Zootaxa 3736 (1): 001–053 ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2013 Magnolia Press ISSN 1175-5334 (online edition) http://dx.doi.org/10.11646/zootaxa.3736.1.1 http://zoobank.org/urn:lsid:zoobank.org:pub:BC992886-EDAB-4D3E-A1F7-DEA34DB06A10 Review of the hymenopteran fauna of New Caledonia with a checklist of species

JOHN T. JENNINGS1, LARS KROGMANN2 & CHRIS BURWELL3,4 1Australian Centre for Evolutionary Biology and Biodiversity, and School of Earth and Environmental Sciences, The University of Adelaide, SA 5005, Australia. E-mail: [email protected] 2State Museum of Natural History Stuttgart, Entomology, Germany. E-mail: [email protected] 3Natural Environments Program, Queensland Museum, PO Box 3300, South Brisbane, QLD 4101, Australia. E-mail: chris.bur- [email protected] 4Environmental Futures Centre and Griffith School of Environment, Griffith University, Nathan, QLD 4111, Australia

Table of contents

Abstract ...... 1 Introduction ...... 2 Material and methods ...... 4 The Hymenopteran Fauna of New Caledonia ...... 4 ...... 4 ...... 4 Xiphydrioidea ...... 33 Orussoidea ...... 33 Stephanoidea ...... 33 Megalyroidea ...... 33 Trigonaloidea ...... 33 ...... 34 ...... 34 ...... 35 Diaprioidea ...... 35 ...... 36 ...... 36 Chalcidoidea ...... 37 ...... 39 ...... 39 ...... 40 ...... 40 ...... 42 Apiformes ()...... 42 ...... 43 Conclusions ...... 43 Acknowledgements ...... 45 References ...... 45

Abstract

The hymenopteran fauna of New Caledonia is reviewed and compared with that of Australia and New Zealand, as well as other islands in the south-west Pacific. In conclusion, several different scenarios (e.g., recent dispersal events and radia- tions) can be used to explain the extant distribution of New Caledonian . A detailed checklist of 409 species and subspecies of Hymenoptera of New Caledonia is provided, along with estimates of the undescribed fauna, and a sum- mary of the general biology of the families represented in the region.

Key words: Hymenoptera, , , , Australia, New Zealand, south-west Pacific, biogeography

Accepted by M. Buffington: 1 Oct. 2013; published: 11 Nov. 2013 1 Introduction

The French Overseas Territory of New Caledonia is located in the south-west Pacific about 1,200 km east of Queensland, Australia, and 1,500 km north-north-west of New Zealand. The land area is about 19,103 km2, much smaller than that of New Zealand (269,057 km2) (Pauly & Munzinger 2003) and Australia (7,659,861 km2) (Geoscience Australia 2010). New Caledonia comprises the main island of Grande Terre which is dominated by a chain of high mountains which run along its entire length, and several smaller islands; the Belep archipelago to the north, Loyalty Islands to the east, Île des Pins (Isle of Pines) to the south, and the Chesterfield Islands and Bellona Reefs to the west.

FIGURE 1. Map showing position of New Caledonia relative to other land masses in the south-west Pacific.

New Caledonia separated from the margin of northeast Australia about 65 mya and reached its present position around 50 mya (see, for example, Coleman 1980; Raven 1980; Yan & Kroenke 1993). New Caledonia has until recently been considered as a continental island because of its geological origin and the presence of phylogenetic relicts (see, for example, Holloway 1979; Morat 1993), and as a consequence, its biota were considered to date from, at least partly, Gondwanan times. Some workers, such as Diamond (1984), have argued for relatively recent dispersal and subsequent radiation to account for some of the New Caledonian biota, although Bauer & Sadlier (2000) suggested that endemism of herpetofauna is extremely high in New Caledonia. Bauer (1988) concluded that “the long independent history of the island has accounted for the [herpeto]faunal composition”, and “there is little

2 · Zootaxa 3736 (1) © 2013 Magnolia Press JENNINGS ET AL. evidence of “recent” immigration via dispersal.” More recently, Bauer & Jackman (2006) suggested that “microendemism [of lizards] is the result of geologically and climatically-mediated fragmentation of habitats throughout the second half of the Tertiary”. Sharma & Giribet (2009) in their multi-gene phylogenetic analyses of the family Troglosironidae (Opiliones) found support for evolutionary relicts in the New Caledonian opilione fauna, and suggested a scenario of an ancient (> 200 Ma) origin of the family, with subsequent diversification of extant lineages in the Eocene. Geological evidence indicates Eocene ophiolitic obduction (39-36 mya; Lowry 1999; Lee et al. 2001), which resulted in the overthrusting of peridotite sheets, which today dominate the southern one-third of Grande Terre, followed by marine submersion(s) in the Oligocene and by Miocene marine regression and mountain building (see Bauer and Jackman 2006). This evidence and recent biogeographic and phylogenetic studies, suggest that at least some of the biota of New Caledonia are no older than the Oligocene (Grandcolas et al. 2008). An example supporting this is the study by Murienne et al. (2005) of the cockroach genus Angustonicus (subfamily Tryonicinae). Both geological and molecular dating showed that this group diversified less than two million years ago and that local species richness and endemism are not necessarily indicative of an old Gondwanan origin. This and other examples cited in Grandcolas et al. (2008), suggest that the biota is not that of a continental island which has retained many relict groups, but is that of an oceanic island with a rich local biota dominated by recent radiations following many dispersal events (Gillespie & Roderick 2002; Grandcolas et al. 2008). Murienne (2009) recently summarised three different models for explaining the biota of New Caledonia. The first of these is the ‘museum model’ or ‘Noah’s Ark model’ which implies that the island has acted as a ‘museum’ from Gondwanan times and the origin and diversification of New Caledonian lineages predates any submersion event(s). The other two models involve diversification after the land re-emerged, either with a series of mountain refugia and/or island refugia and long-distance dispersal, or without refugia and the current diversity being explained by long-distance dispersal alone. Murienne (2009) concluded that all three models could exhibit identical phylogenetic patterns. Molecular dating, such as with Angustonicus cockroaches (Murienne et al. 2005), could at least discern between the first and last models. Crisp and Cook (2009) recently proposed that cryptic mass extinction would result in a phylogenetic pattern indistinguishable from that of an evolutionary radiation. Interestingly, these scenarios might equally apply to New Zealand (Giribet & Boyer 2010). The geography and climate of New Caledonia have resulted in a wide range of vegetation types on Grand Terre. The major types are: maquis vegetation, a low, sclerophyllous, evergreen, heath-like formation largely restricted to ultrabasic (ultramafic) substrates which cover about one-third of Grand Terre (see, for example, Murienne et al. 2008); moist evergreen rainforest found principally in the east and the mountainous centre of the island where rainfall is highest; and remnant stands of sclerophyllous forest found between sea level and about 300 m (Lowry 1999). New Caledonia is renowned for its remarkably diverse flora, with more than 75% of the approximately 3,200 species of angiosperms and gymnosperms endemic (Myers 1988; Morat 1993; Lowry 1999). By contrast, the native vertebrate fauna of New Caledonia lacks amphibians and non-volant mammals (see Balgooy 1969). Of the 78 freshwater fishes, nine species are endemic to New Caledonia (Marquet et al. 1997), and there are few endemic genera and only one endemic family of birds (Diamond 1984). However, the New Caledonian terrestrial reptiles show a high degree of endemicity, with 61 of the 71 terrestrial reptiles strictly endemic (Bauer & Sadlier 2000). Some invertebrate groups have high levels of endemism (Chazeau 1993). Among these are land snails, with 99% endemism on New Caledonia (Tillier & Clarke 1983), cockroaches, especially the speciose genus Lauraesilpha (Blattidae: Tryonicinae) (Grandcolas 1997; Murienne et al. 2008), Trichoptera, with all but two of 132 described species endemic (Swedish Museum of Natural History 2003), and Agnotecous crickets (Orthoptera: Eneopterinae) (Robillard & Desutter-Grandcolas 2004; Robillard et al. 2007). Other groups, however, display lower endemicity. For example, only 38% of the more than 70 native species of butterflies and 300 species of moths are endemic (Chazeau 1993). Phytophagous diversity shows only modest endemism as well as species richness lower than might be expected for the land area (Holloway 1993). For the vast majority of insect groups there is, however, no precise information available on species numbers in New Caledonia. Chazeau (1993) estimated about 3,500 insect species for New Caledonia, including about 1,500 Coleoptera, 350 Hemiptera and 250 Hymenoptera. However, the hymenopteran fauna of New Caledonia has been relatively poorly studied. Recent papers by Aguiar and Jennings (2005, 2007), Bohart (1999), Baptiste and Kimsey (2000), Jennings and Austin (2002, 2005), Jennings et al. (2004, 2007), Jourdan & Mille (2006) and Smith (2008),

REVIEW OF HYMENOPTERAN FAUNA OF NEW CALEDONIA Zootaxa 3736 (1) © 2013 Magnolia Press · 3 and a survey by Monteith et al. (2006) of four conservation reserves at the southern tip of New Caledonia, prompted us to examine the hymenopteran fauna of New Caledonia and compare it with that of Australia and New Zealand, as well as other islands in the south-west Pacific.

