Overview of Australian Cynipoidea (Hymenoptera)

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Australian Journal of Entomology (2013) 52, 73–86

Overview of Australian Cynipoidea (Hymenoptera) Jordi Paretas-Martínez,1* Mattias Forshage,2 Matthew Buffington,3 Nicole Fisher,4 John La Salle5 and Juli Pujade-Villar1 1

Department of Animal Biology, Faculty of Biology, University of Barcelona, Avda. Diagonal 645, 08028 Barcelona, Spain. 2 Department of Entomology, Swedish Museum of Natural History, Box 50007, SE-104 05 Stockholm, Sweden. 3 Systematic Entomology Laboratory, USDA, c/o NMNH, Smithsonian Institution, 10th & Constitution Avenue NW. PO Box 37012 MRC-168, Washington, DC 20013, USA. 4 Australian 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 for the following genera of Eucoilinae: Aganaspis Lin, Areaspis Lin, Chrestosema Förster, Didyctium Riley, Endecameris Yoshimoto, Ganaspis Förster, Leptolamina Yoshimoto, Micreriodes Yoshimoto, Pseudodiranchis Yoshimoto, Sinochresta Lin and Weldia Yoshimoto. Nine new combinations, two new synonymies and one 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 marguerita comb. 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. (Thrasorus rieki Paretas-Martínez & Pujade-Villar 2011 syn. n.). Aspects on the systematics, distribution, biology and morphology of all cynipoid families and figitid subfamilies in Australia are given. A multi-character online key 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.

I NTRODUCT IO N Cynipoid wasps (Hymenoptera: Apocrita) form a worldwide group that includes ~3000 species (Ronquist 1999). Cynipoids can be divided into two major groups, the so-called macro- and microcynipoids. Macrocynipoids (up to 20 mm in size) have relatively low species richness when compared with the microcynipoids and include the rare Austrocynipidae, as well as Ibaliidae and Liopteridae. The biology of Austrocynipidae is not fully known, but the only species of this family was reared from larvae of an undescribed oecophorid moth dissected from seeds in infested cones of Araucaria (Araucariaceae) (Riek 1971); thus, their host niche is similar to that of the other macrocynipoids, which are koinobiont endoparasites of woodboring or cone-boring insects (Liu 1998). Ibaliids are parasitoids of siricid woodwasp larvae in conifers and hardwoods (Ronquist 1999). Liopterids are putative parasitoids of buprestid, cerambycid and curculionid beetle larvae boring in twigs and stems of deciduous trees and bushes (Ronquist 1995b; Liu 1998), though no definitive rearing records have surfaced (Buffington et al. 2012).

*[email protected] © 2012 The Authors Australian Journal of Entomology © 2012 Australian Entomological Society

The microcynipoids (0.7–8 mm in size), consisting of Cynipidae and Figitidae, are the most species-rich clade of Cynipoidea, with ~1500 described species each. Most species of Cynipidae are gall inducers on woody dicots, but the family also includes inquilines (guests of gall inducers, incapable of gall induction themselves) in galls of other cynipids. Figitids, for which the biology is known, are almost exclusively koinobiont endoparasitoids of endopterygote insect larvae, with most species being primary parasitoids of the muscomorph flies (Diptera: Schizophora) in habitats from leaf-mines to algae to dung and carrion (Buffington 2002; Fontal-Cazalla et al. 2002; Buffington et al. 2007). Figitidae is the most species diverse family within Cynipoidea (Ronquist 1999) and is separated into 12 subfamilies: Anacharitinae, Aspicerinae, Charipinae, Emargininae, Euceroptrinae, Eucoilinae, Figitinae, Mikeiinae, Parnipinae, Plectocynipinae, Pycnostigminae and Thrasorinae (Paretas-Martínez et al. 2011). The cynipoids include economically important species that are 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 many figitids (Anacharitinae, Aspicerinae and Charipinae) conflict doi:10.1111/j.1440-6055.2012.00877.x

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with human interests when parasitising predators or parasitoids of aphid pests. On the other side, entomophagous cynipoids 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 been used as biological control agents of the woodwasp Sirex noctilio F. (Taylor 1976, 1981). As for many small-bodied Hymenoptera, cynipoid biodiversity is certainly underestimated, and much remains to be learned of their systematics, fundamental biology and the ecosystem services that they provide. Cynipoids are especially poorly known outside the Holarctic and tropical Africa. The vast majority of described species are North American, European and sub-Saharan African, whereas many areas of the Eastern Palaearctic, the tropical regions and the Southern Hemisphere (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 an example, only 5% of species of figitids have been described, with 1400 described species (Buffington et al. 2007) compared with an estimated global diversity of 24 000 species (Nordlander 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 authors have contributed to the description of the Australian cynipoid fauna (Ashmead 1900; Kieffer 1911; Weld 1944, 1952; Kerrich & Quinlan 1960; Riek 1970, 1971; New 1979; Carver 1992, 1993; Paretas-Martínez & Pujade-Villar 2006, 2010; Buffington 2008; Paretas-Martínez et al. 2011). The Australian species of three figitid groups have recently been revised: the Charipinae (Carver 1992; Paretas-Martínez & Pujade-Villar 2006), the Figitinae (Paretas-Martínez & Pujade-Villar 2010) and the Thrasorinae + Mikeiinae (Paretas-Martínez et al. 2011). Biological studies on Australian cynipoids have been focused on Ibalia control of siricid wasps (Taylor 1976, 1981 and references therein). A taxonomic overview of all families, subfamilies, genera and species of Cynipoidea present in Australia is presented here, including new records. An interactive and multicharacter LucID key to all Australian cynipoids can be accessed at http://www.ces.csiro.au/keys/Hymenoptera/ Australian_Cynipoidea/Australian-Cynipoidea-Keys.html. Aspects of the taxonomy, distribution, biology and morphology of all cynipoid families and figitid subfamilies in Australia are given.

