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First Australian record of Aleiodes (Hemigyroneuron) (Hymenoptera: Braconidae: Rogadinae) with the description of a new species from Tasmania
Journal of Asia-Pacific Entomology 19 (2016) 977–980
Contents lists available at ScienceDirect
Journal of Asia-Pacific Entomology journal homepage: www.elsevier.com/locate/jape
First Australian record of Aleiodes (Hemigyroneuron) (Hymenoptera: Braconidae: Rogadinae) with the description of a new species from Tasmania Buntika A. Butcher ⁎,1, Donald L.J. Quicke 2 Department of Biology, Faculty of Science, Chulalongkorn University, Phayathai Road, Pathumwan, BKK 10330, Thailand
a r t i c l e
i n f o
Article history: Received 12 May 2016 Accepted 30 August 2016 Available online 01 September 2016 Keywords: Wing venation New species Australasia Host Parasitoid wasp
a b s t r a c t A new species of Aleiodes (Hemigyroneuron) from Tasmania is described and illustrated. This is the first species of the subgenus known from Australia. The type and only known specimen was reared from a mummified caterpillar collected on Coastal Wattle, Acacia longifolia subsp. sophorae (Labill.) Court, and appears to belong to the Geometridae in agreement with most previous host records for the subgenus. The new species, A. (H.) ellingsenae, has a color pattern that is typical of many larger Australian braconids, with the head and anterior mesosoma orange red, the rest of the body black except for a bright white tergites 2 + 3. © 2016 Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant Protection Society. Published by Elsevier B.V. All rights reserved.
Introduction Parasitic Hymenoptera are an enormously diverse but under-studied group, with some 125,000 species described worldwide. The family Braconidae alone has some 17,000 described species (Quicke, 2015), but it is estimated that between 5 and 20 times as many await description (Dolphin and Quicke, 2001; Jones et al., 2009; Smith et al., 2012; Rodriguez et al., 2013), primarily outside of the Holarctic. The Australian fauna is perhaps somewhat depauperate but is nevertheless poorly known for many groups, many of which have received little taxonomic attention since early in the 20th century. The situation regarding the subfamily Rogadinae (excluding Betylobraconini) is particularly anomalous. As of 2013, only 21 species have been recorded from Australia (Australian faunal directory, 2016), 11 in the distinctive genus Yelicones, five in Teresirogas Quicke & Shaw, two in Spinaria Brullé, one in Batotheca Enderlein, and two in Aleiodes Wesmael. The last of these is an extraordinarily low number since this cosmopolitan genus is one of the largest in the World, and a recent study on the fauna of Thailand revealed 184 species of which only five had been previously described, and based on the sampling, it was estimated that the true fauna would be in excess of 400 (Butcher et al., 2012) species. It is certain ⁎ Corresponding author. E-mail address: [email protected] (B.A. Butcher). http://zoobank.org/urn:lsid:zoobank.org:author:3AC486D1-9045-44FB-9FE4FF71A0C2909C. 2 http://zoobank.org/urn:lsid:zoobank.org:author:4FAA3C17-13D3-467B-AAA582B27FE783EF. http://zoobank.org/urn:lsid:zoobank.org:pub:24B0CAFC-6344-45A5A784-1B6965AC9839. 1
that the Australian Aleiodes fauna is far larger than the two described species, and it is simply the case that no taxonomic revision has been attempted. Aleiodes species, and other rogadines, are koinobiont endoparasitoids exclusively of Lepidoptera caterpillars, and mummify their hosts prior to pupation. This makes them an especially useful group for the study of host ranges and their evolution, since if specimens can be reared from field-collected mummies, an absolutely definite association can be made between the parasitoid and its host, which can usually be identified at least to family. This is important since a very large proportion of parasitic wasp/host associations are erroneous (Shaw, 1994, 2003; Noyes, 1994). The species described here is of interest because it is reared and its host indicates a high level of taxonomic conservation, and also extends the known range of the subgenus Hemigyroneuron Baker by approximately 4000 km. Aleiodes (Hemigyroneuron) comprises species with highly modified fore wing venation, the distal part of the subbasal cell being expanded and glabrous. Hemigyroneuron species are relatively large, with body lengths between 5.3 and 10.0 mm (Butcher and Quicke, 2011), and based on molecular studies (Zaldívar-Riverón et al., 2009, Quicke et al., in prep.) are derived rather basally within Aleiodes. The males of all but one of the species for which males are known, possess glands that open at a sub-posterior pores on the 5th–7th metasomal tergites. The prior known distribution of the subgenus ranged from South Africa and Madagascar through Arabia, China, South-East Asia through IndoAustralia to Papua New Guinea (Butcher and Quicke, 2011, 2015). The new species of Aleiodes (Hemigyroneuron) from Tasmania represents the first record of the group from Australia. It differs considerably
http://dx.doi.org/10.1016/j.aspen.2016.08.014 1226-8615/© 2016 Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant Protection Society. Published by Elsevier B.V. All rights reserved.