Material and methods

The checklist is mainly based on published records which we sourced from the extensive literature and also various electronic data bases (including Noyes 2011 for Chalcidoidea, Yu et al. 2011 for Ichneumonoidea, and Hymenoptera Online Database (HOD)). New records were added from museum specimens in the following collections: MNHN = Muséum national d'Histoire naturelle, Paris; NC = Institu de recherche por le développement insect collection, New Caledonia, Nouméa; QM = Queensland Museum, Brisbane; SMNS = State Museum of Natural History, Stuttgart; WINC = Waite Insect and Nematode Collection, Adelaide.

The Hymenopteran Fauna of New Caledonia

Traditionally the order has been divided into two major groups with the status of suborder (“Symphtya” and ); however, recent phylogenetic analyses have demonstrated that only Apocrita is monophyletic and Symphyta is paraphyletic (Vilhelmsen et al. 2010 and references therein). Also, we have adopted the classification of Melo & Gonçalves (2005) whereby the Apoidea along with Apidae s.l. (i.e. all bees) is the sister group to the , the latter previously being accommodated in a broader, paraphyletic s.l. (see below for further details). Here we provide a summary of the families of Hymenoptera occurring in Australia, New Caledonia and New Zealand, along with brief comments on their biology and distribution. We also provide the first detailed checklist of the Hymenoptera of New Caledonia.

Tenthredinoidea

Argidae. The second largest symphytan family with about 800 species worldwide and with most diversity found in tropical regions, is poorly represented in Australia, with 12 species (Stevens et al. 2007). They are not known from either New Zealand or New Caledonia. . This is the dominant family in Australia (mostly Perga species), probably as a result of the diversity of myrtaceous host plants such as Angophora and Eucalyptus (Schmidt 2006). They show a Gondwanan distribution, with the majority of species occurring in Australia and South America but no species are known from Africa (Schmidt 2006). In New Caledonia and New Zealand, only the introduced leafblister sawfly (Phylacteophaga froggatti Riek) is known (Schmidt 2006). . This is the largest sawfly family with more than 5,000 species, most of which occur in the northern hemisphere. As with other sawfly families in Australasia (with the exception of the Pergidae), tenthredinids are poorly represented. There are only nine described species recorded for Australia, and four for New Zealand, with most having been introduced accidentally (Stevens et al. 2007; Table 1). Only two genera are endemic to Australia, Cheilophleps and Senoclidea (Naumann et al. 2002). The family is not represented in New Caledonia.

Siricoidea

Siricidae. The sirex woodwasp (Sirex noctilio F.) is a native of Europe, Asia, and northern Africa. It has been introduced accidentally into many forestry regions of the world, and now occurs in south-eastern Australia and New Zealand where it is a pest of introduced conifers, especially Pinus radiata (Jennings & Smith 2006). A recent incursion into New South Wales is Tremex fuscicornis (F.), a species of Palaearctic origin (Jennings & Smith 2006). To date, there have been no reports of siricid wasps in New Caledonia.

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Xiphydriidae. Xiphydriid woodwasps are worldwide in distribution, except sub-Saharan Africa, and well represented in Australasia (Australia, Fiji, New Caledonia, New Guinea and New Zealand) (Jennings et al 2007; Smith 2008), with at least 12 undescribed species in eastern Australia (Jennings, unpublished data). Smith (2008) recently revised the New Caledonian fauna. Lissoxiphyda, comprising seven species, is distributed in a small area from Papua New Guinea to Aru and Sulawesi in Indonesia, with one species in New Caledonia, Lissoxiphyda tripotini Smith (Smith 2008). Calexiphyda is known only from New Caledonia, and is represented by four described species (Smith 2008; Table 1), although there are a number of unassociated males from New Caledonia known in collections. Generally the hosts of the wood-boring xiphydriid larvae are unknown, but they can usually be found in weakened or dying limbs of various woody plants (see, for example, Smith 2008; Jennings et al. 2009). Nothing is known about the biology of any New Caledonian species.

Orussoidea

Orussidae. This small family of about 85 species worldwide has 12 described species from Australia (Riek 1955a; Schmidt & Vilhelmsen 2002; ABRS 2013). No native orussids are known from New Zealand, although Guiglia schauinslandi (Ashmead) has been introduced as a parasitoid of sirex woodwasps (Rawlings 1957). The family is not known from New Caledonia. Orussids are the only parasitoids among the and are considered to be the sister group to the Apocrita (Schmidt & Gibson 2001; Schmidt & Vilhelmsen 2002).

Stephanoidea

Stephanidae. Twenty-one species in three genera are recorded for Australia (Aguiar 2001), with most species belonging to the genus Parastephanellus. Schlettererius is represented only by S. cinctipes (Cresson), which has been introduced into Australia as a biocontrol agent for the sirex woodwasp Sirex noctilio. Stephanids are absent from New Zealand, but four species of Parastephanellus are known from New Caledonia (Aguiar & Jennings 2005, Table 1). No host records are available for any Australasian species but elsewhere stephanids are ectoparasitoids, mostly of wood-boring coleopteran larvae and, much more rarely, sawflies and solitary bees.

Megalyroidea

Megalyridae. Megalyrids are more diverse in the Australasian region than anywhere else in the world (Shaw 1990b), apparently because they have adapted to the dry conditions of Eucalyptus woodlands and Acacia scrub habitats. The Australian fauna comprises 22 described species of Megalyra, with the genus also represented by a few species in New Guinea and South-east Asia, and one species in New Caledonia (Shaw 1990a,b; Table 1). A single species of Carminator also occurs in New Britain and the Bismarck Archipelago. The family does not occur in New Zealand. Megalyrids are thought to be ectoparasitoids mostly of immature stages of wood-boring Coleoptera, although one Australian species is a parasitoid of a mud-nesting pemphredonine (Crabronidae) (Naumann 1991).

Trigonaloidea

Trigonalidae. This is a small family of about 120 species worldwide, and with about 10 species from Australia (Weinstein & Austin 1991; ABRS 2013; Table 1). Trigonalids have a cosmopolitan distribution, with the greatest species diversity in the tropics (Carmean & Kimsey 1998). Trigonalids have not been recorded from New Zealand and New Caledonia. Most species show an unusual biology. Female trigonalids lay several very small eggs into incisions in a leaf; each female laying up to several thousand eggs over a season. The eggs are eaten by

REVIEW OF HYMENOPTERAN FAUNA OF NEW CALEDONIA Zootaxa 3736 (1) © 2013 Magnolia Press · 33 phytophagous lepidopteran or sawfly larvae and hatch inside the larvae where they may then develop as primary parasitoids, but most develop as hyperparasitoids if the primary phytophagous larva is parasitised by an ichneumonid wasp or tachinid fly.