M ATE RIAL AND ME T HO D S This study was made through an exhaustive literature search on Australian cynipoids, as well as the study of Australian material from the Australian National Insect Collection (ANIC, CSIRO Ecosystem Sciences, Canberra, Australia) and the Queensland Museum (QM, Brisbane, Australia). For © 2012 The Authors Australian Journal of Entomology © 2012 Australian Entomological Society

Eucoilinae, no exhaustive studies were made on the abundant material from Australia present in Australian collections. The treatment of the Eucoilinae in this work is based on literature records, and mainly on studies of relevant types, as well as studies of Australian material in the Natural History Museum, London (BMNH) and The United States National Museum, Washington (USNM). Abbreviations of other holotype repositories are: ANSP (Academy of Natural Sciences of Philadelphia); 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, the following sections are given: List of species with valid name of species present in Australia, author and year of description, original combination, relevant synonymies in Australia (if any), repository of holotype (in cases where the holotype is not deposited in an Australian institution, we also indicate if there are available specimens for study in an Australian institution); Systematics of the family/subfamily worldwide, with number of subfamilies/tribes (where this applies), and genera included (names of genera are given except when the number is too high); World distribution of genera present in Australia to indicate if the genera present in Australia are endemic or also present in other regions; Biology with a brief summary of the biology of the family/subfamily and details of the Australian taxa, if known; Morphology with a brief diagnosis of the subfamily. Overall, the classification of cynipoid wasps presented by Ronquist (1999) is followed; however, several new groups have been described in the Figitidae since that time. For the higher level classification of Figitidae, we follow ParetasMartínez et al. (2011); for eucoiline classification, we follow Forshage and Nordlander (2008). Two special sections have been added: at the end of the Austrocynipidae, a brief summary of its phylogenetic position is given, since this endemic Australian taxon putatively holds a key position in the evolutionary origin of Cynipoidea; inside the biology section of Ibaliidae, a detailed summary of the introduction of Ibalia in Australia is given, since this is an important example of the use of cynipoids in biological control in Australia.

RES U L T S All genera and family-group taxa of Cynipoidea described in Australia are in Table 1, including the new genus records given herein.

Family Austrocynipidae (Fig. 1a) Species list Austrocynips mirabilis Riek 1971 (Fig. 1a) holotype in QM,

specimens available in ANIC

Australian Cynipoidea

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Table 1 Cynipoidea cited in Australia Family: Subfamily Austrocynipidae Ibaliidae Liopteridae: Dallatorrellinae Cynipidae Figitidae: Anacharitinae Figitidae: Charipinae Figitidae: Emargininae Figitidae: Eucoilinae

Figitidae: Figitinae Figitidae: Mikeiinae Figitidae: Thrasorinae

Tribe: Genus Austrocynips Ibalia Dallatorrella Aylacini: Phanacis; Cynipini: Andricus Anacharis, Xyalaspis Alloxysta, Phaenoglyphis, Thoreauana, Dilapothor Thoreauella 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 Xyalophora Mikeius Cicatrix, Palmiriella, Thrasorus

†New record.

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

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

World distribution of genera present in Australia. Austrocynips is endemic to Australia.

Family Ibaliidae (Fig. 1b)

Biology. Austrocynips is only known from three female specimens 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, consistent with the hypothesis that the larvae were phytophagous. However, the collector reported that the specimens were actually reared from larvae of an undescribed oecophorid moth dissected from seeds in infested cones (NW Heather pers. comm. 1971). The oecophorid moths attack the cones when they are still attached to the tree, and the larvae bore through three to six seeds before pupating (Heather 1970). Thus, the host niche of Austrocynips is similar to that of the other macrocynipoids, the Ibaliidae and Liopteridae (Ronquist 1995b). Morphology. Austrocynips has a number of traits that are unique among cynipoids: pterostigma in forewing present; last flagellomere as long or shorter than penultimate flagellomere; radicle indistinctly delineated, but with a line indicating it; 13 flagellomeres in female antenna; posterior margin of pronotum projecting over anterior margin of mesopleuron, not abutting, covering mesothoracic spiracle which is not visible laterally; lateral bars of scutellum absent; nucha absent; annulus with tergal and sternal parts well defined but separate, not fused. Phylogenetic position. Austrocynips is the sister group of all other cynipoids, the latter being supported as a monophyletic group by several character states (Ronquist 1999). This sistergroup relationship is based on the presence in Austrocynips of a number of character states that are more plesiomorphic than other cynipoids (Ronquist 1995b). Some of these traits are unique among cynipoids but are commonly found in the proctotrupoid complex and probably belong to the cynipoid ground

Species list Ibalia leucospoides (Hochenwarth 1785) (Ichneumon leucospoides) (Fig. 1b) holotype presumably lost, specimens avail-

able in ANIC Ibalia rufipes Cresson 1879 holotype in ANSP, specimens

available in ANIC Systematics. The Ibaliidae include three genera: Ibalia Latreille, Heteribalia Sakagami and Eileenella Fergusson. World distribution of genera present in Australia. Ibalia is cosmopolitan. Biology. Ibaliids are parasitoids of siricid woodwasp larvae (Liu & Nordlander 1994). Ibalia leucospoides Hochenwarth attacks eggs and first-instar larvae of Sirex noctilio F. and is endoparasitic until its third instar, a stage at which it becomes an ectoparasite. The Ibalia life cycle is closely related with the development of woodwasp larvae, usually lasting one year (but see Corley et al. 2004). Introduction of Ibalia into Australia. The introduction of Ibalia (and other Sirex parasitoids) into Australia is reported by Taylor (1967, 1976, 1981). Ibalia leucospoides was first successfully introduced to control Sirex noctilio (after early unsuccessful attempts) in New Zealand in 1950, from stocks obtained in England some years earlier. After the discovery of S. noctilio in a private plantation of Pinus radiata D. Don. at Pittwater near Hobart, Tasmania, in 1952, and an initial period of unsuccessful attempts to eradicate this woodwasp in Tasmania, the Tasmanian Department of Agriculture approached the New Zealand authorities for shipments of I. leucospoides and Rhyssa persuasoria (L.) (Hymenoptera: Ichneumonidae, © 2012 The Authors Australian Journal of Entomology © 2012 Australian Entomological Society

