museum, London SW7 5BD, UK 3 Department
of Integrative Biology & the Biodiversity Institute of Ontario, University of Guelph, Guelph, Ontario N1G
2W1, Canada 4 National Museum of Natural History, Smithsonian Institution, PO Box 37012, MRC 105
Washington, DC 20013-7012, USA 5Faculty of Science, University of South Bohemia and Biology Center,
Czech Academy of Sciences, Branisovska 31, 37005 Ceske Budejovice, Czech Republic 6 Department of Bio-
logy, Faculty of Science, Chulalongkorn University, BKK 10330, ailand
† urn:lsid:zoobank.org:author:4FAA3C17-13D3-467B-AAA5-82B27FE783EF
‡ urn:lsid:zoobank.org:author:E46EE6EB-E096-4FCD-BF5A-F91D4A8294EE
§ urn:lsid:zoobank.org:author:D70079F1-5135-46BA-A790-CB833B5892EF
| urn:lsid:zoobank.org:author:31771CB4-A88A-4BF6-ACF0-28F9564D2692
¶ urn:lsid:zoobank.org:author:3AC486D1-9045-44FB-9FE4-FF71A0C2909C
Corresponding author: Dr. Donald L. J. Quicke (d.quicke@imperial.ac.uk)
Academic editor: Gavin Broad|Received 7 July 2012|Accepted 13 August 2012|Published 24 August2012
urn:lsid:zoobank.org:pub:1352252C-D511-44A1-A2D0-B74E00C7D616
Citation:
Quicke DLJ, Smith MA, Miller SE, Hrcek J, Butcher B (2012) Colastomion Baker (Braconidae, Rogadinae):
nine new species from Papua New Guinea reared from Crambidae. Journal of Hymenoptera Research 28: 85–121. doi:
10.3897/JHR.28.3484
Abstract
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 eld col-
lected 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. e 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
Keywords
cytochrome oxidase I, DNA barcoding, Lepidoptera, hosts, Rogadini, Crambidae
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 Cysto-
mastacoides van Achterberg, Macrostomion Szépligeti, Megarhogas Szépligeti and Myo-
cron 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 mummied host
larva (Shaw 2002, Maeto and Arakaki 2005).
To date, only six species of Colastomion have been described, viz. C. abdominale
Baker (1917) from the Philippines, C. bicoloricorne (Granger 1949) from Madagas-
car, C. concolor (Szépligeti 1911) originally described from Tanzania and subsequently
reported from Democratic Republic of Congo, Madagascar, Malawi and Togo (Yu et
al. 2005), C. formosanum Watanabe (1932) originally described from Taiwan, but re-
cently recorded from Japan (Tenma 2002), C. nigricorne (Granger 1949) and C. tristis
(Granger 1949) both from Madagascar.
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 Pap-
ua New Guinea. ese were obtained during an extensive caterpillar collecting and
rearing programme which has additionally yielded a number of other interesting roga-
dine parasitoids including the recently described, highly distinctive genus Vojtechirogas
Quicke & van Achterberg (Quicke et al. 2012).
e host records for some of the species include a degree of fuzziness because iden-
tications are based on caterpillar morphology. e 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 rear-
ing 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 identication. Two rearing records for C.
masalaii sp. n. and C. parotiphagus sp. n. were conrmed by sequencing host remnants
(Hrcek et al. 2011).
New reared rogadine wasp species from New Guinea 87
e 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. us 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 (1979, 1988). Measurements of the height and
horizontal length of the eye are approximations because the very bulbous face and very
large eyes make it dicult to measure consistently; in our attempts to do so, the head
was orientated so that the face (dened as running from the anterior edge of the toruli
to the dorsal margin of the clypeus) was horizontal or perpendicular to the measure-
ment axis.
Abbreviations: NHM (e 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 max-
imum 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 Biodiver-
sity 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 Light-
room software and edited using Adobe Photoshop CS4.
Results
DNA barcodes were generated from all but one of the species described below, the ex-
ception 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 intraspecic
variation was limited to two individuals of C. masalaii sp. n. (vouchers USNM ENT
00503254 and 00643295) diering from the remainder at a single base position.
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
88
Figure 1. Maximum likelihood tree (using GTR+G parameter model) of the reared and barcoded species
of Colastomion (nal Gamma –based likelihood -2578.386754).
Systematics
Key to species
1 Fore wing vein cu-a antefurcal (Figs 5, 15, 21); propodeum with short mid-
longitudinal carina anteriorly, giving rise to a midlongitudinal depression
with transverse crenulae or transverse rugosity (Fig. 7); antenna with fewer
than 42 [31–39] agellomeres (Fig. 2); hind wing vein M+CU shorter [0.6–
0.9 ×] than 1-M; largely yellow or yellow brown, metasoma the same colour
as mesosoma ...............................................................................................2
– Fore wing vein cu-a postfurcal; propodeum with a complete midlongitudinal
carina or if incomplete, replaced by rugosity on posterior half; antenna with
more than 43 [45–57]; agellomeres; hind wing vein M+CU longer [1.1–2.5
×] than 1-M; colour variable, sometimes metasoma black and cream contrast-
ing with yellow-brown mesosoma ...............................................................4
2 Antenna with fewer than 35 [32] agellomeres (Fig. 2); 3
rd
segment of female
maxillary palp distinctly swollen, approximately 4.5 × longer than maximally
New reared rogadine wasp species from New Guinea 89
wide; pterostigma entirely pale yellow (Fig. 2); mesosoma brown yellow dor-
sally with posterior of pronotum, mesopleuron, mesosternum and metapleu-
ron whitish (Fig. 4) ..........................................................C. gregarius sp. n.
– Antenna with more than 35 [37–40] agellomeres; 3
rd
segment of female
maxillary palp slender, approximately 7 × longer than maximally wide; pter-
ostigma either bicolorous (Fig. 21) or entirely dark grey (Fig. 15); mesosoma
unicolorous, either entirely pale yellow or brown yellow (Fig. 13) ............... 3
3 Pterostigma largely dark brown; antenna largely dark brown (Fig. 15); fore
wing vein cu-a strongly inclivous (Fig. 15) ........................C. masalaii sp. n.
– Pterostigma bicolorous, cream on basal 0.3 and on anterior margin, remain-
der brown (Fig. 21); antenna cream coloured; fore wing vein cu-a vertical
(Fig. 21) .....................................................................C. madangensis sp. n.
4 Metasoma almost entirely brown-yellow, sometimes with basal grooves of ter-
gites brown medially (Figs 29, 30); 5
th
segment of female maxillary palp less
than 0.6 [0.5 ×] 6
th
segment (Fig. 32) .........................................................5
– Metasomal tergites 3–4 largely black (Figs 39, 47, 51, 52); 5
th
segment of
female maxillary palp more than 0.6 [0.7–0.9 ×] 6
th
segment (Fig. 54) .......6
5 Flagellum entirely yellow; metasomal tergites 3–5 pale yellow with distinctly
brown yellow mark narrowly mediobasally, tergite 6 somewhat more exten-
sively brown yellow medially (Fig. 29) ................. C. crambidiphagus sp. n.
– Flagellum black on basal half becoming paler, sometimes yellow, from about
middle to shortly before tip, contrasting strongly with pale yellow scapus and
pedicellus (Fig. 31); metasomal tergites more or less evenly coloured pale yel-
low (Fig. 30) ............................................................. C. parotiphagus sp. n.