Ceraphronoidea

Ceraphronidae. The Australasian ceraphronid fauna is poorly known, although they are commonly collected from soil and litter (Stevens et al. 2007). Fifty-one species are known from Australia (Johnson et al. 2009) but the New Zealand fauna has not been described. Only a few ceraphronid specimens have been collected from New Caledonia, representing about five species (see Table 1). Ceraphronid biology is largely inferred from studies of northern hemisphere species, where they have been reared as primary endoparasitoids of Diptera, Lepidoptera, Thysanoptera and Neuroptera, and with some species hyperparasitoids of these host groups. . One species, Dendrocerus (Macrostigma) noumeae Dessart, is endemic to New Caledonia (Dessart 1967), compared with 24 species known from Australia, and a New Zealand fauna known only from introduced taxa (Stevens et al. 2007; Table 1). The Australasian fauna is poorly studied, and their biology largely unknown. Northern hemisphere species have been reared as primary parasitoids of scale , lacewings, or fly puparia, and some species are hyperparasitoids of aphids (Stevens et al. 2007).

Evanioidea

Aulacidae. The Australian fauna comprises 41 described species but current research is expected to more than treble this number (Jennings, unpublished data). The family is absent from New Zealand but is known from New Caledonia (Jennings et al. 2004; see Table 1) and New Guinea (Jennings & Austin 2006). Based on a number of characters, particularly those associated with fore wing venation and hind coxal morphology, the three species of Aulacus from New Caledonia appear unrelated to Australian members of the family (Jennings et al. 2004). It should be noted, however, that Aulacus is paraphyletic (Turrisi et al. 2009). Also, one undescribed Pristaulacus is known from New Caledonia (Jennings, unpublished data; see Table 1). Host records for Australasian species are scarce but records from other continents indicate that they are endoparasitoids of wood-boring wasp (Xiphydriidae) and beetle (Cerambycidae and Buprestidae) larvae (see, for example, Carlson 1979; Smith 2001; Jennings & Austin 2004). . Hatchet or ensign wasps develop as predators of cockroach eggs within oothecae (Deans 2005). In Australia there are 36 described species (Jennings 2007; Table 1) although more than 50 species await description. The Australian fauna is mostly endemic although a few species, including Acanthinevania mediana Schletterer and A. princeps (Westwood), are found in both Australia and New Guinea. The family is unknown from New Zealand. Apart from the cosmopolitan Evania appendigaster (L.), found in both Australia and New Caledonia (e.g. ANIC, QM, WINC), there are three endemic Szepligetella species known to occur on New Caledonia (Balhoff et al. 2013). . Female gasteruptiids oviposit in the nests of solitary bees and vespid wasps, where the larvae are predator-inquilines, eating the host egg or larvae and consuming the pollen store (Jennings & Austin 2002, 2004). One hundred and eighty-eight described species of gasteruptiid wasps occur in Australia: 113 species in the subfamily Gasteruptiinae (Gasteruption spp.) and 75 in the subfamily Hyptiogastrinae (two genera – Hyptiogaster and Pseudofoenus) (Jennings & Austin 1997, 2002; Jennings & Deans 2006; see Table 1). However, more than 100 undescribed Gasteruption and Pseudofoenus species are known from Australia (Jennings, unpublished data). The Gasteruptiinae is worldwide in distribution with Australian species comprising about 25% of the world’s fauna. Three undescribed species of Gasteruption are known from New Caledonia (Jennings, unpublished data), the same number as known from New Zealand, but we expect further New Caledonian species await discovery. Hyptiogastrine wasps are largely confined to Australia although there are two species from New Zealand, several species from Fiji, New Guinea and Vanuatu and two species in South America (Jennings & Austin 2002). Pseudofoenus caledonicus Jennings & Austin is endemic to New Caledonia and P. ritae (Cheesman) is found in

34 · Zootaxa 3736 (1) © 2013 Magnolia Press JENNINGS ET AL. both New Caledonia and Vanuatu (Jennings & Austin 2002, 2005; Table 1). Jennings & Austin (2002) found a close relationship between these Pseudofoenus species, as well as others from Fiji and New Guinea, although they formed a trichotomy with strictly Australian taxa. Pseudofoenus crassipes (Smith) and P. unguiculatus (Westwood) from New Zealand, however, were placed apically in a different clade but which also contained Australian species. Jennings and Austin (2002) suggested that the fauna from New Caledonia and New Zealand may have resulted from independent recent dispersal events from mainland Australia and/or, in the case of New Caledonia, from New Guinea.

Cynipoidea

Austrocynipidae. This rare endemic Australian family is known from a single species, Austrocynips mirabilis (Riek), reared from larvae of the family Oecophoridae (Lepidoptera) in the cones of Araucaria in Queensland (Riek 1971; Paretas-Martínez et al. 2013). Cynipidae. Cynipids are gall inducers or phytophagous inquilines in galls and are particularly diverse in the northern hemisphere (Richie 1993). The Cynipidae are poorly represented in Australia and New Zealand. The sole described Australian species, Phanacis hypochoeridis (Kieffer, 1887) is introduced (ABRS 2013; Table 1), although specimens of Andricus sp. are known (Paretas-Martinez et al. 2013). The family is unknown from New Caledonia. . This is the largest cynipoid family in the Australasian region with 47 described species in Australia and six in New Zealand (Buffington 2008; Paretas-Martínez & Pujade-Villar 2010; Paretas-Martínez et al. 2011, 2013; ABRS 2013; Table 1). There are also a significant number of undescribed genera and species in various collections (Paretas-Martinez et al. 2013). The most familiar members of the group are phytophagous, especially as gall-formers, though the majority of included species are parasitoids or hyperparasitoids. Figitids have not been formally recorded from New Caledonia but unidentified specimens exist in several collections (Matt Buffington, pers. comm.). . This family is represented naturally in Australasia only by a single very rare genus, Eileenella, from New Guinea (Liu & Nordlander 1994). The dominant genus Ibalia is known from 15 species distributed in the northern hemisphere (Liu & Nordlander 1994). Two of these species have been introduced into Australia and New Zealand as biological control agents of sirex woodwasps (Siricidae) (Stevens et al. 2007; Table 1). The family is not known from New Caledonia. . In Australasia this family is known from a single genus, Dallatorrella, with one species described from Australia (Liu 2001; Paretas-Martinez et al. 2013; Table 1) and one from New Guinea (Liu 2001). Limited host records from the northern hemisphere indicate that members of the family parasitise larvae of cerambycids and other wood-boring beetles.

Diaprioidea

Diapriidae. In Australia the largely endemic diapriid fauna consists of about 149 described species (Stevens et al. 2007), and in New Zealand, about 52 species (Table 1). Naumann (1982, 1988) revised the Gondwanan subfamily Ambositrinae for Australasia but the species of the two larger subfamilies, Belytinae and Diapriinae, are mostly undescribed. Diapriids parasitise a wide variety of fly larvae, particularly those occurring in low vegetation and leaf-litter. Species have also been recorded from termite nests. Elsewhere in the world, host records also include beetles, ants and , attacking the latter group as hyperparasitoids (Stevens et al. 2007). The New Caledonian fauna is represented by more than 20 undescribed species (HOD; Table 1). . Ismarinae, formerly a subfamily of , was raised to family status by Sharkey et al. (2011). Globally, there are only 33 species described which are distributed worldwide and of which at least some develop as hyperparasitoids of (Liu et al. 2011). Ismaridae have not yet been identified from New Caledonia, while two species occur in Australia (Naumann 1982). The family is not represented in New Zealand (Naumann 1988). Maamingidae. The family is endemic to New Zealand and is known from a single genus, Maaminga, comprising two species (Early et al. 2001; Table 1). The biology of the family is unknown.

REVIEW OF HYMENOPTERAN FAUNA OF NEW CALEDONIA Zootaxa 3736 (1) © 2013 Magnolia Press · 35 Monomachidae. Monomachids occur in Australia (3 species), New Guinea (2 species) and the Neotropical region from Mexico to Chile (Naumann 1985; Johnson & Musetti 2012; Table 1). However, they have not been recorded from either New Zealand or New Caledonia. Little is known of their biology but one Australian species is known to parasitise the eggs of a soldier fly, Boreoides (Diptera: Stratiomyidae) (Naumann 1985).