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(a)

(b)

Following the discovery in 1961 that S. noctilio was established in Victoria, and the establishment of the National Sirex Fund, a worldwide search for natural enemies of siricids was instituted. Further stocks of I. leucospoides and R. persuasoria were introduced from throughout their range in the northern hemisphere (both are Holarctic) between 1962 and 1973. Nineteen other parasitoid species were introduced to control S. noctilio, including other species of Ibalia. In total, the origin and number of Ibalia females imported to Tasmania in this period 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 and Turkey (1469 specimens), Morocco (40 specimens), Japan (= suprunenkoi 243 specimens). Of all these, only the last three were successfully reared for release and are now established in Australia. Both subspecies of I. leucospoides emerge in summer and autumn to attack siricid larvae hatching soon after oviposition; Ibalia rufipes drewseni emerge in spring to attack siricid larvae in trees where the hatching of the eggs has been delayed over winter.

(c)

Morphology. Ibaliids are characterised by being of much larger size than other cynipoids, with an elongate yet strongly laterally flattened metasoma, and the reduction of wing venation is less strong than other cynipoids. Other diagnostic character states include: marginal cell of forewing very long and narrow; median notch in pronotal crest; pair of posterior scutellar processes; short metafemur; enlarged seventh tergum in female metasoma; apical tubular process present on second metatarsomere.

Family Liopteridae (Fig. 1c) Species list Dallatorrellinae Dallatorrella rubriventris Kieffer 1911 (Fig. 1c) holotype in

BMNH, specimens available in ANIC Fig. 1. Macrocynipoids. (a) Austrocynips (Austrocynipidae); (b) Ibalia (Ibaliidae); (c) Dallatorrella (Liopteridae).

another siricid parasitoid). In 1959 and 1960, shipments of I. leucospoides were received and released in the same plantation; the specimens of I. leucospoides from New Zealand were 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. © 2012 The Authors Australian Journal of Entomology © 2012 Australian Entomological Society

Systematics. The Liopteridae include 10 genera classified into four subfamilies: Dallatorrellinae (2), Mayrellinae (2), Oberthuerellinae (3) and Liopterinae (3). World distribution of genera present in Australia. Dallatorrella is distributed in the Oriental and Australian regions. Biology. The biology of the subfamily Dallatorrellinae is unknown. Females of one species of Dallatorrella have been

Australian Cynipoidea collected on logs of Syzygium (Myrtaceae) in Papua New Guinea, suggesting an association with wood-boring insects on broad-leaved trees (Liu 2001). Species of Mayrellinae have been 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 all these records only report collections on trees attacked by these beetles, lacking direct evidence of a parasitoid-host association. Thus, although all records suggest an association of Liopteridae with wood-boring Coleoptera, further studies are needed to confirm the hosts of this cynipoid family. Morphology. According to Ronquist (1995a,b, 1999), the Liopteridae share at least 12 synapomorphies including: lateral part of cranium expanded, resulting in a swollen gena; deeply foveate pronotal sculpture; presence of mesopleural impression; dorsolateral scutellar processes; metapleural sulcus reaching anterior metapectal margin far above midheight; propodeum with two distinct median carinae; relatively long nucha; and short metatibia. However, there are exceptions to nearly all of these features in form of partial reduction or total loss of the characteristic in some species of Liopteridae; also, many of these morphological structures show some homoplasy within the Cynipoidea due to independent, parallel gains in other cynipoids (Ronquist 1995a).

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(a)

(b)

Family Cynipidae (Fig. 2a) Species list Tribe Aylacini

(c)

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

ANIC Tribe Cynipini Andricus sp. specimens available in ANIC

Systematics. The Cynipidae include about 77 genera and 1400 species classified into eight tribes: the gall inducers Aylacini (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 (Nothofagaceae), two genera), Pediaspidini (gallwasps on Acer (Sapindaceae), three genera), and Qwaqwaiini (gallwasps on Scolopia (Salicaceae), one genus); and the inquilines Synergini (seven genera) (Liljeblad & Ronquist 1998; Liljeblad et al. 2008, 2011; Nieves-Aldrey et al. 2009). World distribution of genera present in Australia. Andricus is an oak gallwasp genus that has more than 300 species (Pujade-Villar et al. 2001), but it is restricted to the Holartic region together with the natural distribution of its host Quercus. The presence of Andricus specimens in Australia

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

must be due to an accidental and isolated introduction. Phanacis hypochoeridis is a Western Palaearctic species that has been introduced in the Nearctic, Neotropical, Australasian and Afrotropical regions (Melika & Prinsloo 2007). Biology. Riek (1970) mentioned for Australian cynipids: ‘Andricus sp. forms galls on introduced oaks. Aylax hypochoeridis, gall-former in the flower stems of the introduced © 2012 The Authors Australian Journal of Entomology © 2012 Australian Entomological Society

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

(a)

Morphology. A combination of ‘loss’ character states can help to identify the Cynipidae (these character states can be found separately in some figitids but never all together): vein Rs+M directed towards middle area of basal vein, pronotal carinae absent, ring of setae at base of metasoma absent and club in the antenna absent.