6 Fore wing vein M+CU thickened up to near its middle and sharply narrow-
ing and curved beyond this (Figs 41, 42) ...........................C. wanang sp. n.
– Fore wing vein M+CU more or less evenly thick and straight or only weakly
curved (Figs46, 50, 52) ...............................................................................7
7 Anterior of propodeum strongly sculptured, rugose (Fig. 47, see also Fig. 38);
2
nd
subdiscal cell rather widened distally, maximum length of cell membrane
< 3.5 [3.1] × maximum width (see Fig. 41) ................. C. cheesmanae sp. n.
– Anterior of propodeum on either side of midlongitudinal carina largely
smooth with punctures or at most with narrow crenulated groove next to
carina (Figs 49, 54); 2
nd
subdiscal cell long and narrow, maximum length of
cell membrane < 3.7 [4.0] × maximum width (Fig. 50) ..............................8
8 1
st
tergite in lateral prole distinctly deeper near midlength (Fig. 55) ............
.......................................................................................... C. pukpuk sp. n.
– 1
st
tergite in lateral prole deepest on anterior third and behind this at or
weakly depressed (Fig. 56) .................................................C. maclayi sp. n.
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
90
Descriptions of new species
Colastomion gregarius Quicke, sp. n.
urn:lsid:zoobank.org:act:6C64E6C5-8CB9-48B3-9A92-1475E7779B24
http://species-id.net/wiki/Colastomion_gregarius
Figs 2–8
Material examined. Holotype female, Papua New Guinea, Madang Province, Wa-
nang, 24-May-07, 145°10.910'E, 5°13.853'S, 100m, ex caterpillar on Ficus variegata
Blume (Moraceae) (voucher USNM ENT 00680021; BOLD ASQSP084-08; Gen-
bank JF963128) [furthest specimen from pin on topmost card] (USNM)
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).
Morphology. Length of body 4.0–4.8 mm, of fore wing 3.0–3.6 mm and of an-
tenna 5.1–5.2 mm.
Head. Antenna with 31–32 agellomeres. Terminal agellomere acuminate. Me-
dian agellomeres 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. Dis-
tance 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 ne transverse striation laterally above level of clypeus. Short-
est 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 lon-
gitudinally striate-rugulose at conuence 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.
Fore wing. Vein 1-CU1 antefurcal. Vein 2-CU1 6.0 × longer than 1-CU1. Vein
1-M weakly curved. Lengths of veins SR1: 3-SR: r = 6.0: 2.7 : 1.0. Vein 2-SR 1.0 ×
2-SR+M.
Hind wing. Vein M+CU 0.9 × length of 1-M. Vein 2-SC+R thickened and dis-
tinctly inclivous. Vein SR moderately strongly curved on basal half, almost reaching
wing margin approximately half distance to wing tip, then running more or less paral-
lel to wing margin.
Metasoma. 1
st
metasomal tergite 1.7 × longer than posteriorly wide. 2nd meta-
somal tergite 1.6 × wider than long, with wide midbasal triangular area giving rise
New reared rogadine wasp species from New Guinea 91
Figures 2–5. Colastomion gregarius sp. n. light photomicrographs. 2 Habitus, lateral view 3 face 4 head
and mesosoma, lateral view 5 basal venation of fore wing.
to weak midlongitudinal carina; rather weakly irregularly longitudinally striate with
approximately 7 striate lateral to midlongitudinal carina. 3rd metasomal tergite 1.9 ×
wider than long; weakly irregularly longitudinally rugulose-striate basally and medially.
4
th
- 6
th
tergite s smooth.
Coloration. Body, legs, wing venation and antennae largely pale ochreous yellow
with malar region, posteroventral part of pronotum, propleuron, mesopleuron and
mesosternum, metapleuron, sides of tergites 2–6 and fore and mid coxa, trochanter
and trochantellus whitish; terminal 6 or 7 agellomeres dark grey-black.
Biology. Gregarious parasitoid of Glyphodes near stolalis.
Etymology. Based on gregarious biology.
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
92
Colastomion masalaii Quicke, sp. n.
urn:lsid:zoobank.org:act:0C6B94FD-975C-435A-A554-171939C4D138
http://species-id.net/wiki/Colastomion_masalaii
Figs 9–16
Material examined. Holotype female, East Sepik Province, Wamangu, 143°39.125’E,
03° 47.228'S, 100m, 30-Mar-05, ex caterpillar on Ficus bernaysii King (Moraceae),.
(voucher USNM ENT 00503254; BOLD ASQSP055-08; Genbank JF271305) [fur-
Figures 6–8. Colastomion gregarius sp. n. light photomicrographs. 6 Head and anterior mesosoma, dor-
sal view 7 propodeum 8 metasoma.
New reared rogadine wasp species from New Guinea 93
thest individual from pin on card]. Host remnants were sequenced (BOLD ASPN766-
09, Genbank JF271356) and the sequence identied as Glyphodes margaritaria.
Paratypes. 2 additional females on same card and from same brood as holotype; see
Table 1 for all data on 30 additional rearings totalling 33 paratypes. Specimens USNM
ENT 00491800, USNM ENT 00491811 and USNM ENT 00648414 are deposited
in the BMNH.
Figures 9–12. Colastomion masalaii sp. n. holotype and paratypes, Cell^D® light photomicrographs.
9 Brood including holotype (furthest individual from pin on card) and two female paratypes 10 habitus,
dorsal view 11 face, oblique view 12 head and mesosoma, dorsal view.
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
94
Table 1. Collection data, sex, host and host plant associations of paratypes of Colastomion masalaii sp. n.
USNM ENT
voucher
number
BOLD process
ID
Genbank
accession
number
No.
and
sex
Province Locality
a
Latitude and longitude Collection
date
Host identication Host plant
00209066 - - 4F Madang Baitabag 145°47'E, 5°08'S 20-Nov-02 Crambidae Ficus molior F. Meull. ex
Benth. (Moraceae)
00211702 - - 1F East Sepik Elem 143°55'E, 4°49'S 16-Apr-03 Crambidae Ficus conocephalifolia Ridley
(Moraceae)
00491773 - - 1M Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00491774 - - 1F Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00491775 - - 1F Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00491776 - - 1F Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00491790 - - 1F Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00491794 ASQSP126-08 JF271307 1F Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00491795 ASQSP125-08 JF271308 1M Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00491799 - - 1F Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00491800 ASQSP128-08 JF271306 1F Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00491801 - - 1F Sandaun Utai 141°35.153'E, 3°23.043'S 30-Jul-04 Talanga sexpunctalis
complex
— ditto —
00491807 - - 1F Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 no host information — ditto —
00491808 - - 1F Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00491809 - - 1F Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00491810 - - 1F Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00491811 - - 1F Sandaun Utai 141°35.153'E, 3°23.043'S 28-Jul-04 Crambidae — ditto —
00506254 - - 4F Madang Ohu 145°41'E, 5°14'S 12-Jul-04 Glyphodes
margaritaria
(Cramer)
Ficus dammaropsis Diels
00643279 - - 1F Madang Wanang 145°10.910'E, 5°13.853'S 26-Apr-05 Crambidae Ficus conocephalifolia Ridley
00643281 - - 1F Madang Wanang 145°10.910'E, 5°13.853'S 26-Apr-05 Crambidae — ditto —
00643282 - - 1F Madang Wanang 145°10.910'E, 5°13.853'S 26-Apr-05 Crambidae — ditto —
New reared rogadine wasp species from New Guinea 95
USNM ENT
voucher
number
BOLD process
ID
Genbank
accession
number
No.