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from other species both in its coloration and morphology. In common with several medium-sized to large Australian braconids it belongs to a homeochromatic complex characterized by an orange head and anterior mesosoma, a black body with a conspicuous white median zone. Also in common with many Australian species, the body is largely smooth and shiny, whereas nearly all other species of the subgenus it is largely coriaceous or granular.
Materials and methods Photography The specimen was imaged using an Olympus SXZ16 microscope with automated multiple image capture at preset focal levels using an Olympus DP72 camera, and image combination using the Cell^D image processing system.
Terminology Terminology follows van Achterberg (1988) except for wing venation nomenclature which follows Sharkey and Wharton (1997); see also Fig. 2.2 in Quicke (2015) for comparison of wing venation naming systems.
Systematics The holotype is deposited in the Tasmanian Museum and Art Gallery (TMAG), Hobart Australia. Genus Aleiodes Wesmael, 1838 Subgenus Hemigyroneuron Baker, 1917 Aleiodes (Hemigyroneuron) ellingsenae sp. nov. (Figs. 1–2) Holotype female: AUSTRALIA (TASMANIA), Denison Beach, 41 49′ 21″S, 148 15′57″E, mummified host collected from plant of Coastal Wattle, Acacia longifolia subsp. sophorae (Labill.) Court, on 12.iii.2016, wasp emerged 16.iii.2016, Coll. Kristy Ellingsen. (TMAG; registration no. F19515). Length of body (corrected for bent metasoma) 8.0 mm, of fore wing 6.8 mm, of antennae 9.4 mm. Antenna with 64 flagellomeres; terminal flagellomere acuminate; median flagellomeres quadrate; 1st to 3rd flagellomeres approximately equally long. Eyes large, sharply and deeply emarginated opposite antennal sockets; width of head:width of face:height of eye = 1.0:0.67:0.8. Face smooth and shiny with moderately dense setiferous punctures. Frons smooth and shiny, moderately depressed, with a fine ridge running sub parallel to the eyes. Ocelli large; shortest distance between posterior ocelli:transverse diameter of posterior ocellus:shortest distance between posterior occellus and eye = 1.0:3.0:1.7. Occiput smooth and shiny. Occipital carina completely absent dorsally from
Fig. 1. Aleiodes (Hemigyroneuron) ellingsenae sp. n. holotype female. (a) Habitus, lateral view, (b) head frontal view, (c) mesosoma and head lateral view, (d) head, dorsal view.
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Fig. 2. Aleiodes (Hemigyroneuron) ellingsenae sp. n. holotype female. (a) Fore wing, (b) metasomal tergites 2–6, dorsal view, (c) apex of metasoma, lateral view, (d) mummified host remains (Geometridae) showing postero-dorsal emergence hole.
