Amy Carmichael Thesis (PDF 17MB)

Taxonomy and Diagnostics of Fruit Fly infesting Opiine Braconids in Australia and the South Pacific

Amy Elizabeth Carmichael

BSc., BFA (Hons)

Submitted in fulfilment of the requirements for the Master of Applied Science

School of Natural Resource Sciences Queensland University of Technology September, 2009

Frontis piece: An insect cabinet drawer of opine braconids

i ii Abstract Opiine wasps (Hymenoptera: Braconidae: Opiinae) are parasitoids of dacine fruit flies (Diptera: Tephritidae: Dacinae), the primary horticultural pests of Australia and the South Pacific. Effective use of opiines for biological control of fruit flies is limited by poor taxonomy and identification difficulties. To overcome these problems, this thesis had two aims: (i) to carry out traditional taxonomic research on the fruit fly infesting opine braconids of Australia and the South Pacific; and (ii) to transfer the results of the taxonomic research into user friendly diagnostic tools. Curated wasp material was borrowed from all major Australian museum collections holding specimens. This was supplemented by a large body of material gathered as part of a major fruit fly project in Papua New Guinea: nearly 4000 specimens were examined and identified. Each wasp species was illustrated using traditional scientific drawings, full colour photomicroscopy and scanning electron microscopy. An electronic identification key was developed using Lucid software and diagnostic images were loaded on the web-based Pest and Diseases Image Library (PaDIL). A taxonomic synopsis and distribution and host records for each of the 15 species of dacine-parasitising opiine braconids found in the South Pacific is presented. Biosteres illusorius Fischer (1971) was formally transferred to the genus Fopius and a new species, Fopius ferrari Carmichael and Wharton (2005), was described. Other species dealt with were Diachasmimorpha hageni (Fullaway, 1952), D. kraussii (Fullaway, 1951), D. longicaudata (Ashmead, 1905), D. tryoni (Cameron, 1911), Fopius arisanus (Sonan, 1932), F. deeralensis (Fullaway, 1950), F. schlingeri Wharton (1999), Opius froggatti Fullaway (195), Psyttalia fijiensis (Fullaway, 1936), P. muesebecki (Fischer, 1963), P. novaguineensis (Szépliget, 1900i) and Utetes perkinsi (Fullaway, 1950). This taxonomic component of the thesis has been formally published in the scientific literature. An interactive diagnostics package (“OpiineID”) was developed, the centre of which is a Lucid based multi-access key. Because the diagnostics package is computer based, without the space limitations of the journal publication, there is no pictorial limit in OpiineID and so it is comprehensively illustrated with SEM photographs, full colour photographs, line drawings and fully rendered illustrations. The identification key is only one small component of OpiineID and the key is supported by fact sheets with morphological descriptions, host associations, geographical information and images. Each species contained within the OpiineID package has also been uploaded onto the PaDIL website (www.padil.gov.au). Because the identification of fruit fly parasitoids is largely of concern to fruit fly workers, rather than braconid specialists, this thesis deals directly with an area of growing importance to many areas of pure and applied

iii biology; the nexus between taxonomy and diagnostics. The Discussion chapter focuses on this area, particularly the opportunities offered by new communication and information tools as new ways delivering the outputs of taxonomic science.

Keywords Opiinae, Braconidae, Parasitoids, Taxonomy, Identification, Diagnostics.

iv Table of Contents

Abstract ...... iii Keywords...... iv Table of Contents ...... v List of Figures ...... vii List of Tables ...... ix Statement of original authorship...... xi Acknowledgments ...... xiii Chapter 1 Introduction ...... 1 General Introduction ...... 1 Introduction to the Family Braconidae, Sub-Family Opiinae...... 2 Taxonomic History of the Opiinae ...... 3 Opiinae as biological control agents of Fruit flies ...... 7 Background to current project ...... 8 Aims and Objectives of Thesis ...... 10 Chapter 2 Materials and Methods...... 12 Geographic scope of project...... 12 Material examined ...... 12 Identification Methodology...... 13 Terminology...... 15 Illustration techniques...... 16 Scanning Electron Microscopy ...... 19 Lucid 3 Key Builder...... 19 Chapter 3 Taxonomy ...... 21 Introduction...... 21 Key to the Opiinae parasitising Tephritidae of the South Pacific region...... 25 Diachasmimorpha hageni (Fullaway, 1952) ...... 29 Diachasmimorpha kraussii (Fullaway, 1951)...... 32 Diachasmimorpha longicaudata (Ashmead, 1905)...... 40 Diachasmimorpha tryoni (Cameron, 1911)...... 46 Fopius arisanus (Sonan, 1932) ...... 50 Fopius deeralensis (Fullaway, 1950)...... 62 Fopius ferrari Carmichael & Wharton ...... 66 Fopius illusorius (Fischer, 1971)...... 69 Fopius schlingeri Wharton, 1999 ...... 76

v Fopius cf. vandenboschi...... 80 Opius froggatti (Fullaway, 1950)...... 83 Psyttalia fijiensis (Fullaway, 1936)...... 86 Psyttalia muesebecki (Fischer, 1963)...... 90 Psyttalia novaguineensis (Szépligeti, 1900)...... 91 Utetes cf. albimanus...... 97 Utetes perkinsi (Fullaway, 1950) ...... 99 Chapter 4 Identification and Diagnostics ...... 102 Introduction...... 102 Materials and Methods ...... 107 Results...... 112 Discussion ...... 124 Chapter 5 Taxonomy, Identification and Diagnostics ...... 127 Introduction & thesis summary ...... 127 Taxonomy, identification and diagnostics...... 127 Changes in Taxonomy...... 128 Progression of Identification ...... 130 Diagnostics in a digital age...... 131 Communication and Collaboration ...... 132 Chapter 6 Appendices...... 134 Appendix 1 ...... 134 Appendix 2 ...... 160 Appendix 3 ...... 190 Chapter 7 Bibliography...... 212 CD Envelope ...... 220

vi List of Figures

Figure 2.1. Geographic region covered by study...... 12 Figure 2.2. Opiine wing venation and terminology (following Sharkey & Wharton 1997)...... 15 Figure 2.3. Opiine body morphology and terminology (following Sharkey & Wharton 1997)...... 15 Figure 3.1. Diachasmimorpha hageni. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view...... 31 Figure 3.2. Diachasmimorpha kraussii. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view...... 39 Figure 3.3. Diachasmimorpha longicaudata. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view...... 45 Figure 3.4. Diachasmimorpha tryoni. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view...... 49 Figure 3.5. Fopius arisanus. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view...... 61 Figure 3.6. Fopius deeralensis. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view...... 65 Figure 3.7. Fopius ferrari. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view...... 68 Figure 3.8. Fopius illusorius. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view...... 75 Figure 3.9. Fopius schlingeri. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view...... 79

vii Figure 3.10. Fopius cf. vandenboschi. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view...... 82 Figure 3.11. Opius froggatti. a, head, anterior view; b, mesonotum, dorsal view; c, wing...... 85 Figure 3.12. Psyttalia novaguineensis. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view...... 96 Figure 3.13. Utetes cf. albimanus. Habitus...... 98 Figure 3.14. Utetes perkinsi. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view...... 101 Figure 3.15. Hind femur and tibia showing tibial carina (arrow) typical of Utetes..101 Figure 4.1. An example of a character page with OpiineID...... 120 Figure 4.2. Screen capture of the species menu of the Braconidae within the PaDIL website...... 123 Figure 4.3. A comparative image table illustrating three morphological characters of three Fopius species; Fopius arisanus Fopius deeralensis and Fopius schlingeri. 124

viii List of Tables

Table 1.1 An overview of the changes for each genus within the subfamily Opiinae. This information has been modified from Wharton (2008) ...... 6 Table 2.1. Source and amount of Opiinae material examined in this thesis. [ANIC, Australian National Insect Collection; ASCU, Agricultural Scientific Collections Unit; BPBM, Bernice P. Bishop Museum; GU, Griffith University; QDPIF, Queensland Department of Primary Industries and Fisheries; SPC, Secretariat of the Pacific Community; TAMU, Texas A&M University]...... 14 Table 3.1. Host Records. For each host association record, the source of the record is given in the table using the following code: 1 (label data); 2 Wharton and Gilstrap (1983); 3 Waterhouse (1993); 4 Quimio & Walter (2001); 5 Wharton (1999); 6 Fullaway (1952); 7 Fischer (1963a)...... 27 Table 4.1. Character state matrix required for creating the dichotomous fruit identification key above. “1” indicates that the character state was scored positively for the particular fruit type. Blank cells are not negative scores, but illustrate where a character state did not need to be scored for the key to be developed...... 105 Table 4.2. Character state matrix required for creating a multi-access fruit identification key. The presence or absence of a character state is indicated by the following code: 0 absence; 1 presence; 2 possibly misinterpreted...... 105 Table 4.3. Character state matrix required for creating the multi-entry or matrix key to opiine fruit fly parasitoids of the South Pacific as presented in OpiineID. The presence or absence of a character state is indicated by the following code: 0 absent; 1 present; 2 rarely present; 3 uncertain...... 109 Table 4.4. An example species page within OpiineID: Fopius ferrari Carmichael and Wharton...... 114 Table 4.5. Static version of morphological tutorial of the face...... 121

ix x Statement of original authorship

The work contained in this thesis has not been previously submitted to meet the requirements for an award at this or any other higher education institution. To the best of my knowledge and belief, the thesis contains no material previously published or written by another person except where due reference is made.

Signed: ………………………………………. Date:

xi xii Acknowledgments

I would like to thank the following people for their contributions to this thesis:

My supervisors Dr. Tony Clarke and Dr. Andrew Baker for their help, guidance and patience throughout this study. Tony has been exceptionally supportive in his two, often conflicting, roles as both my employer and my supervisor over the past few years.

Prof. Bob Wharton for his help with confirming my identifications, and his support and assistance when it came to describing my first species.

My parents, friends and family who have endlessly supported and encouraged me throughout this research.

Lastly, my dear husband James, who has been there for me every step of the way.

xiii

Chapter 1

Introduction

General Introduction Tephritid fruit flies (Diptera: Tephritidae) are the focus of extensive taxonomic and diagnostic research, with description, recategorisation and genetic analysis regularly changing the taxonomy and systematics of the Tephritidae (Norrbom et al. 1998). The biosecurity importance and global distribution of these pest flies helps to maintain the momentum of continuing taxonomic revision and the ongoing search for control options. In contrast, significantly less taxonomic effort has been put into the natural enemies of fruit flies, particularly in Australasia, yet this is an area of primary importance for the development of sustainable management strategies for fruit flies.

Interest in the use of natural enemies for fruit fly control dates back to the early 1900’s (Silvestri 1913; Silvestri 1914; Lever 1938). Opiine fruit fly parasitoids (Hymenoptera: Braconidae: Opiinae) have been introduced and released as classical biological control agents of fruit flies in many regions, although often with poor success (Waterhouse 1993). One likely reason for the lack of success in some areas is the shortage of both taxonomic and ecological understanding of the parasitoids themselves (Wharton 1997a). Effective biocontrol is hindered by the inability to accurately identify opiines, with inadequate species descriptions, continuously changing taxonomy and poor identification tools. Despite their importance in applied entomology the identification of opiine parasitoids remains difficult, with reliance on complex characters, and identification keys being taxonomically dated and restricted in the taxa covered. Specialist morphological terminology and composite couplets make identification remarkably complicated even when suitable keys do exist. Problems of parasitoid identification are additionally exacerbated by the fact that regional faunas are now mixed as a result of biological control liberations. The difficulties of identification of fruit fly parasitoids will continue to hamper efforts to use the agents effectively in fruit fly biological control.

This thesis addresses the issues of poor opiine taxonomy and identification tools for Australia and the South Pacific by carrying out a taxonomic revision of the regional fruit fly attacking opiine fauna and the development of different identification tools. As an introduction to the thesis, this chapter will look broadly at the Braconidae and then focus on the generic classification and reclassification within the Opiinae.

1 Following this taxonomic section, I introduce briefly fruit flies as pests and the biological control potential of opiines. I finish the Introduction with an over-view of the thesis structure and questions.

Introduction to the Family Braconidae, Sub-Family Opiinae The Hymenoptera is one of the largest insect orders (Naumann 1991). It contains a diverse assortment of phytophagous insects, social insects, solitary bees, wasps and parasitoids. Among the parasitoids are the two largest families of Hymenoptera, the Ichneumonidae and Braconidae, which together with the small (and non-Australian) Apozygidae constitute the super-family Ichneumonoidea (Naumann 1991). Superficial recognition of the families is usually by the generally smaller size of braconids as compared with their ichneumonid relatives; however, examination of the forewing venation will more definitively identify each family (Matthews 1974). The braconids possess only one recurrent vein, whereas ichneumonids have two, resulting in a more complete venation. Tendency towards reduction in wing veins is a characteristic of braconids, as is the relative stability of venation for ichneumonids (Tobias 1967; Capek 1970; Matthews 1974). The Ichneumonidae contains over 60 000 species most of which are parasitoids of larvae and pupae of Coleoptera, Hymenoptera and Lepidoptera (Naumann 1991). Approximately 12 000 species have been described from the Braconidae, however, it is estimated that between 40 000 and 50 000 species occur within the family (Sharkey & Wahl 1992). The most common hosts of braconid parasitoids are the larvae of Lepidoptera, Coleoptera and Diptera. Adult braconids oviposit almost exclusively in, on or near other insects, with the immature stages completing their development at the host’s expense. Only a few species are known to be phytophagous (Macedo & Monteiro 1989; Wharton 1993; Infante et al. 1995).

The Braconidae contains both ectoparasitoids (i.e. some immature development stage of the parasitoid is external to the host) and endoparasitoids (i.e. all immature development stages are internal to the host) (Wharton 1993). Although both are well represented amongst the described species, there are more endoparasitoids than ectoparasitoids (Matthews 1974). Braconid ectoparasitoids are represented by members of the subfamilies Braconinae, Hormiinae and Doryctinae (Matthews 1974). The endoparasitoids of the Braconidae belong to the families Alysiinae, Microgastrinae and Opiinae (Wharton 1997b). The Microgastrinae are the most important single group of parasitoids of Lepidoptera in the world (Whitfield 1997). The Alysiinae and Opiinae are closely related, being the two braconid subfamilies

2 that are exclusively endoparasitic on cyclorrhaphous Diptera (Matthews 1974). The Alysiinae are parasitoids of, and have been used in the biological control of, calliphorids, muscids and leaf-mining agromyzids (Wharton 1980). Biologically the subfamilies Alysiinae and Opiinae are very similar. Morphologically, as adults, they can be separated easily on mandibular structure. Alysiines have outwardly directed, non-overlapping (=exodont) mandibles, while opiines, with the exception of the very rarely encountered Exodontiella, do not (Wharton 1988).

Species of the subfamily Opiinae have been recorded attacking leaf-mining Agromyzidae and fruit infesting Tephritidae (Naumann 1991). Confirmed host records are available for about 275 (~29%) opiine species (Fischer 1971a, 1972, 1977, 1987). With the exception of Opius, there is some degree of host specificity at the generic level for most of the opiines. Those species that attack fruit infesting tephritids, even though they have a reasonably broad host range, have not been recorded from tephritids infesting flower heads. Over 100 species of Braconidae have been reared from fruit-infesting tephritids throughout the world and the vast majority of these are from the subfamily Opiinae (Wharton 1993).

Taxonomic History of the Opiinae The Opiinae is one of the larger subfamilies of the Braconidae, containing over 1300 described species. In a long series of papers beginning in 1956, Fischer revised the Opiinae on a global basis, significantly expanding our knowledge of the diversity of this important group of dipterous parasitoids. During the first half of the twentieth century, much of the research on the Opiinae was based on the conservative classification of Gahan (1915), in which most species were placed, ungrouped, in the genus Opius Wesmael. The most recent comprehensive classification of the Opiinae is the three volume monograph published by Fischer (1972; 1977; 1987). This work established a basis for more intense examination of the Opiinae, resulting in several subsequent modifications and additions to the classification. Gahan’s (1915) classification could not adequately accommodate the large number of species added by Fischer and so was modified. The resulting classification (Fischer 1972) still had the vast majority of species in Opius, but segregated into 36 subgenera. Other subgenera were subsequently added (Fischer 1984, 1986).