and
sex
Province Locality
a
Latitude and longitude Collection
date
Host identication Host plant
00643285 - - 1F Madang Wanang 145°10.910'E, 5°13.853'S 26-Apr-05 Crambidae — ditto —
00643293 - - 1F Madang Wanang 145°10.910'E, 5°13.853'S 26-Apr-05 Crambidae — ditto —
00643294 - - 1F Madang Wanang 145°10.910'E, 5°13.853'S 26-Apr-05 Crambidae — ditto —
00643295 ASQSP059-08 JF271309 1M Madang Wanang 145°10.910'E, 5°13.853'S 26-Apr-05 Crambidae — ditto —
00643298 - - 1F Madang Wanang 145°10.910'E, 5°13.853'S 26-Apr-05 Crambidae — ditto —
00643299 - - 1F Madang Wanang 145°10.910'E, 5°13.853'S 26-Apr-05 Crambidae — ditto —
00648414 - - 1F Madang Wanang 145°10.910'E, 5°13.853'S 26-Apr-05 Crambidae — ditto —
00648415 - - 1F Madang Wanang 145°10.910'E, 5°13.853'S 26-Apr-05 Crambidae — ditto —
00648419 - - 1F Madang Wanang 145°10.910'E, 5°13.853'S 26-Apr-05 Crambidae — ditto —
a
Ohu is 200 m above mean sea level, other localities are 100 m above mean sea level
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
96
Figures 13–16. Colastomion masalaii sp. n. Cell^D® light photomicrographs. 13 Mesosoma, lateral view
14 propodeum and 1
st
tergite 15 fore wing 16 metasoma.
Morphology. Length of body 5.5–6.5 mm, of fore wing 48–5.0 mm and of an-
tenna 7.0 mm.
Head. Antenna with 35–37 agellomeres. Terminal agellomere pointed, not or
hardly acuminate. Median agellomeres 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, clos-
est point approximately 1.2 distance from middle of anterior tentorial pit and eye. Dis-
tance between anterior tentorial pits approximately 3.0 × shortest distance between pit
New reared rogadine wasp species from New Guinea 97
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
conuence. 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.
Fore wing. Vein 1-CU1 antefurcal. Vein 2-CU1 9 × longer than 1-CU1. Lengths
of veins SR1: 3-SR: r = 5.5: 3.5: 1.0. Vein 2-SR 2.4 × 2-SR+M.
Hind wing. Vein M+CU 0.7 × length of 1-M.
Metasoma. 1
st
metasomal tergite 1.6–1.9 × longer than posteriorly wide. 2nd metaso-
mal tergite 1.25 × wider than long, with well developed midbasal triangular area giving rise
to weak midlongitudinal carina; irregularly longitudinally striate with approximately 15
striae lateral to midlongitudinal carina. 3rd metasomal tergite 1.9 × wider than long; lon-
gitudinally striate without distinct midlongitudinal carina. 4
th
- 6
th
tergites almost smooth.
Coloration. Body largely pale brown yellow; scapus and pedicellus yellow, agel-
lum 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.
Biology. Solitary and gregarious (3 or 4 individuals per brood) endoparasitoids of
Talanga sexpunctalis complex and Glyphodes margaritaria (Cramer) (both Crambidae)
feeding on Ficus spp (Moraceae). T. sexpunctalis is usually considered a widespread spe-
cies but appears to be a complex of species, including several in New Guinea (Craft et
al. 2010: S2). G. margaritaria is a widespread species (Craft et al. 2010: 5043 and S2).
Colastomion madangensis Quicke, sp. n.
urn:lsid:zoobank.org:act:F30DD23D-34AE-4663-AC71-0A520491E039
http://species-id.net/wiki/Colastomion_madangensis
Figures 17–22
Material examined. Holotype female, Papua New Guinea, Madang Province, Wanang,
24-May-07, 145°10.910'E, 5°13.853'S, 100m, ex caterpillar on Ficus variegata Blume
(Moraceae) (vouchers USNM ENT 00680132; BOLD ASQSP177-08; Genbank
JX034716) [individual furthest from pin on upper card] [host Glyphodes near stolalis].
Paratypes: 5 females, 2 males, from same brood as holotype, mounted on 3 cards on
same pin; 6 females, 3 males mounted on 2 cards on same pin, Madang Province, Wanang,
10-May-06, 145°10.910'E, 5°13.853'S, 100m, ex caterpillar on Ficus variegata Blume,
(voucher USNM ENT 00680061; BOLD ASQSP081-08; Genbank JF271301); 9 fe-
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
98
Figures 17–20. Colastomion madangesis sp. n. holotype, Cell^D® light photomicrographs. 17 Habitus,
dorsal view 18 face, showing slender 3
rd
segment of maxillary palp 19 propodeum 20 head and anterior
mesosoma, dorsal view.
males, 3 males, mounted on three cards on same pin, Madang Province, Wanang, 24-May-
07, 145°10.910'E, 5°13.853'S, 100m, ex caterpillar on Ficus variegata Blume, Morace-
ae, (voucher USNM ENT 00680102; BOLD ASQSP083-08; Genbank JX034721)
[host Glyphodes near stolalis]; 10 individals, Madang Province, Wanang, 10-May-06,
145°10.910'E, 5°13.853'S, 100m, ex caterpillar on Ficus variegata Blume, Moraceae,
(voucher USNM ENT 00680111; BOLD ASQSP139-08; Genbank JX034722).
New reared rogadine wasp species from New Guinea 99
Morphology. Length of body 4.3–5.5 mm, of fore wing 3.0–4.1 mm and of an-
tenna 4.5–5.0 mm.
Head. Antenna with 35–39 agellomeres. Terminal agellomere acuminate. Me-
dian agellomeres 2.3 × longer than wide. 3
rd
segment of maxillary palp of female
slender, approximately 7 longer than maximally wide. 5
th
segment slender and approxi-
mately same length of 6
th
. Base of mandible well separated from eye, closest point
approximately 1.1 distance from middle of anterior tentorial pit and eye. Distance be-
Figures 21–22. Colastomion madangesis sp. n. holotype, Cell^D® light photomicrographs. 21 Wings
22 metasoma.
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
100
tween anterior tentorial pits approximately 5.2 × shortest distance between pit and eye.
Width of face: width of head across eyes: height of eye = 1.0 : 2.25 : 1.36. Face later-
ally with well developed transverse rugosity. Shortest distance between posterior ocelli:
transverse diameter of posterior ocellus: shortest distance between posterior ocellus
and eye = 1.0: 1.0 : 1.0. Occipital carina complete but distinctly weak mediodorsally
Mesosoma. Notauli more or less strongly impressed and crenulated. Mesoscutum
rugulose at conuence of notauli. Mesopleuron largely smooth and shiny. Precoxal
sulcus strongly impressed, weakly sinuate, strongly nely crenulated. Propodeum with
short midlongitudinal carina anteriorly that splits giving rise to a widening midlongi-
tudinal depression with moderately well developed transverse striation.