Family Figitidae (Figs 2b,c,3,4) Systematics. Figitids include about 132 genera and 1400 species classified into 12 subfamilies: Anacharitinae, Aspicerinae, Charipinae, Emargininae, Euceroptrinae, Eucoilinae (the most diverse and species-rich), Figitinae, Mikeiinae, Parnipinae, Plectocynipinae, Pycnostigminae and Thrasorinae (Paretas-Martínez et al. 2011).

(a)

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

Subfamily Anacharitinae (Fig. 2b) Species list Anacharis zealandica Ashmead 1900 (syn Anacharis australiensis Ashmead 1900) (Fig. 2b) holotype in USNM, specimens

available in ANIC Xyalaspis victoriensis New 1979 holotype in ANIC, specimens

(c)

available in ANIC Systematics. The Anacharitinae include nine genera: Acanthaegilips Ashmead, Acothyreus Ashmead, Aegilips Walker, Anacharis Dalman, Calofigites Kieffer, Petricynips Belizin, Proanacharis Kovalev, Solenofigites Díaz and Xyalaspis Hartig (Ronquist 1999). World distribution of genera present in Australia. Anacharis and Xyalaspis are cosmopolitan. Biology. Anacharitines are primary parasitoids of aphidfeeding lacewing larvae (Neuroptera: Hemerobiidae and Chrysopidae) (Ronquist 1999). Australian species are found in association with larvae of Micromus tasmaniae (Walker) and Drepanacra binocula (Newman); these are the two most abundant Hemerobiidae around Melbourne and both are more frequently parasitised by A. zealandica. Xyalaspis victoriensis appears to be relatively rare, but several specimens have been captured by sweeping and beating Acacia trees (New 1979, 1982). Morphology. The Anacharitinae (excluding Petricynips Belizin) are well characterised by the triangular shape of the © 2012 The Authors Australian Journal of Entomology © 2012 Australian Entomological Society

Fig. 3. Figitidae: Emargininae-Eucoilinae. The yellow arrows point to the diagnostic structure of the Eucoilinae, the scutellar cup. (a) Thoreauella (Emargininae); (b) Trybliographa (Eucoilinae); (c) dorsal mesosoma of Trybliographa.

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

Australian Cynipoidea (a)

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

ANIC Alloxysta darci (Girault 1933) (Allotria d⬘arci) holotype in QM Alloxysta fuscicornis (Hartig 1841) (Xystus fuscicornis) (syn Hypodiranchis aphidae Froggatt 1904) holotype in ZSM, speci-

mens available in ANIC Alloxysta victrix (Westwood 1833) (Allotria victrix) (syn Sarothrus io Girault 1932) holotype in OUMNH, holotype of

S. io in QM Dilapothor carverae Paretas-Martínez & Pujade-Villar 2006

holotype in ANIC (b)

Phaenoglyphis villosa (Hartig 1841) (Xystus villosus) (syn Glyptoxysta bifoveata Girault 1931) holotype in ZSM,

holotype of G. bifoveata in QM, specimens available in ANIC Thoreauana giraulti Paretas-Martínez & Pujade-Villar 2006

holotype 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 in

ANIC (c)

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

available in ANIC Systematics. The Charipinae include eight genera: Alloxysta Förster, Apocharips Fergusson, Dilapothor Paretas-Martínez & Pujade-Villar, Dilyta Förster, Lytoxysta Kieffer, Lobopterocharips Paretas-Martínez & Pujade-Villar, Phaenoglyphis Förster and Thoreauana Girault (Paretas-Martínez et al. 2008). World distribution of genera present in Australia. Alloxysta and Phaenoglyphis are cosmopolitan. Dilapothor and Thoreauana are endemic to Australia.

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

Subfamily Charipinae (Fig. 2c) Species list Alloxysta australiae (Ashmead 1900) (Allotria australiae) holo-

type in USNM, specimens available in ANIC

Biology. Alloxysta, Phaenoglyphis and Lytoxysta are hyperparasitoids of Aphidiinae (Hymenoptera: Braconidae) and Aphelinidae (Hymenoptera: Chalcidoidea) parasitising Aphididae (Hemiptera); Dilyta and Apocharips are hyperparasitoids of Encyrtidae (Hymenoptera: Chalcidoidea) parasitising Psyllidae (Hemiptera) (Menke & Evenhuis 1991). The hosts of the remaining genera are unknown. Detailed host records and distribution of the Alloxysta species and P. villosa found in Australia are given in Carver (1992). Morphology. The Charipinae are the only cynipoid group with a smooth scutellum without distinct sculpture or © 2012 The Authors Australian Journal of Entomology © 2012 Australian Entomological Society

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structures (A. carinata has some carinae at the apex of scutellum, but these are difficult to see and the overall aspect of the scutellum is smooth and shiny). The rest of the body is also smooth and shiny (except for Lytoxysta, with a very fine reticulate 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 and Weldiola 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 this record actually refers to Thoreauella. Fontal-Cazalla et al. (2002) reported two undescribed species of Emargininae from Africa (Kenya) and Central America (Belize), which were ‘provisionally classified’ as belonging to Thoreauella. This generic assignment was justified using the oldest available generic name within the Emargininae (Fontal-Cazalla et al. 2002), because Ronquist (1999) considered the subfamily ‘rather’ homogeneous and with unclear generic circumscriptions. Biology. Although the biology of the Emargininae is unknown, they have been found associated with ants. Adults have been obtained through Berlese funnel extraction of refuse deposits of army ants (Weld 1960), and they have been collected in Camponotus nests (Díaz 1978). They are presumably parasitoids of myrmecophilous Diptera larvae. Morphology. The Emargininae are well characterised by having a deeply bilobed forewing (only the eucoiline genus Kleidotoma has a similar forewing among all Cynipoidea). Also, many emarginines have a narrow, rhomboid or elongate area dorsally on the scutellum defined by prominent lateral carinae, but this structure is apparently not universally present in the group (Weld 1960).