approximately the mid-height of the eye, weak ventrally, joining hypostomal carina remote from base of mandible. Back of head rather strongly narrowing behind the eye. Mesosoma 1.7 × longer than high. Mesoscutum smooth and shiny; notauli narrow, punctate to crenulate, only present on anterior half of mesoscutum. Scutellar sulcus wide with one strong medial carina and a short, weaker submedial pair. Mesopleuron and mesosternum largely shiny with punctures at base of setae anteriorly; precoxal sulcus short, comma-shaped, moderately impressed, with fine vertically-orientated carination anteriorly. Propodeum smooth and shiny except for setiferous punctures, with complete midlongitudinal carina and with several shorter carinae arising from the posterior margin submedially. Fore wing. Lengths of veins r-rs:3RSa:3RSb = 1.0:1.85:5.1. Vein 1-M strongly sloping, forming an angle of 10° with C + SC + R, 2.5 longer than both m + cu and (RS + M)b separately. Vein (RS + M)b evenly curved. Subbasal cell with strongly enlarged, ovoid apical part, without a sclerome, with distal part of M + CU and 1CUa forming an ‘S’-shaped curve, 1CUa strongly arched towards fore wing margin. Subbasal cell glabrous distally and sparsely setose for its whole narrow basal part. Vein 1cu-a moderately curved, reclivous, not distally expanded. Hind wing. Vein M + CU 1.27 × 1 M. Vein m-cu entirely absent. Vein R short, longitudinal. Legs long, hind leg longer than fore wing. Lengths of fore femur:tibia:tarsus = 1.05:1.02:1.0. Lengths of hind femur:tibia:basitarsus = 1.85:2.3:1.0. Hind femur 5.5 × longer than maximally deep. Hind basitarsus 7.5 × longer than wide. Claws with 4 or 5 setiform pectin spines. 1st and 2nd metasomal tergites finely longitudinally striate, the remainder smooth and shiny with dense, small, setiferous punctures. Tergite 1 with strong, posteriorly uniting, dorsal carinae that give rise to a strong midlongitudinal carina; strongly widening posteriorly; 1.4 × wider posteriorly than long; lateral margins concave. 2nd tergite with small midbasal triangular area and complete strong midlongitudinal carina; 1.85 × wider posteriorly than medially long, 1.0 × longer than 3rd tergite. Ovipositor sheaths with squared end. Coloration. Head, prothorax and fore legs orange; antennae, remains of mesosoma, mid and hind legs, 1st metasomal tergites and 4th–6th backwards black; posterior half of 1st metasomal tergite, and
whole of 2nd tergite, white. Wing membrane dark brown-grey, venation dark brown to black. Host. The mummified caterpillar from which the new species emerged clearly belongs to a member of the Geometridae having only 2 pairs of prolegs. Etymology. Named after Kristy Ellingsen, the collector. Notes. The new species keys easily in Butcher and Quicke (2011) to couplet 27 which leads to two species (A. (H.) bakeri Butcher & Quicke, from Java, and nigricans (Chen & He, 1997), from China). The new species differs markedly from both of these in coloration and morphology. A. (H.) nigricans is largely brown-black with red-brown markings on the mesopleuron and mesosternum, red-yellow markings on the 1st metasomal tergite, and red-yellow legs; A. (H.) bakeri differs from both of these also in the shape of the distal expansion of the subbasal cell which narrows almost immediately after the junction of vein 1-M. A. (H.) ellingsenae is the only known species of its subgenus with forewing vein 1-CU1 strongly curved and running anteriorly distal to junction 1-M. It is also the only species with the 1st metasomal tergite strongly widening distally with concave sides. Discussion Only four of the 27 known species of A. (Hemigyroneuron) have associated host records. Three species have been reared from Geometridae, and one has associated label data indicating it as having been reared from a pierid, and whilst the latter record is not impossible, there are no host remains with the type specimen and so it cannot be verified (Butcher and Quicke, 2011). The new species was also reared from a geometrid, strongly suggesting that the subgenus is specialized on parasitizing members of that family of moths. There is another issue concerning whether A. (Hemigyroneuron), as presently constituted, is monophyletic. The distally swollen subbasal cell is not a perfect diagnostic character, having evolved in other rogadines including some other, non-Hemigyroneuron, Aleiodes (Quicke and Shaw, 2005; Butcher et al., 2012; Butcher and Quicke, 2015). Among those Hemigyroneuron recognized by Butcher and