Fore wing. Vein 1-CU1 antefurcal. Vein 2-CU1 7 × longer than 1-CU1. Vein 1-M
weakly curved. Lengths of veins SR1: 3-SR: r = 4.5:2.7:1.0. Vein 2-SR 2.8 × 2-SR+M.
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 approxi-
mately 0.6 distance to wing tip, then running more or less parallel to wing margin.
Metasoma. 1
st
metasomal tergite 1.4 × longer than posteriorly wide. 2nd metaso-
mal tergite 1.35 × wider than medially long, with well developed midbasal triangular
area giving rise to complete midlongitudinal carina; irregularly longitudinally striate
with approximately 8 striate lateral to midlongitudinal carina. 3rd metasomal tergite
1.9 × wider than long; longitudinally striate, without distinct midlongitudinal carina.
4
th
tergite distinctly irregularly longitudinally striate, 5
th
tergite more weakly striate and
largely only on basal 0.5; 6
th
tergite smooth.
Coloration. Body largely brown yellow, metasoma somewhat more reddish; anten-
nae 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.
Biology. Gregarious parasitoid of Glyphodes near stolalis Guenee 1854, caterpillars:
the broods reared comprised members of both sexes. Glyphodes near stolalis is a member
of a species complex needing revision (Munroe 1996: 280), complicated by the type of
stolalis being lost (Viette 1987) (Hrcek et al. 2011: supplement).
Colastomion crambidiphagus Quicke, sp. n.
urn:lsid:zoobank.org:act:ACC78520-7B06-49C3-83D1-2701CC861913
http://species-id.net/wiki/Colastomion_crambidiphagus
Figs 23–29
Material examined. Holotype male, Papua New Guinea, Madang Province, Wanang,
12-Apr-07, 145°10.910'E, 5°13.853'S, 100m, WP-5E-2952, ex caterpillar on Mer-
remia peltata (L.) Merr. (Convolvulaceae) (voucher USNM ENT 00680146; BOLD
ASQSP086-08; Genbank JX034720).
New reared rogadine wasp species from New Guinea 101
Figures 23–26. Colastomion crambidiphagus sp. n. holotype, Cell^D® light photomicrographs. 23 Habi-
tus, lateral view 24 face 25 head and anterior mesosoma, dorsal view 26 propodeum.
Paratypes: 1 female, Papua New Guinea, Madang Province, Wanang, 16-Feb-2007,
145°10.910'E, 5°13.853'S, 100m, ex caterpillar feeding on Merremia peltata (L.) Merr.
(Convolvulaceae) (voucher USNM ENT 00680134; BOLD ASQSP135-08; Gen-
bank JF963127); 1 female, Papua New Guinea, Madang Province, Wanang, 20-Jun-
06, 145°10.910'E, 5°13.853'S, 100m, ex caterpillar feeding on Merremia peltata (L.)
Merr. (Convolvulaceae) (voucher USNM ENT 00680173, no sequence data).
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
102
Figures 27–29. Colastomion crambidiphagus sp. n. holotype, Cell^D® light photomicrographs. 27 Wings
28 head and mesosoma, lateral view 29 metasoma.
New reared rogadine wasp species from New Guinea 103
Morphology. Length of body 6.0–8.5 mm, of fore wing 5.0–6.5 mm and of an-
tenna 8.0–8.5 mm.
Head. Antenna with 45 agellomeres. Terminal agellomere strongly acuminate.
Median agellomeres 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 ap-
proximately 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 poste-
rior 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 pos-
teriorly. Propodeum with midlongitudinal carina at least on anterior 0.5, posteriorly ei-
ther complete with strong transverse rugae lateral to it, or replaced by transverse rugosity.
Fore wing. Vein 1-CU1 postfurcal. Vein 2-CU1 18–19 × longer than 1-CU1.
Lengths of veins SR1: 3-SR: r = 4.5 : 2.7 : 1.0. Vein 2-SR 2.1 × 2-SR+M.
Hind wing. Vein M+CU 1.1 × length of 1-M. Vein 1-SC+R slightly thickened, al-
most transverse. Vein SR strongly curved on basal half, almost reaching wing margin ap-
proximately half distance to wing tip, then running more or less parallel to wing margin.
Metasoma. 1
st
metasomal tergite 2.0 × longer than posteriorly wide. 2nd metaso-
mal tergite 1.1 × longer than maximally wide, with well developed midbasal triangular
area giving rise to complete midlongitudinal carina; irregularly longitudinally striate
with approximately 6 striate lateral to midlongitudinal carina. 3rd metasomal tergite
1.1 × longer than maximally wide; longitudinally striate. 4
th
and 5
th
tergites longitudi-
nally striate. 6
th
tergite smooth.
Coloration. Body.
Etymology. Named because of its parasitism of Crambidae.
Biology. Solitary endoparasitoids of Crambidae feeding on Merremia peltata (L.)
Merr. (Convolvulaceae). e holotype was reared from Hydriris guadealis Rothschild.
Paratype USNM ENT 00680134 was reared from Tabidia insanalis Snellen, and para-
type USNM ENT 00680173 was reared from.a further unidentied crambid.
Colastomion parotiphagus Quicke, sp. n.
urn:lsid:zoobank.org:act:BE5DD9B9-5DB2-4C52-A95C-ED2FE0E76FD0
http://species-id.net/wiki/Colastomion_parotiphagus
Figs 30–32
Material examined. Holotype male, Papua New Guinea, Madang Province, Wanang,
30-May-07, 145°10.910'E, 5°13.853'S, 100m, WS-2D-0932, ex caterpillar on Sar-
cocephalus coadunatus (Sm.) Druce (Rubiaceae), CATX043 (voucher USNM ENT
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
104
00680014; BOLD ASQSP026-08; Genbank JX034709) [Host – Parotis tricoloralis
(Pagenstecher)].
For details of 34 paratypes see Table 2.
Figures 30–32. Colastomion parotiphagus sp. n. holotype, Cell^D® light photomicrographs. 30 Habitus,
dorsal view 31 head and anterior mesosoma, dorsal view 32 head and mesosoma, lateral view.
New reared rogadine wasp species from New Guinea 105
For details of 34 paratypes see Table 2. Specimens USNM ENT 00680067, USNM
ENT 00680154 and USNM ENT 00690187 are deposited in the BMNH.
Morphology. Length of body 6.2–9.5 mm, of fore wing 5.8–7.5 mm and of an-
tenna 7.5–9.5 mm.
Head. Antenna with 50 agellomeres. Terminal agellomere acuminate. Median
agellomeres 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 ap-
proximately half length of 6
th
. Base of mandible very close to eye, closest point approxi-
mately 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, nely 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 fu-
sion. Propodeum with complete midlongitudinal carina. Mesopleuron largely smooth.
Precoxal sulcus narrow and deep on posterior half of mesopleuron, nely crenulated.
Propodeum with complete midlongitudinal carina.
Fore wing. Vein 1-CU1 postfurcal. Vein 2-CU1 6.4 × longer than 1-CU1. Lengths
of veins SR1: 3-SR: r = 4.0: 1.6 :1.0. Vein 2-SR 2.0 × 2-SR+M.
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.