The generic concepts of Fischer have been critically examined by Wharton & Marsh (1978), Wharton (1987) and Wharton & Van Achterberg (2000), resulting in several changes. Most notable of these changes was the elevation of Psyttalia Walker and

3 Utetes Förster to generic rank by Wharton (1987). Many Opius species were not formally reassigned following these changes until revision by Wharton (1997a). Lack of assignment was, in part, because of the lack of agreement regarding the limits of the definition of two other key opiine genera, Diachasmimorpha Viereck and Fopius Wharton (Wharton 1987, 1988; Van Achterberg & Maeto 1990). To facilitate the placement of parasitoids, the major genera were described in detail by Wharton (1997a), which has subsequently provided some taxonomic stability to genera that had previously been split and clumped on numerous occasions for various reasons. An overview of former and currently recognised genera within the Opiinae is detailed in Table 1.1 [modified from Wharton (2008)]. The taxonomic history of four currently valid genera within the subfamily Opiinae, are described in further detail below: these genera are detailed as they are directly relevant to the research conducted within this thesis.

Diachasmimorpha Viereck (1913) In publications prior to Wharton (1988), the species of Diachasmimorpha were placed in either the genera Opius or Biosteres. Wharton (1997a) defined the genus Diachasmimorpha on the basis of the forewing m-cu arising from a shortened second submarginal cell, well developed hindwing m-cu and 2M, loss of hind wind RS (at least basally), absence of oblique carina ventrolaterally on propleuron, unsculptured notauli, and the long ovipositor with strongly attenuate hypopygium.

Fopius Wharton (1987) Prior to Wharton (1987), most of the species now placed in the genus Fopius were placed either in Opius or Biosteres, and in some cases in Diachasma or Parasteres. The genus Fopius was defined by Wharton (1997a) on the basis of more heavily sculptured body (especially frons, notauli and propleuron) and the largely unmodified mandibles and clypeus. Additionally, most species of Fopius have the hind wing m-cu slightly recurved near the wing margin.

Psyttalia Walker (1860) Prior to 1987, most of the species of Psyttalia were placed in Opius. Psyttalia was treated as a subgenus of Opius by Fischer in 1972 and then elevated to generic rank by Wharton in 1987. Wharton (1997a) confirmed the separation of the genus Psyttalia from Opius by the combination of a relatively short second metasomal tergum, short clypeus, strongly attenuate hypopygium, and absence of both hind

4 wing m-cu and at least the basal half of hind wing RS. The large second submarginal cell and the absence of a mesonotal pit are other useful distinguishing characters.

5 Table 1.1 An overview of the changes for each genus within the subfamily Opiinae. This information has been modified from Wharton (2008) Genus Current Status Comments Austroopius Szépligeti, Synonym of Psyttalia (Wharton 1987). Tephritid parasitoids formerly included here have usually been referred to as Opius 1900 in the biological control literature. Biosteres Förster, 1862 Valid genus Tephritid parasitoids formerly included here have now been transferred mostly to either Diachasmimorpha or Fopius. See Wharton (1988). Bracanastrepha Brèthes Synonym of Utetes (Wharton 1987, 1988). Tephritid parasitoids formerly included here were referred to as Opius in most of the 1924 biological control literature prior to 1979. Desmiostoma Förster, Synonym of Opius (Wharton 1983, 1988) 1862 Diachasma Förster, 1862 Valid genus (but see Tobias 1977). Diachasmimorpha Viereck, Valid genus which contains some of the most Prior to 1987, most of these species were placed in either Biosteres or Opius, and 1913 widely used parasitoids in tephritid biocontrol. more rarely Parasteres (Wharton 1987). Doryctobracon Enderlein Valid genus. Prior to 1978, these species were placed either in Opius or Parachasma (Fischer 1920 1977, Wharton and Marsh 1978). Fopius Wharton 1987. Valid genus, originally described as a Includes some of the most significant parasitoids used in tephritid biocontrol. Most subgenus of Rhynchosteres (Wharton 1987, of these species have been placed in Biosteres in recent literature, and in Opius van Achterberg and Maetô 1990). prior to about 1970 Hedylus Marshall 1891 A junior homonym of Hedylus Förster, 1862, One species (giffardii) was originally described in Hedylus Marshall, 1891. an ichneumonid. Opius Wesmael 1835 Valid genus. Most of the tephritid parasitoids formerly included in Opius have been transferred elsewhere. Parachasma Fischer, 1967 Junior subjective synonym of Doryctobracon (Fischer 1972, Wharton and Marsh 1978) Parasteres Fischer, 1967 Junior subjective synonym of Diachasmimorpha (Wharton and Marsh 1978, Wharton 1987). Psyttalia Walker, 1860 Valid genus which contains some of the most Most of these species have been referred to as Opius in the biological control widely used parasitoids in tephritid biocontrol literature. (Wharton 1987, 1988). Utetes Förster, 1862 Valid genus. Tephritid parasitoids have been referred to as Opius (Wharton 1988). Treated by Fischer as a subgenus of Opius.

6 Utetes Förster (1862) The species of Utetes were generally included in the genus Opius prior to Wharton (1988), although Tobias (1977) and Tobias and Jakimavicius (1986) included Utetes as a subgenus of Xynobius. Wharton (1997a) characterised the genus Utetes on the carina located basomedially on the hind tibia, a character previously overlooked in the segregation of this genus from Opius by other authors.

Within each of the genera described above, there have been many species specific changes. Each of which are described in detail in Chapter 3.

Opiinae as biological control agents of Fruit flies Fruit flies (Diptera: Tephritidae) are among the most economically important pests of edible fruits worldwide (White & Elson-Harris 1992), representing a threat to the successful establishment of horticultural industries and trade. Different fruit fly genera occur in every continent except Antartica and hundreds of millions of dollars are spent annually on their control and eradication. As fruit flies can be transported readily as eggs or maggots in fruit and fruiting vegetables, the existence of fruit fly for which the crop in question is a host in a cropping district means that commodity access to markets may be restricted due to quarantine regulations of the importing area. Of the more than 4 400 tephritid species known worldwide, about 70 species are considered important agricultural pests, while many others are minor or potential pests (White & Elson-Harris 1992). However, it should be recognised that fruit flies are part of natural ecosystems, hence in an area of origin are a normal part of that ecosystem.

The family Tephritidae has a number of subfamilies, with the genus Bactrocera being placed in the Dacinae. The genus Bactrocera is one of the larger tephritid genera with nearly 500 described species (Norrbom 2004) and is of worldwide notoriety for its destructive impact on agriculture, particularly in South East Asia, Australia and Pacific. Development of biological control methods for dacine fruit flies has been a focus of research for nearly a century, with renewed emphasis in recent years as chemical control methods becoming less favoured or banned altogether. The legacy of pesticides such as DDT, lindane and similar compounds used against fruit flies (e.g. Steiner 1952; Keiser 1968) is still with us many years after their use has ceased, and is often found in ecosystems far removed from those where they were applied (Duan & Messing 1997). Less disruptive methods of fruit fly control, such as post-harvest treatment, orchard sanitation, male annihilation and

7 sterile insect technique (SIT), may be applied in the context of integrated fruit fly pest management programs (Duan & Messing 1997). Interest in the biological control of fruit flies, utilising parasitic (parasitoid) Hymenoptera, dates back to the early 1900’s (Silvestri 1913; Silvestri 1914; Lever 1938) and has been a focus of fruit fly research ever since (Knipling 1992; Waterhouse 1993).

Opiines are koinobiont endoparasites of the egg or larval stages of Diptera and have been introduced and released as classical biological control agents of fruit flies in many regions (Wharton & Gilstrap 1983; Waterhouse 1993; Ovruski et al. 2000). More recently, opiines have been used in manipulative1 and inundative2 release programmes against fruit flies (Messing et al. 1993; Purcell et al. 1998; Ramadan 2004), with foreign exploration continuing (Wharton et al. 2000).

Generally, tropical fruit flies have not proved to be good targets for biological control (Waterhouse 1993), however, some biological control programs have achieved significant results. One outstanding success has been achieved in Hawaii against two introduced species, the oriental fruit fly, Bactrocera dorsalis Hendel and the Mediterranean fruit fly, Ceratitis capitata Weidemann (Clausen et al. 1965; Waterhouse 1993). Some crops have become practically free from attack with infestations reduced by up to 95% (Waterhouse 1993). This is mainly due to the establishment of Fopius arisanus, an opiine parasitoid of which the targeted fly species (Bactrocera dorsalis and Ceratitis capitata) are hosts. In contrast, in Fiji F. arisanus has provided only moderately effective but still inadequate control of the two native pest species, Bactrocera passiflorae (Froggatt) and B. xanthodes (Broun) (O'Connor 1960). In Australia, Fopius arisanus is established, but provides only negligible impacts on the Queensland fruit fly, Bactrocera tryoni (Froggatt), populations (Snowball & Lukins 1964).

Background to current project Effective use of opiines for all types of biological control (ie classical, inundative, manipulative) of fruit flies is limited by poor taxonomy and identification difficulties. The most recent opiine identification key (Wharton and Gilstrap 1983) is now taxonomically dated and highly restricted in the taxa covered. As acknowledged by

1 Manipulative biological control: The manipulation of elements in the environment to enhance the numbers and/or actions of natural enemies (Frank & Gillett 2006).

2 Inundative biological control: Release of large numbers of a biocontrol agents relative to the numbers of a target species, in expectation of a rapid effect. There is no implication that the released biological control agent will establish a permanent population (Frank & Gillett 2006).

8 Wharton (pers comm.), Wharton and Gilstrap is a poor taxonomic document because of the largely arbitrary nature of taxa included, being restricted to those wasps used in tephritid biological control programmes to that time. Because it was neither taxonomically comprehensive for a particular genus (or genera), nor comprehensive for a geographic region, it was always likely to be “stop-gap” of limited use, even for applied workers. However, because the Opiinae is a large group with over 1500 described species, using primary taxonomic literature to identify these wasps is a time-consuming and difficult task, especially if no host records are available, and beyond the scope of all, except opiine authorities.

For fruit fly workers of the South-Pacific region, problems of parasitoid identification are exacerbated due to the fact that regional faunas are now mixed as a result of biological control liberations, the geographic area covered is large, and much of the literature focussing on the region is in difficult to access foreign language (non- English) journals (eg. Fischer 1963a, 1967a, 1978a, 1988a). The need for quick and accurate identification of the parasitoids is, however, a common problem as they are frequently encountered in host rearing surveys of pest fruit flies.

My research stems directly from the Papua New Guinea Fruit Fly Project (PNGFFP). The PNGFFP was a collaborative program between the National Agricultural Research Institute (PNG), the Department of Agricultural and Livestock (PNG), the National Agricultural Quarantine and Inspection Agency (PNG), Griffith University (Australia) and the Plant Protection Service of the Secretariat of the Pacific Community (Fiji). As an outcome of rearing fruit flies directly from host fruit, a large series of opiine parasitoid specimens were collected from throughout Papua New Guinea. I was given the task of identifying these wasps as part of post-project activities. After initial examination of the specimens it became evident that specialist identification skills were required in order to accurately use those keys that were available, as specialist morphological terminology and composite couplets made identification complicated. Wharton and Gilstrap (1983) have a disclaimer at the beginning of their key: “It is impossible to correctly identify every specimen on key characters alone, because of extensive morphological variation, especially in material reared under nonuniform conditions. Unassociated males are particularly difficult to identify. This should be kept in mind when using the following key.” Such a disclaimer does little to comfort one new to the discipline. Extensive study of the literature and borrowing of identified specimens enabled familiarisation with the group. Over time experience led to confidence, which allowed variations to be

9 recognized and new species identified. This experience demonstrated the need for user friendly diagnostic toold to expedite the identification process, while highlighting the poor taxonomic knowledge of our regional opiine fauna. This learning process evolved to be the foundation of this thesis.

Aims and Objectives of Thesis

In this thesis I explore the taxonomy and identification of opine parasitoids of the South Pacific region with a view of enabling quick and accurate identification for competent non-expert workers. In so doing I explore the issue of taxonomy versus diagnostics and the issue of how diagnostic capacity will be maintained in the face of ongoing loss of taxonomic expertise.

The thesis contains an expanded Materials and Methods (Chapter 2) in which I detail a variety of scientific illustration techniques used in this thesis. Illustration is a core component of both accurate taxonomy and diagnostics, but not all means of illustration, be they traditional or modern, are suitable for all purposes. Because I use different illustration techniques for different purposes in this thesis, and these illustrations form a key element of the work, I considered it appropriate to provide technical detail on the methods used.

In Chapter 3 I present the core taxonomic component of this thesis: a taxonomic revision of the fruit fly infesting Opiinae of Australia and the South Pacific. In museum collections around Australia there are, thousands of specimens of fruit fly infesting Opiinae. In order to conduct a taxonomic revision of the parasitoids of the South Pacific region I borrowed museum material from all major collections holding material. These supplemented material gathered as part of the PNGFFP. Nearly 4000 specimens were examined and identified using the keys available (Szépligeti 1900; Fullaway 1950b; Wharton & Gilstrap 1983). Some specimens were previously identified; the identification of those specimens was either confirmed or corrected in this study. The results were published already by Carmichael et al. (2005).

Traditional dichotomous keys (such as that presented in Carmichael et al. 2005) are often very difficult to use and have been largely ignored by general field workers in favour of field guides and interactive identification methods (Stevenson et al. 2001). Diagnostic tools instead of dichotomous keys are rapidly becoming the new public face of scientific diagnostics. Thus while my taxonomic work as presented in

10 Chapter 3 and Carmichael et al. 2005 makes identification of the regional fruit fly infesting opiine fauna possible for anyone willing to invest time in learning opiine taxonomy, it is still likely to fail to meet the needs of fruit fly researchers who “just want to know what the little wasp is”. To meet this need, in Chapter 4, I present both an interactive multi-access key and a diagnostic package to the wasps dealt with in Chapter 3. Multi-access keys have been demonstrated to have many advantages over traditional dichotomous keys (White & Sandlant 1998). These include a reduced chance of error, a reduction in the number of questions required to arrive at a solution and the elimination of the ‘unanswerable couplet’ problem, to name a few. As the diagnostics package is computer based there is no limitation to illustrations, unlike printed keys where illustrations are often limited by cost and space. The key and diagnostics package are electronic and hence presented via a CD inserted inside the backcover of this thesis. The text of Chapter 4 simply introduces this electronic component.

Taxonomy by definition is the science of naming and classifying organisms; Identification is the process of ascertaining the taxonomic identity of an organism; and Diagnostics is the distinguishing or differentiating the species, genus, etc from others which are similar; this can be through morphological, molecular, biological (ie. using the characters provided by the taxonomist) or a combined methodology (Walter & Winterton 2007). Using the opiines as a case study, in Chapter 5, I discuss the changing roles of taxonomy, identification and diagnostics in the modern, “user driven” world.

Chapter 2

Materials and Methods

This section details all the materials and methods used throughout this thesis.

Geographic scope of project This project covers the countries and territories of the South Pacific, including Polynesia, Melanesia and Australia (Figure 2.1). It does not extend north of the equator and therefore excludes the Federated States of Micronesia. LeBlanc and Allwood (1997) failed to collect any fruit fly parasitoids in 1100 fruit and vegetable samples weighing >800kg from Micronesia and considered that there are no fruit fly parasitoids in that island group.

Figure 2.1. Geographic region covered by study.