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Quicke (2011) is the only other species with an Australasian distribution, A. (H.) dubiosus (Fullaway) from Papua New Guinea. This species is only known from a male and unlike other male Hemigyroneuron, it lacks the gland pore that opens near the midposterior margins of 4th– 6th metasomal tergites, and instead has 4th and 5th tergites of male with large subbasal depressions with fine pores and distinctive pattern of setae (see Fig. 36C in Butcher et al. (2012)). It seems likely that the pores of dubiosus originate from paired glands that are homologous to the glands of the other species. Glands opening at a submarginal medial pore on the posterior metasomal tergites of males are not restricted to Hemigyroneuron but occur in some Aleiodes with normal wing venation, including A. cameronii (de Dalla Torre, 1898) from Mexico, the Palaearctic A. fortipes (Reinhard, 1863) (and several other species of the A. pulchripes Wesmael, 1838, group to which it belongs) (Shaw et al., 1997; Delfín-González and Wharton, 2002), some undescribed species from Madagascar, and in the genus Arcaleiodes Chen and He which appears to be the sister group to Aleiodes. These mentioned Aleiodes species have often been treated as members of the subgenus Chelonorhogas, but this is shown by molecular analyses to be a basal grade taxon. The male metasomal glands opening into a medial pore is therefore a plesiomorphic character within Aleiodes, and which may have been lost repeatedly. Acknowledgements Kirrily Moore (TMAG) kindly facilitated study of the type specimen. We thank Senior Postdoctoral Fellowship under Rachadapisek Sompote Fund, Graduate School, Chulalongkorn University to DLJQ, and the Animal Systematics Research Unit and the Integrative Ecology Lab, Department of Biology, Faculty of Science, Chulalongkorn University for allowing us to use their Cell^D imaging facility. References van Achterberg, C., 1988. Revision of the subfamily Blacinae Foerster (Hymenoptera: Braconidae). Zool. Verhandel. 249, 1–324. Australian Faunal Directory, 2016. https://biodiversity.org.au/afd/taxa/BRACONIDAE? Accessed 23 August . Butcher, B.A., Quicke, D.L.J., 2011. Revision of Aleiodes (Hemigyroneuron) parasitic wasps (Hymenoptera: Braconidae: Rogadinae) with reappraisal of subgeneric limits, descriptions of new species and phylogenetic analysis. J. Nat. Hist. 45, 1403–1476. http://dx.doi.org/10.1080/00222933.2011.557557. Butcher, B.A., Quicke, D.L.J., 2015. First record of Aleiodes (Hemigyroneuron) (Hymenoptera: Braconidae: Rogadinae) from the Arabian Peninsula: description of new species with remarkable wing venation convergence to Gyroneuron and Gyroneuronella. Zootaxa 4033, 275–279. http://dx.doi.org/10.11646/zootaxa.4033.2.7. Butcher, B.A., Smith, M.A., Sharkey, M.J., Quicke, D.L.J., 2012. A turbo-taxonomic study of Thai Aleiodes (Aleiodes) and Aleiodes (Arcaleiodes) (Hymenoptera: Braconidae:
Rogadinae) based largely on COI bar-coded specimens, with rapid descriptions of 179 new species. Zootaxa 3457, 1–232. Chen, X., He, J., 1997. Revision of the subfamily Rogadinae (Hymenoptera: Braconidae) from China. Zool. Verhandel. 308, 1–187. de Dalla Torre, C.G., 1898. Catalogus Hymenopterorum. Volumen IV. Braconidae. Guilelmi Engelmann, Lipsiae. Delfín-González, H., Wharton, R.A., 2002. Distribution of species and species-groups of Aleiodes (Hymenoptera: Braconidae) in Mexico. Folia Entomol. Mex. 41, 215–227. Dolphin, K.P., Quicke, D.L.J., 2001. Estimating the global species-richness of incompletely described taxa: an example using parasitoid wasps (Hymenoptera: Braconidae). Biol. J. Linn. Soc. 73, 279–286. http://dx.doi.org/10.1006/bij1.2001.0537. Jones, O.R., Purvis, A., Baumgart, E., Quicke, D.L.J., 2009. Using taxonomic revision data to estimate the geographic and taxonomic distribution of undescribed species richness in the Braconidae (Hymenoptera: Ichneumonoidea). Ins. Cons. Diver. 