Subfamily Eucoilinae (Fig. 3b,c) Species list

(many undetermined specimens of Eucoilinae are available in ANIC) Diglyphosematini: Gronotoma domestica Girault 1932 holotype in QM Gronotoma spp. (unidentified species) Eucoilini: Leptopilina boulardi (Barbotin, Carton & Kelner-Pillault, 1979) new record (Cothonaspis boulardi) (syn Charips © 2012 The Authors Australian Journal of Entomology © 2012 Australian Entomological Society

mahensis Kieffer 1911: the combination Leptopilina mahensis cannot be used because it is a secondary homonym, there is another Leptopilina mahensis (Kieffer 1911), originally described as Erisphagia mahensis (Nordlander 1980)) Holotype in MNHN Leptopilina heterotoma (Thomson 1862) new record (Eucoila heterotoma) (syn. Pseudeucoila bochei Weld 1944) Holotype in MZLU Leptopilina lonchaeae (Cameron 1912) comb. n. (Heptamerocera lonchaeae) holotype in BMNH examined by MF Leptopilina maria (Girault 1930) comb. n. (Hexaplasta maria) holotype in QM examined by MF Leptopilina spp. (several unidentified species and undescribed species) Maacynips distincta Yoshimoto 1963 holotype in BPBM Maacynips papuana Yoshimoto 1963 new record holotype in BPBM Maacynips spp. (several undescribed species) Trybliographa australiensis Ashmead 1900 stat. rev. (syn Rhoptromeris unimaculus Girault 1931 syn. n.) holotype of australiensis in USNM, holotype of unimaculus in QM checked by Susan Wright & Chris Burwell for MF Trybliographa spp. (at least one undescribed species) (Fig. 3b,c) Ganaspini: Aganaspis spp. (few unidentified and/or undescribed species) new record Areaspis spp. (several undescribed species) new record Chrestosema sp (unidentified species) new record Didyctium spp. (several unidentified and undescribed species) new record Endecameris cf striata Yoshimoto 1962 new record Ganaspis spp. (several unidentified and undescribed species) new record Hexacola aemilia (Girault 1930) comb. n. (Hexaplasta aemilia) holotype in QM examined by MF Hexacola florentia (Girault 1930) comb. n. (Hexaplasta florentia) holotype in QM examined by MF Hexacola julia (Girault 1930) comb. n. (Hexaplasta julia) holotype in QM examined by MF Hexacola mozarti (Girault 1930) comb. n. (Hexaplasta mozarti) holotype in QM examined by MF Hexacola thoreauini (Girault 1930) comb. n. (Hexaplasta thoreauini) holotype in QM examined by MF Hexacola spp. (unidentified and undescribed species) Leptolamina spp. (several unidentified and undescribed species) new record Micreriodes sp. (at least one undescribed species) new record Pseudodiranchis sp. (unidentified species) new record Sinochresta sp. (unidentified species) new record

Australian Cynipoidea Striatovertex occipitalis (Kerrich & Quinlan 1960) (Eucoila occipitalis) holotype and paratypes in ANIC, additional paratypes in BMNH and USNM Weldia sp. (unidentified species) new record Kleidotomini: Cothonaspis atricornis Ashmead 1896 holotype in BMNH, specimens available in ANIC Kleidotoma carlylei Girault 1932 holotype in QM Kleidotoma marguerita (Girault 1931) comb. n. (Pentracrita marguerita) holotype in QM Kleidotoma melancholica Girault 1932 holotype in QM Kleidotoma spp. (several undescribed species) Trichoplastini: Rhoptromeris operarius Girault 1934 holotype in QM Rhoptromeris spp. (several undescribed species) Systematics. Eucoilinae is the largest figitid subfamily with at least 85 recognisable genera that either have valid names or are currently being described. They are classified into six tribes (Forshage & Nordlander 2008; Buffington 2009): Diglyphosematini (12), Zaueucoilini (13), Kleidotomini (5), Trichoplastini (>6), Eucoilini (>8), Ganaspini (>36) (number of genera in these figures include many unpublished assignments to tribes of genera, and some yet undescribed genera, but a certain number of genera of uncertain placement are not included in the count). World distribution of genera present in Australia. Of the 20 genera so far recorded from Australia, eight genera are cosmopolitan or near-cosmopolitan: Chrestosema, Didyctium, Ganaspis, Gronotoma, Hexacola, Kleidotoma, Leptopilina and Rhoptromeris; eight genera are largely eastern Palaeotropical – some, as far as currently known, restricted to the eastern Oriental and Oceanic regions (Areaspis, Pseudodiranchis, Sinochresta and Weldia) and some also present but rare in western Oriental, Southeast Palaearctic and usually also Afrotropics (Endecameris, Leptolamina, Maacynips and Micreriodes). Although the distribution of Trybliographa species is mainly Holarctic, with a few tropical species, Aganaspis species are typically Neotropical or Oriental, with a few unique widespread representatives; and the distribution of Cothonaspis species is mainly Holarctic with one widespread species. Finally, Striatovertex is a Nearctic and Neotropical genus, introduced into Hawaii, and probably accidentally introduced into Australia (Schick et al. 2011). None are Australian endemics.

