Overview of Australian Cynipoidea (Hymenoptera)

Overview of Australian Cynipoidea (Hymenoptera)

Jordi Paretas-Martínez,1* Mattias Forshage,2 Matthew Buffington,3 Nicole Fisher,4 John La Salle5 andJuli Pujade-Villar1

1Department of Animal Biology, Faculty of Biology, University of Barcelona, Avda. Diagonal 645, 08028 Barcelona,Spain.2Department of Entomology, Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm, Sweden.3Systematic Entomology Laboratory, USDA, c/o NMNH, Smithsonian Institution, 10th & Constitution Avenue NW. POBox 37012 MRC-168, Washington, DC 20013, USA.4Australian National Insect Collection and 5Atlas of Living Australia, CSIRO Ecosystem Sciences, GPO Box 1700,Canberra, ACT 2601, Australia.

Abstract An overview of all families, subfamilies, genera and species of Cynipoidea present in Australia is presented.The Australian cynipoid fauna is very poorly known, with 37 genera recorded: one each for Austrocynipidae,Ibaliidae and Liopteridae; two for Cynipidae; and 32 for Figitidae. The first Australian records are given forthe following genera of Eucoilinae: Aganaspis Lin, Areaspis Lin, Chrestosema Förster, Didyctium Riley,Endecameris Yoshimoto, Ganaspis Förster, Leptolamina Yoshimoto, Micreriodes Yoshimoto, Pseudodiran-chis Yoshimoto, Sinochresta Lin and Weldia Yoshimoto. Nine new combinations, two new synonymies andone reinstatement are made: Eucoilinae (Figitidae): Hexacola aemilia comb. n., Hexacola florentia comb. n.,Hexacola julia comb. n., Hexacola mozarti comb. n., Hexacola thoreauini comb. n., Kleidotoma margueritacomb. n., Leptopilina lonchaeae comb. n., Leptopilina maria comb. n., Trybliographa australiensis stat. rev.(Rhoptromeris unimaculus Girault 1931 syn. n.); Thrasorinae (Figitidae): Thrasorus berlesi comb. n. (Thra-sorus rieki Paretas-Martínez & Pujade-Villar 2011 syn. n.). Aspects on the systematics, distribution, biologyand morphology of all cynipoid families and figitid subfamilies in Australia are given. A multi-character onlinekey to the genera of Australian Cynipoidea can be found at http://www.ces.csiro.au/keys/Hymenoptera/Australian_Cynipoidea/Australian-Cynipoidea-Keys.html.

Key words Australia, Austrocynipidae, Cynipidae, Figitidae, Ibaliidae, Liopteridae.

INTRODUCTION

Cynipoid wasps (Hymenoptera: Apocrita) form a worldwidegroup that includes ~3000 species (Ronquist 1999). Cynipoidscan be divided into two major groups, the so-called macro- andmicrocynipoids. Macrocynipoids (up to 20 mm in size) haverelatively low species richness when compared with the micro-cynipoids and include the rare Austrocynipidae, as well asIbaliidae and Liopteridae. The biology of Austrocynipidae isnot fully known, but the only species of this family was rearedfrom larvae of an undescribed oecophorid moth dissected fromseeds in infested cones of Araucaria (Araucariaceae) (Riek1971); thus, their host niche is similar to that of the othermacrocynipoids, which are koinobiont endoparasites of wood-boring or cone-boring insects (Liu 1998). Ibaliids are parasi-toids of siricid woodwasp larvae in conifers and hardwoods(Ronquist 1999). Liopterids are putative parasitoids ofbuprestid, cerambycid and curculionid beetle larvae boring intwigs and stems of deciduous trees and bushes (Ronquist1995b; Liu 1998), though no definitive rearing records havesurfaced (Buffington et al. 2012).

The microcynipoids (0.7–8 mm in size), consisting ofCynipidae and Figitidae, are the most species-rich clade ofCynipoidea, with ~1500 described species each. Mostspecies of Cynipidae are gall inducers on woody dicots, butthe family also includes inquilines (guests of gall inducers,incapable of gall induction themselves) in galls of othercynipids. Figitids, for which the biology is known, are almostexclusively koinobiont endoparasitoids of endopterygoteinsect larvae, with most species being primary parasitoids ofthe muscomorph flies (Diptera: Schizophora) in habitats fromleaf-mines to algae to dung and carrion (Buffington 2002;Fontal-Cazalla et al. 2002; Buffington et al. 2007). Figitidaeis the most species diverse family within Cynipoidea (Ron-quist 1999) and is separated into 12 subfamilies: Anachari-tinae, Aspicerinae, Charipinae, Emargininae, Euceroptrinae,Eucoilinae, Figitinae, Mikeiinae, Parnipinae, Plectocynipi-nae, Pycnostigminae and Thrasorinae (Paretas-Martínez et al.2011).

The cynipoids include economically important species thatare either harmful (the herbivorous cynipids) or beneficial(entomophagous figitids) to human interests. On one side,cynipids can become important plagues of several trees (i.e.chestnut and cork oak gallwasps, and many others), and manyfigitids (Anacharitinae, Aspicerinae and Charipinae) conflict*[email protected]

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with human interests when parasitising predators or parasi-toids of aphid pests. On the other side, entomophagouscynipoids have been used in biological control programs:eucoiline (Figitidae) species are used to control pest Diptera(Buffington 2002; Fontal-Cazalla et al. 2002; Buffington et al.2007), and species of Ibalia Latreille (Ibaliidae) have beenused as biological control agents of the woodwasp Sirex noc-tilio F. (Taylor 1976, 1981).

As for many small-bodied Hymenoptera, cynipoid biodiver-sity is certainly underestimated, and much remains to belearned of their systematics, fundamental biology and the eco-system services that they provide. Cynipoids are especiallypoorly known outside the Holarctic and tropical Africa. Thevast majority of described species are North American, Euro-pean and sub-Saharan African, whereas many areas of theEastern Palaearctic, the tropical regions and the SouthernHemisphere (including Australia) remain almost unstudied,although being extremely species rich (Nieves-Aldrey &Fontal-Cazalla 1997; Fontal-Cazalla & Nieves-Aldrey 1999;Buffington & Ronquist 2006; Ronquist et al. 2006). As anexample, only 5% of species of figitids have been described,with 1400 described species (Buffington et al. 2007) comparedwith an estimated global diversity of 24 000 species (Nord-lander 1984).

