Papers and Proceedingsof the RoyalSociety of Tasmania, Volume 144, 20 IO 15

TRICHOMALOPSIS S/SYRAE ASKEW, SP. NOV. (HYMENOPTERA: PTEROMALIDAE), A PARASITOID OF S/SYRA (: ) IN TASMANIA, AUSTRALIA

by G. N. R. Forteath, A. W. Osborn and R. R. Askew

(with one text-figure and one plate)

Forteath, G.N.R., Osborn, AW. & Askew, R.R. 2010 (30:xi): Trichomalopsis sisyraeAskew, sp. nov. (Hymenoptera: Pteromalidae}, a parasitoid of Sisyra (Neuroptera: Sisyridae) in Tasmania, Australia. Papers and Proceedings of the Royal Society of Tasmania 144: 15-19. https://doi.org/10.26749/rstpp.144.15 ISSN 0080-4703. Queen Victoria Museum and Art Gallery, PO Box 403, Launceston, Tasmania 7250, Australia {GNRF*, AWO); 5 Beeston Hall Mews, Beeston, Tarpodey, Cheshire CW6 9TZ, UK (RRA). *Author for correspondence. Email: [email protected]

A new species ofhymenopteran parasitoid, Trichomalopsissisyrae Askew (H ymenoptera: Pteromalidae) is described fromTasmania, Aus­ tralia. Adults emerge solitarily from pupal cocoons of the endemic neuropteran Sisyrapedderensis S mithers. The firstrecord of a pteromalid from Tasmania, the species is compared with other pteromalid parasitoids of Sisyridae (Neuroptera) and Gyrinidae (Coleoptera) with similar life history strategies. KeyWords: Trichomalopsis, new species, Sisyra, Gyrinophagus, Sisyridivora, Chalcidoidea, Pteromalidae, Sisyridae, Gyrinidae, ectoparasitoid.

INTRODUCTION hosts may be physically sufficiently alike to be located and accepted as hosts by G. aper. Species of Sisyridae (Neuroptera), the spongilla flies, have Sisyridivoracavigena was described by Gahan ( 1951) from aquaticlarvae that feedon freshwater sponges. S. pedderensis specimens obtained from "nymphs" of the sisyrid Smithers, 2008 larvae feed on the endemic freshwater sponge areolaris (Hagen, 1861) in Ohio, USA, and probably the Radiospongillap edkrensisOsborn, Forteath & Stanisic, 2008 same parasitoid species was foundin 1952 in Michigan by which abounds in Lake Pedder, Tasmania (Northings 5264680 Spangler attacking cocooned prepupae or pupae of Sisyra and 5235000 and Eastings 416550 and 448280 (GDA94)) vicaria (Walker, 1853) at the tops of sedge stems, sometimes (Osborn et al. 2008). adjacent to gyrinid cocoons (Parfin & Gurney 1956). When fullygrown, sisyrid larvae leave the water andmove, Brown (1951) gave a detailed account of the biology of 5. often a considerable distance, to a drier situation where cavigena as a parasitoid of C areolaris. Only cocooned stages they spin two-layered silken cocoons. Pupation sites often of C. areolaris are attacked, exposed pupae and prepupae used by members of this genus include walls of bridges, being ignored by host-searching females. S. cavigena is a crevices in tree trunks and spaces beneath loose bark. It solitary ectoparasitoid, ovipositing on the surface of host is this cocooned terrestrial phase in the sisyrid life cycle prepupae after these have been paralysed by an injection that is vulnerable to attack from parasitic Chalcidoidea of vebom. Brown observed a female S. cavigena applying (Hymenoptera). its mouthparts to the puncture made by the ovipositor; Previously reported chalcid parasitoids of Sisyridae are this was probably an instance of host-feeding. In order to the pteromalids Gyrinophagus aper (Walker, 1839) and reach the host surface, the ovipositor of S. cavigena must Sisyridivora cavigena Gahan, 1951. G. aper has been reared penetrate the two-layered host cocoon, and sometimes its in Britain from cocoons of Sisyrafuscata (Fabricius, 1793) abdomen becomes entangled in the meshes of the outer at Oxford in 1932 (Killington 1933 as Eupteromalus sp., net-like covering so that it may die before laying an egg. Graham 1969) and at Speybank, Inverness, in 1997 &om The total duration from egg to adult in this pteromalid is a cocoon found under loose alder bark (Hancock 1999). 9-15 days while adult S. cavigena are thought to be active Other European records from the same host are from Austria from spring to autumn. (Weissmair 1994) and former Czechoslovakia (Peck et al. Parasitoids of Sisyridae have previously not been recorded 1964). G. aper has been found in Canada (Boui'ek 1993) from Australia. Adults of a parasitoid emerged solitarily from where it was reared from cocoons ofGyrinidae (Coleoptera) pupal cocoons of Sisyra pedderensis (Smithers et al. 2008), (Bou eek & Heydon 1997). an endemic sisyrid found at Lake Pedder in southwestern A second species of Gyrinophagus, G. !uteipes Ruschka, Tasmania. The undescribed parasitoid exhibited similar 1914, is considered by Graham (1969) and Boucek (1990) biology to G. aper and S. cavigena and is placed in the probably to be a small variant of G. aper. However, neither genus Trichomalopsis Crawford, all three genera placed in author proposed formal synonymy, chiefly because of Pteromalidae: Pteromalinae. Other pteromalids apparently the difference in hosts of the two species, Gyrinidae and related to these three sisyrid parasitoids,or whichshare with Sisyridae respectively. Polyphagy, however, is very common them certain morphological features, have been reared from in Pteromalidae which are sometimes known to utilise cocoons of Gyrinidae (Coleoptera); G. aper is possibly a a range of unrelated hosts sharing particular ecological parasitoid of both Sisyridae and Gyrinidae. features (e.g., Askew 1962), and the small, terrestrial or Here we describe the newly discovered species of semi-terrestrial cocooned stages of two unrelated, aquatic Trichomalopsis. 16 Forteath, G.NR, Osborn, A W. and Askew, RR