2, 204–212. http://dx.doi.org/10.1111/j.1752-4598.2009.00057.x. Noyes, J.S., 1994. The reliability of published host parasitoid records: a taxonomist's view. Norw. J. Agric. Sci. Suppl. 16, 59–69. Quicke, D.L.J., 2015. Biology, Systematics, Evolution and Ecology of Braconid and Ichneumonid Parasitoid Wasps. Wiley Blackwell, Chichester, UK. Quicke, D.L.J., Shaw, M.R., 2005. First host record for the rogadine genus Pholichora van Achterberg (Hymenoptera: Braconidae) with description of a new species and notes on convergent wing venation features. J. Nat. Hist. 39, 531–537. http://dx.doi. org/10.1080/00222930410001708678. Reinhard, H., 1863. Beitrage zur Kenntniss einiger Braconiden-Gattungen. Dtsc. Entomol. Z. 7, 248–274. Rodriguez, J.J., Fernández-Triana, J.L., Smith, M.A., Janzen, D.H., Hallwachs, W., Erwin, T.L., Whitfield, J.B., 2013. Extrapolations from field studies and known faunas converge on dramatically increased estimates of global microgastrine parasitoid wasp species richness (Hymenoptera: Braconidae). Ins. Cons. Diver 6, 530–536. http://dx.doi.org/ 10.1111/icad.12003. Sharkey, M.J., Wharton, R.A., 1997. Morphology & terminology. In: Wharton, R.A., Marsh, P.M., Sharkey, M.J. (Eds.), Identification Manual to the New World Genera of Braconidae. Special Publication of the International Society of Hymenopterists vol. 1, pp. 19–37 (Washington D.C.). Shaw, M.R., 1994. Parasitoid host ranges. In: Hawkins, B.A., Sheehan, W. (Eds.), Parasitoid Community Ecology. Oxford University Press, Oxford, pp. 111–144. Shaw, M.R., 2003. Host ranges of Aleiodes species (Hymenoptera: Braconidae), and an evolutionary hypothesis. In: Melika, G., Thuróczy, C. (Eds.), Parasitic Wasps: Evolution, Systematics, Biodiversity and Biological Control. Agroinform Kiadó, Budapest, pp. 321–327 (2002). Shaw, S.R., Marsh, P.M., Fortier, J.C., 1997. Revision of North American Aleiodes (part 1): the pulchripes Wesmael species-group in the New World (Hymenoptera: Braconidae: Rogadinae). J. Hymenopt. Res. 6, 10–35. Smith, M.A., Fernandez-Triana, J.J., Eveleigh, E., Gomez, J., Guclu, C., Hallwachs, W., Hebert, P.D.N., Hrcek, J., Huber, J.T., Janzen, D., Mason, P.G., Miller, S., Quicke, D.L.J., Rodriguez, J.J., Rougerie, R., Shaw, M.R., Várkonyi, G., Ward, D.F., Whitfield, J.B., Zaldívar-Riverón, A., 2012. DNA barcoding and the taxonomy of Microgastrinae wasps (Hymenoptera, Braconidae): impacts after 8 years and nearly 20 000 sequences. Mol. Ecol. Resour. 13, 168–176. http://dx.doi.org/10.1111/1755-0998.12038. Wesmael, C., 1838. Monographie des Braconides de Belgique. 4. Nouv. Mém. Acad. Roy. Sci. Belles-Lettres, Bruxelles 11, 1–166. Zaldívar-Riverón, A., Shaw, M.R., Saez, A.G., Mori, M., Belokobylskij, S.A., Shaw, S.R., Quicke, D.L.J., 2009. Evolution of the parasitic wasp subfamily Rogadinae (Braconidae): phylogeny and evolution of lepidopteran host ranges and mummy characteristics. BMC Evol. Biol. 8, 329.
Contents lists available at ScienceDirect
Journal of Asia-Pacific Entomology journal homepage: www.elsevier.com/locate/jape
First Australian record of Aleiodes (Hemigyroneuron) (Hymenoptera: Braconidae: Rogadinae) with the description of a new species from Tasmania Buntika A. Butcher ⁎,1, Donald L.J. Quicke 2 Department of Biology, Faculty of Science, Chulalongkorn University, Phayathai Road, Pathumwan, BKK 10330, Thailand
a r t i c l e
i n f o
Article history: Received 12 May 2016 Accepted 30 August 2016 Available online 01 September 2016 Keywords: Wing venation New species Australasia Host Parasitoid wasp
a b s t r a c t A new species of Aleiodes (Hemigyroneuron) from Tasmania is described and illustrated. This is the first species of the subgenus known from Australia. The type and only known specimen was reared from a mummified caterpillar collected on Coastal Wattle, Acacia longifolia subsp. sophorae (Labill.) Court, and appears to belong to the Geometridae in agreement with most previous host records for the subgenus. The new species, A. (H.) ellingsenae, has a color pattern that is typical of many larger Australian braconids, with the head and anterior mesosoma orange red, the rest of the body black except for a bright white tergites 2 + 3. © 2016 Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant Protection Society. Published by Elsevier B.V. All rights reserved.