Australian cynipoid biodiversity is very poorly known.Most species descriptions are found in Girault’s works (1930,1931, 1932, 1933, 1934a,b, 1935, 1937), but other authorshave contributed to the description of the Australian cynipoidfauna (Ashmead 1900; Kieffer 1911; Weld 1944, 1952;Kerrich & Quinlan 1960; Riek 1970, 1971; New 1979; Carver1992, 1993; Paretas-Martínez & Pujade-Villar 2006, 2010;Buffington 2008; Paretas-Martínez et al. 2011). The Australianspecies of three figitid groups have recently been revised: theCharipinae (Carver 1992; Paretas-Martínez & Pujade-Villar2006), the Figitinae (Paretas-Martínez & Pujade-Villar 2010)and the Thrasorinae + Mikeiinae (Paretas-Martínez et al.2011). Biological studies on Australian cynipoids have beenfocused on Ibalia control of siricid wasps (Taylor 1976, 1981and references therein).

A taxonomic overview of all families, subfamilies, generaand species of Cynipoidea present in Australia is presentedhere, including new records. An interactive and multi-character LucID key to all Australian cynipoids canbe accessed at http://www.ces.csiro.au/keys/Hymenoptera/Australian_Cynipoidea/Australian-Cynipoidea-Keys.html.Aspects of the taxonomy, distribution, biology and morphol-ogy of all cynipoid families and figitid subfamilies in Australiaare given.

MATERIAL AND METHODS

This study was made through an exhaustive literature searchon Australian cynipoids, as well as the study of Australianmaterial from the Australian National Insect Collection(ANIC, CSIRO Ecosystem Sciences, Canberra, Australia) andthe Queensland Museum (QM, Brisbane, Australia). For

Eucoilinae, no exhaustive studies were made on the abundantmaterial from Australia present in Australian collections. Thetreatment of the Eucoilinae in this work is based on literaturerecords, and mainly on studies of relevant types, as well asstudies of Australian material in the Natural History Museum,London (BMNH) and The United States National Museum,Washington (USNM). Abbreviations of other holotype reposi-tories are: ANSP (Academy of Natural Sciences of Philadel-phia); BPBM (Bernice P. Bishop Museum, Honolulu, Hawaii);MNHN (Muséum National d′Histoire Naturelle, Paris,France); MZLU (Lund Museum of Zoology, Lund, Sweden);OUMNH (Hope Department of Entomology, Oxford,England); and ZSM (Zoologische Staatssammlung Museum,München, Germany).

The text below is divided into families, and within Figitidae,into subfamilies. For each family and figitid subfamily, thefollowing sections are given: List of species with valid nameof species present in Australia, author and year of description,original combination, relevant synonymies in Australia (ifany), repository of holotype (in cases where the holotype is notdeposited in an Australian institution, we also indicate if thereare available specimens for study in an Australian institution);Systematics of the family/subfamily worldwide, with numberof subfamilies/tribes (where this applies), and genera included(names of genera are given except when the number is toohigh); World distribution of genera present in Australia toindicate if the genera present in Australia are endemic or alsopresent in other regions; Biology with a brief summary of thebiology of the family/subfamily and details of the Australiantaxa, if known; Morphology with a brief diagnosis of thesubfamily. Overall, the classification of cynipoid wasps pre-sented by Ronquist (1999) is followed; however, several newgroups have been described in the Figitidae since that time. Forthe higher level classification of Figitidae, we follow Paretas-Martínez et al. (2011); for eucoiline classification, we followForshage and Nordlander (2008).

Two special sections have been added: at the end of theAustrocynipidae, a brief summary of its phylogenetic positionis given, since this endemic Australian taxon putatively holdsa key position in the evolutionary origin of Cynipoidea; insidethe biology section of Ibaliidae, a detailed summary of theintroduction of Ibalia in Australia is given, since this is animportant example of the use of cynipoids in biological controlin Australia.

RESULTS

All genera and family-group taxa of Cynipoidea described inAustralia are in Table 1, including the new genus records givenherein.

Family Austrocynipidae (Fig. 1a)

Species list

Austrocynips mirabilis Riek 1971 (Fig. 1a) holotype in QM,specimens available in ANIC

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Systematics. The Austrocynipidae include a single species:Austrocynips mirabilis Riek.

World distribution of genera present in Australia. Austro-cynips is endemic to Australia.

Biology. Austrocynips is only known from three female speci-mens collected in Queensland, reared from cones of hoop pine(Araucaria cunninghamii Aiton ex D.Don) (Riek 1971). Riek(1971) stated that the wasps emerged from seeds, consistentwith the hypothesis that the larvae were phytophagous.However, the collector reported that the specimens were actu-ally reared from larvae of an undescribed oecophorid mothdissected from seeds in infested cones (NW Heather pers.comm. 1971). The oecophorid moths attack the cones whenthey are still attached to the tree, and the larvae bore throughthree to six seeds before pupating (Heather 1970). Thus, thehost niche of Austrocynips is similar to that of the othermacrocynipoids, the Ibaliidae and Liopteridae (Ronquist1995b).

Morphology. Austrocynips has a number of traits that areunique among cynipoids: pterostigma in forewing present;last flagellomere as long or shorter than penultimate flagel-lomere; radicle indistinctly delineated, but with a lineindicating it; 13 flagellomeres in female antenna; posteriormargin of pronotum projecting over anterior margin of meso-pleuron, not abutting, covering mesothoracic spiracle whichis not visible laterally; lateral bars of scutellum absent; nuchaabsent; annulus with tergal and sternal parts well defined butseparate, not fused.

Phylogenetic position. Austrocynips is the sister group of allother cynipoids, the latter being supported as a monophyleticgroup by several character states (Ronquist 1999). This sister-group relationship is based on the presence in Austrocynips ofa number of character states that are more plesiomorphic thanother cynipoids (Ronquist 1995b). Some of these traits areunique among cynipoids but are commonly found in the proc-totrupoid complex and probably belong to the cynipoid ground

plan. Thus, Austrocynips is a key taxon in linking cynipoids toother apocritan wasps (Ronquist 1999).

Family Ibaliidae (Fig. 1b)

Species list

Ibalia leucospoides (Hochenwarth 1785) (Ichneumon leuco-spoides) (Fig. 1b) holotype presumably lost, specimens avail-able in ANIC

Ibalia rufipes Cresson 1879 holotype in ANSP, specimensavailable in ANIC

Systematics. The Ibaliidae include three genera: IbaliaLatreille, Heteribalia Sakagami and Eileenella Fergusson.

World distribution of genera present in Australia. Ibalia iscosmopolitan.

Biology. Ibaliids are parasitoids of siricid woodwasp larvae(Liu & Nordlander 1994). Ibalia leucospoides Hochenwarthattacks eggs and first-instar larvae of Sirex noctilio F. and isendoparasitic until its third instar, a stage at which it becomesan ectoparasite. The Ibalia life cycle is closely related with thedevelopment of woodwasp larvae, usually lasting one year (butsee Corley et al. 2004).