Proctotrupoidea

Austroniidae. This endemic Australian family is only known from three species of Austronia from wet forest habitats of south-eastern Australia and Tasmania (Riek 1955c; Table 1). Nothing is known of their biology. . This family comprises a single genus, Helorus, with 13 species, most in the Holarctic (Townes 1977b; Achterberg, 2006). Two species are known from the Australasian region: H. australiensis New, from Australia and H. niuginiae Naumann, from Papua New Guinea (Achterberg 2006; Table 1). They are not present in either New Zealand or New Caledonia. Helorids are solitary endoparasitoids of chrysopid lacewings (Neuroptera: Chrysopidae), ovipositing into the larvae but emerging from the host pupae. Peradeniidae. This endemic Australian family comprises one genus, Peradenia, with two described species (Naumann & Masner 1985; Table 1). They are only known from a few specimens collected in forested habitats in south-eastern Australia and Tasmania. Their biology is unknown. . Seven described species of proctotrupids are known from New Zealand and 23 from Australia (ABRS 2013; Table 1), although for Australia this is probably less than a quarter of the true species number. In Australasia, species are probably endoparasitoids of beetle larvae, although some northern hemisphere taxa are reported to parasitise fungus gnats (Diptera: Mycetophilidae) (Masner 1993). The New Caledonian proctotrupid fauna is undescribed (See Table 1).

Platygastroidea

Platygastridae. Sharkey (2007) combined the families Scelionidae and under the single name Platygastridae. About 666 species are described from Australia and 27 from New Zealand (ABRS 2013; Table 1) but there are probably around 4,000 platygastrid species in Australasia (Stevens et al. 2007). Within Telenominae, three described species of Trissolcus occur in New Caledonia: T. basalis (Wollaston), introduced to control green vegetable bug, Nezara viridula (L.), T. flaviscapus Dodd (also found in Australia), and T. personatus (also New Guinea, Fiji & Tahiti) (Johnson 1992). Telenomus lucellus (Nixon) has been introduced to New Caledonia (Sands & Liebregts 2005; Pintureau et al. 2010) and at least one additional species is known (QM; HOD). One species of Psix Kozlov & Lê has been collected from the Riviere Bleue Territorial Park (HOD; see Table 1). Several species of Teleas Latreille, Trimorus Foerster and Xenomerus Walker from the Teleasinae are known from New Caledonia (HOD; see Table 1), each represented by only a few specimens. Over 30 genera of are known to occur in New Caledonia (QM; HOD; see Table 1), many of which are known from fewer than 10 specimens, and in most instances probably represent undescribed species. Described taxa include the endemic spider egg parasitoids Ceratobaeus umari Iqbal & Austin and C. wattora Iqbal & Austin (Iqbal & Austin 2000) and the more widely distributed Baeus maryae Stevens (also Norfolk Island and throughout northeastern Queensland – Stevens & Austin 2007). In New Caledonia, three grasshopper egg parasitoids occur: Scelio bipartitus Kieffer (also Australia, New Guinea and Sarawak), S. javanicus Roepke (widespread – including Vietnam, Indonesia, New Guinea and Solomon Islands), and S. planithorax Dodd (also from Australia). Platygastrid wasps have a very diverse host range, however, they exhibit “little overlap in host groups used by different taxa within the family” (Galloway & Austin 1984). They parasitise the eggs of Coleoptera, Orthoptera and Hemiptera, larvae of Cecidomyiidae (Diptera) in their galls (see Austin et al. 2005), as well as spider eggs (Iqbal & Austin 2000).

36 · Zootaxa 3736 (1) © 2013 Magnolia Press JENNINGS ET AL. Chalcidoidea

This superfamily represents one of the largest species radiations in Hymenoptera. Most families are ubiquitous and most likely occur on New Caledonia in much higher species numbers than indicated by the published record. Large numbers of unidentified representatives of various chalcidoid families from throughout the south-west Pacific region, including Australia, exist in collections (e.g., ANIC, MNHN, QM, SMNS). . Agaonids are associated with figs (Ficus; Moraceae), acting as pollinators where the relationship is an obligate mutualism (Weiblen 2002). Two agaonid species are known from New Caledonia (Noyes 2011; Table 1): Dolichoris boschmai (Wiebes) and Pleistodontes greenwoodi (Grandi), with the latter having a wide distribution including Australia, American Samoa and Fiji. New Caledonia has 31 Ficus spp., most of which are endemic and found in rainforest (Jaffré et al. 2001), compared with 42 spp. of Ficus in Australia (ABRS 2011). Given the large number of agaonid species associated with figs in Australia (ABRS 2013; see Table 1) it is likely that there is a larger undiscovered agaonid fauna in New Caledonia. . Most aphelinids are primary ecto- or endoparasitoids of Hemiptera, especially Coccoidea, Aleyrodoidea and Aphidoidea (Viggiani 1984). However, others are egg parasitoids of these hemipteran families or of grasshoppers and crickets, and some are hyperparasitoids (Noyes 2011). There are many examples of aphelinids being used successfully in classical biological control (see Noyes 2011). About 286 species are known from Australia, 23 from New Zealand and 11 from New Caledonia (Table 1). Of the few recorded New Caledonian species, only five are apparently endemic, four species of Aphytis and Encarsia neocala Heraty & Polaszek (Noyes 2011; Table 1). . Most chalcidids are primary larval and pupal endoparasitoids of various Lepidoptera and Diptera, although Hymenoptera, Coleoptera and Neuroptera are also parasitised (see Noyes 2011). The family also contains hyperparasitoids and gregarious species. Keys to Australasian genera are included in Bouček (1988). About 178 species are described from Australia with just three species known from New Zealand (Noyes 2011; ABRS 2013; Table 1). In New Caledonia, three species of Brachymeria have been recorded and one of these, B. ucalegon (Walker), also occurs in Australia and New Zealand (Noyes 2011; Table 1). . The larvae of the majority of encyrtid wasps are primary parasitoids of Hemiptera, mostly scale insects (Coccoidea), although many other hosts are attacked. Life histories can be variable, for example, some attack eggs or larvae, and others are hyperparasitoids. They are found worldwide in virtually all habitats and are extremely important as biological control agents of many economic pests, especially scale insects (see summary in Noyes & Hayat 1994). Over 520 encyrtid species are known from Australia and over 70 from New Zealand (Noyes 2011; ABRS 2013; Table 1). However, apart from the endemic Habrolepis neocaledonensis Fabres, which has been found to parasitise the trilobite scale Pseudaonidia trilobitiformis (Green), the remaining 10 encyrtid species recorded from New Caledonia are introduced (Noyes 2011; Pintureau et al. 2010). . Eucharitid wasps are exclusively parasitoids and are particularly diverse in tropical and subtropical regions (Heraty 2002). Ninety-one species are recorded from Australia but none from New Zealand (Noyes 2011; ABRS 2013; Table 1). Orasema koghisiana Heraty is currently the only recorded representative of the family from New Caledonia and is also found in Vanuatu (Heraty 1994). The genus Orasema is circumtropical and occurs widely in the Americas, Africa, Madagascar, southern Asia, New Guinea and Australia. . Eulophids are the largest chalcidoid family and are speciose in both Australia (840 spp.) and New Zealand (54 spp.). Only seven introduced species have been formally recorded from New Caledonia (Noyes 2011; ABRS 2013; Table 1) but many more exist. Several genera are represented in collections with Renaniana and Gallowayia known previously only from Australia (Bouček 1988). Various eulophid species are known to attack leaf-mining dipterans but virtually all groups are used, including mites, gall forming nematodes, spider egg cases (hyperparasitoids on ichneumonid egg predators), a few scale insects (Coccidae, Diaspididae), Thysanoptera, and numerous coleopteran, lepidopteran, dipteran, and hymenopteran families (Askew 1971, Burks et al. 2011). Phytophagous eulophids are known from the subfamilies Opheliminae and Tetrastichinae and are especially diverse in Australia where they develop as gall-formers on Eucalyptus and various other plants (Bouček 1988, Burks et al. 2011). One species, Tetrastichus brontispae Ferriere, has been widely introduced in the Pacific Islands, including New Caledonia, for the control of the coconut hispine beetle, Brontispa longissima Gestro (Chrysomelidae) (Cochereau 1969).