Metasoma. 1
st
metasomal tergite 2.2 × longer than posteriorly wide. 2nd metasomal
tergite as long as maximally wide, with well developed midbasal triangular area giving
rise to complete midlongitudinal carina; irregularly longitudinally striate with approxi-
mately 8 striate lateral to midlongitudinal carina. 3rd metasomal tergite 1.4 × wider
than long; longitudinally striate and with midlongitudinal carina more or less dier-
entiated. 4
th
and 5
th
tergites longitudinally striate. 6
th
tergite smooth (faintly aciculate).
Coloration. Head and body ochreous yellow, stemmaticum black, hypopygium
usually brown. Scapus and pedicellus bright yellow, contrasting with agellum which
is usually largely black but with variable paler brown-yellow to yellow zone from ap-
proximately 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.
Biology. Solitary larval endoparasitoids of Parotis tricoloralis (Pagenstecher)
(Crambidae), Parotis hilaralis (Walker) (Crambidae) and Haritalodes adjunctalis Leraut
(Crambidae).
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
106
Table 2. Collection data, sex, host and host plant associations of paratypes of Colastomion parotiphagus sp. n.
USNM ENT
voucher
number
BOLD
process ID
Genbank
accession
number
Sex Province Locality
a
Latitude and longitude Collection
date
Host identication Host plant
00206476 - - F Madang Baitabag 145°47'E, 5°08'S 04-Jun-02 Haritalodes adjunctalis
Leraut
Hibiscus tiliaceus L.
(Malvaceae)
00680001 - - M Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) Sarcocephalus coadunatus
(Sm.) Druce (Rubiaceae)
00680023 - - F Madang Wanang 145°10.910'E, 5°13.853'S 23-May-07 Parotis tricoloralis (Pag.) — ditto —
00680026 ASQSP094-08 JX034711 M Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680031 ASQSP090-08 JX034723 M Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680035 - - F Madang Wanang 145°10.910'E, 5°13.853'S 24-Feb-06 Parotis tricoloralis (Pag.) — ditto —
00680040 ASQSP137-08 JX034713 M Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680056 ASQSP018-08 JF271317 M Madang Wanang 145°10.910'E, 5°13.853'S 23-May-07 Parotis tricoloralis (Pag.) — ditto —
00680057 - - F Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680060 - - M Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680067 - - M Madang Wanang 145°10.910'E, 5°13.853'S 23-Feb-06 Parotis tricoloralis (Pag.) — ditto —
00680075 - - F Madang Wanang 145°10.910'E, 5°13.853'S 12-May-06 Parotis tricoloralis (Pag.) — ditto —
00680082 ASQSP091-0 JX034710 M Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680085 - - ? Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680086 ASQSP089-08 JX034714 F Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680088 ASQSP092-08 JX034718 F Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680093 - - M Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680097 ASQSP020-08 JF271315 F Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680103 - - M Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680106 - - Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680108 ASQSP136-08 JX034717 M Madang Wanang 145°10.910'E, 5°13.853'S 13-May-06 most likely Parotis — ditto —
00680117 ASQSP082-08 JF963129 F Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680124 - - M Madang Wanang 145°10.910'E, 5°13.853'S 12-May-06 Parotis tricoloralis (Pag.) — ditto —
New reared rogadine wasp species from New Guinea 107
USNM ENT
voucher
number
BOLD
process ID
Genbank
accession
number
Sex Province Locality
a
Latitude and longitude Collection
date
Host identication Host plant
00680128 ASQSP025-08 JF271314 ? Madang Wanang 145°10.910'E, 5°13.853'S 23-May-07 Parotis tricoloralis (Pag.) — ditto —
00680130 ASQSP027-08 JX034715 F Madang Wanang 145°10.910'E, 5°13.853'S 21-Jun-07 no host information Actinodaphne nitida
Teschn. (Lauraceae)
00680131 ASQSP138-08 JF271318 F Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) Sarcocephalus coadunatus
(Sm.) Druce (Rubiaceae)
00680142 - - ? Madang Wanang 145°10.910'E, 5°13.853'S 31-May-07 No host information Trema orientalis (L.) Blume
(Ulmaceae)
00680143 ASQSP019-08 JF271316 M Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) Sarcocephalus coadunatus
(Sm.) Druce (Rubiaceae)
00680145 ASQSP032-08 JX034708 M Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680148 - - M Madang Wanang 145°10.910'E, 5°13.853'S 30-May-07 Parotis tricoloralis (Pag.) — ditto —
00680154 ASQSP093-08 JX034712 F Madang Wanang 145°10.910'E, 5°13.853'S 12-May-06 Parotis tricoloralis (Pag.) — ditto —
00680176 - - ? Madang Wanang 145°10.910'E, 5°13.853'S 7-Mar-07 most likely Parotis — ditto —
00680187 ASQSP130-08 JX034719 M Madang Wanang 145°10.910'E, 5°13.853'S 18-Apr-06 Parotis tricoloralis (Pag.) — ditto —
00680194 ASQSP181-08 JF271319 F Madang Wanang 145°10.910'E, 5°13.853'S 13-May-06 Parotis hilaralis (Walker)
(host sequence: ASPN768-
09; Genbank JF271498)
— ditto —
a
Both localities are 100m above mean sea level
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
108
Colastomion wanang Quicke, sp. n.
urn:lsid:zoobank.org:act:EDAACAC2-0645-45D7-860F-2C959EFE4D0F
http://species-id.net/wiki/Colastomion_wanang
Figs 33–41
Material examined. Holotype female, Papua New Guinea, Madang Province, Wa-
nang, 29-Apr-05, 145°10.910'E, 5°13.853'S, 100m, SSW, ex caterpillar on Syzigium
longipes (Warb.) Merrill & Perry (Myrtaceae), Mark (voucher USNM ENT 00643283;
BOLD ASQSP063-08; Genbank JF271302). [see Notes on possible conspecics ex-
cluded from type series].
Morphology. Length of body 12 mm, of fore wing 8.8 mm.
Head. Antenna with 56–57 agellomeres. Terminal agellomere acuminate. Me-
dian agellomeres 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 ap-
proximately 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 ner 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: trans-
verse 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.
Mesosoma. Notauli weakly impressed anteriorly, dorsally crenulated to foveate as
impressions get weaker. Mesoscutum smooth where notauli converge. Mesopleuron
smooth and shiny. Precoxal sulcus strongly impressed, rather narrow, smooth. Propo-
deum with complete midlongitudinal carina.
Fore wing. Vein 1-CU1 postfurcal. Vein 2-CU1 5.8 × longer than 1-CU1. Vein
1-M weakly curved near base. Lengths of veins SR1: 3-SR: r = 3.5:2.7:1.0. Vein 2-SR
4 × 2-SR+M.
Hind wing. Vein M+CU 2.5 × length of 1-M. Vein 2-SC+R very thick, short trans-
verse. Vein SR gently curving basally and gradually getting closer to anterior margin
towards wing tip.
Metasoma. 1
st
metasomal tergite 2.0 × longer than posteriorly wide. 2nd metaso-
mal tergite 1.1 × wider than long, with well developed midbasal triangular area giving
rise to complete midlongitudinal carina; irregularly longitudinally striate with approxi-
mately 7 striate lateral to midlongitudinal carina. 3rd metasomal tergite 1.3 × longer
than maximally wide; longitudinally striate. 4
th
and 5
th
tergites entirely coarsely longi-
tudinally striate. 6
th
tergite faintly aciculate.