SYSTEMATICS developed. Head in profile about 1.5x as bigh as long with lower face rather protuberant below level of eyes; eye length pteromalidae Dalman, 1820 about 1.5x malar space. Antenna (fig. ID) with scape (fig. Trichoma/apsis Crawford, 1913 IC) 7x as long (excluding radicula) as broad, reaching Trichoma/apsis sisyrae Askew sp. n. distinctly above level of vertex; combined length of pedicel and flagellum O.84x breadth of head; pedicel in dorsal view Description about 2.25x as long as broad, scarcely longer than anelli plus Female: Head and mesosoma dark green with extensive first funicle segment (PI); both anelli transverse; flagellum coppery tints, especially dorsally; metasoma black with weakly clavate, funicle proximally slightly broader than coppery reflections. Antenna with scape testaceous, darkened pedicel, PI subquadrate to slightly transverse, somewhat at very apex; pedicel brown, paler ventrally; flagellum dark shorter than F2, F2-F5 subquatlrate, F6 slightly broader brown. Legs with coxae and thorax concolorous, metallic, legs than long, clava hardly longer than combined lengths of below coxae reddish testaceous, only protarsus, mesotarsus F5 and F6 (18:17), about 1.8x as long as broad collapsing distally and fifth segment of metatarsus pale brown and in air-dried specimens; sensilla in a single transverse row pretarsi dark brown. Tegulae testaceous; wings almost dear on funicle and claval segments. to faintly infumate, venation brown. Length 2.2-3.0 mm. Mesosoma I.4x as long as broad. Pronotal collar with Head in dorsal view (fig. lA) slightly more than twice as anterior margin raised but without a distinct carina. broad as long; temple OAx as long as eye, weakly curved; Mesoscutum 2.1x as broad as long, only O.87x as broad as POL only slightly greater than OOL, OOL about 4.5x head; notauli extending over anterior two-thirds. Scutellum ocellar diameter; occiput (fig. IB) moderately emarginate slightly longer than mesoscutum, frenum demarcated by with strong, highly arched occipital carina. Head in front finer reticulate sculpture than elsewhere on scutellar dorsum view (fig. I C) l.4x as broad as high, vertex elevated; eye but frenal groove absent. Dorsellurn very short, smooth. height 1.6-1.7x malar space, gena moderately buccate; Propodeum medially about O.7x as long as scutellum; lower margin of torulus above ocular line but nearer to median area about I.4x as broad as long, margined by anterior margin of clypeus than to lower edge of median strong, sinuate lateral plicae which extend down sides of ocellus (13:16); clypeus with radiating striae which extend nucha, median carina well-raised, straight and extending only a short distance onto lower face, its anterior margin over transverse groove in front of nucha, sculpture of median weaklyemarginate. Mouth opening 1.8x malar space; both area strong and quite evenly reticulate but finer and weaker mandibles with four teeth, the left with inner tooth poorly in shallow antero-Iateral foveae; nucha occupying almost