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others parts of plants infested by dipteran larvae (FontalCazalla et al. 2002; Buffington et al. 2012). Few actual host records exist from Australia. Striatovertex occipitalis was noted in the original description as reared from Sarcophagidae in dung but also taken on carrion (Kerrich & Quinlan 1960). Representatives of this genus elsewhere attack other flies (Calliphoridae, Muscidae) in these habitats (Schick et al. 2011). Leptopilina lonchaeae is stated in the original description as reared from the cucumber fly Lonchaea splendida (currently Lamprolonchaea brouniana (Bezzi 1919)) (Cameron 1912). However, all other host records of wellknown wasps of this genus, kept in laboratories worldwide, pertain to Drosophilidae, and unidentified species of Leptopilina have been recorded from Drosophilidae also in Australia (i.e. Tribe 1991; Spinner et al. 2011); therefore, unless more extensive documentation of this rearing can be found, this record should be regarded as either questionable or untypical. Among the other genera, some host associations are more or less well established from elsewhere. Many of these associations are reviewed, and references given, in Buffington (2007) and Buffington et al. (2012), but a lot of the information remains difficult to access on label data of museum specimens all over the world, as well as in obscure publications of applied entomology; a critical review of the sum of records is forthcoming. Gronotoma and Weldia are parasitoids of leaf-mining Agromyzidae, Cothonaspis attack Sepsidae in dung and carrion, Rhoptromeris attack Chloropidae and Aganaspis attack Tephritidae and Lonchaeidae in fruit. Trybliographa are usually but not always parasitoids of Anthomyiidae, Ganaspis are known from several fly families but often Drosophilidae or Chloropidae, Didyctium from several families but often Phoridae and Hexacola from several families but often Chloropidae or Ephydridae. Kleidotoma has a broad range of host fly families worldwide. Chrestosema and Leptolamina have been recorded from Drosophilidae, but rearings are not well documented and data are scarce. For the remaining genera, Areaspis, Endecameris, Maacynips, Micreriodes, Pseudodiranchis and Sinochresta, there are still no host records. Morphology. Eucoilines are easily identified by the presence of an elevated plate dorsally on the scutellum (Fig. 3c), referred to as the scutellar plate. The plate has a glandular release pit in the posterior or central part, the function of which is unknown. The scutellar plate is present in all eucoilines and is unique to them among parasitic wasps. The majority of species are shining black to brown in colour, and the body is largely polished, without distinct surface sculpture (FontalCazalla et al. 2002; Buffington et al. 2007).

Subfamily Figitinae (Fig. 4a) Biology. Eucoilines are solitary koinobiont endoparasitoids that attack first-instar larvae of cyclorrhaphous Diptera in various microhabitats. The adults frequent the sites where their hosts develop, such as fungi, bird nests, cow manure, rotten wood, rotting vegetation, rotting carcasses, or leaves and

Species list Xyalophora australiana Paretas-Martínez and Pujade-Villar 2010 (Fig. 4a) holotype in ANIC examined by JP-M, speci-

mens available in ANIC © 2012 The Authors Australian Journal of Entomology © 2012 Australian Entomological Society

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Xyalophora mauri Paretas-Martínez and Pujade-Villar 2010

holotype in ANIC examined by JP-M Systematics. The Figitinae comprise 14 genera: Amphithectus Hartig, Seitneria Tavares, Sarothrus Hartig, Figites Latreille, Foersthomorus Pujade-Villar & Petersen-Silva (= Homorus Förster), Neralsia Cameron, Xyalophora Kieffer, Xyalophoroides Jiménez & Pujade-Villar, Lonchidia Thomson, Trischiza Förster, Paraschiza Weld, Sarothrioides, Nebulovena Pujade-Villar & Paretas Martinez and Zygosis Förster (Ronquist 1999; Paretas-Martínez et al. 2012; PujadeVillar et al. 2011). World distribution of genera present in Australia. Xyalophora is cosmopolitan. Biology. The Figitinae are primary parasitoids of the ‘higher’ flies (Diptera: Schizophora) (Buffington et al. 2007). There are no verified host records for species of Xyalophora, but their phylogenetic position suggests they are parasitoids of muscomorph flies in habitats such as cow dung (Jiménez et al. 2008).

Biology. Mikeius hartigi (Girault) emerged from Ophelimus sp. (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 of females and males (J La Salle & I-K Kim pers. comm. 1971) (Buffington 2008). Morphology. Mikeiinae have two carinae in the median area of the pronotum that do not form a projected pronotal plate.

Subfamily Thrasorinae (Fig. 4c) Species list Cicatrix pilosiscutum (Girault 1929) (Amblynotus pilosiscutum)

holotype in QM examined by JP-M Cicatrix neumannoides Paretas-Martínez & Restrepo-Ortiz 2011 holotype in ANIC examined by JP-M Cicatrix schauffi (Buffington 2008) (Mikeius schauffi) holotype

in ANIC examined by MB and JP-M Morphology. The Figitinae has been defined usually by the lack of derived character states present in other figitid subfamilies and has been an obvious ‘classificatory wastebasket’ (Ronquist 1999). However, the second abdominal tergite in the form of a ring or collar, sclerotised, sometimes large, usually longitudinally furrowed or carinate, can help to distinguish the Figitinae among the Figitidae.

Subfamily Mikeiinae (Fig. 4b)

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 available

in ANIC Species list Mikeius berryi Buffington 2008 holotype in ANIC examined by

Thrasorus schmidtae Buffington 2008 holotype in ANIC exam-

MB and JP-M, specimens available in ANIC

ined by MB and JP-M

Mikeius clavatus Pujade-Villar & Restrepo-Ortiz 2011 holotype

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

in ANIC examined by JP-M, specimens available in ANIC Mikeius gatesi Buffington 2008 holotype in ANIC examined by MB 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 Girault 1930: replacement name for Amblynotus parvus Girault 1929 (not Hartig 1840)) (Fig. 4b) holotype in QM examined by MB

and JP-M, specimens available in ANIC Systematics. The Mikeiinae include a single genus: Mikeius Buffington. World distribution of genera present in Australia. Mikeius is endemic to Australia. © 2012 The Authors Australian Journal of Entomology © 2012 Australian Entomological Society

World distribution of genera present in Australia. Cicatrix, Palmiriella and Thrasorus are endemic to Australia. Biology. All records to date indicate that species of Thrasorinae in Australia are associated with chalcidoid hosts that induce galls on species of Acacia and Eucalyptus, although most of these host records await verification (Paretas-Martínez et al. 2011). Morphology. The Thrasorinae are morphologically defined by the presence of the circumtorular impression (Ros-Farré & Pujade-Villar 2007).