Coloration. Head, hind trochanter and trochantellus and metasomal tergites 1–4
(largely) black, tergite 5 black basally and laterally, hind tibia dark brown to black,
New reared rogadine wasp species from New Guinea 109
Figures 33–38. Colastomion wanang sp. n. 33, 34 Holotype female and putatively conspecic individ-
ual (USNM ENT 00206919) habitus respectively. 35 face of holotype 36 head and anterior mesosoma
of holotype, dorsal view 37 head and anterior mesosoma of holotype, lateral view 38 propodeum of
putatively conspecic individual (USNM ENT 00452009).
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
110
Figures 39–41. Colastomion wanang sp. n. 39 Metasomal tergites of holotype 40 base of fore wing of
holotype showing distinctive thickening and sudden curving of vein M+CU 41 wings of putatively con-
specic individual (USNM ENT 00452009).
mesosoma and rest of legs orange red, metasomal tergites 5 medioposteriorly and ter-
gite 6 largely cream white. Wings largely pale brown with dark brown venation, pter-
ostigma entirely dark brown.
Etymology. Named after collection locality as a noun in apposition.
Biology. Host unknown for holotype.
New reared rogadine wasp species from New Guinea 111
Notes. Four specimens with host data from a dierent locality and host tree but
lacking barcoding data probably belong to this species but are excluded for the above
reason from the type series. We can nd no convincing morphological dierence be-
tween them and the holotype of C. wanang sp. n. though they do have the 1
st
metaso-
mal tergite largely orange. eir collection data are:
Papua New Guinea, Madang Province, Baitabag, 23-Jul-02, 145°47'E, 5°08'S,
100m, Ulai ex caterpillar on Leea indica Merrill (Vitaceae) (voucher USNM ENT
00206919); 1 male, Madang, Baitabag, 23-Jan-04, 145°47'E, 5°08'S, 100m, ex cat-
erpillar on Leea indica Merrill (Vitaceae) (voucher USNM ENT 00452009); 1 male,
Madang Province, Mis, 1-Aug-02, 145°47'E, 5°11'S, 50m, ex caterpillar on Leea in-
dica Merrill, Vitaceae, David (voucher USNM ENT 00206716); 1 female [metasoma
missing but palps not swollen], Madang Province, Mis, 23-Jul-02, 145°47'E, 5°11'S,
50m (voucher USNM ENT 00206736). ese four specimens were solitary larval
endoparasitoids of one or both of two similar species which are supercially similar to
“Syllepte” crotonalis Walker (CRAM078, Genbank GU695707 and CRAM092, Gen-
bank GU695702). e genus Syllepte has accumulated many unrelated brown moths,
and needs revision, so the correct generic name is also unclear.
Colastomion cheesmanae Quicke, sp. n.
urn:lsid:zoobank.org:act:757380FD-963E-4583-A8AC-44FD96ABCB19
http://species-id.net/wiki/Colastomion_cheesmanae
Figs 42–47
Material examined. Holotype female, Papua New Guinea, Oro Province, Kokoda,
iv.1933, 1200ft [= 365 m], L. E. Cheesman (NHM).
Morphology. Length of body 11 mm, of fore wing 8.5 mm and of antenna 9.2 mm.
Head. Antenna with 55 agellomeres. Terminal agellomere acuminate. Median
agellomeres approximately 2 × longer than wide. 3
rd
segment of maxillary palp of
female rather slender, approximately 5 longer than maximally wide. 5
th
segment 1.5 ×
length of 6
th
. Base of mandible well very close to eye, closest point approximately same
distance as 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 : 2.2. Face rather shiny, with quite
strong diagonal striation laterally above level of clypeus. Shortest distance between
posterior ocelli: transverse diameter of posterior ocellus: shortest distance between pos-
terior ocellus and eye = 1.0: 2.5 : 1.4. Occipital carina complete mediodorsally
Mesosoma. Notauli narrow, weakly impressed, nely crenulated anteriorly becom-
ing more foveate posteriorly. Mesoscutum smooth and broadly depressed at conuence
of notauli. Mesopleuron largely smooth and shiny. Precoxal sulcus deeply, narrow,
weakly crenulate. Propodeum with complete midlongitudinal carina and largely ru-
gose with only narrow smooth and punctate area submedially anteriorly. Propodeum
evenly rounded, without mediolateral projections.
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
112
Fore wing. Vein M+CU distinctly though weakly curved on distal half, but not
thickened medially. Vein 1-CU1 postfurcal. Vein 2-CU1 8.5 × longer than 1-CU1.
Vein 1-M moderately strongly curved posteriorly. Vein 1-SR+M strongly bisinuate.
Lengths of veins SR1: 3-SR: r = 5.0: 4.3: 1.0. Vein 2-SR 3.5 × 2-SR+M.
Hind wing. Vein M+CU 1.75 × length of 1-M. Vein 2-SC+R strongly thickened,
vertical. Vein SR gently curving basally and gradually getting closer to anterior margin
towards wing tip.
Figures 42–47. Colastomion cheesmanae sp. n. light photomicrographs of holotype 42 Habitus, lateral
view 43 head dorsal view 44 mesosoma, lateral view 45 propodeum 46 basal half of fore wing 47 meta-
somal tergites 2 and 3.
New reared rogadine wasp species from New Guinea 113
Metasoma. 1
st
metasomal tergite 2.1 × longer than posteriorly wide. 2nd meta-
somal tergite 1.1 × wider than long, with wide midbasal triangular area giving rise
to weak midlongitudinal carina; rather weakly irregularly longitudinally striate with
approximately 9 striate lateral to midlongitudinal carina. 3rd metasomal tergite 1.2 ×
wider than long; longitudinally striate. 4
th
and basal 0.6 of 5th tergites coarsely longi-
tudinally striate, posterior of tergite 5 and all of tergite 6 smooth.
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 pos-
teriorly and 6 largely cream-white. Antenna largely black with a distinct brown-yellow
zone occupying agellomeres 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 inter-
esting 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.
Colastomion pukpuk Quicke, sp. n.
urn:lsid:zoobank.org:act:28EBB931-2A14-4544-AB0E-6A88C757CD21
http://species-id.net/wiki/Colastomion_pukpuk
Figs 48–51, 55
Material examined. Holotype female, Papua New Guinea, East Sepik Province, Wa-
mangu, 11-Mar-05, 143°39.125'E, 03°47.228'S, 100m, PSS, ex caterpillar on Psycho-
tria micrococca (Laut. & Schum.) Val. (Rubiaceae) (voucher USNM ENT 00505491;
BOLD ASQSP060-08; Genbank JF271303)
Morphology. Length of body 10.0 mm, of fore wing 8.5–9.5 mm and of antenna
9.5–11.2 mm.
Head. Antenna with 56 agellomeres. Terminal agellomere strongly acuminate.
Median agellomeres 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 approxi-
mately 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 ocel-
lus and eye = 1.0: 2.0: 1.2. Occipital carina complete, rather wavy when viewed laterally.
Mesosoma. Notauli weakly impressed anteriorly, dorsally represented by converg-
ing lines of discrete pits. Mesoscutum smooth where notauli converge. . Mesopleuron
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
114
almost entirely smooth ad shiny, precoxal sulcus weakly impressed, unsculptured. Pro-
podeum with complete strong midlongitudinal carina.
Fore wing. Vein 1-CU1 postfurcal. Vein 2-CU1 9.0 × longer than 1-CU1. Lengths
of veins SR1: 3-SR: r = 4.8 : 3.5 : 1.0. Vein 2-SR 2.0 × 2-SR+M.