6 J a o

A

B

FIG. 1-(:4) Head offemale Tricbomalopsis sisyrae sp. n. in dorsal view. (B) Posterior view ofhead offemale T. sisyrae sp. n. with right mandible. (C) Front view of head of female T. sisyrae sp. n. with scape of right antenna. (D) Antennalpedicel andflagellum offemale T. sisyrae sp. n.. (E) T. sisyrae sp. o n. petiole in dorsal view and anterior part of basal segment ofmetasoma offrmale T. sisyrae p. n. (F) Antennal pedicel and flagellum of male T sisyrae sp. n. Scale bar = 0.5 mm. c Trichomalopsis sisyrae Askew, sp. nov. in Tasmania, Australia 17

PLATE 1 Dorsal view ofan adult male jewel wasp, Trichomalopsis sisyrae sp. n. Actual size: 2.4mm.

half of propodeallength, with strong, reticulate sculpture; Type Material spiracles small, separated from metanotum by less than a major diameter; callus rather densely pilose. HOLOTYPE: I' QVM (Queen Victoria Museum & Art Forewing 2.4x as long as broad, reaching just beyond Gallery, Launceston): 12:53189, Tasmania: Lake Pedder, apex of gaster in fluid preserved specimens but well beyond Starfish Inlet, ex pupal cocoon of Sisyra pedderensis Smithers, in air-dried material, bare proximally to level of base of emerged 24.iii.2008 (G.N.R. Forteath &A. W. Osborn). marginal vein except for setae on submarginal vein and ALLOTYPE: 0 QVM: 12:53190, same data as holotype costal cell pilosity; costal cell undersurface with a complete except emerged 6.iii.200B. hair row and additional hairs on apical quarter; speculum PARATYPES: 91'1' llOO. Same data as holotype except open below, moderately large; wing membrane distal to emergence dates 27.ii - 23.xi.2008, although mostly in March speculum densely pilose; lengths costal cell: marginal vein: 300 QVM: 12: 53191-53193; 21'1' QVM: 12: 53194-- stigmal vein: postmarginal vein as 30:13:B:11; venation 53195;300 AustralianNationallnsectCollection(ANIC) slender, especially stigmal vein which is slightly curved 32:057477-{)57479,2 1'1' ANIC 32:05748(}-()57481; 2 with stigma occupying only about one-third of its length. 1'1',200 collection ofR.R. Askew. Metasoma ovate, in air-dried specimens slightly longer than mesosoma, l.5x as long as broad, 1.2x broader than Additional material: A further 461' I' and 2600 were thorax; basal tergite occupying almost one-third (one-fifth reared from S. pedderensis cocoons from the type locality in alcohol-preserved specimens) of metasomallength with in 2008 giving an overall 1':0 ratio of 3:2. posterior margin almost straight and a forwardly-directed flange on each side of the petiole (fig. IE), a few setae Etymology laterally in anterior half; basal sternite also with a flange which ventrally half surrounds the posterior expansion of The only species of Trichomalopsis currently known to the petiole, and a basal longitudinal carina flanked by two parasitise pupal cocoons of Sisyra (Neuroptera: Sisyridae). foveae; apex of hypopygium at about half gaster length. Petiole in dorsal view (fig. IE) campaniform, broadest posteriorly, about as broad as long and slightly shorter than COMMENTS propodeal nucha which mostly conceals it, its surface with raised, irregular sculpture. Trichomalopsis sisyrae is referable to Trichomalopsis Male: Resembles female bur differs as follows: head and Crawford, 1913 (= Eupteromalus Kurdjumov, 1913) in mesosoma with coppery tints usually less extensive; forewing Pteromalinae having an occipital carina, antennal clava a little more strongly infumate. Length 1.9-2.6 mm. Eye that is only slightly broader than the funicle, a reticulate somewhat larger, eye height in front view 1.Bx malar space. propodeum with a large subglobose nucha that forms Antenna longer, combined length of pedicel and flagellum a hood over the small, vertical metasomal petiole, and almost as long as breadth of head (18:19); flagellum (fig. forewing with slender venation and bare basal cell, basal IF) hardly expanded distally (pI. 1); all funicle segments vein and speculum. longer than broad. Forewing slightly narrower, 2.5x as long There are at least 35 species of Trichomalopsis in Europe as broad. Metasoma (air-dried) dorsoventrally flattened, 1.2x (Boueek & Rasplus 1991), 15 in the Nearctic region as long as broad, slightly shorter than mesosoma (43:46), (Gibson & Floate 2001), four species are found in rice basal tergite occupying two-fifths of metasomallength. paddy in Japan and elsewhere in southeast Asia and India (Kamijo & Grissell 1982). Boui'ek (1988) notes that at least 10 species are known from New Zealand and five are listed for Australasia. 18 Forteath, G.NR, Osborn, A. W and Askew, RR