Introduction Parasitic Hymenoptera are an enormously diverse but under-studied group, with some 125,000 species described worldwide. The family Braconidae alone has some 17,000 described species (Quicke, 2015), but it is estimated that between 5 and 20 times as many await description (Dolphin and Quicke, 2001; Jones et al., 2009; Smith et al., 2012; Rodriguez et al., 2013), primarily outside of the Holarctic. The Australian fauna is perhaps somewhat depauperate but is nevertheless poorly known for many groups, many of which have received little taxonomic attention since early in the 20th century. The situation regarding the subfamily Rogadinae (excluding Betylobraconini) is particularly anomalous. As of 2013, only 21 species have been recorded from Australia (Australian faunal directory, 2016), 11 in the distinctive genus Yelicones, five in Teresirogas Quicke & Shaw, two in Spinaria Brullé, one in Batotheca Enderlein, and two in Aleiodes Wesmael. The last of these is an extraordinarily low number since this cosmopolitan genus is one of the largest in the World, and a recent study on the fauna of Thailand revealed 184 species of which only five had been previously described, and based on the sampling, it was estimated that the true fauna would be in excess of 400 (Butcher et al., 2012) species. It is certain ⁎ Corresponding author. E-mail address: [email protected] (B.A. Butcher). http://zoobank.org/urn:lsid:zoobank.org:author:3AC486D1-9045-44FB-9FE4FF71A0C2909C. 2 http://zoobank.org/urn:lsid:zoobank.org:author:4FAA3C17-13D3-467B-AAA582B27FE783EF. http://zoobank.org/urn:lsid:zoobank.org:pub:24B0CAFC-6344-45A5A784-1B6965AC9839. 1
that the Australian Aleiodes fauna is far larger than the two described species, and it is simply the case that no taxonomic revision has been attempted. Aleiodes species, and other rogadines, are koinobiont endoparasitoids exclusively of Lepidoptera caterpillars, and mummify their hosts prior to pupation. This makes them an especially useful group for the study of host ranges and their evolution, since if specimens can be reared from field-collected mummies, an absolutely definite association can be made between the parasitoid and its host, which can usually be identified at least to family. This is important since a very large proportion of parasitic wasp/host associations are erroneous (Shaw, 1994, 2003; Noyes, 1994). The species described here is of interest because it is reared and its host indicates a high level of taxonomic conservation, and also extends the known range of the subgenus Hemigyroneuron Baker by approximately 4000 km. Aleiodes (Hemigyroneuron) comprises species with highly modified fore wing venation, the distal part of the subbasal cell being expanded and glabrous. Hemigyroneuron species are relatively large, with body lengths between 5.3 and 10.0 mm (Butcher and Quicke, 2011), and based on molecular studies (Zaldívar-Riverón et al., 2009, Quicke et al., in prep.) are derived rather basally within Aleiodes. The males of all but one of the species for which males are known, possess glands that open at a sub-posterior pores on the 5th–7th metasomal tergites. The prior known distribution of the subgenus ranged from South Africa and Madagascar through Arabia, China, South-East Asia through IndoAustralia to Papua New Guinea (Butcher and Quicke, 2011, 2015). The new species of Aleiodes (Hemigyroneuron) from Tasmania represents the first record of the group from Australia. It differs considerably
http://dx.doi.org/10.1016/j.aspen.2016.08.014 1226-8615/© 2016 Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant Protection Society. Published by Elsevier B.V. All rights reserved.
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B.A. Butcher, D.L.J. Quicke / Journal of Asia-Pacific Entomology 19 (2016) 977–980
from other species both in its coloration and morphology. In common with several medium-sized to large Australian braconids it belongs to a homeochromatic complex characterized by an orange head and anterior mesosoma, a black body with a conspicuous white median zone. Also in common with many Australian species, the body is largely smooth and shiny, whereas nearly all other species of the subgenus it is largely coriaceous or granular.
Materials and methods Photography The specimen was imaged using an Olympus SXZ16 microscope with automated multiple image capture at preset focal levels using an Olympus DP72 camera, and image combination using the Cell^D image processing system.
Terminology Terminology follows van Achterberg (1988) except for wing venation nomenclature which follows Sharkey and Wharton (1997); see also Fig. 2.2 in Quicke (2015) for comparison of wing venation naming systems.