Figures 48–51. Colastomion pukpuk sp. n. holotype, Cell^D® light photomicrographs. 48 Face 49 pro-
podeum showing largely smooth anterior 50 wings 51 metasomal tergites 2–6.
New reared rogadine wasp species from New Guinea 115
Hind wing. Vein M+CU 1.7 × length of 1-M. Vein SR gently curving basally and
gradually getting closer to anterior margin towards wing tip.
Metasoma. 1
st
metasomal tergite 2.1 × longer than posteriorly wide. 2nd metaso-
mal tergite 1.1 × longer than maximally wide, with well developed midbasal triangular
area giving rise to complete midlongitudinal carina; irregularly longitudinally striate
with approximately 11 striate lateral to midlongitudinal carina. 3rd metasomal tergite
1.2 × wider than long; longitudinally striate. 4
th
tergite largely longitudinally striate.
5
th
tergite longitudinally striate medially becoming smooth laterally with punctures. 6
th
tergite entirely smooth and punctate.
Coloration. Head, and tergites 2–4 entirely, anterior half of tergite 5, and extreme
base of tergite 6 black, hind tibia dark brown, hind trochanter and trochantellus dark
brown, mesosoma, 1
st
tergite and rest of legs orange red, metasomal tergites 4 and 5
posteriorly cream-white, tergites 6 and 7 largely cream-white. Antenna largely black
with paler brown zone approximately occupying agellomeres 22–39. Wing mem-
brane largely pale brown, venation largely (except more basal veins, dark brown; pter-
ostigma dark brown.
Etymology. Pukpuk in Melanesian Pidgin means crocodile, a favourite of the in-
digenous people living around Sepik river.
Biology. Unknown.
Colastomion maclayi Quicke, sp. n.
urn:lsid:zoobank.org:act:DC1DB59F-2C17-45B6-9254-6DBD0A890EE8
http://species-id.net/wiki/Colastomion_maclayi
Figs 52–54, 56
Material examined. Holotype female, Papua New Guinea, East Sepik Province, Yap-
siei, 27-Jan-04, 141°05.839'E, 4°37.695'S, 100m, (voucher USNM ENT 00454207;
BOLD ASQSP061-08; Genbank JF271312) [mounted on same pin is a pupal case,
presumably of the host species, though not the host of this individual]
Paratypes. 1 male, East Sepik Province, Yapsiei, 27-Feb-04, 141°05.839'E,
4°37.695'S, 100m (voucher USNM ENT 00454275; BOLD ASQSP064-08; Gen-
bank JF271310); 1 male, East Sepik Province, Yapsiei, 18-Feb-04, 141°05.839'E,
4°37.695'S, 100m (voucher USNM ENT 00454148; BOLD ASQSP062-08; Gen-
bank JF271311)
Morphology. Length of body 11 mm, of fore wing 8.7 mm and of antenna 11 mm.
Head. Antenna with 57 agellomeres. Terminal agellomere distinctly acuminate.
Median agellomeres 1.8 × longer laterally than wide. 3
rd
segment of maxillary palp
of female very weakly swollen medially swollen, approximately ?7 × longer than maxi-
mally 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
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
116
Figures 52–54. Colastomion maclayi sp. n. holotype, Cell^D® light photomicrographs. 52 Habitus,
dorsal view 53 head and anterior mesosoma, dorsal view 54 head and mesosoma, lateral view.
: 1.9. Face largely nely transversely striate. Shortest distance between posterior ocelli:
transverse diameter of posterior ocellus: shortest distance between posterior ocellus and
eye = 1.0 : 3.0 : 1.1. Occipital carina well developed, complete mediodorsally.
Mesosoma. Notauli weakly impressed anteriorly, dorsally represented by converging
lines of discrete pits. Mesoscutum smooth where notauli converge. Propodeum with
complete midlongitudinal carina.
New reared rogadine wasp species from New Guinea 117
Fore wing. Vein 1-CU1 postfurcal. Vein 2-CU1 8.5 × longer than 1-CU1. Lengths
of veins SR1: 3-SR: r = 4.6: 3.2:1.0. Vein 2-SR 2.4 × 2-SR+M.
Hind wing. Vein M+CU 2.0 × length of 1-M. Hind wing vein 2-SC+R very thick-
ened, almost quadrate. Vein SR gently curving basally and gradually getting closer to
anterior margin towards wing tip.
Metasoma. 1
st
metasomal tergite 2 × longer than posteriorly wide. 2nd metasomal
tergite 1.1 × longer than maximally wide, with well developed midbasal triangular
area giving rise to complete midlongitudinal carina; irregularly longitudinally striate
with approximately 12 striae lateral to midlongitudinal carina. 3rd metasomal tergite
Figures 55–56. Lateral views of 1
st
metasomal tergite. 55 Colastomion pukpuk sp. n. 56 Colastomion
maclayi sp. n.
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
118
1.4 × wider than long; longitudinally striate, with midlongitudinal carina distinct on
anterior 0.5. 4
th
and 5
th
tergites nely longitudinally striate medially becoming weakly
punctate-striate laterally
Coloration. Head, and tergites 2–4 entirely, and anterior half of tergite 5 black,
hind tibia dark brown, mesosoma, 1
st
tergite and legs orange red, metasomal tergite 5
posteriorly cream-white, tergites 6 and 7 largely cream-white. Antenna largely black
with paler brown zone approximately occupying agellomeres 27–42. Wing mem-
brane largely pale brown, venation largely (except more basal veins, dark brown; pter-
ostigma dark brown.
Biology. Solitary endoparasitoid of Meekiaria (Crambidae) caterpillars on Psycho-
tria sp. (Rubiaceae). Meekiaria have distinctive male genitalia with numerous closely
related species in PNG (Munroe 1974, Craft et al. 2010: S2).
Etymology. Named after Nicholai Miklucho-Maklaj, a Russian explorer of New
Guinea who lived around the Madang area for some time.
Discussion
e 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. cram-
bidiphagus 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. e
amount of morphological variation observed between these three groups of species is
considerable, and might in the past have been deemed sucient by some taxonomists
to warrant division into dierent genera or subgenera, and the relatively long internal
branches on the molecular phylogeny support their high level of dierentiation. Ad-
dition 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 de-
scribed here are gregarious. is biology is unusual among the Rogadinae. A few
species of the cosmopolitan genus Aleiodes Wesmael are gregarious (specically A.
pallescens on various Notodontidae, A. stigmator (Say) on Noctuidae, and A. leptoca-
rina Fortier on an unidentied hairy host caterpillar). Another aspect of the gregari-
ousness 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. us they
may make interesting study organisms for investigating the evolutionary transition
New reared rogadine wasp species from New Guinea 119
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 de-
gree 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.