Material examined Material examined (Table 2.1) came from: the Australian National Insect Collection, Canberra, Australia (ANIC); the Agricultural Scientific Collections Unit, NSW Dept of Primary Industries, Orange, Australia (ASCU); the Bernice P. Bishop Museum, Honolulu, Hawaii (BPBM); the Griffith University Fruit Fly Research Group, Nathan, Australia (GU); the Queensland Department of Primary Industries and Fisheries Insect Collection, Indooroopilly, Australia (QDPIF); the Secretariat of the Pacific Community, Suva, Fiji (SPC); and Texas A&M University Insect Collection (TAMU). Holotype specimens from international collections were loaned and examined by Dr.

Robert Wharton. Collections were strongly biased towards Australian and PNG material, with the number of specimens examined by country as follows: American Samoa 1, Australia 1297, Cook Islands (NZ) 123, Fiji 713, Papua New Guinea 1296, Samoa 36, Solomon Islands 5, Tonga 373, and Vanuatu 27.

Identification Methodology Specimens were examined under a Leica MZ6 stereo microscope using a Volpi intralux 4000-1 cold light source and a Planapo 1.0x objective. The vast majority of specimens were unidentified prior to this study. For those specimens that were identified, their identification was either confirmed or corrected. Various separate identification keys were utilised from the existing opiine literature (Szépligeti 1900; Fullaway 1950b; Wharton & Gilstrap 1983). No single key was available to identify all specimens encountered. Identifications were confirmed by Prof. R. A. Wharton, Texas A&M University.

Table 2.1. Source and amount of Opiinae material examined in this thesis. [ANIC, Australian National Insect Collection; ASCU, Agricultural Scientific Collections Unit; BPBM, Bernice P. Bishop Museum; GU, Griffith University; QDPIF, Queensland Department of Primary Industries and Fisheries; SPC, Secretariat of the Pacific Community; TAMU, Texas A&M University]. ANIC ASCU BPBM GU QDPIF SPC TAMU ♂ ♀ ? ♂ ♀ ? ♂ ♀ ? ♂ ♀ ? ♂ ♀ ? ♂ ♀ ? ♂ ♀ ? Diachasmimorpha 5 4 1 hageni D. kraussi 20 32 2 4 6 1 89 107 7 176 159 3 D. longicaudata 1 2 8 16 1 70 86 1 3 D. tryoni 20 23 1 15 3 5 11 Fopius arisanus 86 105 18 3 4 416 711 34 2 3 F. deeralensis 17 13 3 1 1 30 60 F. ferrari 9 16 10 F. illusorius 1 227 504 65 3 2 15 20 F. schlingeri 6 10 2 20 65 13 34 F. cf. vandenboschii 10 25 1

Opius froggatti 1 1 2 Psytallia fijiensis 24 20 5 4 7 77 81 P. novoguineensis 38 57 3 25 75 Utetes cf. albimanus 1 U. perkinsi 1 2 1 5 Totals 178 207 30 30 28 3 1 402 790 86 816 1224 37 3 6 15 21 Institution totals 415 61 1 1278 2077 9 36 Grand Total 3877

Terminology The morphological terminology follows that of Sharkey & Wharton (1997) and is illustrated in Figure 2.2 and Figure 2.3.

Figure 2.2. Opiine wing venation and terminology (following Sharkey & Wharton 1997).

Figure 2.3. Opiine body morphology and terminology (following Sharkey & Wharton 1997).

Illustration techniques Species were illustrated using a variety of techniques (Scientific Illustration, Scanning Electron Microscopy and colour photomicroscopy) as detailed below. As such, all figures within presented in this thesis are my own.

Scientific Illustration A specimen outline was traced using a Leica camera lucida (or drawing tube) attached to the microscope, which allows the specimen on the microscope stage and the drawing surface to be seen simultaneously. Each body part was drawn separately, to ensure each part was drawn in exactly the same plane and magnification to avoid body distortion. Colour patterns, sculpturing or other morphological details were noted for each body region. To ensure bilateral symmetry for dorsal view images, each outline was folded in half and symmetry corrected if required. Once all body regions were complete, the individual drawings were positioned along a ruled central axis and combined, starting with the central body region (thorax). Once all body regions were in place, the complete outline was traced over a light box onto another A4 page. The outline was then transferred in pencil to an A4 piece of EssDee ‘British’ scraper board. The outline was then traced and the illustration rendered using Rotring Black ink and Rotring 0.25 and 0.35 size pens.

For predominantly white regions, the illustration was drawn using a variety of stippling techniques. Stippling is the creation of graduations of light and shade through a series of small, discrete dots. The closer the dots the darker the shade, the farther the area is from the viewer. Conversely, the closer the area is to the viewer, the lighter the shading, depicted by dots which are farther apart. Dots were drawn in a or pattern, which allowed for repetition and flexibility without distinct lines forming to distract the viewers eye. If required, these patterns were then over-stippled to increase the darkness and depth of an area. To create hairs, the pen was pressed firmly at the hair insertion point and rapidly drawn away in the direction of the hair whilst lifting the pen from the scraper board. If a hair was placed in a dark region, the hair was drawn using the scraping method.

For predominantly black regions, the outline was painted black using black rotring ink and a fine camel hair brush. Once dry, this was ‘scraped’ back leaving crisp, clear white marks. Scraping is the removal of black ink to reveal the white underlying

clay coating of the paper and involves use of a scraper tool (a sharpened triangular piece of metal inserted into a quill or handle). Nibs of varying shapes and sizes gave a different stipple texture or line thickness. A similar technique to stippling was used in ‘lighter’ areas, but in this case more stippling using the scraper tool was required. Darker, more distant areas were left predominantly black (ie unscraped). To create hairs, the scraping tool was pressed firmly at the hair follicle and rapidly drawn away in direction of hair whilst lifting the blade from the scraper board.

For both techniques; scraping and stippling, each of the body regions was clearly defined using solid scraped lines to separate each body region as per the original outline. Hairs that overlap the body edge were drawn using a combination of techniques. The portion of the hair within the body outline was scraped, while the remainder was drawn in. Hairs were ‘sharpened’ by scraping beside the hair.

Colour Photomicroscopy

Equipment A Leica MZ16A microscope with planapo 0.5x, 1.0x and 2.0x objectives, Leica CLS150X cold light source and a DC500 (16 megapixel) digital camera, attached to a computer was used to capture images of the specimens. The Leica MZ16A is a stereo microscope with a motorized column to allow for very small incremental movements of the Z axis. Magnification is provided to one decimal place. The 0.5x, 1.0x and 2.0x objectives were used to increase magnification capability. The DC500 digital camera is a fixed camera which takes 16 pictures of 1 megapixel each in a Z- grid pattern (below) resulting in a combined picture 16 megapixels in size.

Automontage The software package Syncroscopy Automontage was used to help eliminate the problems of limited depth of field by automatically capturing the in-focus regions from a range of focal planes and combining them into a single fully-focused, high resolution image.

The process Each specimen was positioned on the microscope stage and illuminated by cold light sources. The microscope was focused on the point of the specimen closest to the microscope lens. The first image was captured and then the focus knob was turned between 1/8th-1/16th of a turn (depending on magnification) before the second picture was taken. It was critical for each image to have a region in focus overlapping with the previous image. This process was continued until the entire specimen was photographed. This usually consisted of between 40-90 images per specimen. These images were then imported into Automontage which analysed each image sequentially, retaining only those regions in focus. The final image was created by stitching all of the in-focus regions together creating a composite (or montage) of the entire photographic series. The final image is entirely in focus with a larger depth of field than that achieved by SLR cameras. Where required (ie out of focus areas still present), the final image was edited by selecting regions in the original image ‘slices’ and overriding those areas in the final image. If required, images were then touched up in Adobe Photoshop, to create an even background, removing artifacts and adjusting brightness and contrast.

For each wasp species colour photomicrographs were taken of: Head – frontal view showing clypeus Head – lateral view showing occipital carina Thorax – dorsal view showing notauli sculpture Thorax – lateral view showing sternaulus Thorax – dorsal view showing metasomal sculpturing Abdomen – dorsal view showing sculpturing on petiole and tergum 2 Whole body – dorsal Whole body – lateral Wing – showing forewing venation

I was fully responsible for purchasing, physically setting up and getting running the photomicrograph system used to produce the photos in this thesis. This system has subsequently been successfully used as a research tool by myself and colleagues. As part of the establishment process a comprehensive user guide was developed for the use of Automontage and the microscopic equipment: this user guide is presented as Appendix 2.

Scanning Electron Microscopy One to three curated specimens from each parasitoid species were selected for study and photography under a scanning electrom microscope (SEM). Prior to examination, specimens were gently cleaned using a camel hair brush to remove dust and debris. Specimens did not require drying as they were already dry preserved. For SEM, specimens were removed from the pin and/or point and attached with double-sided sticky dots onto steel stubs. The specimens and stubs were gold-coated in a BioRad SC500 gold coater for 30-45 seconds. Specimens were examined in a FEI Quanta 200 Environmental Scanning Electronic Microscope at magnifications between 150x and 400x. Digital photographs were taken at varying magnifications and scale bars were added.

For each wasp species SEM photographs were taken of: Head – frontal view showing clypeus Head – lateral view showing occipital carina Thorax – dorsal view showing notauli sculpture Thorax – lateral view showing sternaulus Thorax – Dorsal view showing metasomal sculpturing Abdomen – dorsal view showing sculpturing on petiole and tergum 2

Lucid 3 Key Builder The Lucid Builder is a software product that allows construction of multimedia matrix or multi-access keys. Lucid version 3.3 was used to create the OpiineID key presented in Chapter 4. Lucid consists of two components, a “builder” and “player”. The key developer works in the builder component, while the key user only runs the player.

The key builder opens with the screen divided into three main panels, named Properties, Features and Entities. The features panel contains the features or character sets; the entity panel contains the taxon or species names to be used in

the key; the properties panel allows the addition of web pages and images to support the key. After all of the features and entities are entered into the builder, each species is ‘described’ to the builder by checking boxes of characters that are present for that species. Once all of the species are scored in this way, the key is compiled and tested for accuracy and refined as required. Images and web pages were added for each species and each character state to assist users of the key and supply additional information. The web pages were created using Dreamweaver 8. The Lucid Builder and scoring of character states will be further detailed in Chapter 4.

Chapter 3

Taxonomy

Note: The information presented here has been formally published by Carmichael AE, Wharton RA & Clarke AR. (2005). Opiine parasitoids (Hymenoptera: Braconidae) of tropical fruit flies (Diptera: Tephritidae) of the Australia and South Pacific region. Bulletin of Entomological Research 95: 545-569. The published paper is presented in Appendix 1. This chapter is modified from Carmichael et al (2005) by the addition of “material examined” and recognition of the taxonomic changes published by Carmichael et al. (2005), e.g. Fopius ferrari sp.n. is now Fopius ferrari Carmichael and Wharton.

Contributions of authors: Associate Professor Clarke supervised the study and provided feedback on early drafts of the manuscript. Professor Wharton confirmed my identifications, provided guidance on the species description of Fopius ferrari; borrowed and examined the type specimen of Fopius illusorius from the Bernice P. Bishop Museum; and provided comments on the draft versions of the manuscript. I examined and identified all other material, completed all of the illustrations and SEM photomicrographs, wrote the first draft of the manuscript and saw it through the publication process.

Introduction Fruit flies (Diptera: Tephritidae) are among the most economically important pests of edible fruits worldwide (White & Elson-Harris 1992). Interest in the biological control of fruit flies, utilising parasitic Hymenoptera, dates back to the early 1900s (Silvestri 1913; Silvestri 1914; Lever 1938) and has been a focus of fruit fly research ever since (Knipling 1992; Waterhouse 1993). Among the most commonly used biological control agents against fruit flies are members of the Opiinae (Hymenoptera: Braconidae). Opiines are koinobiont endoparasites of the egg or larval stages of Diptera and have been introduced and released as classical biological control agents of fruit flies in many regions (Wharton & Gilstrap 1983; Waterhouse 1993; Ovruski et al. 2000). More recently, opiines have been used in manipulative and inundative release programmes against fruit flies (Messing et al. 1993; Purcell et al. 1998; Ramadan 2004), with foreign exploration continuing (Wharton et al. 2000).

Despite their importance in applied entomology, the identification of opiine fruit fly parasitoids remains difficult. A previously published key to opiine parasitoids of dacine tephritid pests (Wharton & Gilstrap 1983) is now taxonomically dated and restricted in the taxa covered. Because the Opiinae is a large group with over 1500 described species, using primary taxonomic literature to identify these wasps is a time-consuming and difficult task, especially if no host records are available. For fruit fly workers of the South-Pacific region, problems of parasitoid identification are exacerbated due to the fact that regional faunas are now mixed as a result of biological control liberations, the geographic area covered is large and much of the primary regional literature is in difficult to access foreign language journals (eg (Fischer 1963a, 1967a, 1978b, 1988b).

Fruit fly parasitoids are best collected through the rearing of fruit flies from their host fruit, rather than as collections of individual adults. Such host rearing work led, for example, to the first comprehensive survey of opiine fruit fly parasitoids from Thailand and Malaysia (Chinajariyawong et al. 2000). From the mid-1990s, a series of fruit fly projects in the South Pacific region resulted in substantial, new parasitoid collections being generated as the indirect result of host fruit studies. These projects included the Bactrocera papayae Drew & Hancock eradication programme in far north Queensland (Cantrell et al. 2001), the [Pacific] Regional Fruit Fly Project (Ferrar 1997) and the Papua New Guinea Fruit Fly Project (AusAid/ACIAR Project C32/1996/225). These new parasitoid collections provided an opportunity hitherto unavailable to examine the taxonomy, distribution and host associations of the regional fruit fly parasitoid fauna.

Based on an examination of nearly 4000 specimens, the greater percentage collected in the last decade, we present a synopsis of the South Pacific tephritid- infesting opiine fauna. Information provided includes a key to taxa and synopses of each species (including illustrations, distributions and host records). The document is targeted at fruit fly workers and inclusion of taxa is based on the parasitoids having been reared from a dacine tephritid, as evidenced by label or previously published data. Opiines reared from other dipterans (e.g. Belokobylskij et al. 2004) are not included in this review. In line with standard taxonomic manuscripts, each species is discussed individually and as such there is no final discussion section for this chapter.

Materials and Methods Geographic scope This paper covers the countries and territories of the South Pacific, including Polynesia, Melanesia and Australia. It does not extend north of the equator, therefore excludes Hawaii, the Federated States of Micronesia and the Marshall Islands.

Material examined Material examined came from: the Australian National Insect Collection, Canberra, Australia (ANIC); the Agricultural Scientific Collections Unit, NSW Dept of Primary Industries, Orange, Australia (ASCU); the Bernice P. Bishop Museum, Honolulu, Hawaii (BPBM); the Griffith University Fruit Fly Research Group, Nathan, Australia (GU); the Queensland Department of Primary Industries and Fisheries Insect Collection, Indooroopilly, Australia (QDPIF); the Secretariat of the Pacific Community, Suva, Fiji (SPC); Texas A&M University Insect Collection (TAMU). Holotype specimens from international collections were examined by R.A.W. Collections were strongly biased towards Australian and PNG material, with the number of specimens examined by country as follows: American Samoa 1, Australia 1297, Cook Islands (NZ) 123, Fiji 713, Papua New Guinea 1296, Samoa 36, Solomon Islands 5, Tonga 373, Vanuatu 27. The holotype of the newly described species is housed at the Queensland Museum, Brisbane, Australia (QM).

Terminology The morphological terminology follows that of Sharkey & Wharton (1997) and is illustrated in Figure 2.2 and Figure 2.3. The mesosoma is the thorax plus propodeum and the metasoma is the petiole plus the gaster.

Distribution and host records The distribution and host fly associations have been gathered from the literature or from specimen label data. For each host association record, the source of the record is given, in brackets, after the fly species using the following code: 1 (label data); 2 (Wharton & Gilstrap, 1983); 3 (Waterhouse, 1993); 4 (Quimio & Walter 2001); 5 (Wharton, 1999); 6 (Fullaway, 1952); 7 (Fischer, 1963a). One important caveat has been placed on the host association data to minimise error. When host fruit-rearing is being carried out, multiple fly species may be reared from the one fruit sample. If parasitoids also emerge from such samples, then ambiguity exists over

the fly species with which the parasitoid was associated (Clausen et al. 1965). Where label data for an individual specimen showed that the parasitoid was associated with two or more host fruit flies, then that label data has not been reported in this paper. Host data is given individually for each species and as a comparative summary table (Table 3.1).