Introduction of Ibalia into Australia. The introduction ofIbalia (and other Sirex parasitoids) into Australia is reportedby Taylor (1967, 1976, 1981). Ibalia leucospoides was firstsuccessfully introduced to control Sirex noctilio (after earlyunsuccessful attempts) in New Zealand in 1950, from stocksobtained in England some years earlier. After the discovery ofS. noctilio in a private plantation of Pinus radiata D. Don. atPittwater near Hobart, Tasmania, in 1952, and an initial periodof unsuccessful attempts to eradicate this woodwasp in Tas-mania, the Tasmanian Department of Agriculture approachedthe New Zealand authorities for shipments of I. leucospoidesand Rhyssa persuasoria (L.) (Hymenoptera: Ichneumonidae,

Table 1 Cynipoidea cited in Australia

Family: Subfamily Tribe: GenusAustrocynipidae AustrocynipsIbaliidae IbaliaLiopteridae: Dallatorrellinae DallatorrellaCynipidae Aylacini: Phanacis; Cynipini: AndricusFigitidae: Anacharitinae Anacharis, XyalaspisFigitidae: Charipinae Alloxysta, Phaenoglyphis, Thoreauana, DilapothorFigitidae: Emargininae ThoreauellaFigitidae: Eucoilinae Diglyphosematini: Gronotoma; Eucoilini: Leptopilina, Maacynips, Trybliographa; Ganaspini: Aganaspis†, Areaspis†,

Chrestosema†, Didyctium†, Endecameris†, Ganaspis†, Hexacola, Leptolamina†, Micreriodes†, Pseudodiranchis†,Sinochresta†, Striatovertex, Weldia†; Kleidotomini: Cothonaspis, Kleidotoma; Trichoplastini: Rhoptromeris

Figitidae: Figitinae XyalophoraFigitidae: Mikeiinae MikeiusFigitidae: Thrasorinae Cicatrix, Palmiriella, Thrasorus

†New record.

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another siricid parasitoid). In 1959 and 1960, shipments ofI. leucospoides were received and released in the same plan-tation; the specimens of I. leucospoides from New Zealandwere found to be asynchronous with S. noctilio in Tasmania.Colonisation of a Mediterranean strain of I. leucospoides,together with I. leucospoides ensiger from North America,brought about a much more satisfactory synchronisation.

Following the discovery in 1961 that S. noctilio was estab-lished in Victoria, and the establishment of the National SirexFund, a worldwide search for natural enemies of siricids wasinstituted. Further stocks of I. leucospoides and R. persuasoriawere introduced from throughout their range in the northernhemisphere (both are Holarctic) between 1962 and 1973.Nineteen other parasitoid species were introduced to controlS. noctilio, including other species of Ibalia. In total, the originand number of Ibalia females imported to Tasmania in thisperiod are:

Ibalia aprilina Kerrich: Japan (54 specimens)I. montana Cresson: USA (66 specimens)I. ruficollis Cameron: USA (97 specimens)I. rufipes rufipes Cresson: Canada (three specimens), USA(89)I. rufipes drewseni Borries: Europe (161 specimens), Turkey(four specimens )I. leucospoides ensiger Norton: USA (68 specimens), Canada(two specimens)I. leucospoides leucospoides Hochenwarth: Europe andTurkey (1469 specimens), Morocco (40 specimens), Japan(= suprunenkoi 243 specimens).

Of all these, only the last three were successfully reared forrelease and are now established in Australia. Both subspeciesof I. leucospoides emerge in summer and autumn to attacksiricid larvae hatching soon after oviposition; Ibalia rufipesdrewseni emerge in spring to attack siricid larvae in treeswhere the hatching of the eggs has been delayed over winter.

Morphology. Ibaliids are characterised by being of muchlarger size than other cynipoids, with an elongate yet stronglylaterally flattened metasoma, and the reduction of wing vena-tion is less strong than other cynipoids. Other diagnostic char-acter states include: marginal cell of forewing very long andnarrow; median notch in pronotal crest; pair of posteriorscutellar processes; short metafemur; enlarged seventh tergumin female metasoma; apical tubular process present on secondmetatarsomere.

Family Liopteridae (Fig. 1c)

Species list

Dallatorrellinae

Dallatorrella rubriventris Kieffer 1911 (Fig. 1c) holotype inBMNH, specimens available in ANIC

Systematics. The Liopteridae include 10 genera classifiedinto four subfamilies: Dallatorrellinae (2), Mayrellinae (2),Oberthuerellinae (3) and Liopterinae (3).

World distribution of genera present in Australia. Dalla-torrella is distributed in the Oriental and Australian regions.

Biology. The biology of the subfamily Dallatorrellinae isunknown. Females of one species of Dallatorrella have been

(a)

(b)

(c)

Fig. 1. Macrocynipoids. (a) Austrocynips (Austrocynipidae);(b) Ibalia (Ibaliidae); (c) Dallatorrella (Liopteridae).

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collected on logs of Syzygium (Myrtaceae) in Papua NewGuinea, suggesting an association with wood-boring insectson broad-leaved trees (Liu 2001). Species of Mayrellinae havebeen reported as being biologically associated with buprestid,cerambycid and curculionid beetles (Weld 1956; Díaz 1973;Yang & Gu 1994; Ronquist 1995b; Liu et al. 2007), but allthese records only report collections on trees attacked by thesebeetles, lacking direct evidence of a parasitoid-host associa-tion. Thus, although all records suggest an association ofLiopteridae with wood-boring Coleoptera, further studies areneeded to confirm the hosts of this cynipoid family.

Morphology. According to Ronquist (1995a,b, 1999), theLiopteridae share at least 12 synapomorphies including:lateral part of cranium expanded, resulting in a swollen gena;deeply foveate pronotal sculpture; presence of mesopleuralimpression; dorsolateral scutellar processes; metapleuralsulcus reaching anterior metapectal margin far above mid-height; propodeum with two distinct median carinae; rela-tively long nucha; and short metatibia. However, there areexceptions to nearly all of these features in form of partialreduction or total loss of the characteristic in some speciesof Liopteridae; also, many of these morphological structuresshow some homoplasy within the Cynipoidea due toindependent, parallel gains in other cynipoids (Ronquist1995a).

Family Cynipidae (Fig. 2a)

Species list

Tribe Aylacini

Phanacis hypochoeridis (Kieffer 1887) (Aylax hypochoeridis)(Fig. 2a) holotype presumably lost, specimens available inANIC

Tribe Cynipini

Andricus sp. specimens available in ANIC

Systematics. The Cynipidae include about 77 genera and1400 species classified into eight tribes: the gall inducersAylacini (herb gallwasps, 21 genera), Cynipini (oak gallwasps,43 genera), Diplolepidini (rose gallwasps, two genera),Eschatocerini (gallwasps on Acacia and Prosopis (Fabaceae),one genus), Paraulacini (gallwasps on Nothofagus (Nothof-agaceae), two genera), Pediaspidini (gallwasps on Acer (Sap-indaceae), three genera), and Qwaqwaiini (gallwasps onScolopia (Salicaceae), one genus); and the inquilines Syn-ergini (seven genera) (Liljeblad & Ronquist 1998; Liljebladet al. 2008, 2011; Nieves-Aldrey et al. 2009).