REVIEW OF HYMENOPTERAN FAUNA OF NEW CALEDONIA Zootaxa 3736 (1) © 2013 Magnolia Press · 37 . Eupelmids are primary parasitoids or hyperparasitoids of a variety of insects, often associated with hosts concealed in plant tissue. Calosotinae and Neanastatinae develop primarily on wood-boring beetles, although some species attack gall-midges (Diptera: Cecidomyiidae). In contrast, Eupelminae display a much wider range of hosts, among them species of Lepidoptera, Coleoptera, Hemiptera (Coccoidea), Orthoptera, Blattodea, Mantodea, and Diptera (Gibson 1995; Stevens et al. 2007). Keys to the Australasian genera are included in Bouček (1988). In Australia, some 176 species are known, although there are just four known in New Zealand (Noyes 2011; Table 1). Only Tineobius columbi (Girault) has been formally recorded from New Caledonia; it also occurs in Queensland (Noyes 2011). . Many eurytomid species develop as solitary ecto- or endoparasitoids of phytophagous insects (e.g., gall formers) or they became secondarily phytophagous and develop as seed or stem borers (Noyes 2011). There are about 135 species of Eurytomidae recorded from Australia and 7 from New Zealand (ABRS 2013; Table 1). Only the introduced species Systole foeniculi Otten has been recorded from New Caledonia but many additional species exist in collections. . This is a rarely collected family with 11 described species of Leucospis in Australia (Bouček 1988; ABRS 2013; Table 1), which are parasitoids of solitary aculeate Hymenoptera, mostly bees. They do not occur in New Zealand. Leucospsis antiqua Walker occurs in both the North and Southern provinces of New Caledonia (Noyes 2011; HOD; MNHN). Mymaridae. Virtually all mymarids are egg parasitoids of numerous arthropod groups, with hemipterans, coleopterans, psocopterans, dipterans, and orthopterans being the most frequently utilised orders (Huber 1986; Noyes 2011). There are 276 described species in Australia (Lin et al. 2007; ABRS 2013) and 30 in New Zealand (Table 1). The only recorded New Caledonian species, Anagroidea dubia (Girault), is also known from Queensland and NSW (Triapitsyn & Berezovskiy 2002). . Most species are parasitoids of gall-inducing insects, including other chalcidoids, cynipids and Diptera. This family is represented in Australia by 12 described species in a single genus, Ormyrus (Bouček 1988; ABRS 2013; Table 1). There is a single species, Ormyrus hebridensis Narendran, known from New Caledonia and Vanuatu. Ormyridae has not been recorded from New Zealand. . Thirty-three described species are known from Australia, only one from New Zealand and they have not been recorded from New Caledonia (Noyes 2011; Table 1), although Perilampus specimens exist in collections (e.g., SMNS). Many perilampids are hyperparasitoids of Lepidoptera, developing through ichneumonid or braconid wasps, or tachinid flies. Other species are primary parasitoids of Hymenoptera, Coleoptera or Neuroptera (Bouček 1988). . Pteromalidae is morphologically extremely diverse and has been identified as a polyphyletic assemblage (e.g., Gibson et al. 1999; Krogmann & Vilhelmsen 2006, Heraty et al. 2013). The biology and host- associations of pteromalids are extremely varied and some groups are even secondarily phytophagous. Most species are idiobiont ectoparasitoids and common hosts are concealed larvae and pupae of Lepidoptera, cyclorrhaphous Diptera, Coleoptera and Hymenoptera, the latter often used by hyperparasitoids (see Bouček 1988). Fig associated pteromalids are known from four subfamilies (Epichrysomallinae, Otitesellinae, Sycoecinae, Sycoryctinae). Currently 565 pteromalid species are described from Australia and 30 from New Zealand (Noyes 2011; ABRS 2013; Table 1). Only seven species have been formally recorded from New Caledonia (Noyes 2011; Table 1), which is just a fraction of the actual species diversity (Krogmann, unpublished observations; QM). Three species are introduced (see Table 1), Grasseiana boschmai (Wiebes) and Pseudoceraphron burwelli Desjardins are endemic to New Caledonia, whilst two species, Chalcedectus cuprescens (Westwood) and Toxeumorpha nigra Girault, are also found in Australia (Noyes 2011). New Caledonian species of Neapterolelaps are known, a genus that occurs in Eastern Australia and New Guinea (Desjardins 2007; QM). . This family was originally thought to be endemic to New Zealand but has now also been recorded from Chile (Gibson & Huber 2000). In New Zealand it is known from a single genus Rotoita comprising at least two species, only one of which is described (Gibson pers. comm.). Their biology is unknown. . Signiphorids are often reared from the hemipteran families Diaspididae and Pseudococcidae and may be primary or hyperparasitoids (Noyes 2011). Aleyrodidae (Hemiptera) and dipteran puparia are also attacked, as are lepidopteran eggs (Noyes 2011). Sixteen species are known from Australia and three from New Zealand, with just the introduced Signiphora flavopalliata Ashmead known from New Caledonia (Noyes 2011; ABRS 2013; Table 1).

38 · Zootaxa 3736 (1) © 2013 Magnolia Press JENNINGS ET AL. . The Tanaostigmatidae are known from 11 described species in Australia and are apparently absent from New Zealand and New Caledonia (Noyes 2011; ABRS 2013; Table 1). Species for which the biology is known are mostly phytophagous and always associated with galls, either as gall-inducers or inquilines (Noyes 2011). . Only seven described species occur in Australia and the family has not been recorded from New Zealand or New Caledonia (Burwell 1998; ABRS 2103; Table 1). The biology of very few species is known but most appear to be parasitoids of leaf-mining larvae or insect eggs (Bouček 1988). . The Torymidae are known from 138 described species from Australia, five from New Zealand, and four from New Caledonia (Noyes 2011; ABRS 2013; Table 1). Two species of the torymine genus Podagrion, which are egg parasitoids of Mantodea, are apparently endemic to New Caledonia and one, Podagrion abbreviatum Cockerell, is known from both New Caledonia and Australia (Noyes 2011). The megastigmine Bootania neocaledonica (Milliron) is known from both New Caledonia and Fiji (Noyes 2011). Although many species are associated with plants, either parasitising hosts hidden within plant tissue or through secondarily developed phytophagy, there are a range of other biologies exhibited within the family (see Bouček 1988). . Trichogrammatids are primary (solitary or gregarious) egg parasitoids (Noyes 2011). They are often overlooked as they are among some of the smallest known insects (0.2–1 mm). One-hundred and forty-five species have been described from Australia and seven from New Zealand (ABRS 2013; Table 1). At least three species are known from New Caledonia, all introduced. They comprise two species of Trichogramma, T. achaeae Nagarkatti & Nagaraja (Pintureau et al. 2010) and T. chilonis Ishii (Sands & Liebregts 2005; Pintureau et al. 2010), and the Australian Trichogrammatoidea bactrae Nagaraja (Pintureau et al. 2010). In addition, undetermined Trichogramma and Trichogrammatoidea have been collected (Pintureau et al. 2010; Table 1).

Mymarommatoidea

Mymarommatidae. Within Hymenoptera, mymarommatids are among the most distinct wasp groups and are characterized by several uniquely derived morphological features, including a pleated membrane that allows the head to expand and contract in an accordion-like manner (Vilhelmsen & Krogmann 2006). Nothing is known about their host associations. The family is known from ten species in three genera (Gibson et al. 2007). Three mymarommatid species are known from Australia and one from New Zealand (Gibson et al. 2007; ABRS 2013; Table 1). One undescribed Mymaromma species has been found on New Caledonia (Gibson pers. comm.).