Australian Cynipoidea DISCUSSION The taxonomy of Australian Cynipoidea is still quite immature. The Austrocynipidae, the Mikeiinae (Figitidae), some genera of Thrasorinae (Figitidae) and a few species of other groups (Liopteridae and some subfamilies of Figitidae) are endemic to Australia. Other cynipoid groups present in this region have been introduced intentionally for biological control purposes (Ibaliidae), or accidentally (Cynipidae). Further studies are needed to understand the evolutionary and biogeographical history of the Cynipoidea around the world and in Australia. The historical biogeography of macrocynipoids has been explored through a series of studies (Ronquist 1995b; Nordlander et al. 1996; Liu 1998). Several cross-Beringian vicariance events that presumably date back at least to the terminal Eocene, about 33 million years ago (Ma), have been identified in ibaliids and liopterids associated with broad-leaved forests. At the end of Eocene, previously continuous Asian and American broad-leaved forests became permanently separated by other habitats in the Beringian area through climatic deterioration (Nordlander et al. 1996 and references cited therein). Both the Ibaliidae and the Liopteridae show a basal split between Gondwanian and Laurasian groups, suggesting that their earliest diversification goes back to the Jurassic (about 145 Ma) (Ronquist 1995b). This date agrees roughly with an estimate based on the amount of morphological character change in the phylogeny of the Ibaliidae before and after the cross-Beringian vicariance in Ibalia (Tremibalia) (Nordlander et al. 1996; Liu 1998), as well as more recent non-clock divergence estimates utilising molecular data (Buffington et al. 2012). The natural distributions of species of Ibalia appear to be confined to the Holarctic region and Oriental China. Some species of Ibalia have been introduced around the world, Australia included. Ibalia leucospoides is now found throughout most pine-tree plantations in the Southern Hemisphere where there are woodwasps (Fernández-Arhex & Corley 2005). This is because I. leucospoides has been distributed together with S. noctilio by transportation of infested wood (Corley 2001), or by the deliberate introduction as described in the Biology section. Four of the five known species of Heteribalia are from Oriental China or northern Vietnam; the remaining species is from northern Japan. The only known species of Eileenella is from New Guinea, and further intensive fieldwork may reveal this genus to inhabit north-eastern Australia. Dallatorrellinae are divided between Southeast Asia and the Australian Region. Seven of the nine species of Dallatorrellinae are distributed in southeast and eastern Asia (Liu 2001). The two species of the subfamily that do not occur in this region are from Australia and Papua New Guinea. The Dallatorrellinae are believed to have originated in the Oriental region and subsequently dispersed to the Australian region (Liu 2001), contrary to an earlier hypothesis by Ronquist (1995a), who suggested the Dallatorrellinae originated in the Australian region and subsequently dispersed to the Oriental region. The split of the Dallatorrellinae from the stem species of the two Gondwanian subfamilies Liopterinae and

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Oberthuerellinae probably corresponds to the break up of Pangea into Gondwana and Laurasia in the middle to late Jurassic (180–145 Ma) (Liu 2001). The Mayrellinae predominantly occur in the Northern Hemisphere. The relatively high diversity of Paramblynotus in Southeast Asia is considered to be caused partly by the frequent sea level changes since late Oligocene (29 Ma), which drastically changed the land configuration of this area (Liu et al. 2007). The two species of Cynipidae found in Australia, Andricus sp. and Phanacis hypochoeridis, have been introduced to this region. Species richness of Cynipidae is distributed unequally around the world (Liljeblad & Ronquist 1998; Ronquist 1999). The Aylacini, Diplolepidini are Holarctic (Liljeblad et al. 2008); the Cynipini (the most species-rich group of cynipid gallwasps) are Holarctic and Neotropical (Liljeblad et al. 2008; Pujade-Villar 2008; Medianero & Nieves-Aldrey 2010); the Pediaspidini are Palaearctic (Liljeblad et al. 2008), but some species are introduced to South America (Pujade-Villar & Díaz 2001); the Paraulacini and Eschatocerini are restricted to the Neotropics (Nieves-Aldrey et al. 2009); the Qwaqwaiini are endemic to South Africa (Liljeblad et al. 2011); the inquilinous Synergini are mainly Holarctic, but some species are present in the Oriental and Neotropical regions, and the genus Rhoophilus Mayr is restricted to the Afrotropics (Melika et al. 2005; Abe et al. 2007; Nieves-Aldrey & Medianero 2010). Ronquist and Liljeblad (2001) hypothesised that the Cynipidae arose in Europe, around the Black Sea, and that the Eschatocerini and Rhoophilus arose by dispersal events to South America and South Africa, respectively. However, phylogenetic findings contradict this hypothesis; the Paraulacini, Eschatocerini and Rhoophilus seem to belong to older, more basal cynipid lineages than the groups present in the Holarctic region (Nylander 2004). Nieves-Aldrey et al. (2009) gives an alternative hypothesis to explain the distribution of these cynipid groups in the Southern Hemisphere: they suggest that it may ‘strangely’ indicate a common Gondwanan origin, since that also would explain the distribution of the genus Himalocynips (in Pediaspidini, closely related to Paraulacini) as a Gondwanian relict on the Indian continent. This hypothesis (Nieves-Aldrey et al. 2009) could also explain the finding of microcynipoids associated with galls induced by chalcids also in the Australasian region, with members of the figitid subfamily Thrasorinae being reported from chalcid galls on Eucalyptus and possibly also Acacia (Buffington 2008). If we follow this hypothesis, it is important to remark that the nonAustralian members of Thrasorinae are found only in the Neotropical area (except one species found in the southern Nearctic) (Ros-Farré & Pujade-Villar 2007, 2009). Furthermore, species of another ‘basal’ figitid subfamily, the Plectocynipinae, also are found only in the Neotropics and have been reared from cynipid and chalcid galls on Nothofagus (RosFarré & Pujade-Villar 2002, 2007; Buffington & NievesAldrey 2011). Further research on Cynipidae + Figitidae must be done to test this hypothesis and to elucidate the biogeographical history of the microcynipoids and the evolutionary process between entomophagy and phytophagy in Cynipoidea. © 2012 The Authors Australian Journal of Entomology © 2012 Australian Entomological Society