World distribution of genera present in Australia. Andri-cus is an oak gallwasp genus that has more than 300 species(Pujade-Villar et al. 2001), but it is restricted to the Holarticregion together with the natural distribution of its hostQuercus. The presence of Andricus specimens in Australia

must be due to an accidental and isolated introduction. Phana-cis hypochoeridis is a Western Palaearctic species that hasbeen introduced in the Nearctic, Neotropical, Australasian andAfrotropical regions (Melika & Prinsloo 2007).

Biology. Riek (1970) mentioned for Australian cynipids:‘Andricus sp. forms galls on introduced oaks. Aylax hypoch-oeridis, gall-former in the flower stems of the introduced

(a)

(b)

(c)

Fig. 2. Cynipidae-Figitidae. (a) Phanacis (Cynipidae: Ayla-cini); (b) Anacharis (Figitidae: Anacharitinae); (c) Thoreauana(Figitidae: Charipinae).

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“dandelion”, Hypochoeris radicata, is possibly the mostcommon cynipoid in Australia’.

Morphology. A combination of ‘loss’ character states canhelp to identify the Cynipidae (these character states can befound separately in some figitids but never all together): veinRs+M directed towards middle area of basal vein, pronotalcarinae absent, ring of setae at base of metasoma absent andclub in the antenna absent.

Family Figitidae (Figs 2b,c,3,4)

Systematics. Figitids include about 132 genera and 1400species classified into 12 subfamilies: Anacharitinae, Aspiceri-nae, Charipinae, Emargininae, Euceroptrinae, Eucoilinae (themost diverse and species-rich), Figitinae, Mikeiinae, Parnipi-nae, Plectocynipinae, Pycnostigminae and Thrasorinae(Paretas-Martínez et al. 2011).

Australia. Seven figitid subfamilies are recorded in Australia:Anacharitinae, Charipinae, Emargininae, Eucoilinae, Figiti-nae, Mikeiinae and Thrasorinae. See data below for eachsubfamily. A key to figitid subfamilies can be found inParetas-Martínez et al. (2011).

Subfamily Anacharitinae (Fig. 2b)

Species list

Anacharis zealandica Ashmead 1900 (syn Anacharis austra-liensis Ashmead 1900) (Fig. 2b) holotype in USNM, specimensavailable in ANIC

Xyalaspis victoriensis New 1979 holotype in ANIC, specimensavailable in ANIC

Systematics. The Anacharitinae include nine genera: Acan-thaegilips Ashmead, Acothyreus Ashmead, Aegilips Walker,Anacharis Dalman, Calofigites Kieffer, Petricynips Belizin,Proanacharis Kovalev, Solenofigites Díaz and XyalaspisHartig (Ronquist 1999).

World distribution of genera present in Australia. Anach-aris and Xyalaspis are cosmopolitan.

Biology. Anacharitines are primary parasitoids of aphid-feeding lacewing larvae (Neuroptera: Hemerobiidae andChrysopidae) (Ronquist 1999). Australian species are found inassociation with larvae of Micromus tasmaniae (Walker) andDrepanacra binocula (Newman); these are the two most abun-dant Hemerobiidae around Melbourne and both are more fre-quently parasitised by A. zealandica. Xyalaspis victoriensisappears to be relatively rare, but several specimens have beencaptured by sweeping and beating Acacia trees (New 1979,1982).

Morphology. The Anacharitinae (excluding PetricynipsBelizin) are well characterised by the triangular shape of the

head, protruding mandibles, pronotal plate strongly developedand a more or less elongate petiole. The two Australian speciescan be easily differentiated, because A. zealandica has a veryelongate petiole (not in X. victoriensis), and X. victoriensishas a prominent spine at the apex of the scutellum (not inA. zealandica).

(a)

(a)

(c)

Fig. 3. Figitidae: Emargininae-Eucoilinae. The yellow arrowspoint to the diagnostic structure of the Eucoilinae, the scutellarcup. (a) Thoreauella (Emargininae); (b) Trybliographa (Eucoili-nae); (c) dorsal mesosoma of Trybliographa.

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Subfamily Charipinae (Fig. 2c)

Species list

Alloxysta australiae (Ashmead 1900) (Allotria australiae) holo-type in USNM, specimens available in ANIC

Alloxysta carinata Carver 1992 (Carvercharips carinata inKovalev (1995)) holotype in ANIC, specimens available inANIC

Alloxysta darci (Girault 1933) (Allotria d�arci) holotype in QM

Alloxysta fuscicornis (Hartig 1841) (Xystus fuscicornis) (synHypodiranchis aphidae Froggatt 1904) holotype in ZSM, speci-mens available in ANIC

Alloxysta victrix (Westwood 1833) (Allotria victrix) (synSarothrus io Girault 1932) holotype in OUMNH, holotype ofS. io in QM

Dilapothor carverae Paretas-Martínez & Pujade-Villar 2006holotype in ANIC

Phaenoglyphis villosa (Hartig 1841) (Xystus villosus)(syn Glyptoxysta bifoveata Girault 1931) holotype in ZSM,holotype of G. bifoveata in QM, specimens available inANIC

Thoreauana giraulti Paretas-Martínez & Pujade-Villar 2006holotype in ANIC, specimens available in ANIC

Thoreauana mascagnini (Girault 1935) (Dilyta mascagnini)(Alloxysta mascagnini in Weld (1952)) holotype in QM, speci-mens available in ANIC

Thoreauana nativa Girault 1930 (Alloxysta nativa in Weld(1952)) (Fig. 2c) holotype in QM, specimens available inANIC

Thoreauana thoreauini (Girault 1935) (Alloxysta thoreauini)(Dilyta thoreauini in Weld (1952)) holotype in QM, specimensavailable in ANIC

Systematics. The Charipinae include eight genera: AlloxystaFörster, Apocharips Fergusson, Dilapothor Paretas-Martínez &Pujade-Villar, Dilyta Förster, Lytoxysta Kieffer, Loboptero-charips Paretas-Martínez & Pujade-Villar, PhaenoglyphisFörster and Thoreauana Girault (Paretas-Martínez et al. 2008).

World distribution of genera present in Australia. Al-loxysta and Phaenoglyphis are cosmopolitan. Dilapothor andThoreauana are endemic to Australia.