A number of the parasitoids dealt with in this paper are known to parasitise fruit fly species that do not occur in the South Pacific region. These records are not included here so as to avoid possible confusion over the geographical distribution of those flies.

Key to the Opiinae parasitising Tephritidae of the South Pacific region 1 Hind tibia dorso-posteriorly without basal carina...... 3 - Hind tibia dorso-posteriorly with distinct basal carina (Figure 3.15)...... 2 2 Body uniformly dark brown to black, tarsi white...... Utetes cf. albimanus - Head and at least apical metasomal terga dark brown to black, mesosoma yellow/orange, hind tarsi dark…...... Utetes perkinsi (Fullaway) 3 Notauli smooth (Figure 3.2d) to absent or nearly so (Figure 3.12d)...... 4 - Notauli crenulate (Figure 3.5d)...... 13 4 Labrum exposed, not concealed by clypeus (Figure 3.12a)...... 5 - Labrum concealed by clypeus (Figure 3.4a)...... 9 5 Midpit of mesoscutum absent (Figure 3.12d)...... 6 - Midpit of mesoscutum deep and circular (Figure 3.11b)...... Opius froggatti Fullaway 6 Propodeal carina present (Figure 3.12e)...... 7 - Propodeal carina absent...... Psyttalia muesebecki (Fischer) 7 Forewing vein (RS+M)b present ( Figure 2.2); vein 2RS narrow throughout, not thickened medially...... 8 - Forewing vein (RS+M)b absent; junction of 2RS and m-cu expanded into a large, irregular thickening; 2RS also thickened medially in most specimens…...... Psyttalia fijiensis/novaguineensis 8 Forewing vein 2RS about twice the length of r...... Psyttalia concolor (Szépligeti)* - Forewing vein 2RS about equal in length to r…...... Psyttalia incisi (Silvestri)* 9 Ventral margin of clypeus sinuate or evenly convex (Figure 3.3a)...... 10 - Clypeus with two small teeth along midline at apex of clypeus...... Diachasmimorpha fullawayi (Silvestri)* 10 Metasomal tergum 2 smooth and polished (Figure 3.4f) (or occasionally with weak striae at extreme base)...... 11 - Metasomal tergum 2 extensively longitudinally striate...... Diachasmimorpha longicaudata (Ashmead) 11 Metasoma at least partly black...... 12 - Metasoma entirely yellow/orange...... Diachasmimorpha kraussii (Fullaway) 12 Occipital carina well developed, extending dorsally to level of mid-eye...... Diachasmimorpha hageni (Fullaway)

- Occipital carina nearly absent, or present ventrally only near base of mandible...... Diachasmimorpha tryoni (Cameron) 13 Tergum 2 densely striate (Figure 3.5f)...... 14 - Tergum 2 either without striae (Figure 3.8f) or with only a few weak striae basal-medially...... 17 14 Frons and vertex rugose-punctate...... 16 - Frons densely punctate, vertex more sparsely punctate, neither frons nor vertex rugose-punctate...... 15 15 Metasomal terga 1-5 yellow to orange...... Fopius deeralensis (Fullaway) - At least metasomal terga 2+3 black...... Fopius cf. vandenboschi 16 Metasoma dark brown to black. Apex of ovipositor lacking dorsal ridges, distinctly constricted subapically...... Fopius arisanus (Sonan) - Petiole dark brown, remainder of metasoma usually yellow/orange, at least in female. Apex of ovipositor with weak dorsal ridge and without subapical constriction...... Fopius vandenboschi (Fullaway)* 17 Metasomal terga 1-4 entirely yellow/orange (apical metasomal terga sometimes black in males)...... 18 - Metasomal terga entirely dark brown to black...... 19 18 Clypeus with ventral margin very obtusely angulate (Figure 3.6a)……………...... Fopius deeralensis (Fullaway) - Clypeus with rounded ventral margin (Figure 3.9a)...... Fopius schlingeri Wharton 19 Clypeus with ventral margin very obtusely angulate, body densely setose (Figure 3.7a)...... Fopius ferrari Carmichael & Wharton - Clypeus with rounded ventral margin (Figure 3.8a)...... Fopius illusorius (Fischer) * Indicates species which have been released but not yet recovered.

Table 3.1 Host Records. For each host association record, the source of the record is given in the table using the following code: 1 (label data); 2 Wharton and Gilstrap (1983); 3 Waterhouse (1993); 4 Quimio & Walter (2001); 5 Wharton (1999); 6 Fullaway (1952); 7 Fischer (1963a).

(Szépligeti) (Ashmead) (Fischer) (Fullaway) Wharton (Fullaway) (Fischer) (Fullaway) (Fullaway) (Fullaway) (Fullaway) (Fullaway) (Fullaway) Carmichael & Wharton (Cameron) D. hageni D. kraussii P. muesebecki P. novaguineensis D. longicaudata D. tryoni F. arisanus (Sonan) F. deeralensis F. ferrari illusorius F. F. schlingeri F. cf. vandenboschi O. froggatti P. fijiensis U. cf. albimanus U. perkinsi B. aberrans 1 (Hardy) B. aglaiae 1 1 1 (Hardy) B. bancrofti 1 (Tryon) B. barringtoniae 2 1,4 1, 2 1 (Tryon) 2 B. bidentata 1 (May) B. cacuminata 2, 1,2 2 (Hering) 3 ,4 B. calophylli 1 (Perkins & May) B. caryeae 2 Kapoor B. cucurbitae 1 1,2 (Coquillett) B. curvipennis 2,3 2 (Froggatt) B. dissidens 1 Drew B. distincta 1 (Malloch) B. dorsalis 2,3 2 2,3 Hendel ,4 B. endiandrae 1 1, 1 1 (Perkins & May) 2 B. facialis 1 1 (Coquillett) B. fagraea 2 2 1 (Tryon) B. frauenfeldi 1 1,2 1, 1 1 1 1 2,3 1 (Schiner) 2, 3 B. halfordiae 1 1,5 (Tryon) B. incisus Walker B. jarvisi 1, 1,4 2 1,2 2 (Tryon) 2, 3 B. kirki 1 1 (Froggatt) B. kraussi 1, 1,4 2 1 2 1,2 1 1, (Hardy) 2, 2 3 B. laticaudus 2 2 2 2 (Hardy) B. latifrons 2 2,4 Hendel B. limbiferus Bezzi

B. manskii 1 (Perkins & May) B. melanotus 1 (Coquillett) B. murrayi 2 (Perkins) B. musae 2,1 1 1 2,3 1 (Tryon) 3 B. neohumeralis 1, 1,4 1,5 1 1 (Hardy) 3 B. oleae (Gmelin) B. pallida 2 1, 1,2 (Perkins & May) 2 B. passiflorae 2, 1 2 1,2 1,2 (Froggatt) 3, ,4 ,3 6 B. pedestris 2 (Bezzi) B. penefurva 1 1 1 Drew B. peninsularis 5 (Drew & Hardy) B. psidii 2 2, 2 7 (Froggatt) 3 B. redunca 1 (Drew) B. rufofuscula 5 (Drew & Hancock) B. tinomiscii 1 1 Drew B. trivialis 1 (Drew) B. tryoni 1 2 1, 1 1, 5 1,2 2, (Froggatt) 2, 2, ,3 3 3, 3 7 B. umbrosa 1 1 (Fabricius) B. visenda 1 1 (Hardy) B. vulgaris 1 (Drew) B. xanthodes 1 2 1 1,2 (Broun) ,3 B. zonata 2 (Saunders) D. ciliatus 2 Loew B (Bulladacus) sp. 1 1 1

B.(Gymnodacus) sp. 1

Ceratitis capitata 2 2, 2 Wiedemann 7

Diachasmimorpha hageni (Fullaway, 1952) Opius hageni: (Fullaway 1952): 412. Opius (Biosteres) hageni: (Fischer 1963b): 231. Biosteres hageni: (Fischer 1971a): 25. Biosteres (Chilotrichia) hageni: (Fischer 1978a): 386. Diachasmimorpha hageni: (Wharton 1987): 62; (Waterhouse 1993): 38. Taxonomic History This species was originally named Opius hageni by Fullaway (1952), it then underwent subgeneric reclassification by Fischer (1963b) to be Opius (Biosteres) hageni. Fischer (1971a) formally gave the subgenus, Biosteres, generic ranking to be Biosteres hageni. After further review, it was again subgenerically reclassified, [Biosteres (Chilotrichia) hageni] (Fischer 1978a). Wharton (1987) formally removed Diachasmimorpha from Biosteres due to sufficiently different clypeal morphology and forewing venation, thus the species was renamed Diachasmimorpha hageni. Synopsis (Figure 3.1a-f) Frons and vertex sparsely setose and punctate. Frons polished and weakly punctate, the punctures anteriad ocellar field sparse and shallow; ocellar triangle arranged to form an obtuse triangle with a slight depression anteriad the median ocellus. Occipital carina well developed; clypeus in profile slightly bulged medially; ventral margin of clypeus slightly thickened and sufficiently convex to completely conceal labrum when mandibles closed. Notauli deep, broad, unsculptured; propodeum rugose laterally, with a median longitudinal carina stemming a pentagonal areola, the base of which coincides with the posterior margin. Sternaulus broad, deep, crenulate. Wings strongly infumate. Forewing r vein arising slightly basad midpoint of stigma, 2RS slightly longer than 3RSa, (RS+M)a weakly sinuate; (RS+M)b absent, m-cu arising distad 2RS. Hindwing RS absent; m-cu well developed and pigmented to wing margin. Petiole with dorsal lateral carinae strongly elevated over basal half, weaker posteriorly, but extending to posterior margin. Metasomal tergum 2 smooth and polished. Ovipositor sheath about equal in length to body including head. Ovipositor subapically sinuate. Head, mesosoma and petiole yellow/orange, remainder of metasoma dark brown; legs yellow/orange. Distribution Fiji: Viti Levu. Type Locality Ndeumba, Viti Levu, Fiji.

Material Examined Fiji: Koronivia, Viti Levu, 4.i.1955, T.G. Campbell, (ANIC), 5♂, 4♀, 1 sex undetermined. Host Associations Bactrocera passiflorae (Froggatt) (2,3,6). Discussion Morphologically similar to D. tryoni and D. kraussii. Distinguished from D. tryoni by paler colouration and D. kraussii by dark metasoma. This species is native to Fiji (Wharton & Gilstrap 1983; Waterhouse 1993) and was described from specimens collected by N.L.H. Krauss in May 1951.

Figure 3.1. Diachasmimorpha hageni. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view.

Diachasmimorpha kraussii (Fullaway, 1951) Opius kraussii (Fullaway 1951): 249. Opius (Biosteres) kraussii: (Fischer 1963b): 233. Biosteres kraussi: (Fischer 1971a): 26. Biosteres (Chilotrichia) kraussi: (Fischer 1978a): 386. Diachasmimorpha kraussii: (Wharton 1987): 62; (Waterhouse 1993): 40. Taxonomic History Diachasmimorpha kraussi was originally named Opius kraussii by Fullaway (1951), it then underwent subgeneric reclassification by Fischer (1963b) to be named Opius (Biosteres) kraussii. Fischer (1971a) formally gave the subgenus, Biosteres, generic ranking [Biosteres kraussi]. The Biosteres genus was further reviewed by Fischer (1978a) and again subgenerically divided [Biosteres (Chilotrichia) kraussi]. Wharton (1987) reviewed the genus and found sufficient differences in clypeal morphology and forewing venation to formally remove Diachasmimorpha from Biosteres, thus the species was named Diachasmimorpha kraussii. Synopsis (Figure 3.2a-f) Frons and vertex punctate and setose. Ocellar triangle arranged to form an obtuse triangle with a slight depression anteriad the median ocellus. Occipital carina well developed; clypeus semicircular, ventral margin completely concealing labrum when mandibles closed. Notauli deep, broad, unsculptured; propodeum rugose laterally, with a median longitudinal carina stemming a pentagonal areola, the base of which coincides with the posterior margin. Wings hyaline. Forewing r arising from midpoint of stigma, 2RS slightly longer than 3RSa, (RS+M)a weakly sinuate; (RS+M)b absent, m-cu arising distad 2RS. Hindwing RS absent; m-cu well developed and pigmented to wing margin. Petiole weakly longitudinally striate, dorsal carinae strongly elevated and weakly convergent to posterior margin. Metasomal tergum 2 smooth; Ovipositor sheath about equal in length to body, including head. Ovipositor apically sinuate. Entire body yellow orange. Distribution Australia: Collected from Biogu Is., Darnley Is., Dauan Is., Sabai Is. and mainland Australia, from Cape Tribulation in the north to Sydney in the south. Papua New Guinea: Collected from both high and low altitude regions of the main island of New Guinea and the Gazelle Peninsular of East New Britain. Solomon Islands: Visale. Vanuatu: Efate. Type Locality Deeral, Australia.

Material examined Australia: Atherton Tableland, Queensland, .viii.1949, Solanum mauritianum, N.L.H. Krauss, (ASCU), 1♂; Bauple, Queensland, 6.iii.1960, ex yellow guava, T.G. Campbell, (ANIC), 1♂; Biniguy, N.S.W., 9.iii.1976, ex green husks of pecan nut, Dacus tryoni, D.A. Herps, (ASCU), 2♂, 2♀; Brisbane, Queensland, .iii.1981, on guava, Dacus tryoni, G. Fitt, (ANIC), 2♂, 3♀; Caboolture, Queensland, 6.ii.1959, Solanum mauritianum, (ANIC), 1♀; Cairns, Queensland, .iii.1981, Solanum seaforth, Dacus neohumeralis, G. Fitt, (ANIC), 1♀; Cairns, Queensland, ii.1962, Nauclea orientalis, G.J. Snowball, (ANIC), 1♀; Cairns, Queensland, 9.i.1982, Planchonella careya, Dacus tryoni, G. Fitt, (ANIC), 1♂; Cairns, Queensland, 9.ii.1954, Terminalia catappa, Dacus tryoni, A.W.S. May, (ANIC), 1♂; Coffs Harbour, N.S.W., 5.iii.1957, ex Mango, G.J. Snowball, (ANIC), 1♀; Currumbin, Queensland, 12.ii.1959, Solanum mauritianum, (ANIC), 2♂, 3♀; Duranbah, Queensland, 21.ii.1960, Solanum mauritianum, T.G. Campbell, (ANIC), 3♀, 2 sex undetermined; East Palmerston, Queensland, .ii.1962, Solanum mauritianum, G.J. Snowball, (ANIC), 2♀, 2♂; Ellis Beach, Cairns, Queensland, 19.iii.1982, Terminalia catappa, Dacus jarvisi, G. Fitt, (ANIC), 1♂, 1♀; Ellis Beach, Cairns, Queensland, 15.i.1982, Terminalia catappa, Dacus jarvisi, G. Fitt, (ANIC), 1♂, 1♀; Pinjin Hill, near Innisfail, Queensland, 9.vii.1959, Solanum, mauritianum, T.G. Campbell, (ANIC), 3♀, 1 sex undetermined; Trinity Beach, Cairns, Queensland, 18.iii.1982, Terminalia catappa, Dacus, jarvisi, G. Fitt, (ANIC), 2♀; Freshwater, Queensland, .i.1963, ex guava, G.J. Snowball, (ANIC), 1♀; Freshwater, Queensland, 31.vii.1959, on comquat, T.G. Campbell, (ANIC), 1♂; Mt. Gravatt, Queensland, 28.ii.1955, M.A. Bateman, (ANIC), 1♂, 1♀; Murwillumbah, N.S.W., xii.1964, Solanum mauritianum, R.G. Lukins, (ANIC), 1♀, 2♂; Nerang, Queensland, 15.ii.1961, ex china pear, G.J. Snowball, (ANIC), 1♂, 4♀; North Queensland, Cape Kimberley, 5.viii.1996, Fagraea cambagei, Bactrocera bidentata, CK655, (QDPIF), 1♀; North Queensland, Cape Kimberley, 3.viii.1996, Acmena graveoleus, Bactrocera kraussi, CK675B, (QDPIF), 1♀; North Queensland, Cape Kimberley, 7.viii.1996, Syzygium sp., Bactrocera kraussi & Bactrocera neohumeralis CK682, (QDPIF), 1♂, 1♀; North Queensland, Cape Tribulation at RFCT41, 15.viii.1996, Ficus virgata, Trypetines, CT1002, (QDPIF), 6♂, 2♀; North Queensland, Cape Tribulation at RFCT 2, 20.iii.1996, Bactrocera kraussi & Bactrocera manskii, CT271, (QDPIF), 1♂, 2♀; North Queensland, Cape Tribulation at RFCT1, 3.iv.1996, Acmena sp, Bactrocera kraussi, CT373, (QDPIF), 2♂; North Queensland, Cape Tribulation at RFCT1, 28.v.1996, Acmena sp, Bactrocera kraussi, CT917, (QDPIF), 1♂, 1♀; North Queensland, Josephine Falls at RFJF8,