Acknowledgements
We thank the New Guinea Binatang Research Center parataxonomists, village assis-
tants and Lauren Helgen for technical assistance and George Weiblen for plant iden-
tication. is paper stems from a rearing campaign led by Vojtech Novotny, George
Weiblen, SEM, and Yves Basset and supported by the US National Science Foundation
(DEB0841885, 0816749, 0515678), Grant Agency of the Czech Republic (206/08/
H044, 206/09/0115 and P505/10/0673), Czech Ministry of Education (LC06073,
ME9082, and MSM6007665801) and project CZ.1.07/2.3.00/20.0064 co-nanced
by the European Social Fund and the state budget of the Czech Republic. Labora-
tory reagents and Barcode of Life Datasystem (BOLD; HYPERLINK http://www.
boldsystems.org) infrastructure were funded by Genome Canada through the Ontario
Genomics Institute. e Animal Systematic Research Unit and Integrated Ecology
Lab, Department of Biology, Faculty of Science, Chulalongkorn University kindly al-
lowed use of their Cell^D® imaging facility.
References
Baker CF (1917) Ichneumonoid parasites of the Philippines. i. Rhogadinae. Philippines Jour-
nal of Science D12: 281–327.
Butcher BA, Smith MA, Quicke DLJ (2011) A new derived species group of Aleiodes parasitoid
wasps (Hymenoptera, Braconidae, Rogadinae) from Asia with descriptions of three new
species. Journal of Hymenoptera Research 23: 35–42.
Cameron P (1907) Hymenoptera of the Dutch expedition to New Guinea in 1904 an 1905.
Part II: Parasitic Hymenoptera. Tijdschrift voor Entomologie 50: 27–57.
Chen X-X, He J-H (1997) Revision of the subfamily Rogadinae (Hymenoptera: Braconidae)
from China. Zoologische Verhandelingen, Leiden 308: 1–187.
Craft KJ, Pauls SU, Darrow K, Miller SE, Hebert PDN, Helgen LE, Novotny V, Weiblen GD
(2010) Population genetics of ecological communities with DNA barcodes: an example
from New Guinea Lepidoptera. Proceedings of the National Academy of Sciences of the
United States of America 107: 5041–5046.
Donald L.J. Quicke et al. / Journal of Hymenoptera Research 28: 85–121 (2012)
120
Fortier JC (2000) Description of a new gregarious species of Aleiodes Wesmael (Hymenoptera:
Braconidae: Rogadinae). Journal of Hymenoptera Research 9: 288–291.
Granger C (1949) Braconides de Madagascar. Memoires de l’Institut Scientique de Madagas-
car (A) 2: 1–428.
Guinnee MA, Bernal JS, Bezemer TM, Fidgen JG, Hardy ICW, Mayhew PJ, Mills NJ, West SA
(2005) Testing predictions of small brood models using parasitoid wasps. Evolution and
Ecology Research 7: 779–794.
Hrcek J, Miller SE, Quicke DLJ, Smith MA (2011) Molecular detection of trophic links in a
complex insect host-parasitoid food web. Molecular Ecology Resources 11: 786–794.
Kimmins DE (1962) Miss L.E. Cheesman’s expeditions to New Guinea. Trichoptera. Bulletin
of the British Museum (Natural History) Entomology 11: 99–187.
Maeto K, Arakaki N (2005) Gregarious emergence of Macrostomion sumatranum (Hyme-
noptera: Braconidae; Rogadinae) from the mummied, full-grown larvae of eretra sil-
hetensis (Lepidoptera: Sphingidae). Entomol Science 8: 131–132 doi: 10.1111/j.1479-
8298.2005.00107.x
Mayhew PJ, Glaizot O (2001) Integrating theory of clutch size and body size evolution for
parasitoids. Oikos 92: 372–376.
Mayhew PJ, van Alphen JJM (1999) Gregarious development in alysiine parasitoids evolved
through a reduction in larval aggression. Animal Behaviour 58: 131–141.
Munroe EG (1974) New Pyralidae (Lepidoptera) from the Papuan region. III. Canadian En-
tomologist 106: 21–30.
Munroe EG (1996) Distributional patterns of Lepidoptera in the Pacic Islands. In: Keast A,
Miller SE (Eds) e Origin and Evolution of Pacic Island Biotas, New Guinea to Eastern
Polynesia: Patterns and Processes. Academic Publishers, Amsterdam, 275–295.
Quicke DLJ, Shaw MR (2005) First host record for the rogadine genus Pholichora van Achter-
berg (Hymenoptera: Braconidae) with description of a new species and notes on conver-
gent wing venation features. Journal of Natural History 39: 531–537.
Quicke DLJ, Yen S-H, Mori M, Shaw MR (2003) First host records for the rogadine genus
Conspinaria (Hymenoptera: Braconidae), and notes on Rogadinae as parasitoids of Zygae-
nidae (Lepidoptera). Journal of Natural History 38: 1437–1442.
Shaw MR (2002) A new species of Macrostomion Szépligeti (Hymenoptera: Braconidae: Ro-
gadinae) from Papua New Guinea, with notes on the biology of the genus. Zoologische
Mededelingen, Leiden 76: 133–140.
Shaw MR [2002] (2003) Parasitic Hymenoptera: Evolution, Systematics, Biodiversity and
Biological Control. Melika G, uroczy C (Eds) Budapest: Agroinform. Host ranges of
Aleiodes species (Hymenoptera: Braconidae), and an evolutionary hypothesis, 321–327.
Stamatakis A (2006) RAxML-VI-HPC: Maximum likelihood-based phylogenetic analyses with
thousands of taxa and mixed models. Bioinformatics 22: 2688–2690.
Szépligeti G (1911) Braconidae der I. Zentral-Afrika-Expedition. Wissenschaftliche Ergebnisse
Deutschen Zentral-Afrika Expedition 3: 393–418.
Tenma K (2002) Newly Recorded Genera, Conspinaria Schulz, 1906 and Colastomion Baker,
1917 (Hymenoptera, Braconidae, Rogadinae) from Japan. Japanese Journal of Systematic
Entomology 8: 185–191.
New reared rogadine wasp species from New Guinea 121
van Achterberg C (1979) A revision of the subfamily Zelinae auct. (Hymenoptera, Braconidae).
Tidjschrift voor Entomologie 122: 241–479.
van Achterberg C (1988) Revision of the subfamily Blacinae Foerster (Hymenoptera, Braconi-
dae). Zoologische Verhandelingen, Leiden 249: 1–324.
van Achterberg C (1989) Four new genera of Braconinae and Rogadinae from the Oriental
Region (Hymenoptera: Braconidae). Zoologische Mededelingen, Leiden 63: 79–85.
van Achterberg C (1991) Revision of the genera of the Afrotropical and W. Palaeartica Ro-
gadinae Foerster (Hymenoptera: Braconidae). Zoologische Verhandelingen, Leiden 273:
1–102.
van Achterberg C, Chen X (1996) Canalirogas, a new genus of the subfamily Rogadinae Foer-
ster (Hymenoptera: Braconidae) from the Indo-Australian region. Zoologische Mededelin-
gen, Leiden 70: 63–92.
Viette P (1987) Le cas de Glyphodes stolalis Guenée (Lep. Crambidae Spilomelinae). Lambil-
lionea 87: 108–110.
Yu DS, van Achterberg C, Horstmann K (2005) World Ichneumonoidea 2004 – Taxonomy,
Biology, Morphology and Distribution. Taxapad 2005. Vancouver.
Zaldivar-Riverón A, Shaw MR, Saez AG, Mori M, Belokobylskij SA, Shaw SR, Quicke DLJ
(2009) Evolution of the parasitic wasp subfamily Rogadinae (Braconidae): phylogeny and
evolution of lepidopteran host ranges and mummy characteristics. BMC Evolutionary Bi-
ology 8: 329.