Biology. Alloxysta, Phaenoglyphis and Lytoxysta are hyper-parasitoids of Aphidiinae (Hymenoptera: Braconidae)and Aphelinidae (Hymenoptera: Chalcidoidea) parasitisingAphididae (Hemiptera); Dilyta and Apocharips are hyperpara-sitoids of Encyrtidae (Hymenoptera: Chalcidoidea) parasitis-ing Psyllidae (Hemiptera) (Menke & Evenhuis 1991). Thehosts of the remaining genera are unknown. Detailed hostrecords and distribution of the Alloxysta species and P. villosafound in Australia are given in Carver (1992).

Morphology. The Charipinae are the only cynipoid groupwith a smooth scutellum without distinct sculpture or

(a)

(b)

(c)

Fig. 4. Figitidae. (a) Xyalophora (Figitinae); (b) Mikeius(Mikeiinae); (c) Thrasorus (Thrasorinae).

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structures (A. carinata has some carinae at the apex of scutel-lum, but these are difficult to see and the overall aspect of thescutellum is smooth and shiny). The rest of the body is alsosmooth and shiny (except for Lytoxysta, with a very fine reticu-late sculpture in head and mesosoma).

Subfamily Emargininae (Fig. 3a)

Species list

Thoreauella amatrix Girault 1930 (Fig. 3a) holotype in QM,specimens available in ANIC

Systematics. The Emargininae currently include four genera:Emargo Weld, Quinlania Kovalev, Thoreauella Girault andWeldiola Kovalev (Ronquist 1999; Pujade-Villar et al. 2002).

World distribution of genera present in Australia.Thoreauella is originally described from Australia. Quinlan(1988) reported Emargo Weld 1960 from Australia, but thisrecord actually refers to Thoreauella. Fontal-Cazalla et al.(2002) reported two undescribed species of Emargininaefrom Africa (Kenya) and Central America (Belize), whichwere ‘provisionally classified’ as belonging to Thoreauella.This generic assignment was justified using the oldest avail-able generic name within the Emargininae (Fontal-Cazallaet al. 2002), because Ronquist (1999) considered the sub-family ‘rather’ homogeneous and with unclear genericcircumscriptions.

Biology. Although the biology of the Emargininae isunknown, they have been found associated with ants. Adultshave been obtained through Berlese funnel extraction of refusedeposits of army ants (Weld 1960), and they have been col-lected in Camponotus nests (Díaz 1978). They are presumablyparasitoids of myrmecophilous Diptera larvae.

Morphology. The Emargininae are well characterised byhaving a deeply bilobed forewing (only the eucoiline genusKleidotoma has a similar forewing among all Cynipoidea).Also, many emarginines have a narrow, rhomboid or elongatearea dorsally on the scutellum defined by prominent lateralcarinae, but this structure is apparently not universally presentin the group (Weld 1960).

Subfamily Eucoilinae (Fig. 3b,c)

Species list

(many undetermined specimens of Eucoilinae are available inANIC)

Diglyphosematini:Gronotoma domestica Girault 1932holotype in QMGronotoma spp. (unidentified species)Eucoilini:Leptopilina boulardi (Barbotin, Carton & Kelner-Pillault,1979) new record (Cothonaspis boulardi) (syn Charips

mahensis Kieffer 1911: the combination Leptopilina mahensiscannot be used because it is a secondary homonym, there isanother Leptopilina mahensis (Kieffer 1911), originallydescribed as Erisphagia mahensis (Nordlander 1980))Holotype in MNHNLeptopilina heterotoma (Thomson 1862) new record(Eucoila heterotoma) (syn. Pseudeucoila bochei Weld 1944)Holotype in MZLULeptopilina lonchaeae (Cameron 1912) comb. n. (Heptam-erocera lonchaeae)holotype in BMNH examined by MF

Leptopilina maria (Girault 1930) comb. n. (Hexaplasta maria)holotype in QM examined by MFLeptopilina spp. (several unidentified species and undescribedspecies)Maacynips distincta Yoshimoto 1963holotype in BPBMMaacynips papuana Yoshimoto 1963 new recordholotype in BPBMMaacynips spp. (several undescribed species)Trybliographa australiensis Ashmead 1900 stat. rev. (synRhoptromeris unimaculus Girault 1931 syn. n.)holotype of australiensis in USNM, holotype of unimaculus inQM checked by Susan Wright & Chris Burwell for MFTrybliographa spp. (at least one undescribed species)(Fig. 3b,c)Ganaspini:Aganaspis spp. (few unidentified and/or undescribed species)new recordAreaspis spp. (several undescribed species) new recordChrestosema sp (unidentified species) new recordDidyctium spp. (several unidentified and undescribed species)new recordEndecameris cf striata Yoshimoto 1962 new recordGanaspis spp. (several unidentified and undescribed species)new recordHexacola aemilia (Girault 1930) comb. n. (Hexaplastaaemilia)holotype in QM examined by MFHexacola florentia (Girault 1930) comb. n. (Hexaplastaflorentia)holotype in QM examined by MFHexacola julia (Girault 1930) comb. n. (Hexaplasta julia)holotype in QM examined by MFHexacola mozarti (Girault 1930) comb. n. (Hexaplastamozarti)holotype in QM examined by MFHexacola thoreauini (Girault 1930) comb. n. (Hexaplastathoreauini)holotype in QM examined by MFHexacola spp. (unidentified and undescribed species)Leptolamina spp. (several unidentified and undescribedspecies) new recordMicreriodes sp. (at least one undescribed species) new recordPseudodiranchis sp. (unidentified species) new recordSinochresta sp. (unidentified species) new record

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Striatovertex occipitalis (Kerrich & Quinlan 1960) (Eucoilaoccipitalis)holotype and paratypes in ANIC, additional paratypes inBMNH and USNMWeldia sp. (unidentified species) new recordKleidotomini:Cothonaspis atricornis Ashmead 1896holotype in BMNH, specimens available in ANICKleidotoma carlylei Girault 1932holotype in QMKleidotoma marguerita (Girault 1931) comb. n. (Pentracritamarguerita)holotype in QMKleidotoma melancholica Girault 1932holotype in QMKleidotoma spp. (several undescribed species)Trichoplastini:Rhoptromeris operarius Girault 1934holotype in QMRhoptromeris spp. (several undescribed species)

Systematics. Eucoilinae is the largest figitid subfamily withat least 85 recognisable genera that either have valid names orare currently being described. They are classified into sixtribes (Forshage & Nordlander 2008; Buffington 2009): Dig-lyphosematini (12), Zaueucoilini (13), Kleidotomini (5), Tri-choplastini (>6), Eucoilini (>8), Ganaspini (>36) (number ofgenera in these figures include many unpublished assignmentsto tribes of genera, and some yet undescribed genera, but acertain number of genera of uncertain placement are notincluded in the count).