4.ii.1996, Acmena graveolens, Bactrocera kraussi, JF29, (QDPIF), 2♂; AUSTRALIA, North Queensland, Mulgrave R, 14.ii.1996, Bactrocera neohumeralis, MR8, (QDPIF), 1♂, 1♀; North Queensland, Tully Heads, 27.xi.1995, Mangifera indica, Bactrocera tryoni, NQ139, (QDPIF), 2♀; North Queensland, Port Douglas, 31.i.1996, Terminalia catappa, Bactrocera frauenfeldi, Bactrocera jarvisi, Bactrocera mayi, Bactrocera neohumeralis, Bactrocera tryoni, NQ452, (QDPIF), 4♂, 4♀; North Queensland, Palm Cove, 1.iii.1996, Terminalia catappa, Bactrocera tryoni, Bactrocera papayae, Bactrocera frauenfeldi, Bactrocera jarvisi, Bactrocera neohumeralis, NQ552, (QDPIF), 8♂, 7♀; North Queensland, Wangetti Beach, 5.iii.1996, Terminalia muelleri, Bactrocera tryoni & Bactrocera pallida, NQ607B, (QDPIF), 1♀; North Queensland, Wangetti Beach, 15.iii.1996, Terminalia muelleri, Bactrocera neohumeralis & Bactrocera tryoni, NQ630, (QDPIF), 3♀; North Queensland, Rocky Point, 20.iii.1996, Terminalia arenicola, Bactrocera tryoni & Bactrocera neohumeralis, NQ655, (QDPIF), 1♂; North Queensland, Charters Towers, 9.xi.1995, Prunus persica var nectarina, Bactrocera tryoni, NQ69, (QDPIF), 3♂, 1♀; Queensland, Nambour, ii.1993, Feijoa, Bactrocera tryoni & Bactrocera neohumeralis, Q67, M.M. Elson-Harris, (QDPIF), 2♂, 1♀; Queensland, Redlands H.R.C., xi.1994, peaches, Bactrocera tryoni & Bactrocera neohumeralis, H. Lahey, (QDPIF), 2♂, 8♀; Queensland, Rockhampton, i.1995, ? Planchonia careya, B. (Afrodacus) jarvisi, P. Leach, (QDPIF), 77♂, 57♀; Queensland, Rockhampton, 3.ii.1982, Planchonella careya, Dacus jarvisi, G. Fitt, (ANIC), 1♂, 1♀; Sydney, N.S.W., 28.vi.1954, G.J. Snowball, (ANIC), 1♂; Tamworth, N.S.W., 20.iii.1963, ex Boxthorn berries, L.C. Williamson, (ASCU), 1♀; Tamworth, N.S.W., 10.iii.1964, ex Boxthorn berries, L.C. Williamson, (ASCU), 1♀; Tamworth, N.S.W., 13-14.iii.1963, ex Boxthorn berries, NSW AG, L.C. Williamson, (ASCU), 1♂, 1♀; The Pocket, Maryborough, Queensland, 6.iv.1950, on guava, T.G. Campbell, (ANIC), 1♀; Toowoomba, Queensland, .xii.1948, ex walnut, Strumeta spp., ANIC, A.W.S. May, (ANIC), 1♂, 2♀; Yarrahappini, N.S.W., 31.i.1962, ex peach, R.G. Lukins, (ANIC), 1♀. Boigu Island (Australia): Village, 28.iii.1994, Terminalia catappa, Bactrocera frauenfeldi & Bactrocera neohumeralis, BG 060, E. Hamacek, (QDPIF), 3♀, 2, sex undetermined. Dauan Island (Australia): 25.x.1993, Terminalia catappa, Bactrocera frauenfeldi, DU019, B. Sabine & T. Jawai, (QDPIF), 3♂; School, 7.xi.1993, Terminalia catappa, Bactrocera frauenfeldi, DU 024, Thomas Jawai, (QDPIF), 1♂, 2♀; School, 13.ii.1994, Terminalia catappa, Bactrocera frauenfeldi & Bactrocera neohumeralis, DU043, B. Sabine, (QDPIF), 5♂, 15♀; Council flat, 24.x.1993, Terminalia catappa,

Bactrocera frauenfeldi, DU019, B. Sabine & T. Jawai, (QDPIF), 7♂, 3♀; Council Office, 7.xi.1993, Terminalia sp., Bactrocera neohumeralis, DU 028, Thomas Jawai, (QDPIF), 2♂, 2♀; Village, 12.iii.1994, Terminalia catappa, Bactrocera frauenfeldi, DU 060, E. Hamacek & T. Jawai, (QDPIF), 1♀; Village, 14.ii.1994, Terminalia catappa, Bactrocera frauenfeldi & Bactrocera neohumeralis, DU048, B. Sabine, (QDPIF), 1♀, 1, sex undetermined; Village, 14.ii.1994, Terminalia catappa, Bactrocera frauenfeldi, Bactrocera neohumeralis & Bactrocera turneri, DU049, B. Sabine & T. Jawai, (QDPIF), 1♂, 9♀; Village, 13.v.1993, Terminalia catappa, DU5, E. Hamacek, (QDPIF), 2♀. Papua New Guinea: E. New Britain, Kerevat: L.A.E.S., 12-Dec-97, MYRTACEAE, Syzygium, Bactrocera frauenfeldi & Bactrocera obliqua, I 118-122, (GU), 3♂, 1♀; E. New Britain, Kerevat: L.A.E.S., Michael Wan house, 18-Dec-97, MYRTACEAE, Syzygium, Bactrocera frauenfeldi & Bactrocera obliqua, I 143-148, (GU), 3♂, 5♀; E. New Britain, Kerevat: L.A.E.S., Kavieng compound, 19-Dec-97, MYRTACEAE, Syzygium malaccense, Bactrocera frauenfeldi & Bactrocera obliqua, I 156-158, (GU), 9♂, 8♀; E. New Britain, Sonoma: S.D.A. school, mechanics workshop, 22- Dec-97, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, I 162-163, (GU), 1♂; E. New Britain, Sonoma: S.D.A. school, mechanics workshop, 22-Dec-97, MYRTACEAE, Syzygium aqueum, Bactrocera frauenfeldi, I 169-170, (GU), 12♂, 9♀, 1, sex undetermined; E. New Britain, Kerevat: L.A.E.S., Orchard nr. Country Club, 24-Nov-97, MYRTACEAE, Psidium cattleianum, Bactrocera frauenfeldi, I 6, (GU), 2♂, 3♀; E. New Britain, Kerevat: L.A.E.S., Labour compound guest hse., 2- Jul-98, MYRTACEAE, Syzygium malaccense, Bactrocera frauenfeldi, I 613, (GU), 1♂, 1♀; E. New Britain, Kerevat: L.A.E.S., M. Powell house, 15-Jul-98, MYRTACEAE, Syzygium malaccense, Bactrocera frauenfeldi & Bactrocera obliqua, I 724-725, (GU), 2♀; E. New Britain, Kerevat: L.A.E.S., Kavieng cpd: CCRI guest h., 28-Oct-98, MYRTACEAE, Syzygium aqueum, Bactrocera frauenfeldi, I 767-769, (GU), 4♂; E. New Britain, Kerevat: L.A.E.S., tree nursery, 28-Oct-98, MYRTACEAE, Syzygium, Bactrocera frauenfeldi & Bactrocera obliqua, I 797, (GU), 17♂, 6♀, PNG, E. New Britain, Sonoma: S.D.A. school, mechanics workshop, 6-Nov-98, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, I 802-804, (GU), 2♀; E. New Britain, Kerevat: L.A.E.S., near Labourer compound, 9-Feb-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, I 904-905, (GU), 1♂; E. New Britain, Kuradui Block, Raluana 4, 4-Aug-99, ANACARDIACEAE, Mangifera indica, frauenfeldi, I 993-1080, (GU), 1♂; Central, Lobulogo village Hiritano Highway, 23-Feb-99, OXALIDACEAE, Averrhoa carambola, Bactrocera frauenfeldi,

L 1301-1400, (GU), 1♀; PNG Central, Laloki block A.Mehuze garden, 19-May-99, MUSACEAE, Musa x paradisiaca, L 2905, (GU), 1♂; W. Highlands, Hagen: Bob Hagraves farm, 18-Nov-98, RUTACE, Citrus sinensis, MH 57, (GU), 1♀; Morobe, Lae: Mutzing Dist, Kaiapit area, 10-Nov-99, CUCURBITACEAE, Momordica charantia, Bactrocera cucurbitae, N 1113, (GU), 1♂; Morobe, Lae: Omsis forest, 07- Feb-00, COMBRETACEAE, Terminalia, Bactrocera penefurva, N 1259, (GU), 1♂, 1♀; Morobe, Lae: Omsis forest, 16-Feb-00, COMBRETACEAE, Terminalia, Bactrocera frauenfeldi & Bactrocera penefurva, N 1273, (GU), 1♀; Morobe, Lae: Bukawa, Tikeling 2 forest, 04-Apr-00, Bactrocera unidentified, N 1377, (GU), 2♀; Morobe, Lae: Gabensis Village area, 06-Apr-00, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 1384, (GU), 5♀; Morobe, Lae: Markham farm, Mararumi, 06-Apr-00, CUCURBITACEAE, Momordica charantia, frauenfeldi, N 1385, (GU), 1♂; Morobe, Lae: Markham, Nasuapum vill, 28-Apr-00, MYRTACEAE, Psidium guajava, frauenfeldi, N 1411, (GU), 1♀; Morobe, Lae: FRI forest/ botanical gard, 09-May-00, COMBRETACEAE, Terminalia catappa, Bactrocera frauenfeldi & Bactrocera penefurva, N 1428, (GU), 9♂, 6♀; Morobe, Lae: FRI forest/ botanical gard, 31-May- 00, COMBRETACEAE, Terminalia dissidens, N 1491, (GU), 2♂; Morobe, Lae: FRI forest/ botanical gard, 31-May-00, COMBRETACEAE, Terminalia, Bactrocera dissidens & Bactrocera frauenfeldi, N 1494, (GU), 1♂; Morobe, Lae: Bubia Agr. Station, 06-Feb-99, COMBRETACEAE, Terminalia catappa, frauenfeldi, N 299, (GU), 3♀; Morobe, Lae: Lae town, FRI office area, 13-Apr-99, COMBRETACEAE, Terminalia catappa, Bactrocera exima & Bactrocera frauenfeldi, N 325, (GU), 3♂, 8♀, 1, sex undetermined; Morobe, Lae: University of Technology, 22-Apr-99, MYRTACEAE, Syzygium, Bactrocera frauenfeldi, N 359, (GU), 1♂, 2♀; Morobe, Lae: University of Technology, 22-Apr-99, MYRTACEAE, Syzygium, Bactrocera trivialis & Bactrocera frauenfeldi, N 361, (GU), 3♂, 4♀, 2, sex undetermined; Morobe, Lae: Bundun conf centre, 23-Apr-99, MORACEAE, Ficus, N 372, (GU), 3♂, 4♀; Morobe, Lae: FRI bota/garden forest, 30-Apr-99, SAPINDACEAE, Pometia pinnata, Bactrocera latissima (sp.nr.) & Bactrocera lineata, N 405, (GU), 2♂, 1♀; Morobe, Lae: FRI bota/garden forest, 30-Apr-99, APOCYNACEAE, Tabernaemontana novoguineensis, Bactrocera latissima (sp.nr.) & Bactrocera lineata, N 414, (GU), 1♀; Morobe, Lae: 11-mile Suambu farm, 03-May-99, MYRTACEAE, Syzygium, Bactrocera frauenfeldi, N 447, (GU), 1♀; Morobe, Lae: Markham farm, Mararumi, 04-May-99, ANACARDIACEAE, Mangifera indica, B. frauenfeldi, N 457, (GU), 2, sex undetermined; Morobe, Lae: Bubia Agri Station, 18- Aug-99, OXALIDACEAE, Averrhoa carambola,B. frauenfeldi, N 755, (GU), 1♀; Morobe, Lae: Bubia Agri Station, 18-Aug-99, MYRTACEAE, Syzygium, Bactrocera

frauenfeldi & Bactrocera trivialis?, N 756, (GU), 6♀; Morobe, Lae: Markham farm, Mararumi, 25-Aug-99, MYRTACEAE, Psidium guajava, Bactrocera trivialis & Bactrocera frauenfeldi N 795, (GU), 1♂; E. New Britain, Kerevat C.I.S. prison, 2- Feb-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 2591-2640, (GU), 1♀; E. New Britain, Kerevat C.I.S. prison, 10-Feb-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua R 2641-2690, (GU), 3♂, 11♀: E. New Britain, Kerevat C.I.S. prison, 24-Feb-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 2752-2801, (GU), 1♂; E. New Britain, Kerevat C.I.S. prison, 17-Mar-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 2852-2901, (GU), 1♂; E. New Britain, Kerevat: L.A.E.S., Orchard nr. Country Club, 30-Aug-99, OXALIDACEAE, Averrhoa carambola, Bactrocera frauenfeldi, R 3606- 3725, (GU), 3♀, 1, sex undetermined; E. New Britain, Kerevat: L.A.E.S., Guava block 408, 1-Sep-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 3726-3775, (GU), 5♀; E. New Britain, Kerevat C.I.S. prison, 3-Mar-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R2802-2851, (GU), 1♂, 1♀. Saibai Island (Australia): 9.iii.1994, Terminalia catappa, Bactrocera frauenfeldi, Bactrocera jarvisi, Bactrocera neohumeralis & Bactrocera papayae SB 076, R. Enosa, (QDPIF), 1♀; Mission House, 9.iii.1994, Terminalia catappa, Bactrocera frauenfeldi, Bactrocera jarvisi, Bactrocera neohumeralis & Bactrocera papayae, SB 078, R. Enosa, (QDPIF), 6♂, 8♀; Village, 27.i.1994, Terminalia catappa, Bactrocera frauenfeldi, Bactrocera neohumeralis & Bactrocera papayae, SB066, R. Enosa, (QDPIF), 10♂, 7♀; Village, 27.i.1994, Terminalia catappa, Bactrocera frauenfeldi, Bactrocera neohumeralis & Bactrocera papayae, SB067, R. Enosa, (QDPIF), 2♀; Village, 27.i.1994, Terminalia catappa, Bactrocera frauenfeldi, Bactrocera neohumeralis & Bactrocera papayae, SB068, R. Enosa, (QDPIF), 13♂, 2♀. Solomon Islands: Visale, N.W., Guadalcanal, 29.v.1996, Terminalia catappa, Bactrocera frauenfeldi & Bactrocera tigrina, SI 1617, (QDPIF), 1♀, Darnley Island (Australia): Paul's Garden, 21.ii.1994, Psidium guajava, Bactrocera frauenfeldi & Bactrocera neohumeralis, TSD056, B. Sabine & P. Saylor, (QDPIF), 2♂, 1♀; By old school, 21.ii.1994, Terminalia catappa, Bactrocera frauenfeldi, TSD068, B. Sabine & P. Saylor, (QDPIF), 2♀. Vanuatu: Efate, 4.ix.1995, Bactrocera umbrosa, QDPI, M. Vagalo, (QDPIF), 15♂, 4♀. Host Associations

Bactrocera barringtoniae (Tryon) (2), B. bidentata (May) (1), B. cacuminata (Hering) (2,3), B. cucurbitae (Coquillett) (1), B. dissidens Drew (1), B. frauenfeldi (Schiner) (1), B. jarvisi (Tryon) (1,2,3), B. kraussi (Hardy) (1,2,3), B. murrayi (Perkins) (2), B. neohumeralis (Hardy) (1,3), B. pallida (Perkins & May) (2), B. penefurva Drew (1), B. tryoni (Froggatt) (1), and B. umbrosa (Fabricius) (1). Discussion Morphologically similar to D. longicaudata and D. hageni, but can be readily distinguished from these species by its pale colouration and smooth metasomal tergum 2. Diachasmimorpha kraussii is considered native to Australia (Wharton & Gilstrap 1983; Waterhouse 1993) and Solomon Islands (Waterhouse 1993); PNG is also likely to be part of its native range.