Trichomalopsis species are often difficult to identifY, good somal petiole inconspicuous under propodeal nucha, about morphological distinguishing characters for many species as long as broad; left mandible with three teeth ...... being difficult to find, and the biologies of some species ...... Gyrinophagus Ruschka2 may be confusingly similar. For instance, four of the five species known to parasitise filth fly puparia in North America 3. Gena with a large hollow extending from base ofmandible include Musca domestica 1. in their host ranges (Gibson almost to eye; ~ flagellum fusiform, broadest at apex & Floare 2001) and the previously known Australian of funicle; notauli indicated only near anterior margin species have been reared only as secondary parasitoids of of mesoscutum; scutellum with distinct frenal groove; Lepidoptera attacking cocoons of microgastrine Braconidae forewing with basal cell bare but speculum absent; (Boucek 1988). abdominal petiole twice as long as apical breadth Boucek (1988) lists four species of Trichomdlopsis from ...... Sisyridivora Gahan3 Australia, but later in the same work one of these, T. pulchra (Girault & Dodd, 1915 in Girault 1915), is put in the genus Gena with at most a small hollow at corner of mouth; S2 Oxyhanna Balleek characterised by an elongated metasomal flagellumsubcylindrical to subclavate, funicle not broader petiole. The type material of T. australiensis (Girault, 1926) than clava; notauli distinct over anterior two-thirds of is lost but it is considered by Balleek probably to be a junior mesoscutum; scutellum with frenum not separated by synonym of T. braconophaga (Cameron, 1912). a frenal groove; forewing with basal cell bare in most Therefore, with the description of T. sisyrae, only three species, speculum present; abdominal petiole very small, species are known with certainty from Australia. concealed beneath nucha... Trichomalopsis Crawford4 We have examined the type of T. braconis (Dodd, 1917), but that of T. braconophaga was unavailable for study. lOne species, D. dineutis (Ashmead), parasitoid in cocoons However, from the original description of Trichoglenes(?) of Dineutes Gyrinidae), USA. braconophagus Cameron, 1912, both it and T. braconis 2 G. aper (= marginatus Thomson), parasitoid in cocoons differ morphologically from T. sisyrae in having a female of Sisyridae, Holarctic; G. luteipes, parasitoid in cocoons of metasoma shorter than the mesosoma (1.25x as long in T. Gyrinidae, central Europe and G. cychreus (Walker, 1850), sisyrae) and at least the front coxae largely testaceous (all host unknown but possibly Ephydridae (Diptera), Dublin, coxae darkened and metallic in T. sisyrae). Also the very may be the same as G. aper, central Europe. different host association distinguishes T. sisyrae from the 3 One species, S. cavigena, parasitoid in cocoons ofSisyridae, rno secondary parasitoids of Lepidoptera. USA. In Graham's (1969) key to females of European and North 4A large genus, formerly knownasEupteromalus Kurdjumov, American Eupteromalus, T. sisyrae approaches "sp. indet. which occurs in all major regions. Mosdysolitary or gregarious, A", but differs from it in its mainly copper colouration, primary or secondary parasitoids of a range of hosts in in having a well-developed propodeal median carina and cocoons or puparia, attacking particularly Lepidopteraor their more transverse head in dorsal view. This species is now braconid (Hymenoptera) or tachinid (Diptera) parasitoids, known as T. arzoneae (Boucek, 1970) which is biologically Diptera and Coleoptera, sometimes aculeate Hymenoptera very different from T. sisyrae; T. arzoneae is gregarious and and egg sacs of spiders. The species described above is a was described from 21','',' and 300 reared in Italy from parasitoid of Sisyridae. Species range from being apparently a prepupa of the Ailanthus Silkmoth, Philosamia cynthia more or less host-specific to very polyphagous. (Drury, 1773) (Saturniidae) parasitised by the tachinid Pales pavic14 (Meigen, 1824) (Boucek 1970). Although species of Trichomalopsis are often polyphagous, ACKNOWLEDGEMENTS their host ranges are limited by ecological factors. The unusual host association of T. sisyrae, together with its We thank Drs LD. Naumann and J. LaSalle for valuable advice combination of morphological characters, provide sufficient in the preliminary stages of this work. We also thank Messrs P. grounds for recognising it as a new species. Harding and C. Livingston for assistance as coxswains which enabled us to conduct our field work on Lake Pedder. The research was made possible by grants from Hydro Tasmania KEY TO GENERA OF PTEROMALIDAE and the W.O. Booth Charitable Trust. Finally invaluable WHICH INCLUDE PARASITOIDS OF support was forthcoming from the Queen Victoria Museum SISYRIDAE AND GYRINIDAE & Art Gallery, Launceston, Tasmania, Australia.

In all genera the propodeal nucha is large, subglobose and has coarse, strongly-raised reticulate sculpture. REFERENCES

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