World distribution of genera present in Australia. Of the20 genera so far recorded from Australia, eight genera arecosmopolitan or near-cosmopolitan: Chrestosema, Didyctium,Ganaspis, Gronotoma, Hexacola, Kleidotoma, Leptopilinaand Rhoptromeris; eight genera are largely eastern Palaeo-tropical – some, as far as currently known, restricted to theeastern Oriental and Oceanic regions (Areaspis, Pseudodiran-chis, Sinochresta and Weldia) and some also present but rare inwestern Oriental, Southeast Palaearctic and usually also Afro-tropics (Endecameris, Leptolamina, Maacynips and Micreri-odes). Although the distribution of Trybliographa species ismainly Holarctic, with a few tropical species, Aganaspisspecies are typically Neotropical or Oriental, with a fewunique widespread representatives; and the distribution of Cot-honaspis species is mainly Holarctic with one widespreadspecies. Finally, Striatovertex is a Nearctic and Neotropicalgenus, introduced into Hawaii, and probably accidentallyintroduced into Australia (Schick et al. 2011). None are Aus-tralian endemics.

Biology. Eucoilines are solitary koinobiont endoparasitoidsthat attack first-instar larvae of cyclorrhaphous Diptera invarious microhabitats. The adults frequent the sites where theirhosts develop, such as fungi, bird nests, cow manure, rottenwood, rotting vegetation, rotting carcasses, or leaves and

others parts of plants infested by dipteran larvae (Fontal-Cazalla et al. 2002; Buffington et al. 2012).

Few actual host records exist from Australia. Striatovertexoccipitalis was noted in the original description as reared fromSarcophagidae in dung but also taken on carrion (Kerrich &Quinlan 1960). Representatives of this genus elsewhere attackother flies (Calliphoridae, Muscidae) in these habitats (Schicket al. 2011). Leptopilina lonchaeae is stated in the originaldescription as reared from the cucumber fly Lonchaea splen-dida (currently Lamprolonchaea brouniana (Bezzi 1919))(Cameron 1912). However, all other host records of well-known wasps of this genus, kept in laboratories worldwide,pertain to Drosophilidae, and unidentified species of Lep-topilina have been recorded from Drosophilidae also in Aus-tralia (i.e. Tribe 1991; Spinner et al. 2011); therefore, unlessmore extensive documentation of this rearing can be found,this record should be regarded as either questionable oruntypical.

Among the other genera, some host associations are more orless well established from elsewhere. Many of these associa-tions are reviewed, and references given, in Buffington (2007)and Buffington et al. (2012), but a lot of the informationremains difficult to access on label data of museum specimensall over the world, as well as in obscure publications of appliedentomology; a critical review of the sum of records is forth-coming. Gronotoma and Weldia are parasitoids of leaf-miningAgromyzidae, Cothonaspis attack Sepsidae in dung andcarrion, Rhoptromeris attack Chloropidae and Aganaspisattack Tephritidae and Lonchaeidae in fruit. Trybliographa areusually but not always parasitoids of Anthomyiidae, Ganaspisare known from several fly families but often Drosophilidae orChloropidae, Didyctium from several families but often Phori-dae and Hexacola from several families but often Chloropidaeor Ephydridae. Kleidotoma has a broad range of host fly fami-lies worldwide. Chrestosema and Leptolamina have beenrecorded from Drosophilidae, but rearings are not well docu-mented and data are scarce. For the remaining genera, Area-spis, Endecameris, Maacynips, Micreriodes, Pseudodiranchisand Sinochresta, there are still no host records.

Morphology. Eucoilines are easily identified by the presenceof an elevated plate dorsally on the scutellum (Fig. 3c),referred to as the scutellar plate. The plate has a glandularrelease pit in the posterior or central part, the function of whichis unknown. The scutellar plate is present in all eucoilines andis unique to them among parasitic wasps. The majority ofspecies are shining black to brown in colour, and the body islargely polished, without distinct surface sculpture (Fontal-Cazalla et al. 2002; Buffington et al. 2007).

Subfamily Figitinae (Fig. 4a)

Species list

Xyalophora australiana Paretas-Martínez and Pujade-Villar2010 (Fig. 4a) holotype in ANIC examined by JP-M, speci-mens available in ANIC

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Xyalophora mauri Paretas-Martínez and Pujade-Villar 2010holotype in ANIC examined by JP-M

Systematics. The Figitinae comprise 14 genera: Amphithec-tus Hartig, Seitneria Tavares, Sarothrus Hartig, FigitesLatreille, Foersthomorus Pujade-Villar & Petersen-Silva(= Homorus Förster), Neralsia Cameron, Xyalophora Kieffer,Xyalophoroides Jiménez & Pujade-Villar, LonchidiaThomson, Trischiza Förster, Paraschiza Weld, Sarothrioides,Nebulovena Pujade-Villar & Paretas Martinez and ZygosisFörster (Ronquist 1999; Paretas-Martínez et al. 2012; Pujade-Villar et al. 2011).

World distribution of genera present in Australia. Xyalo-phora is cosmopolitan.

Biology. The Figitinae are primary parasitoids of the ‘higher’flies (Diptera: Schizophora) (Buffington et al. 2007). Thereare no verified host records for species of Xyalophora, buttheir phylogenetic position suggests they are parasitoids ofmuscomorph flies in habitats such as cow dung (Jiménez et al.2008).

Morphology. The Figitinae has been defined usually by thelack of derived character states present in other figitid sub-families and has been an obvious ‘classificatory wastebasket’(Ronquist 1999). However, the second abdominal tergite in theform of a ring or collar, sclerotised, sometimes large, usuallylongitudinally furrowed or carinate, can help to distinguish theFigitinae among the Figitidae.

Subfamily Mikeiinae (Fig. 4b)

Species list

Mikeius berryi Buffington 2008 holotype in ANIC examined byMB and JP-M, specimens available in ANIC

Mikeius clavatus Pujade-Villar & Restrepo-Ortiz 2011 holotypein ANIC examined by JP-M, specimens available in ANIC

Mikeius gatesi Buffington 2008 holotype in ANIC examined byMB and JP-M, specimens available in ANIC

Mikeius grandawi Buffington 2008 holotype in ANIC exam-ined by MB and JP-M, specimens available in ANIC

Mikeius hartigi (Girault 1930) (Amblynotus hartigi Girault1930: replacement name for Amblynotus parvus Girault 1929(not Hartig 1840)) (Fig. 4b) holotype in QM examined by MBand JP-M, specimens available in ANIC

Systematics. The Mikeiinae include a single genus: MikeiusBuffington.

World distribution of genera present in Australia. Mikeiusis endemic to Australia.