Figure 3.2. Diachasmimorpha kraussii. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view.

Diachasmimorpha longicaudata (Ashmead, 1905) Biosteres longicaudatus (Ashmead 1905): 970. Diachasmimorpha comperei (Viereck 1913): 641; syn. by (Wharton & Gilstrap 1983): 733. Biosteres compensans (Silvestri 1916):168; (Fischer 1963b) 234 as subsp.; (Wharton & Gilstrap 1983): 733 as syn. Biosteres formosanus (Fullaway 1926): 283; syn. by (Fischer 1963b): 234. Opius longicaudatus: (Fullaway 1951): 243. Opius longicaudatus var. chocki (Fullaway 1953): 310. Opius longicaudatus var. novocaledonicus (Fullaway 1953): 311. Opius longicaudatus var. malaiaensis (Fullaway 1953): 312. Opius longicaudatus var. taiensis (Fullaway 1953): 313. Opius (Biosteres) longicaudatus taiensis: (Fischer 1963b): 234. Opius (Biosteres) longicaudatus longicaudatus: (Fischer 1963b): 234. Biosteres longicaudatus: (Fischer 1971a): 27. Biosteres (Chilotrichia) longicaudatus: (Fischer 1978a): 386. Diachasmimorpha longicaudata: (Wharton 1987): 62. Taxonomic History Biosteres longicaudatus was originally described by Ashmead (1905). Viereck (1913) described the same species as Diachasmimorpha comperei; Silvestri (1916) also described the same species under the name Biosteres compensans, both D. comperei and B. compensans were synonymised as B. longicaudatus by Wharton and Gilstrap (1983). Fullaway (1926) described the species as Biosteres formosanus which was then synonymised by Fischer (1963b). Fullaway (1951) reviewed the generic classification and placed the species into the genus Opius [Opius longicaudatus]. In 1953, Fullaway again reviewed the species and divided it into a series of varietal names: Opius longicaudatus var. chocki; Opius longicaudatus var. novocaledonicus; Opius longicaudatus var. malaiaensis; Opius longicaudatus var. taiensis. Fischer (1963b) formally ranked the varietal name taiensis as subspecies in the subgenus Biosteres [Opius (Biosteres) longicaudatus taiensis] and the remaining varietal names were formally named Opius (Biosteres) longicaudatus longicaudatus. Wharton & Marsh (1978) stated that the longicaudatus complex needed to be examined in more detail. After further revision Fischer (1971a) formally renamed the species Biosteres longicaudatus. Another subgeneric reclassification, by Fischer (1978a) named it Biosteres (Chilotrichia) longicaudatus. Wharton (1987) formally removed Diachasmimopha from Biosteres due to sufficient

morphological differences in the clypeus and forewing venation, thus the species was named Diachasmimorpha longicaudata. Synopsis (Figure 3.3a-f) Frons and vertex sparsely punctate and setose. Ocellar triangle arranged to form an obtuse triangle with a slight depression anteriad the median ocellus. Occipital carina well developed; ventral margin of clypeus completely concealing labrum when mandibles closed. Notauli deep, broad, unsculptured; propodeum irregularly areolate with a median longitudinal carina extending from anterior margin to areola. Wings hyaline. Forewing r arising slightly basad midpoint of stigma, 2RS slightly longer than 3RSa, (RS+M)a weakly sinuate; (RS+M)b absent, m-cu arising distad 2RS. Hindwing RS absent; m-cu well developed and pigmented to wing margin. Petiole longitudinally striate, with dorsal lateral carinae strongly elevated and slightly divergent to posterior margin. Metasomal tergum 2 longitudinally striate; Ovipositor sheath about equal in length to body, including head. Body entirely yellow orange, or with black transverse stripe(s) subapically on metasomal terga. Distribution Australia: Collected from the Bloomfield River, far North Queensland and Lord Howe Island. Fiji: Collected from many sites on Viti Levu and Vanua Levu, also collected on islands of the Lau and Yasawa groups. Papua New Guinea: Lae and Sonoma. Vanuatu: Efate. Type Locality Manila, Philippines Material examined Australia: Lord Howe Island, .iii.1961, ex cherry guava, T.G. Campbell, (ASCU), 1♂, 2♀; North Queensland, Bloomfield R, 21.viii.1996, Chionanthus, ramiflora, Euphranta, linocierae, BR43, (QDPIF), 1♂. Fiji: Colo-i-Suva Forest Park, 11.vi.1992, Amaroria soulameiodes, Bactrocera passiflorae, F2876, G. Walker et al, (QDPIF), 2♀; Koronivia, ex lab culture of B. xanthodes, vii.1992, Bactrocera xanthodes, SPC, (SPC), 1♂, 1♀; Lambasa, Vunika, Labasa, Vanua Levu, 16.vii.1992, Inocarpus fagiferus, Bactrocera passiflorae, F3105, A. Allwood et al, (QDPIF), 1♀; Lautoka, Shankar's, 25.i.1991, Averrhoa carambola, Bactrocera passiflorae, F0225, G. Walker & Sada Nand Lal, (QDPIF), 1♂; Malevu, Nailiko Rd Bridge, 19.xii.1991, Ochrosia oppositifolia, Bactrocera passiflorae, F1892, A. Allwood et al, (QDPIF), 8♂, 19♀; Malevu, Nailiko Rd Bridge, 26.ii.1992, Ochrosia oppositifolia, Bactrocera passiflorae, F2322, G. Walker et al, (QDPIF), 9♂, 4♀; Malevu, Nailiko Rd Bridge, 19.xi.1992, Ochrosia oppositifolia, Sp., nov. 2, F3464, Ema Tora, (QDPIF), 1♀; Nadi, Legalega R.S., 15.xi.1993, Psidium

guajava, Bactrocera passiflorae, F049, G. Walker et al, (QDPIF), 1♂; Nadi, Legalega R.S., 11.i.1991, Psidium guajava, Bactrocera passiflorae, F0163, A. Allwood et al, (QDPIF), 2♂; Nadi, Legalega R.S., 11.i.1991, Psidium guajava, Bactrocera passiflorae, F0163, A. Allwood & G. Walker, (QDPIF), 2♂, 6♀; Nadi, Legalega R.S., 26.xi.1991, Artocarpus altilis, Bactrocera xanthodes, F1714, G. Walker et al, (QDPIF), 1♂; Nadi, Legalega R.S., 19.xii.1991, Mangifera indica, Bactrocera passiflorae, F1887, A. Allwood et al, (QDPIF), 1♂, 1♀; Nadi, Legalega R.S., 23.i.1992, Mangifera indica, Bactrocera passiflorae, F1999, Harikesh Lai, (QDPIF), 1♂; Nadi, Legalega R.S., 29.i.1992, Psidium guajava, Bactrocera passiflorae, F2057, G. Walker et al, (QDPIF), 6♂, 8♀; Nadi, Tauleou mango orchard, 13.xi.1992, Mangifera indica, Bactrocera passiflorae, F3457, G. Walker et al, (QDPIF), 1♀; Nausori, Naduruloulou R.S., 6.i.1992, Citrus maxima, Bactrocera passiflorae, F1926, A. Allwood et al, (QDPIF), 5♂, 3♀; Nausori, Naduruloulou R.S., 22.i.1992, Psidium guajava, Bactrocera passiflorae, F1956, A. Allwood et al, (QDPIF), 3♂; Nausori, Naduruloulou R.S., 6.ii.1992, Psidium guajava, Bactrocera passiflorae, F2156, G. Walker et al, (QDPIF), 7♂, 4♀; Nausori, Naduruloulou R.S., 19.ii.1992, Garcinia mangostana, Bactrocera passiflorae, F2185, G. Walker et al, (QDPIF), 1♂; Nausori, Naduruloulou R.S., 22.iv.1992, Fortunella japonica, Bactrocera passiflorae, F2611, Ema Tora et al, (QDPIF), 1♂; Nausori, Naduruloulou R.S., 4.v.1992, Citrus maxima, Bactrocera passiflorae, F2747, G. Walker et al, (QDPIF), 1♂; Nausori, Naduruloulou R.S., 11.vi.1992, Psidium guajava, Bactrocera passiflorae, F2908, G. Walker et al, (QDPIF), 1♀; Nausori, Toga Village, 21.x.1992, Averrhoa carambola, Bactrocera passiflorae, F2624, Ema Tora et al, (QDPIF), 4♂, 3♀; Pacific Harbour, Lomaloma Beach, 14.vi.1992, Bactrocera passiflorae, F2934, G. Walker et al, (QDPIF), 1♂, 1♀; Rakiraki, A. Prasad, 21.iv.1992, Citrus sinensis, Bactrocera passiflorae, F2667, G. Walker et al, (QDPIF), 1♀; Rakiraki, Prassad's, 19.ii.1991, Inocarpus fagiferus, Bactrocera passiflorae, F0398, A. Allwood & G. Walker, (QDPIF), 1♂; Rakiraki, Lal's, 23.iv.1991, Fortunella japonica, Bactrocera passiflorae, F0676, G. Walker & E. Hamacek, (QDPIF), 3♀; Rakiraki, Sat Deo's, 27.xi.1991, Artocarpus altilis, Bactrocera passiflorae & Bactrocera xanthodes, F1755, A. Allwood et al, (QDPIF), 5♂, 11♀; Rakiraki, Sat Deo's, 21.x.1992, Artocarpus altilis, Bactrocera passiflorae & Bactrocera xanthodes, F3383, G. Walker et al, (QDPIF), 5♀; Rakiraki, Waimari, 26.v.1992, Citrus reticulata, Bactrocera passiflorae, F2836, G. Walker et al, (QDPIF), 1♂, 1♀; Sigatoka, Sigatoka R.S, 25.vii.1991, Citrus paradisi, Bactrocera passiflorae, F1147, G. Walker et al, (QDPIF), 1♂, 1♀; Sigatoka, Sigatoka R.S, 26.ii.1992, Psidium guajava, Bactrocera passiflorae, F2310, G. Walker et al, (QDPIF), 1♂; Sigatoka Valley, 1st Deo,

23.iv.1992, Citrus reticulata, Bactrocera passiflorae, F2719, G. Walker et al, (QDPIF), 1♂; Suva, U.S.P. Compound, 12.iv.1992, Terminalia littoralis, Bactrocera passiflorae, F2575, E. Hamacek et al, (QDPIF), 4♂. Papua New Guinea: E. New Britain, Sonoma: S.D.A. school, mechanics workshop, 22-Dec-97, MYRTACEAE, Syzygium aqueum, Bactrocera frauenfeldi, I 169-170, (GU), 3♀; Morobe, Lae: Mutzing Dist, Kaiapit area, 10-Nov-99, CUCURBITACEAE, Momordica charantia, Bactrocera cucurbitae, N 1113, (GU), 2♂, 2♀; Morobe, Lae: Omsis forest, 16-Feb-00, COMBRETACEAE, Terminalia, Bactrocera frauenfeldi & Bactrocera penefurva, N 1273, (GU), 1♀; Morobe, Lae: Markham farm, Mararumi, 28-Jul-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 652, (GU), 1, sex undetermined; Morobe, Lae: Mutzing Dist, Kaiapit area, 10-Nov-99, CUCURBITACEAE, Momordica charantia, cucurbitae, N 1111, (GU), 5♀; Morobe, Lae: Mutzing Dist, Kaiapit area, 10-Nov-99, CUCURBITACEAE, Momordica charantia, cucurbitae, N 1114, (GU), 1♀; Morobe, Lae: Kaipit, Markham valley, 23- Jun-99, CUCURBITACEAE, Momordica charantia, cucurbitae, N 603, (GU), 4♂, 2♀; Morobe, Lae: Bubia Agri Station, 30-Jun-99, CUCURBITACEAE, Cucurbita pepo, N 604, (GU), 1♀; Morobe, Lae: Markham farm, Mararumi, 28-Jul-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 691, (GU), 1♂, 1♀; Morobe, Lae: Erab Agri Station, 4-Aug-99, ANNONACEAE, Annona reticulata, frauenfeldi, N 708, (GU), 1♂. Tonga: Eua, 6.iv.2000, guava, (SPC), 2♀. Vanuatu: Efate, 4.ix.1995, Bactrocera umbrosa, M. Vagalo, (QDPIF), 8♀. Host Associations B. caryeae Kapoor (2), B. cucurbitae (1,2), B. curvipennis (Froggatt) (2,3), B. dorsalis (Hendel) (2,3), B. frauenfeldi (1,2), B. latifrons (Hendel) (2), B. passiflorae (1), B. pedestris (Bezzi) (2), B. psidii (Froggatt) (2), B. tryoni (2), B. umbrosa (1), B. xanthodes (Broun) (1), B. zonata (Saunders) (2), Dacus ciliatus Loew (2), and Ceratitis capitata (Wiedemann) (2). Discussion Morphologically similar to D. kraussii and D. hageni. Distinguished by pale colouration and having striae on metasomal tergum 2. Diachasmimorpha longicaudata is not considered native to the South Pacific region, but was introduced to Australia and elsewhere around the Pacific from Hawaii for the biological control of pest Bactrocera species during the 1950s. Multiple releases were made under a number of different varietal names, the biological status of which was, and is still, unclear (O'Connor 1960; Wharton & Gilstrap 1983; Waterhouse 1993). There are a range of colour variants of D. longicaudata on several of the Pacific islands and since this species, and those related to it, have traditionally been separated only by

colour and the presence or absence of striae on tergum 2, this creates a taxonomic problem. All of the variations have been listed here as D. longicaudata, however, further study is required to accurately differentiate these populations and determine their true biological status.

Figure 3.3. Diachasmimorpha longicaudata. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view.