Ichneumonoidea

Braconidae. Braconids are known from 694 described species for Australia and 91 for New Zealand (Stevens et al. 2007; ABRS 2013; Table 1). This probably represents less than 10% of the true size of the Australian fauna. Although 27 braconids have been described from New Caledonia (see Table 1) the fauna is much larger as several undescribed species are known in collections. Interestingly, the genus Chremyloides, with three species otherwise confined to Australia, is also represented in New Caledonia by the endemic C. tobiasi Papp (Papp 2004). The majority of braconids are larval parasitoids but also oviposit into host eggs, pupae or adults. They develop as either primary endo- or ectoparasitoids on a wide variety of insect hosts (Wahl & Sharkey 1993). . This family probably comprises more than 1,500 species in Australia, however only 433 have been described (ABRS 2013; Table 1). The New Zealand fauna contains around 90 described species (Stevens et al. 2007; Table 1). The family is known from New Caledonia by 45 described species or subspecies (Gupta 1987; Table 1). Of these, Agrypon dozense Cheesman and Netelia (Netelia) shopar Cheesman, are endemic to the Loyalty Islands. Quite a few of the ichneumonid species from New Caledonia (see Table 1) are either cosmopolitan (e.g. Diplazon laetatorius (Fabricius)) or have a broader distribution in the southern tropics (e.g. Echthromorpha agrestoria (Swederus) (Broad, pers. comm.). Adults are commonly seen at flowers or searching for hosts around tree trunks, logs, vegetation, or in litter. Biologically, the group is very diverse. They can be ecto- or endoparasitoids, parasitising the larvae, prepupae or pupae of various endopterygote insects, and more rarely spiders and spider egg sacs. Some species are hyperparasitoids.

REVIEW OF HYMENOPTERAN FAUNA OF NEW CALEDONIA Zootaxa 3736 (1) © 2013 Magnolia Press · 39 Chrysidoidea

Bethylidae. There are some 2,200 species of bethylids worldwide and they are particularly abundant in the tropics (Gordh & Móczár 1990). Fifty-six species have been described from Australia (see Table 1; ABRS 2013) but in New Zealand there is just one described endemic species and several introduced species. The introduced Cephalonomia stephanoderis Betrem is recorded from New Caledonia (Jourdan & Mille 2006) and additional unidentified specimens exist in collections. Bethylids are ectoparasitoids of larvae, and occasionally pupae, of Coleoptera and Lepidoptera (Gordh & Móczár 1990), while a few species parasitise other hosts, such as Hymenoptera (Melo & Evans 1993). Chrysididae. Chrysidid wasps occur in all regions except Antarctica and Kimsey & Bohart (1990) suggest they may have evolved, or at least diversified, after the breakup of Gondwana. There are 67 described Australian species, with the fauna principally related to the Oriental fauna (Kimsey & Bohart 1990); the family is not known from New Zealand. The subfamily Chrysidinae, the cuckoo wasps, are mostly parasitoids or cleptoparasitoids in the nests of solitary Apoidea and (Stevens et al. 2007). The subfamilies Amiseginae and Lobosceliidinae are probably parasitoids of stick insect eggs (Phasmatodea) (Stevens et al. 2007). Only two described species of chrysidids are recorded from New Caledonia, the chrysidine Primeuchroeus caledonicus (Mocsáry) (Bohart 1988) and the amisegine Atoposega decorata Kimsey (Kimsey 1995) (Table 1). Additional undescribed species of New Caledonian Atoposega are known (Monteith et al. 2006). Dryinidae. Around 139 described species of dryinids are known from Australia and two from New Zealand (Olmi 1984; ABRS 2013; Table 1). They are ectoparasitoids of (Hemiptera: Fulgoroidea) nymphs and adults. Females of the vast majority of species use their chelate fore legs to catch and hold their hosts while they sting and temporarily paralyse them. Wingless ant-mimicking females can often be found where ants tend leafhoppers for honeydew. Several described species are known from New Caledonia (Olmi 1984) and the survey of Monteith et al. (2006) revealed one undescribed Dryinus sp. (Table 1). . This small family is known from fewer than 20 species worldwide, including two species described from Australia and one from New Zealand, all belonging to the nominal genus Embolemus (Olmi 1995; ABRS 2013; Table 1). One species, E. notogeicus Olmi, is known from New Caledonia (Olmi 1995; Monteith et al. 2006; Table 1). The biology of the Australasian species is unknown. . This family is known from Australasia by a single genus, Probethylus, with two species from Australia (Olmi 2004), and is apparently absent from New Zealand and New Caledonia. Sclerogibbids are known as solitary or gregarious ectoparasitoids of webspinners (Embioptera). Hosts of the Australian species are unknown. . Scolebythids parasitise the larvae of wood-boring beetles (Brothers 1981) and are known in the Australian region by two species. Of these, Ycaploca evansi Nagy, is also known from South Africa (Nagy 1975) as well as Norfolk Island and New Caledonia (Monteith et al. 2006). Naumann (1991) speculated that Y. evansi may have been introduced accidentally to the region in wooden sailing ships or imported timber.

Vespoidea

Formicidae. New Caledonia has a large, relatively well studied endemic ant fauna, together with a number of native and exotic species (see Taylor 1987, Ward & Wetterer 2006). A total of 49 ant genera are known from the region, compared with 103 from Australia (including one undescribed, CSIRO, 2012) and 24 from New Zealand. None of the New Caledonian genera are endemic and seven are unequivocally represented only by introduced species (Anoplolepis, Brachymyrmex, Cardiocondyla, Paratrechina, Solenopsis, Tapinoma and Wasmannia). Most New Caledonian genera are also known from Australia, with the exception of Brachymyrmex and Rogeria. Several genera are known from the region in the published literature or in collections but are not represented by described species, viz. Adelomyrmex (Taylor 1987), Crematogaster (Taylor 1987), Leptanilla (Monteith et al. 2006), and Metapone As with other isolated islands of the region, such as Fiji and Vanuatu, many ant genera typical of continental Australia are absent, and conversely, other continental genera have successfully radiated on New Caledonia, including Camponotus, Paraparatrechina, Pheidole and particularly Monomorium and Rhytidoponera (see Taylor