Biology. Mikeius hartigi (Girault) emerged from Ophelimussp. (Chalcidoidea: Eulophidae) galls on Eucalyptus cinerea F.Muell. from the middle of October to the middle of November.This species is biparental, producing almost equal numbers offemales and males (J La Salle & I-K Kim pers. comm. 1971)(Buffington 2008).

Morphology. Mikeiinae have two carinae in the medianarea of the pronotum that do not form a projected pronotalplate.

Subfamily Thrasorinae (Fig. 4c)

Species list

Cicatrix pilosiscutum (Girault 1929) (Amblynotus pilosiscutum)holotype in QM examined by JP-M

Cicatrix neumannoides Paretas-Martínez & Restrepo-Ortiz2011 holotype in ANIC examined by JP-M

Cicatrix schauffi (Buffington 2008) (Mikeius schauffi) holotypein ANIC examined by MB and JP-M

Palmiriella neumanni (Buffington 2008) (Mikeius neumanni)holotype in ANIC examined by MB and JP-M

Thrasorus pilosus Weld 1944 (Fig. 4c) holotype in USNM,specimens available in ANIC

Thrasorus berlesi (Girault 1937) comb. n. (Amblynotus berlesi)(Thrasorus rieki Paretas-Martínez & Pujade-Villar 2011 syn.n.) holotype in QM examined by JP-M, specimens availablein ANIC

Thrasorus schmidtae Buffington 2008 holotype in ANIC exam-ined by MB and JP-M

Systematics. The Thrasorinae include five genera: CicatrixParetas-Martínez, Myrtopsen Rübsaamen, PalmiriellaPujade-Villar & Paretas-Martínez, Scutimica Ros-Farré andThrasorus Weld (Ros-Farré & Pujade-Villar 2007;Paretas-Martínez et al. 2011).

World distribution of genera present in Australia.Cicatrix, Palmiriella and Thrasorus are endemic toAustralia.

Biology. All records to date indicate that species of Thrasori-nae in Australia are associated with chalcidoid hosts thatinduce galls on species of Acacia and Eucalyptus, althoughmost of these host records await verification (Paretas-Martínezet al. 2011).

Morphology. The Thrasorinae are morphologically definedby the presence of the circumtorular impression (Ros-Farré &Pujade-Villar 2007).

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DISCUSSION

The taxonomy of Australian Cynipoidea is still quite imma-ture. The Austrocynipidae, the Mikeiinae (Figitidae), somegenera of Thrasorinae (Figitidae) and a few species of othergroups (Liopteridae and some subfamilies of Figitidae) areendemic to Australia. Other cynipoid groups present in thisregion have been introduced intentionally for biologicalcontrol purposes (Ibaliidae), or accidentally (Cynipidae).Further studies are needed to understand the evolutionary andbiogeographical history of the Cynipoidea around the worldand in Australia.

The historical biogeography of macrocynipoids has beenexplored through a series of studies (Ronquist 1995b; Nord-lander et al. 1996; Liu 1998). Several cross-Beringian vicari-ance events that presumably date back at least to the terminalEocene, about 33 million years ago (Ma), have been identifiedin ibaliids and liopterids associated with broad-leaved forests.At the end of Eocene, previously continuous Asian and Ameri-can broad-leaved forests became permanently separated byother habitats in the Beringian area through climatic deteriora-tion (Nordlander et al. 1996 and references cited therein). Boththe Ibaliidae and the Liopteridae show a basal split betweenGondwanian and Laurasian groups, suggesting that their earli-est diversification goes back to the Jurassic (about 145 Ma)(Ronquist 1995b). This date agrees roughly with an estimatebased on the amount of morphological character change in thephylogeny of the Ibaliidae before and after the cross-Beringianvicariance in Ibalia (Tremibalia) (Nordlander et al. 1996; Liu1998), as well as more recent non-clock divergence estimatesutilising molecular data (Buffington et al. 2012).

The natural distributions of species of Ibalia appear to beconfined to the Holarctic region and Oriental China. Somespecies of Ibalia have been introduced around the world, Aus-tralia included. Ibalia leucospoides is now found throughoutmost pine-tree plantations in the Southern Hemisphere wherethere are woodwasps (Fernández-Arhex & Corley 2005). Thisis because I. leucospoides has been distributed together withS. noctilio by transportation of infested wood (Corley 2001),or by the deliberate introduction as described in the Biologysection. Four of the five known species of Heteribalia are fromOriental China or northern Vietnam; the remaining species isfrom northern Japan. The only known species of Eileenella isfrom New Guinea, and further intensive fieldwork may revealthis genus to inhabit north-eastern Australia.

Dallatorrellinae are divided between Southeast Asia and theAustralian Region. Seven of the nine species of Dallatorrelli-nae are distributed in southeast and eastern Asia (Liu 2001).The two species of the subfamily that do not occur in thisregion are from Australia and Papua New Guinea. The Dalla-torrellinae are believed to have originated in the Orientalregion and subsequently dispersed to the Australian region(Liu 2001), contrary to an earlier hypothesis by Ronquist(1995a), who suggested the Dallatorrellinae originated in theAustralian region and subsequently dispersed to the Orientalregion. The split of the Dallatorrellinae from the stem speciesof the two Gondwanian subfamilies Liopterinae and

Oberthuerellinae probably corresponds to the break up ofPangea into Gondwana and Laurasia in the middle to lateJurassic (180–145 Ma) (Liu 2001). The Mayrellinae predomi-nantly occur in the Northern Hemisphere. The relatively highdiversity of Paramblynotus in Southeast Asia is considered tobe caused partly by the frequent sea level changes since lateOligocene (29 Ma), which drastically changed the land con-figuration of this area (Liu et al. 2007).

The two species of Cynipidae found in Australia, Andricussp. and Phanacis hypochoeridis, have been introduced to thisregion. Species richness of Cynipidae is distributed unequallyaround the world (Liljeblad & Ronquist 1998; Ronquist 1999).The Aylacini, Diplolepidini are Holarctic (Liljeblad et al.2008); the Cynipini (the most species-rich group of cynipidgallwasps) are Holarctic and Neotropical (Liljeblad et al.2008; Pujade-Villar 2008; Medianero & Nieves-Aldrey 2010);the Pediaspidini are Palaearctic (Liljeblad et al. 2008), butsome species are introduced to South America (Pujade-Villar& Díaz 2001); the Paraulacini and Eschatocerini are restrictedto the Neotropics (Nieves-Aldrey et al. 2009); the Qwaqwaiiniare endemic to South Africa (Liljeblad et al. 2011); theinquilinous Synergini are mainly Holarctic, but some speciesare present in the Oriental and Neotropical regions, and thegenus Rhoophilus Mayr is restricted to the Afrotropics (Melikaet al. 2005; Abe et al. 2007; Nieves-Aldrey & Medianero2010).