Diachasmimorpha tryoni (Cameron, 1911) Opius tryoni (Cameron 1911): 343. Diachasma tryoni: (Silvestri 1913): 116. Opius (Biosteres) tryoni: (Fischer 1959): 29. Biosteres (Parasteres) acidusae (Fischer 1967b): 3; syn. by (Wharton & Marsh 1978): 157. Biosteres tryoni: (Fischer 1971a): 31. Parasteres tryoni: (Fischer 1978a): 394. Diachasmimorpha tryoni: (Wharton 1987): 62. Taxonomic History This species was originally described as Opius tryoni by Cameron (1911). Silvestri (1913) reviewed the generic classification and placed it in the genus Diachasma [Diachasma tryoni]. Fischer (1959) replaced the species in the genus Opius and placed it in the subgenus Biosteres [Opius (Biosteres) tryoni]. Fischer (1967b) described the same species as Biosteres (Parasteres) acidusae, which was subsequently synonymised by Wharton and Marsh (1978). In 1971a, Fischer gave Biosteres generic ranking, therefore renaming Opius (Biosteres) tryoni as Biosteres tryoni. Fischer (1978a) renamed the species under a new genus, Parasteres tryoni. Wharton (1987) formally removed Diachasmimorpha from Biosteres due to differences in forewing venation and clypeal morphology. Since the type species of Parasteres [acidusae] is a synonym of tryoni (Wharton & Marsh 1978), Parasteres becomes the junior subjective synonym of Diachasmimorpha. This species is now known as Diachasmimorpha tryoni. Synopsis (Figure 3.4a-f) Frons and vertex setose and punctate. Frons polished and weakly punctate, the punctures anteriad the ocellar field sparse and shallow; ocellar triangle arranged to form an obtuse triangle with a slight depression anteriad the median ocellus. Occipital carina weakly developed to almost absent; clypeus in profile slightly bulged medially; ventral margin of clypeus sinuate, ventral margin sufficiently convex to completely conceal labrum when mandibles closed. Notauli deep, unsculptured; propodeum weakly rugose laterally, with a median longitudinal carina stemming from a weakly developed pentagonal areola, the base of which coincides with the posterior margin. Sternaulus broad, shallow, crenulate. Wings strongly infumate. Forewing r arising slightly basad midpoint of stigma, 2RS about equal in length to 3RSa, (RS+M)a weakly sinuate; (RS+M)b absent, m-cu arising distad 2RS. Hindwing RS absent; m-cu well developed and pigmented to wing margin. Petiole

with dorsal lateral carinae strongly elevated over basal half, weaker posteriorly but reaching the posterior margin. Metasomal tergum 2 smooth and polished. Ovipositor sheath about equal in length to body including head. Ovipositor subapically sinuate. Head and mesosoma yellow/orange, petiole and tergum 2 black medially, pale to almost white laterally, remainder of metasoma black dorsally; fore and mid legs yellow/orange except tarsal segments 4-5 brown, hind leg basal half of coxa yellow orange, remainder of leg dark brown. Distribution Australia: Collected along the eastern coast of Australia, from Mossman in the north to Sydney in the south. Type Locality Narara, Australia. Material examined Australia: Coffs Harbour, N.S.W., Rowley Potters Garden, 14.ii.1960, On china pears, T.G. Campbell, (ANIC), 1♂; Coffs Harbour, N.S.W., Thompson's House, 27.ii.1959, on cherry guava, T.G. Campbell, (ASCU), 1♂; Coffs Harbour, N.S.W., 21.ii.1959, ex cherry guava, (ANIC), 2♂, 2♀; Concord, N.S.W., .xii.1958, ex peaches, M.A. Bateman, (ANIC), 5♂, 1♀; Concord, N.S.W., .xii.1958, reared ex peaches, M.A.Bateman, (ASCU), 1♂; CSIRO Orchard, N.Ryde, N.S.W., 14.iv.1955, ex Guava, (ANIC), 2♂, 1♀, 1, sex undetermined; East Palmeston, Queensland, Gregory's Farm, 6.v.1960, T.G. Campbell, (ANIC), 1♀; Freshwater, Queensland, 30.vii.1959, on comquat, T.G. Campbell, (ANIC), 1♂; Gatton, Queensland, 16.i.1953, Pyrus commonis, Strumeta tryoni, A.W.S. May, (ANIC), 1♂, 1♀; Gosford, N.S.W., 25.ii.1981, (ASCU), 1♂; Gosford, N.S.W., 21.iv.1911, Dacus tryoni, W.W. F., (ASCU), 3♂, 1♀; Gregor's Ck, 10.ii.1950, Hemicyclia australasica, Strumeta tryoni, J. Mull, (ANIC), 2♂; Mt. Glorious, Queensland, 1.iii.1953, Rauwenhottia leichardtii, Strumeta halfordiae, A.W.S. May, (ANIC), 1♀; Queensland, Mt. Glorious, 27.x.1991, S.P. Phillips, (QDPIF), 5♂, 10♀; Sydney, N.S.W., .i.1982, on peach, Dacus tryoni, G. Fitt, (ANIC), 1♀; Sydney, N.S.W., 4.i.1934, ex plums, Chaetodacus tryoni, (ASCU), 1♂; Sydney, N.S.W., .ix.1959, Strumeta tryoni, (ASCU), 3♂, 2♀; Mossman, N. Qld, 21.ii.1968, G. Carlirle, (ASCU), 1♂; Narara Res. Stn. Gosford, N.S.W., 25.iii.1959, On Mortons Citrange, T.G. Campbell, (ANIC), 1♀; Narara Res. Stn. Gosford, N.S.W., 25.iii.1959, On Blacks Valencia, T.G. Campbell, (ANIC), 1♂; Narara Res. Stn. Gosford, N.S.W., 25.iii.1959, T.G. Campbell, (ANIC), 1♂, 2♀; Narara Res. Stn. Gosford, N.S.W., 25.iii.1959, On Chapmans Valencia, T.G. Campbell, (ANIC), 4♂; Queensland, Redlands H.R.C., xi.1994, peaches, Bactrocera

tryoni & Bactrocera neohumeralis, H. Lahey, (QDPIF), 1♀; Unknown Australia, (ASCU), 4♂. Hawaii: Kula Maui, 17.viii.1959, reared from peaches, R. Lukins, (ANIC), 12♀. Host Associations B. dorsalis (2), B. halfordiae (Tryon) (1), B. passiflorae (2), B. tryoni (1,2,3,7), B. xanthodes (2), and C. capitata (2,7). Discussion Morphologically similar to D. hageni, D. kraussi and D. longicaudata in having an apically sinuate ovipositor and notauli deep and unsculptured to midpit. Distinguished by dark metasomal colouration, metasomal tergum 2 smooth and polished and having a weakly developed occipital carina. Diachasmimorpha tryoni is native to eastern Australia (Silvestri 1914; Wharton & Gilstrap 1983) and was released, but did not become established, in Fiji, French Polynesia and Western Australia (Waterhouse 1993).

Figure 3.4. Diachasmimorpha tryoni. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view.

Fopius arisanus (Sonan, 1932) Opius arisanus (Sonan 1932): 67-68. Opius oophilus (Fullaway 1951): 248; syn. by (Wharton & Gilstrap 1983): 730. Diachasma arisanum: (Fischer 1967a): 68; (Fischer 1988a): 250. Biosteres arisanus: (Wharton & Gilstrap 1983): 730. Fopius arisanus: (Waterhouse 1993): 41. Taxonomic History Opius arisanus was described by Sonan (1932). Fullaway (1951) described the same species as Opius oophilus which was subsequently synonymised by Wharton & Gilstrap (1983). Fischer (1967a) revised the generic assignment and placed the species in the genus Diachasma [Diachasma arisanum]. Morphological examination by Wharton and Gilstrap (1983) revealed the clypeus of arisanus is of the typical Biosteres form, therefore moving it to the Biosteres genus [Biosteres arisanus]. Wharton (1987) revised the Indo-Pacific species of Biosteres and formed a new genus Fopius, to which this species now belongs [Fopius arisanus] (Waterhouse 1993). Synopsis (Figure 3.5a-f) Frons and vertex densely setose and rugose-punctate. Occipital carina well developed; clypeus in profile not bulging medially, ventral margin almost completely concealing labrum when mandibles closed. Notauli deep and foveolate/crenulate to midpit; propodeum rugose, the sculpture largely without obvious pattern. Sternaulus rugose. Wings weakly infumate. Forewing r arising distad midpoint of stigma, 2RS longer than 3RSa, (RS+M)a weakly sinuate; (RS+M)b sinuate, m-cu arising basad or directly in line with 2RS. Hindwing RS virtually absent, m-cu well developed and at least weakly pigmented to wing margin. Petiole longitudinally striate, dorsal carinae elevated over basal half and reaching posterior margin. Metasomal tergum 2 longitudinally striate. Ovipositor sheath equal to or greater than length of metasoma. Ovipositor narrowed apically, with distinct subapical constriction. Head almost entirely orange-brown, antennae dark; mesosoma varying from orange brown anteriorly, darker brown posteriorly to entirely brown (males commonly darker than females); metasoma dark brown to black. Legs pale yellow, except tarsi brown. Distribution Australia: Collected along the eastern coast of Australia, from Cape Tribulation in the north to Sydney in the south, and Lord Howe Island. Cook Islands (New Zealand): Rarotonga. Fiji: Collected from many sites on Viti Levu and Vanua Levu,

also collected on Mana Island and the islands of the Lau and Yasawa groups. Tonga: Havelotu and Tongatapu. Western Samoa: Savoli and Upolu. Type Locality Funkiko, Arisan, 1400m, Formosa [Taiwan] Material examined American Samoa: Tutuila Malota, 25.viii.1998, mango, L. Ali, (SPC), 1♂. Australia: Atherton, Queensland, 18.iii.1982, on guava, Dacus neohumeralis, G. Fitt, (ANIC), 1♂, 1♀; Brisbane, Queensland, .iii.1981, guava, Dacus tryoni, G. Fitt, (ANIC), 1♀; Cairns, Queensland, 16.iii.1981, guava, Dacus tryoni, G. Fitt, (ANIC), 2♀; Cairns, Queensland, .iii.81, on guava, Dacus tryoni, G. Fitt, (ANIC), 1♂, 1♀; Coffs Harbour, N.S.W., "Orange Trees", 3.iii.1959, T.G. Campbell, (ANIC), 1, sex undetermined; Coffs Harbour, The Orange Trees, Oraro Road, 24.i.1959, captured in flight around ripening mango fruits, T.G. Campbell, (ANIC), 2♀; Coffs Harbour, N.S.W., Watson's house, Watsonia, 2.ii.1959, T.G. Campbell, (ANIC), 1♀; Coffs Harbour, N.S.W., 16.ii.1960, On china pears, T.G. Campbell, (ANIC), 1♀; Coffs Harbour, N.S.W., 14.ii.1960, ex cherry guava, T.G. Campbell, (ANIC), 1♀, Coffs Harbour, N.S.W., 16.ii.1960, ex cherry guava, T.G. Campbell, (ANIC), 1♀; Coffs Harbour, N.S.W., House next to T. Williams Garden, 3.iii.1959, T.G. Campbell, (ANIC), 3♀, 1, sex undetermined; Coffs Harbour, N.S.W., T. Williams Garden, 5.iii.1959, on Mango, T.G. Campbell, (ANIC), 1♂, 2, sex undetermined; Coffs Harbour, N.S.W., 14.ii.1960, ex china pear, T.G. Campbell, (ANIC), 1♂, 1♀, 1, sex undetermined; Coffs Harbour, N.S.W., Rowley's Garden, 14.ii.1960, ex china pear, T.G. Campbell, (ANIC), 2♂, 1♀; Coffs Harbour, N.S.W., T. Williams Garden, 3.iii.1959, T.G. Campbell, (ANIC), 4♂, 8♀; Coffs Harbour, N.S.W., Thompson's Garden, 5.iv.1959, ex cherry guava, T.G. Campbell, (ANIC), 11♂, 2♀, 2, sex undetermined; Currumbin, Queensland, 16.ii.1961, ex mango, G. J. Snowball, (ANIC), 1♂, 2♀; Currumbin, Queensland, .1.1962, Solanum mauritianum, G. J. Snowball, (ANIC), 2♂, 3♀,Duranbah, Queensland, Banana Demo Farm, 7.iv.1959, T.G. Campbell, (ANIC), 1♀; Duranbah, Queensland, Banana Demo Farm, 6.v.1959, on guava, T.G. Campbell, (ANIC), 1♀; Duranbah, Queensland, 21.ii.1960, Solanum mauritianum, T.G. Campbell, (ANIC), 3♂, 1♀; Duranbah, Queensland, 19.ii.1959, ex guava, (ANIC), 3♂, 1♀, 1, sex undetermined; East Palmerston, Queensland, 12.iv.1961, ex wild tobacco sawdust, R.G. Lukins, (ANIC), 3♂, 4♀, 1 sex undetermined; East Palmerston, Queensland, 6.vi.1960, Solanum mauritianum, T.G. Campbell, (ANIC), 2♂, 2♀; Edmonton, Queensland, .i.1962, Averrhoa carambola, G. J. Snowball, (ANIC), 1♂, 1♀; Ellis Beach, Cairns, Queensland, 19.iii.1982, Terminalia catappa, Dacus jarvisi, G. Fitt, (ANIC), 1♂, 1♀; Freshwater, Queensland,

11.vi.1960, Averrhoa carambola, T.G. Campbell, (ANIC), 1♀; Freshwater, Queensland, .i.1962, ex Valencia orange, G.J. Snowball, (ANIC), 2♀; Freshwater, Queensland, 1.vi.1960, Averrhoa carambola, T.G. Campbell, (ANIC), 2♀; Freshwater, Queensland, 17.iv.1961, ex orange, R.G. Lukins, (ANIC), 1♂; Freshwater, Queensland, 17.iv.1961, ex orange, R.G. Lukins, (ASCU), 1♂, 1♀; Freshwater, Queensland, 14.vi.1960, Averrhoa carambola, T.G. Campbell, (ANIC), 1♂; Freshwater, Queensland, 16.iv.1961, ex five corner fig sawdust, R.G. Lukins, (ANIC), 1♂; Gordonvale, Queensland, .i.1962, Averrhoa carambola, G.J. Snowball, (ANIC), 4♂, 2♀; Kairi, Atherton, Queensland, 17.iii.1982, guava, Dacus neohumeralis, G. Fitt, (ANIC), 1♂; Kuranda, Queensland, 14.iv.1961, ex orange, R.G. Lukins, (ANIC), 5♀; Kuranda, Queensland, 14.iv.1961, ex guava, R.G. Lukins, (ANIC), 1♂, 1♀; Kuranda, Queensland, 31.v.1960, Solanum mauritianum, T.G. Campbell, (ANIC), 3♂, 1♀; Lord Howe Island, 5.iii.1961, ex cherry guava, T.G. Campbell, (ANIC), 3♀; Lord Howe Island, 15.iii.1961, ex cherry guava, T.G. Campbell, (ANIC), 4♂, 3♀; Lord Howe Island, 14.iii.1961, ex cherry guava, T.G. Campbell, (ANIC), 1♂, 1♀; Lord Howe Island, 15.iii.1961, ex cherry guava, T.G. Campbell, (ASCU), 1♀; Lord Howe Island, 5.iii.1961, ex cherry guava, T.G. Campbell, (ASCU), 1♂; Mowbray River, Mossman, Queensland, 2.ix.1959, on common lemon, T.G. Campbell, (ANIC), 1 sex undetermined; Murwillumbah, N.S.W., .i.1962, ex mango, G. J. Snowball, (ANIC), 3♂, 5♀; Murwillumbah, N.S.W., .i.1962, ex mango, R.G. Lukins, (ANIC), 6♀, 1, sex undetermined; North Queensland, Cape Tribulation, 29.viii.1996, Niemeyera prunifera, Bactrocera kraussi, QDPI CT1048, (QDPIF), 2♀; North Queensland, Cape Tribulation at RFCT 30, 17.x.1996, Endiandra microneura, QDPI CT1397, (QDPIF), 4♀; North Queensland, Cape Tribulation at RFCT11, 24.x.1996, Endiandra microneura, QDPI CT1437, (QDPIF), 1♂, 2♀; North Queensland, Cape Tribulation at RFCT20, 2.iv.1996, Acmena sp, Bactrocera kraussi, QDPI CT483, (QDPIF), 1♀; North Queensland, Goldsborough Valley at RFGV3, 23.i.1996, Barringtonia calyptrata, Bactrocera barringtoniae, QDPI GV13, (QDPIF), 2♂, 5♀; North Queensland, Goldsborough Valley at RFGV11, 1.vi.1996, Neolitsea dealbata, Bactrocera aberrans, QDPI GV250, (QDPIF), 1♀; North Queensland, Goldsborough Valley at RFGV7, 27.v.1997, Strychnos minor, Bactrocera manskii, QDPI GV401, (QDPIF), 5♀; North Queensland, Lake Morris at RFLM20, 21.v.1996, Solanum mauritianum, Bactrocera cacuminata, QDPI LM1200, (QDPIF), 1♀; North Queensland, Lake Morris at RFLM 25, 21.x.1996, Solanum mauritianum, Bactrocera cacuminata, QDPI LM1354, (QDPIF), 3♂, 9♀; North Queensland, Lake Morris at RFLM28, 23.iv.1996, Solanum mauritianum, Bactrocera cacuminata, QDPI LM951, (QDPIF), 2♀; North