40 · Zootaxa 3736 (1) © 2013 Magnolia Press JENNINGS ET AL. 1987). Lordomyrma, a primarily Melanesian genus known from a few Australian species, has also successfully radiated on New Caledonia, although the majority of species are undescribed (Lucky & Sarnat 2008, Taylor 2009). Intriguingly, the only native extralimital species of the quintessentially Australian ant genus Myrmecia, M. apicalis Emery, is endemic to New Caledonia. The true size of the New Caledonian ant fauna is difficult to ascertain but there are probably numerous undescribed species. There are around 119 described ant species and sub-species known from New Caledonia. They are mostly endemic, or native and occurring elsewhere. Some 22 taxa are exotic, however, distinguishing between native and introduced taxa is problematic for those widespread in the Indo-Pacific region. From only four small botanical reserves at the southern tip of the main island, Monteith et al. (2006) recorded a diverse rainforest fauna of 96 ant species from 38 genera. Numerous pantropical tramp species have been introduced to New Caledonia, including several notorious invasives such as the African big-headed ant, Pheidole megacephala (F.), the yellow crazy ant, Anoplolepis gracilipes (Smith), and the little fire ant Wasmannia auropunctata (Roger). In particular, the latter species, locally referred to as the “fourmi electrique” (electric ant) is widespread, occurring in the lowlands of the main island, the Loyalty Islands, Île des Pins and the remote island of Walpole (Jourdan et al. 2001). Wasmannia auropunctata has become the ecologically dominant ant in a variety of habitats, including highly endangered dry sclerophyll forest, serpentine vegetation communities and pristine rainforest and it has been demonstrated to have negative impacts on native ants (Le Breton et al. 2003, 2005) and reptiles (Jourdan et al. 2001). The biogeographic relationships of the New Caledonian ant fauna have been little studied, although Ward (1985) found that the 17 endemic species of Rhytidoponera formed a monophyletic group relative to extant congeners in Australia and New Guinea and based on this, and limited vagility due to the absence of winged queens, he concluded that these species represent an archaic element in the ant fauna of New Caledonia dating back to the early Tertiary. This contrasts with Lucky’s analysis of Leptomyrmex, an essentially Australo-papuan genus with a monophyletic clade of three New Caledonian species (Lucky 2011). Combined nuclear and mitchondrial datasets suggest that the New Caledonian clade split from a central and eastern Australian clade no more than 10 million years ago. This implies that the New Caledonian fauna originated from a long distance dispersal event, despite the limited mobility of the wingless queens of Leptomyrmex (Lucky 2011). . Mutillids, or ‘velvet ants’, are most diverse in the tropics. About 192 species have been described from Australia but the fauna is probably at least twice this size (Stevens et al. 2007; ABRS 2013; Table 1). Only one introduced species from Australia occurs in New Zealand. Four species of Ancistrotilla Brothers are known from New Caledonia; A. aenigmatica Brothers, A. bleuensis Brothers, A. caledonica (André 1896) and A. nigra Brothers (Brothers 2012); Table 1). Mutillids are ectoparasitoids and can be found on bare ground, sandy areas, tree trunks and walls where they search for nests of sphecid, crabronid and vespid wasps, and bees (Apidae), whose larvae or pupae they parasitise. Pompilidae. Pompilids are represented in Australia by 268 described species with at least this number yet to be recognised, while 11 species are described from New Zealand (Stevens et al. 2007; Table 1). An ongoing generic revision of Australian Pompilidae will lead to generic transfers that affect some of the New Caledonian species (Krogmann et al., in prep.). Twelve described pompilid species occur in New Caledonia (Table 1), but Wahis et al. (2009) estimate that the fauna comprises 15 species. A large proportion of the New Caledonian pompilid fauna seems to be endemic, while some species are very widely distributed: e.g., Anoplius caerulescens (Dalla Torre) in Indonesia, Papua New Guinea, Fiji and New Caledonia, A. opulentus (Smith) in Queensland, Papua New Guinea, Indonesia, the Marianas, Carolines and Solomons, Fiji and New Caledonia, and Cyphononyx vitiensis Turner in Fiji, the Solomon Islands and New Caledonia. Monteith et al. (2006) collected 12 species of pompilid wasps in their New Caledonian survey, tentatively identifing three of them as Ctenostegus vachali (Banks), Monodontonyx parvulus (Banks) and Priocnemis araucariae Williams. Harris (1987) regarded the New Caledonian Priocnemis species as members of Sphictostethus Kohl, but Krogmann & Austin (2011) treated them again as members of Priocnemis Schiødte. One undescribed species of Priocnemis has been found on New Caledonia (Krogmann unpublished data). Pompilids are predatory parasitoids and are often referred to as ‘spider wasps’ because of their obligate use of spiders as food for their developing larva. Only one genus (Epipompilus) is known as a parasitoid of spider eggs. . In Australia, this small family is known from 17 described species that belong to the ant- like, wing-reduced genus Olixon (Krogmann et al. 2009). These species are distributed throughout the Australian

REVIEW OF HYMENOPTERAN FAUNA OF NEW CALEDONIA Zootaxa 3736 (1) © 2013 Magnolia Press · 41 continent, including arid and semi-arid and wet tropical habitats (Krogmann et al. 2009). This genus may also occur in New Guinea but the family is absent from New Zealand and New Caledonia. Although no host records are known from Australia, members of this family are all thought to be ectoparasitoids of crickets (Orthoptera: Gryllidae) (Townes 1977a; Krogmann et al. 2009). . From about 300 species of scoliid wasps worldwide, only about 20 have been described from Australia and there is only one introduced species in New Zealand (Stevens et al. 2007; ABRS 2013; Table 1). Only Campsomeris novocaledonica Turner has been described from New Caledonia. Female scoliids dig into soil or rotting wood where they paralyse scarabaeoid beetle larvae before depositing an egg (Finnamore & Hanson 1995). The solitary larva then develops ectoparasitically. . Flower wasps are parasitoids of beetle larvae in soil and are very diverse in Australia (Naumann 1991; Table 1), although absent from New Zealand. New Caledonia has 18 described tiphiids, all belonging to the genus Eirone (Baptiste & Kimsey 2000; Table 1), which also occurs in Australia. Monteith et al. (2006) collected two additional species of Eirone. Vespidae. Worldwide, there are about 5,000 species of Vespidae comprising solitary (Eumeninae, Euparagiinae, Masarinae, some Stenogastrinae) and eusocial wasps (, , some Stenogastrinae). About 350 described vespid species occur in Australia (ABRS 2013; Table 1). They are naturally absent from New Zealand where they are now represented by five introduced species (Stevens et al. 2007; Table 1). Nine species of Vespidae are known to occur in New Caledonia, four of which have been introduced (Jourdan & Mille 2006). Polistes stigma townsvillensis Giordani Soika, which is native to Australia, is also found in New Caledonia (Jourdan & Mille 2006). Polistes olivaceus (De Geer), which is native to the Oriental region (Harris 1979), is widespread and common in New Caledonia. The yellow jacket species Vespula germanica and V. vulgaris have been introduced to Australia and New Zealand but are not yet known from New Caledonia.

Apoidea

In the past bees and sphecoid wasps have been considered as separate superfamilies, the Apoidea s.str. and Sphecoidea (e.g. Naumann 1991). However, there is growing phylogenetic evidence that the bees, as a robust monophyletic group, are nested inside the sphecoid wasps (e.g. Melo & Gonçalves 2005). This has been confirmed by a recent detailed molecular study by Debevec et al. (2012), which places the bees (‘Anthophila’) within the Crabronidae, which is itself paraphyletic. Here we recognise two informal groups. The Spheciformes (=Sphecoidea) include the , Crabronidae, Heterogynaidae (non-Australasian) and Sphecidae s.str.; and Apiformes including all bees (= Anthophila sensu Debevec et al. 2012) and comprising the subfamilies Apinae, Andreninae, Colletinae, Halictinae, Megachilinae, and Stenotritinae, of which Andreninae and Melittinae do not occur in Australasia (Michener 2000).

Apiformes (Apidae)

In Australia there are more than 1,668 described species of bees, with many more undescribed (ABRS 2013; Ken Walker, pers. comm.; Table 1). This contrasts starkly with 37 species described from New Zealand (Table 1) and 17 from New Caledonia (Pauly & Munzinger 2003), although there are several undescribed species from New Caledonia (Ken Walker, pers. comm.). Pauly and Munziger (2003) compared data from New Caledonia with that of a number of islands in the Pacific and the Indian Oceans, and concluded that “The paucity of [the New Caledonian] fauna is paradoxical compared with the richness and endemicity of the island flora, but may be explained by the fact that the island was isolated before or soon after the emergence of the Apoidea about 130 million years ago”. This equates with the ‘museum model’ or ‘Noah’s Ark model’ of Murienne (2009). Most bees in Australasia are solitary and do not produce honey or wax. The Australian fauna is dominated by the subfamily Colletinae, to which most of the Australian genera and more than half of the described species belong (ABRS 2013). Colletines are solitary nesting bees although some species will nest in aggregations. Brood chambers are made in the ground, termite mounds, or within dead twigs, rotten logs or stumps (e.g., Stevens et al.

42 · Zootaxa 3736 (1) © 2013 Magnolia Press JENNINGS ET AL. 2007). The only Australian native eusocial honeybees are the few species of Austroplebia and Tetragonula which are stingless and restricted to the tropical or subtropical regions of the continent. The introduced European , Apis mellifera L., is widely distributed in the Australasian region including New Caledonia.