Ronquist and Liljeblad (2001) hypothesised that the Cynipi-dae arose in Europe, around the Black Sea, and that theEschatocerini and Rhoophilus arose by dispersal events toSouth America and South Africa, respectively. However, phy-logenetic findings contradict this hypothesis; the Paraulacini,Eschatocerini and Rhoophilus seem to belong to older, morebasal cynipid lineages than the groups present in the Holarcticregion (Nylander 2004). Nieves-Aldrey et al. (2009) gives analternative hypothesis to explain the distribution of thesecynipid groups in the Southern Hemisphere: they suggest thatit may ‘strangely’ indicate a common Gondwanan origin, sincethat also would explain the distribution of the genus Himalo-cynips (in Pediaspidini, closely related to Paraulacini) as aGondwanian relict on the Indian continent. This hypothesis(Nieves-Aldrey et al. 2009) could also explain the finding ofmicrocynipoids associated with galls induced by chalcids alsoin the Australasian region, with members of the figitid sub-family Thrasorinae being reported from chalcid galls on Euca-lyptus and possibly also Acacia (Buffington 2008). If wefollow this hypothesis, it is important to remark that the non-Australian members of Thrasorinae are found only in the Neo-tropical area (except one species found in the southernNearctic) (Ros-Farré & Pujade-Villar 2007, 2009). Further-more, species of another ‘basal’ figitid subfamily, the Plecto-cynipinae, also are found only in the Neotropics and have beenreared from cynipid and chalcid galls on Nothofagus (Ros-Farré & Pujade-Villar 2002, 2007; Buffington & Nieves-Aldrey 2011). Further research on Cynipidae + Figitidae mustbe done to test this hypothesis and to elucidate the biogeo-graphical history of the microcynipoids and the evolutionaryprocess between entomophagy and phytophagy in Cynipoidea.

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The study of the Figitidae phylogeny (Buffington et al.2007) showed that at the base of the figitid tree are groupsassociated with the gall community (at that time included intwo subfamilies: Parnipinae and Thrasorinae). In the lastyears, several taxonomic revisions of the Thrasorinae haveresulted in the split of these groups into several figitid sub-families: Euceroptrinae, Mikeiinae, Plectocynipinae and Thra-sorinae (Ros-Farré & Pujade-Villar 2007; Buffington &Liljeblad 2008; Paretas-Martínez et al. 2011). The phyloge-netic relationships among these ‘basal’ figitid subfamilies werestudied by Paretas-Martínez et al. (2011) but little is knownabout the biology of these figitids associated with chalcid/cynipid galls. Understanding the biology and evolutionaryhistory of these groups (plus Parnipinae) is essential to recon-struct the relationship between gall inducers cynipids andentomophagous figitids. In this sense, a focus should be madeto study the Australian endemic genera Mikeius and Thra-sorus, which also will help in reconstructing the biogeographi-cal evolution of the ‘basal’ Figitidae. The Emargininae(Afrotropical and Australian) may also prove to be critical tounderstanding the figitid biogeographical history.

The second most abundant group in collections of Austral-ian figitids are the Charipinae. The genus most frequentlyfound in Australia is the aphid hyperparasitoid Alloxysta. Theaphid fauna of Australia is relatively meagre and largelyexotic, as is the guild of associated parasitoids and hyperpara-sitoids. Alloxysta fuscicornis is widely distributed in Australiaand is evidently an originally Holarctic, now cosmopolitan,species coexisting with its principal aphid host, the cabbageaphid Brevicoryne brassicae (L.), and the aphidiine (Braconi-dae) host, Diaeretiella rapae (McIntosh), on cabbages.Alloxysta australiae is known only from eastern Australiafrom Queensland to Tasmania, and its host range and distribu-tion does not allow speculation as to its origin, whether Aus-tralasian, or whether Holarctic and accidentally introduced.Alloxysta darci may be an Australasian species because it iswidely spread in Australia, in both cultivated and natural areas,and has been identified also from Tonga. Alloxysta carinatawas collected in areas of native vegetation (Carver 1992). Aparticular case is Phaenoglyphis villosa, the only species ofPhaenoglyphis described in the Southern Hemisphere. Phae-noglyphis villosa has a wide host range within the Aphidiinaeand Aphididae, which explains its cosmopolitan distributionthrough introduction together with its hosts. Phaenoglyphisvillosa is haploid arrhenotokous (females produced biparen-tally) in Europe (Menke & Evenhuis 1991) but deuterotokousin North America (Andrews 1978) and in Australia (Carver1992). A possible explanation, therefore, for the allopatricreproductive behaviour in P. villosa is that this species may bePalaearctic in origin, with one or more individuals of theparthenogenetic component of the species having been intro-duced accidentally from Europe into North America as well asinto Australia (Carver 1992). The two other charipine genera,Dilapothor and Thoreauana, are endemic to Australia; theirhosts are unknown, but their phylogenetic placement indicatesthat their biology may be related to Psyllidae: these two generaform a monophyletic clade together with Dilyta Förster and

Apocharips Fergusson, both psyllid hyperparasitoids (Paretas-Martínez et al. 2007).

The Eucoilinae is the most abundant group within Figitidae,and this is also true in collections of Australian figitids. Theextreme diversification of the Eucoilinae is explained by a shiftfrom parasitisation of hosts associated with the aphid commu-nity (examples of this lineage are Anacharitinae, Aspicerinaeand Charipinae) to parasitisation of the incredibly diversefauna of schizophoran Diptera in exposed and concealed habi-tats (Buffington et al. 2007). The Australian fauna of Eucoili-nae is almost completely unknown (as it happens in mostworld regions), with the identity of the few taxa describedfrom the region very poorly known. Here we have provided afirst attempt at characterising the composition of the Australianfauna, though the fauna remains almost completely unstudiedon the species level, and clearly most species in the region arestill undescribed. All conclusions are highly preliminary, but itcan be noted that there is no endemism on the genus level;most eucoiline genera present are either widespread, Palaeo-tropic or East Palaeotropic. More detailed studies may even-tually find endemic genera new to science, but they are not aconspicuous element. To what extent there is a high degree ofendemism on the species level cannot be assessed at thismoment, since the faunas of both Australia and its neighbour-ing regions are mostly unexplored on this level.

ACKNOWLEDGEMENTS

We are very grateful to Chris Burwell and Susan Wright(QM) for information on type material and to Kathy Schickfor her comments on the type material of Thoreauellaamatrix. We also are grateful to David Notton for hostingMattias Forshage at the BMNH in London. The first author’sstay at ANIC was supported by the Science and EducationMinistry of Spain.

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Accepted for publication 1 August 2012.

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