Systematics The holotype is deposited in the Tasmanian Museum and Art Gallery (TMAG), Hobart Australia. Genus Aleiodes Wesmael, 1838 Subgenus Hemigyroneuron Baker, 1917 Aleiodes (Hemigyroneuron) ellingsenae sp. nov. (Figs. 1–2) Holotype female: AUSTRALIA (TASMANIA), Denison Beach, 41 49′ 21″S, 148 15′57″E, mummified host collected from plant of Coastal Wattle, Acacia longifolia subsp. sophorae (Labill.) Court, on 12.iii.2016, wasp emerged 16.iii.2016, Coll. Kristy Ellingsen. (TMAG; registration no. F19515). Length of body (corrected for bent metasoma) 8.0 mm, of fore wing 6.8 mm, of antennae 9.4 mm. Antenna with 64 flagellomeres; terminal flagellomere acuminate; median flagellomeres quadrate; 1st to 3rd flagellomeres approximately equally long. Eyes large, sharply and deeply emarginated opposite antennal sockets; width of head:width of face:height of eye = 1.0:0.67:0.8. Face smooth and shiny with moderately dense setiferous punctures. Frons smooth and shiny, moderately depressed, with a fine ridge running sub parallel to the eyes. Ocelli large; shortest distance between posterior ocelli:transverse diameter of posterior ocellus:shortest distance between posterior occellus and eye = 1.0:3.0:1.7. Occiput smooth and shiny. Occipital carina completely absent dorsally from
Fig. 1. Aleiodes (Hemigyroneuron) ellingsenae sp. n. holotype female. (a) Habitus, lateral view, (b) head frontal view, (c) mesosoma and head lateral view, (d) head, dorsal view.
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Fig. 2. Aleiodes (Hemigyroneuron) ellingsenae sp. n. holotype female. (a) Fore wing, (b) metasomal tergites 2–6, dorsal view, (c) apex of metasoma, lateral view, (d) mummified host remains (Geometridae) showing postero-dorsal emergence hole.
approximately the mid-height of the eye, weak ventrally, joining hypostomal carina remote from base of mandible. Back of head rather strongly narrowing behind the eye. Mesosoma 1.7 × longer than high. Mesoscutum smooth and shiny; notauli narrow, punctate to crenulate, only present on anterior half of mesoscutum. Scutellar sulcus wide with one strong medial carina and a short, weaker submedial pair. Mesopleuron and mesosternum largely shiny with punctures at base of setae anteriorly; precoxal sulcus short, comma-shaped, moderately impressed, with fine vertically-orientated carination anteriorly. Propodeum smooth and shiny except for setiferous punctures, with complete midlongitudinal carina and with several shorter carinae arising from the posterior margin submedially. Fore wing. Lengths of veins r-rs:3RSa:3RSb = 1.0:1.85:5.1. Vein 1-M strongly sloping, forming an angle of 10° with C + SC + R, 2.5 longer than both m + cu and (RS + M)b separately. Vein (RS + M)b evenly curved. Subbasal cell with strongly enlarged, ovoid apical part, without a sclerome, with distal part of M + CU and 1CUa forming an ‘S’-shaped curve, 1CUa strongly arched towards fore wing margin. Subbasal cell glabrous distally and sparsely setose for its whole narrow basal part. Vein 1cu-a moderately curved, reclivous, not distally expanded. Hind wing. Vein M + CU 1.27 × 1 M. Vein m-cu entirely absent. Vein R short, longitudinal. Legs long, hind leg longer than fore wing. Lengths of fore femur:tibia:tarsus = 1.05:1.02:1.0. Lengths of hind femur:tibia:basitarsus = 1.85:2.3:1.0. Hind femur 5.5 × longer than maximally deep. Hind basitarsus 7.5 × longer than wide. Claws with 4 or 5 setiform pectin spines. 1st and 2nd metasomal tergites finely longitudinally striate, the remainder smooth and shiny with dense, small, setiferous punctures. Tergite 1 with strong, posteriorly uniting, dorsal carinae that give rise to a strong midlongitudinal carina; strongly widening posteriorly; 1.4 × wider posteriorly than long; lateral margins concave. 2nd tergite with small midbasal triangular area and complete strong midlongitudinal carina; 1.85 × wider posteriorly than medially long, 1.0 × longer than 3rd tergite. Ovipositor sheaths with squared end. Coloration. Head, prothorax and fore legs orange; antennae, remains of mesosoma, mid and hind legs, 1st metasomal tergites and 4th–6th backwards black; posterior half of 1st metasomal tergite, and