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The study of the Figitidae phylogeny (Buffington et al. 2007) showed that at the base of the figitid tree are groups associated with the gall community (at that time included in two subfamilies: Parnipinae and Thrasorinae). In the last years, several taxonomic revisions of the Thrasorinae have resulted in the split of these groups into several figitid subfamilies: Euceroptrinae, Mikeiinae, Plectocynipinae and Thrasorinae (Ros-Farré & Pujade-Villar 2007; Buffington & Liljeblad 2008; Paretas-Martínez et al. 2011). The phylogenetic relationships among these ‘basal’ figitid subfamilies were studied by Paretas-Martínez et al. (2011) but little is known about the biology of these figitids associated with chalcid/ cynipid galls. Understanding the biology and evolutionary history of these groups (plus Parnipinae) is essential to reconstruct the relationship between gall inducers cynipids and entomophagous figitids. In this sense, a focus should be made to study the Australian endemic genera Mikeius and Thrasorus, which also will help in reconstructing the biogeographical evolution of the ‘basal’ Figitidae. The Emargininae (Afrotropical and Australian) may also prove to be critical to understanding the figitid biogeographical history. The second most abundant group in collections of Australian figitids are the Charipinae. The genus most frequently found in Australia is the aphid hyperparasitoid Alloxysta. The aphid fauna of Australia is relatively meagre and largely exotic, as is the guild of associated parasitoids and hyperparasitoids. Alloxysta fuscicornis is widely distributed in Australia and is evidently an originally Holarctic, now cosmopolitan, species coexisting with its principal aphid host, the cabbage aphid Brevicoryne brassicae (L.), and the aphidiine (Braconidae) host, Diaeretiella rapae (McIntosh), on cabbages. Alloxysta australiae is known only from eastern Australia from Queensland to Tasmania, and its host range and distribution does not allow speculation as to its origin, whether Australasian, or whether Holarctic and accidentally introduced. Alloxysta darci may be an Australasian species because it is widely spread in Australia, in both cultivated and natural areas, and has been identified also from Tonga. Alloxysta carinata was collected in areas of native vegetation (Carver 1992). A particular case is Phaenoglyphis villosa, the only species of Phaenoglyphis described in the Southern Hemisphere. Phaenoglyphis villosa has a wide host range within the Aphidiinae and Aphididae, which explains its cosmopolitan distribution through introduction together with its hosts. Phaenoglyphis villosa is haploid arrhenotokous (females produced biparentally) in Europe (Menke & Evenhuis 1991) but deuterotokous in North America (Andrews 1978) and in Australia (Carver 1992). A possible explanation, therefore, for the allopatric reproductive behaviour in P. villosa is that this species may be Palaearctic in origin, with one or more individuals of the parthenogenetic component of the species having been introduced accidentally from Europe into North America as well as into Australia (Carver 1992). The two other charipine genera, Dilapothor and Thoreauana, are endemic to Australia; their hosts are unknown, but their phylogenetic placement indicates that their biology may be related to Psyllidae: these two genera form a monophyletic clade together with Dilyta Förster and © 2012 The Authors Australian Journal of Entomology © 2012 Australian Entomological Society

Apocharips Fergusson, both psyllid hyperparasitoids (ParetasMartínez et al. 2007). The Eucoilinae is the most abundant group within Figitidae, and this is also true in collections of Australian figitids. The extreme diversification of the Eucoilinae is explained by a shift from parasitisation of hosts associated with the aphid community (examples of this lineage are Anacharitinae, Aspicerinae and Charipinae) to parasitisation of the incredibly diverse fauna of schizophoran Diptera in exposed and concealed habitats (Buffington et al. 2007). The Australian fauna of Eucoilinae is almost completely unknown (as it happens in most world regions), with the identity of the few taxa described from the region very poorly known. Here we have provided a first attempt at characterising the composition of the Australian fauna, though the fauna remains almost completely unstudied on the species level, and clearly most species in the region are still undescribed. All conclusions are highly preliminary, but it can be noted that there is no endemism on the genus level; most eucoiline genera present are either widespread, Palaeotropic or East Palaeotropic. More detailed studies may eventually find endemic genera new to science, but they are not a conspicuous element. To what extent there is a high degree of endemism on the species level cannot be assessed at this moment, since the faunas of both Australia and its neighbouring regions are mostly unexplored on this level.

ACK N OW L EDGEMEN T S We are very grateful to Chris Burwell and Susan Wright (QM) for information on type material and to Kathy Schick for her comments on the type material of Thoreauella amatrix. We also are grateful to David Notton for hosting Mattias Forshage at the BMNH in London. The first author’s stay at ANIC was supported by the Science and Education Ministry of Spain.

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