Spheciformes

Ampulicidae. The cockroach wasps are a primarily tropical family that are ectoparasitoids of adult and nymphal cockroaches (Riek 1955b). There are 14 species described from Australia (ABRS 2013; Table 1), but they are absent from New Zealand (Table 1). Just one species, Ampulex compressa (Fabr.), is known from New Caledonia. However, this species has a very broad distribution including Ethiopia, Kenya, Tanzania, Arabian Peninsula, Madagascar, Seychelles, Reunion, Mauritius, India, Sri Lanka, Bangladesh, Singapore, south China, Indonesia, Philippines, and Australia (Williams 1942, 1947; Table 1), and has been introduced to Hawaii for the biological control of cockroaches (Williams 1942). Crabronidae. Adult crabronids feed on nectar or honeydew, or at extrafloral nectaries, while females prey on and provision their nests with a range of insect groups as well as spiders. There are 688 species of Crabronidae described from Australia and 18 from New Zealand (ABRS 2013; Table 1). A large number of crabronid species occur in New Guinea and throughout the south-west Pacific. Twenty-six described crabronid species occur in New Caledonia (Table 1), with 17 of them belonging to the genus Arpactophilus. Monteith et al. (2006) also found an additional four species of Arpactophilus in their New Caledonian survey. Only 22 species of Arpactophilus are known from Australia, three from New Guinea and one from Indonesia (Misool Island) (Bohart 1999; Table 1). Within the genus Sericophorus, the vast majority of the approximately 100 described species are Australian endemics (four species in New Guinea and one in Indonesia – Ambon Island), but two species, S. rhinoceros Pulawski (Pulawski 1989) and S. pulawskii Ohl (Ohl 2009), are described from New Caledonia. Ohl (2008) recently described Clitemnestra noumeae Ohl from New Caledonia, which appears to have affinities with the four described species from New Guinea rather than the nine described Australian species. The majority of species (51) are, however, from the New World. Sphecidae s.str. The biology of sphecids is similar to that of crabronid wasps (see above). The New Caledonian sphecid fauna is species poor when compared with that of Australia but is similar to that of New Zealand (see Table 1). Undoubtedly the Australian fauna is much larger than indicated. Sphecids from New Caledonia include Pison ignavum Turner, which is widespread, being also known from Australia, Samoa, Fiji, Marquesas, and Society Islands (Williams 1945). A number of introduced sphecids are established on New Caledonia (Table 1). Three native species of Sceliphron (Callan 1980), and the introduced Madagascan species S. fuscum (Bohart & Menke 1976), are also known.

Conclusions

Sixty-five hymenopteran families are recorded from Australia but only the families Austrocynipidae, Austroniidae and Peradeniidae, and fewer than 10% of subfamilies, are endemic (Naumann, 1991). However, levels of species and generic endemicity are high. Some of the early lineages of Hymenoptera, such as , , , and Xiphydriidae (Xiphydriinae), which otherwise have a relict, worldwide distribution, are well represented in Australia (see Tables 1 & 2). By contrast, only 42 families are recorded from New Zealand, eight of which are represented only by introduced species (Tenthredinidae, Pergidae, Siricidae, Agaonidae, Scolebythidae, Mutillidae, Scoliidae and Vespidae) (see Tables 1 & 2). Only the family Maamingidae is confined to New Zealand (Early et al. 2001), and Rotoitidae, which was originally thought to be endemic to New Zealand, has also been recorded from Chile (Gibson & Huber 2000).

REVIEW OF HYMENOPTERAN FAUNA OF NEW CALEDONIA Zootaxa 3736 (1) © 2013 Magnolia Press · 43 TABLE 2. Comparative land area of Australia, New Caledonia and New Zealand and proportion compared with the Australian land area (in brackets), together with a summary of the numbers of described species of hymenopteran superfamilies (estimated total for New Caledonia in brackets) - data modified from Table 1, after data from ABRS (2013) and Stevens et al. (2007); * introduced species; # known from undetermined species. Australia New Zealand New Caledonia Land area (km2) 7,700,000 269,057 (3.5%) 19,103 (0.25%) Tenthredinoidea 161 5* 1* (1) Siricoidea 2* 1* 0 (0) Xiphydrioidea 7 3 5 (5) Orussoidea 12 1 0 (0) Stephanoidea 21 0 4 (4) Megalyroidea 22 0 1 (1) Trigonaloidea 10 0 0 (0) Ceraphronoidea 75 9 1 (6+) Evanioidea 265 5 9 (14) Cynipoidea 52 9 0 (4+) Diaprioidea 154 52 0 (20) Proctotrupoidea 29 9 0 (1) Platygastroidea 666 27 15 (53) Chalcidoidea 3,499 267 62 (117) Mymarommatoidea 3 1 0 (#1) Ichneumonoidea 1,127 174 72 (92) Chrysidoidea 268 6 10 (17) Vespoidea 2,868 61 164 (202+) Apoidea 2,404 66 65 (70) TOTAL 11,635 748 409 (609+)

New Caledonia is more family-rich than New Zealand with 47 families (see Tables 1 & 2), none of which are confined to New Caledonia, and some (e.g., Pergidae and Signiphoridae) are apparently represented only by introduced species. By way of additional comparison, the fauna of the tiny Lord Howe Island is represented in the Australian Museum collection by 31 families and more than 225 species of parasitoid and predatory wasps (Jennings & Austin, unpublished data). Lord Howe Island covers just 14.6 km2, is an eroded remnant of a 6.9 million-year-old shield volcano (McDougall et al. 1981), and is located approximately 770 km east of Australia and 1,350 km northwest of New Zealand on the submerged Lord Howe Rise. If we examine the New Caledonian Chalcidoidea, for example, many taxa have been introduced for biological control; of the remainder, fewer than half are native, and of these, nearly half are shared with either north-eastern Australia or nearby island groups such as Vanuatu and/or Fiji (see Table 1). This is certainly an artefact as the chalcidoid fauna, apart from few single species descriptions, has not been well collected or studied. Many more chalcidoid species are expected in New Caledonia if one considers the comparative proportions of the land areas based on that of Australia (see Table 2). The numbers of species of several other groups (e.g., Platygastroidea, Vespoidea and Apoidea) are also likely to be underestimates, particularly as a result of under-collecting. In Evanioidea the situation is varied: Gasteruptiidae and Aulacidae are probably undercollected as suitable habitats and a large species diversity of suitable hosts exist. Most cockroaches from New Caledonia are not Blattellidae and lack an exposed ootheca (e.g., Murienne et al. 2005), making them unsuitable hosts for evaniids. Excluding introduced or tramp species, there are a number of taxa shared between New Caledonia and Australia, particularly Queensland and New South Wales. Just one chalcidid, Brachymeria ucalegon (Walker), is shared between all three land masses (and it possibly originated in Australia), but none are shared between New

44 · Zootaxa 3736 (1) © 2013 Magnolia Press JENNINGS ET AL. Zealand and New Caledonia. This is perhaps not surprising given the weather over New Zealand is dominated by the continual eastward passage of anticyclones at about weekly intervals (Robertson 1966), giving virtually no chance for Hymenoptera (or for that matter other insects) to be blown the 1,500 km north-north-west to New Caledonia. More commonly, there are taxa (e.g., Gasteruptiidae, Eucharitidae, Ormyridae, Ichneumonidae, Pompilidae) shared among neighbouring island groups such as Fiji, New Guinea and Vanuatu, and their distributions suggest these groups can readily disperse from island to island. Whilst several different scenarios might explain the origin of the New Caledonian hymenopteran fauna, until various phylogenies are examined using molecular dating, there is little evidence for or against the ‘museum model’, diversification after submersion(s), either with or without refugia and long-distance dispersal (Murienne 2009), or cryptic mass extinction (Crisp & Cook 2009).

Acknowledgements

The authors thank Gavin Broad (Natural History Museum, London) for information on Ichneumonoidea and for postive suggestions for improving the paper; Denis Brothers (University of KwaZulu-Natal, South Africa) for information on Mutillidae; Gary Gibson (Canadian National Collection of Insects) for information on and Rotoitidae; Massimo Olmi (University of Tuscia, Viterbo, Italy) for information on Dryinidae; Ken Walker (Museum Victoria, Melbourne) for information on Apidae; and Gavin Broad (Natural History Museum, London), Susan Wright (Queensland Museum, Brisbane), and Claire Villemant (Muséum national d'Histoire naturelle, Paris) for access to their collections. We especially thank Hervé Jourdan, Institut de Recherche pour le Dévelopment, Noumea for arranging collecting permits and for access to the Institut’s collection in Noumea. This research received support from the SYNTHESYS Project http://www.synthesys.info/ which is financed by European Community Research Infrastructure Action under the FP7 "Capacities" Program (FR-TAF- 341 and GB-TAF-1819).

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