whole of 2nd tergite, white. Wing membrane dark brown-grey, venation dark brown to black. Host. The mummified caterpillar from which the new species emerged clearly belongs to a member of the Geometridae having only 2 pairs of prolegs. Etymology. Named after Kristy Ellingsen, the collector. Notes. The new species keys easily in Butcher and Quicke (2011) to couplet 27 which leads to two species (A. (H.) bakeri Butcher & Quicke, from Java, and nigricans (Chen & He, 1997), from China). The new species differs markedly from both of these in coloration and morphology. A. (H.) nigricans is largely brown-black with red-brown markings on the mesopleuron and mesosternum, red-yellow markings on the 1st metasomal tergite, and red-yellow legs; A. (H.) bakeri differs from both of these also in the shape of the distal expansion of the subbasal cell which narrows almost immediately after the junction of vein 1-M. A. (H.) ellingsenae is the only known species of its subgenus with forewing vein 1-CU1 strongly curved and running anteriorly distal to junction 1-M. It is also the only species with the 1st metasomal tergite strongly widening distally with concave sides. Discussion Only four of the 27 known species of A. (Hemigyroneuron) have associated host records. Three species have been reared from Geometridae, and one has associated label data indicating it as having been reared from a pierid, and whilst the latter record is not impossible, there are no host remains with the type specimen and so it cannot be verified (Butcher and Quicke, 2011). The new species was also reared from a geometrid, strongly suggesting that the subgenus is specialized on parasitizing members of that family of moths. There is another issue concerning whether A. (Hemigyroneuron), as presently constituted, is monophyletic. The distally swollen subbasal cell is not a perfect diagnostic character, having evolved in other rogadines including some other, non-Hemigyroneuron, Aleiodes (Quicke and Shaw, 2005; Butcher et al., 2012; Butcher and Quicke, 2015). Among those Hemigyroneuron recognized by Butcher and
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Quicke (2011) is the only other species with an Australasian distribution, A. (H.) dubiosus (Fullaway) from Papua New Guinea. This species is only known from a male and unlike other male Hemigyroneuron, it lacks the gland pore that opens near the midposterior margins of 4th– 6th metasomal tergites, and instead has 4th and 5th tergites of male with large subbasal depressions with fine pores and distinctive pattern of setae (see Fig. 36C in Butcher et al. (2012)). It seems likely that the pores of dubiosus originate from paired glands that are homologous to the glands of the other species. Glands opening at a submarginal medial pore on the posterior metasomal tergites of males are not restricted to Hemigyroneuron but occur in some Aleiodes with normal wing venation, including A. cameronii (de Dalla Torre, 1898) from Mexico, the Palaearctic A. fortipes (Reinhard, 1863) (and several other species of the A. pulchripes Wesmael, 1838, group to which it belongs) (Shaw et al., 1997; Delfín-González and Wharton, 2002), some undescribed species from Madagascar, and in the genus Arcaleiodes Chen and He which appears to be the sister group to Aleiodes. These mentioned Aleiodes species have often been treated as members of the subgenus Chelonorhogas, but this is shown by molecular analyses to be a basal grade taxon. The male metasomal glands opening into a medial pore is therefore a plesiomorphic character within Aleiodes, and which may have been lost repeatedly. Acknowledgements Kirrily Moore (TMAG) kindly facilitated study of the type specimen. We thank Senior Postdoctoral Fellowship under Rachadapisek Sompote Fund, Graduate School, Chulalongkorn University to DLJQ, and the Animal Systematics Research Unit and the Integrative Ecology Lab, Department of Biology, Faculty of Science, Chulalongkorn University for allowing us to use their Cell^D imaging facility. References van Achterberg, C., 1988. Revision of the subfamily Blacinae Foerster (Hymenoptera: Braconidae). Zool. Verhandel. 249, 1–324. Australian Faunal Directory, 2016. https://biodiversity.org.au/afd/taxa/BRACONIDAE? Accessed 23 August . Butcher, B.A., Quicke, D.L.J., 2011. Revision of Aleiodes (Hemigyroneuron) parasitic wasps (Hymenoptera: Braconidae: Rogadinae) with reappraisal of subgeneric limits, descriptions of new species and phylogenetic analysis. J. Nat. Hist. 45, 1403–1476. http://dx.doi.org/10.1080/00222933.2011.557557. Butcher, B.A., Quicke, D.L.J., 2015. First record of Aleiodes (Hemigyroneuron) (Hymenoptera: Braconidae: Rogadinae) from the Arabian Peninsula: description of new species with remarkable wing venation convergence to Gyroneuron and Gyroneuronella. Zootaxa 4033, 275–279. http://dx.doi.org/10.11646/zootaxa.4033.2.7. Butcher, B.A., Smith, M.A., Sharkey, M.J., Quicke, D.L.J., 2012. A turbo-taxonomic study of Thai Aleiodes (Aleiodes) and Aleiodes (Arcaleiodes) (Hymenoptera: Braconidae:
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