Queensland, Mt. Lewis at RFML33, 13.ii.1996, Solanum, mauritianum Bactrocera cacuminata, QDPI ML108, (QDPIF), 4♀; North Queensland, Mulgrave River at RF MR3, 23.i.1996, Barringtonia calyptrata, Bactrocera barringtoniae, QDPI MR10, (QDPIF), 2♂, 7♀; North Queensland, Mulgrave River at RFMR5, 23.i.1996, Barringtonia calyptrata, Bactrocera barringtoniae, QDPI MR21, (QDPIF), 7♀; North Queensland, Mulgrave River at RFMR8, 23.i.1996, Cryptocarya sp, Bactrocera endiandrae, QDPI MR29A, (QDPIF), 2♂, 3♀; North Queensland, Kamerunga Res. Stn., Cairns, 30.xii.1995, Casimiroa edulis, Bactrocera neohumeralis & Bactrocera tryoni, QDPI NQ 338, (QDPIF), 2♀; North Queensland, Mareeba, 12.v.1996, Averrhoa carambola, Bactrocera tryoni, QDPI NQ1139, (QDPIF), 2♀; North Queensland, Kamerunga Res. Stn., Cairns, 17.xi.1996, Syzygium jambos, Bactrocera tryoni & Bactrocera neohumeralis, QDPI NQ117, (QDPIF), 2♀; North Queensland, Mossman, 16.vi.1996, Averrhoa carambola, Bactrocera tryoni, QDPI NQ1473, (QDPIF), 4♀; North Queensland, Bot. Gardens, Cairns, 19.xii.1995, Barringtonia calyptrata, Bactrocera barringtoniae, QDPI NQ244, (QDPIF), 1♀; North Queensland, Kamerunga Res. Station, Cairns, 9.i.1996, Manilkara achras, Bactrocera tryoni, Bactrocera papayae & Bactrocera neohumeralis, QDPI NQ397, (QDPIF), 1♀; North Queensland, Kamerunga Res. Station, Cairns, 1.ii.1996, Psidium guajava, Bactrocera kraussi, Bactrocera tryoni & Bactrocera jarvisi, QDPI NQ457, (QDPIF), 1♂, 5♀; North Queensland, Kuranda Downs, 21.ii.1996, Musa sp, Bactrocera jarvisi, Bactrocera musae & Bactrocera papayae, QDPI NQ495, (QDPIF), 2♂, 6♀; North Queensland, Kamerunga Res. Station, Cairns, 4.xii.1995, Averrhoa carambola, Bactrocera tryoni, QDPI NQ60, (QDPIF), 3♂, 3♀; North Queensland, Kamerunga Res. Station, Cairns, 9.xi.1996, Mangifera, indica, Bactrocera tryoni & Bacterocera neohumeralis, QDPI NQ68, (QDPIF), 2♂, 4♀; North Queensland, Edmonton, 20.iii.1996, Carica papaya, Bactrocera neohumeralis & Bactrocera tryoni, QDPI NQ662, (QDPIF), 2♂; North Queensland, Julatten, 27.iii.1996, custard apple, Bactrocera neohumeralis, QDPI NQ691C, (QDPIF), 3♀; North Queensland, Cairns, 27.iv.1996, Psidium guajava, Bactrocera calophylli, Bactrocera tryoni & Bactrocera neohumeralis, QDPI NQ917, (QDPIF), 1♂, 2♀; North Ryde, N.S.W., 19.ii.1957, ex peaches, G.J. Snowball, (ANIC), 3♂, 1♀; Pinjin Hill, near Innisfail, Queensland, 16.vi.1960, T.G. Campbell, (ANIC), 1♀; Pinjin Hill, near Innisfail, Queensland, 16.vi.1960, Solanum mauritianum, T.G. Campbell, (ANIC), 1♂, 2♀, 1, sex undetermined; Port Macquarie, N.S.W., .xii.1963, wild tobacco (contaminant?), R.G. Lukins, (ANIC), 1, sex undetermined; Queensland, 26.i.1978, Mango, Dacus spp., G.J. Baker, (ANIC), 3♀; Queensland, Nambour, 16.xii.1992, Sapote, Q20, M.M. Elson-Harris, (QDPIF), 7♂, 3♀; Queensland, Nambour,

14.i.1993, Acerola, Q23, M.M. Elson-Harris, (QDPIF), 6♂, 12♀; Queensland, Nambour, 14.i.1993, Wax Jambu, Q27, M.M. Elson-Harris, (QDPIF), 1♂, 6♀; Queensland, Nambour, 2.ii.1993, Carambola, Bactrocera tryoni & Bactrocera neohumeralis, Q64, M.M. Elson-Harris, (QDPIF), 27, sex undetermined; Queensland, Nambour, ii.1993, Feijoa, Bactrocera neohumeralis & Bactrocera tryoni, Q67, M.M. Elson-Harris, (QDPIF), 8♂, 26♀, 1, sex undetermined; Queensland, Downey Creek, 15.xi.1982, Pulogium galactoxyla, Bactrocera endiandrae, M.M. Elson-Harris, (QDPIF), 1♀; Queensland, Nambour, 18.iii.1993, Carambola, Q99, M.M. Elson-Harris, (QDPIF), 5♂; Sydney, N.S.W., 11.ii.1957, lab reared, Strumeta tryoni, (ANIC), 1♀, Queensland, Redlands H.R.C., xi.1994, peaches, Bactrocera tryoni & Bactrocera neohumeralis, H. Lahey, (QDPIF), 6♂, 12♀; Sydney, N.S.W., 1.vii.1950, G.J. Snowball, (ANIC), 2♀; Sydney, N.S.W., 11.xii.1957, lab reared, Strumeta tryoni, (ANIC), 1♂; Sydney, N.S.W., Exp No. 14, 4.ii.1952, F. Wilson, (ANIC), 2♂, 4♀;Sydney, N.S.W., Botanic Gardens, 6.iii.1958, collected from peach tree, G. J. Snowball, (ANIC), 3♂, 1♀, 1, sex undetermined; Sydney, N.S.W., 4.v.1959, Bactrocera tryoni, (ASCU), 1♂, 2♀; Trinity Beach, Cairns, Queensland, 18.iii.1982, Terminalia catappa, Dacus jarvisi, G. Fitt, (ANIC), 2♂, 1♀; Upper Currumbin, 23.ii.1960, Solanum mauritianum, T.G. Campbell, (ANIC), 1♂, 3♀; W. Gosford, N.S.W., 25.xi.1958, ex plums, T.G. Campbell, (ANIC), 2♂, 2♀; Yarrahappini, N.S.W., 6.iii.1959, on pear, T.G. Campbell, (ANIC), 1♀; Yarrahappini, N.S.W., near Kempsey, 6.iii.1959, on Mango, T.G. Campbell, (ANIC), 1♀, 1, sex undetermined; Yarrahappini, N.S.W., 10.ii.1960, ex peach, T.G. Campbell, (ANIC), 1♂, 1♀; Yarrahappini, N.S.W., 10.ii.1960, On china pears, T.G. Campbell, (ANIC), 2♂, 1♀; Yarrahappini, N.S.W., near Kempsey, 8.iii.1959, ex mango, T.G. Campbell, (ANIC), 3♂, 3♀, 2, sex undetermined; Yarrahappini, N.S.W., near Kempsey, 16.ii.1960, On china pears, T.G. Campbell, (ANIC), 4♂; Yarrahappini, N.S.W., near Kempsey, 10.iii.1959, ex mango, T.G. Campbell, (ANIC), 4♂, 4♀, 1, sex undetermined. Cook Islands: RaroTitikaveka, 12.iii.1991, Psidium guajava, C155, Abdul, 6♂, 12♀; RaroNikao, 14.iii.1991, Psidium guajava, Bactrocera melanotus, C163, Abdul Kassim, 8♂, 3♀; RaroNikao, 18.iii.1991, Psidium guajava, Bactrocera melanotus, C175, Abdul, 1♂, 3♀, 1, sex undetermined; RaroNikao, 18.iii.1991, Inocarpus fagiferus, Bactrocera melanotus, C177, Abdul, 1♂; RaroNgatangiia, 20.iii.1991, Psidium guajava, Bactrocera melanotus, C215, Abdul, 1♀; RaroTitikaveka, 11.iv.1991, Artocarpus altilis, Bactrocera melanotus & Bactrocera xanthodes, C309, Abdul, 1♂; RaroAtupa, 3.v.1991, Psidium guajava, Bactrocera melanotus, C352, Abdul, 4♂, 5♀; RaroT.R.S. Forest, 28.v.1991, Terminalia catappa, Bactrocera

melanotus, C424, Abdul, 5♂, 5♀; RaroT.R.S., 28.v.1991, Terminalia catappa, Bactrocera melanotus, C425, Abdul, 1♂, 2♀; RaroT.R.S., 28.v.1991, Terminalia catappa, Bactrocera melanotus, C426, Abdul, 3♀, 3, sex undetermined; Raro11.vi.1991, Inocarpus, fagiferus, Bactrocera melanotus, C461, Abdul Kassim, 1♂, 1♀; Raro12.vi.1991, Terminalia catappa, Bactrocera melanotus, C483, Abdul Kassim, 1♂; Raro16.vi.1991, Terminalia catappa, Bactrocera melanotus, C495, Abdul Kassim, 7♂, 12♀; Raro27.vi.1991, Artocarpus altilis, Bactrocera xanthodes, C504, Abdul Kassim, 6♂, 2♀; Raro2.vii.1991, Artocarpus altilis, Bactrocera xanthodes, C519, Abdul Kassim, 5♂, 3♀; Raro2.vii.1991, Artocarpus altilis, Bactrocera xanthodes, C520, Abdul Kassim, 4♂, 11♀; Raro2.vii.1991, Artocarpus altilis, Bactrocera xanthodes, C522, Abdul Kassim, 1♂; Raro2.vii.1991, Artocarpus altilis, Bactrocera xanthodes, C523, Abdul Kassim, 4♀; Nadi, Legalega R.S., 11.i.1992, Psidium guajava, Bactrocera passiflorae, F0163, A. Allwood & G. Walter, 19♂, 41♀; Nadi, Legalega R.S., 25.i.1991, Psidium guajava, F0237, G. Walker & Sada Nand Lal, 1♀; Nadi, Legalega R.S., 25.i.1991, Psidium guajava, Bactrocera passiflorae, F0244, G. Walker & Sada Nand Lal, 2♀; Nadi, Legalega R.S., 18.ii.1991, Psidium guajava, Bactrocera passiflorae, F0360, A. Allwood et al, 1♂; Nadi, Legalega R.S., 18.ii.1991, Psidium guajava, Bactrocera passiflorae, F0361, A. Allwood et al, 16♂, 11♀; Nadi, Legalega R.S., 18.ii.1991, Psidium guajava, F0363, A. Allwood et al, 3♂; Nadi, Legalega R.S., 18.ii.1991, Psidium guajava, Bactrocera passiflorae, F0364, A. Allwood & G. Walter, 1♂, 1♀; Nadi, Legalega R.S., 18.ii.1991, Psidium guajava, Bactrocera passiflorae, F0365, A. Allwood & G. Walter, 1♀; Nadi, Legalega R.S., 18.ii.1991, Psidium guajava, Bactrocera passiflorae, F0366, A. Allwood & G. Walter, 1♂, 1♀; Nadi, Legalega R.S., 18.ii.1991, Psidium guajava, F0367, A. Allwood & G. Walter, 1♀; Rakiraki, Prasad's, 19.ii.1991, Inocarpus fagiferus, Bactrocera passiflorae, F0398, A. Allwood & G. Walter, 1♂, 8♀; Nadi, Legalega R.S., 15.xi.1990, Psidium guajava, Bactrocera passiflorae, F049, G. Walker et al, 1♀; Naduruloulou R.S., 22.iii.1991, Psidium guajava, F0508, G. Walker, 1♂, 3♀; Rakiraki, Lal's, 23.iv.1991, Psidium guajava, Bactrocera passiflorae, F0675, G. Walker & E. Hamacek, 2♀; Rakiraki, Lal's, 23.iv.1991, Fortunella japonica, Bactrocera passiflorae, F0676, G. Walker & E. Hamacek, 1♀; Nadi, Garden of the Sleeping Giant, 24.iv.1991, Psidium guajava, Bactrocera passiflorae, F0710, E. Hamacek & G. Walker, 1♂, 1♀; Nadi, Legalega R.S., 12.xii.1990, Psidium guajava, F082, G. Walker et al, 1♂, 1♀; Waimara, Rakiraki, 11.vi.1991, Citrus reticulata, Bactrocera passiflorae, F0889, P. Prassad & G. Walker, 3♀; Nausori, Naduruloulou R.S., 10.vii.1991, Citrus maxima, Bactrocera passiflorae, F1002, Ema Tora et al, 3♀; Nausori, Naduruloulou R.S., 10.vii.1991,

Fortunella japonica, Bactrocera passiflorae, F1006, Ema Tora et al, 1♂, 2♀; Rakiraki, Bari Lal, 23.iv.1991, Fortunella japonica, Bactrocera passiflorae, F1095, G. Walker et al, 1♂, 3♀; Saheed, Lautoka, 24.vii.1991, Citrus paradisi, Bactrocera passiflorae, F1124, G. Walker et al, 1♂, 8♀; Nausori, Naduruloulou R.S., 15.viii.1991, Fortunella japonica, Bactrocera passiflorae, F1205, G. Walker et al, 10♀; Labasa, Seaqaqa R.S. Vanua Levu, 15.viii.1991, Bactrocera passiflorae, F1222, G. Walker et al, 2♂, 1♀; Nausori, Naduruloulou R.S., 8.i.1991, Psidium guajava, Bactrocera passiflorae, F131, G. Walker et al, 14♂, 26♀, 1, sex undetermined; Nausori, Naduruloulou R.S., 8.i.1991, Syzygium malaccense, Bactrocera passiflorae, F138, A. Allwood et al, 6♂, 6♀; Nadi, Matasawaku, 26.xi.1991, Syzygium, jambos, Bactrocera passiflorae, F1682, M. Atur, 20♂, 19♀; Rakiraki, Sat Deo, 27.xi.1991, Artocarpus altilis, Bactrocera passiflorae & Bactrocera xanthodes, F1755, A. Allwood et al, 1♀; 3km W of Tagavere, 28.xi.1991, Inocarpus fagiferus, Bactrocera passiflorae, F1794, A. Allwood et al, 3♀; Nadi, Legalega R.S., 19.xii.1991, Syzygium jambos, Bactrocera passiflorae, F1880, A. Allwood et al, 1♂; Nausori, Naduruloulou R.S., 22.i.1992, Psidium guajava, Bactrocera passiflorae, F1956, A. Allwood et al, 1♂; Nausori, Naduruloulou R.S., 22.i.1992, Psidium guajava, Bactrocera passiflorae, F1966, A. Allwood et al, 1♂, 4♀; Nadi, Legalega R.S., 29.i.1992, Manilkara achras, F2059, G. Walker et al, 6♂, 3♀; Labasa, Seaqaqa R.S. Vanua Levu, 30.i.1992, Psidium guajava, Bactrocera passiflorae, F2122, A. Allwood et al, 8♂, 2♀; Labasa, Seaqaqa R.S. Vanua Levu, 30.i.1992, Syzygium jambos, Bactrocera passiflorae, F2128, A. Allwood et al, 1♀; Nausori, Naduruloulou R.S., 6.ii.1992, Syzygium malaccense, Bactrocera passiflorae, F2153, G. Walker et al, 1♀, 1, sex undetermined; Nausori, Naduruloulou R.S., 6.ii.1992, Psidium guajava, Bactrocera passiflorae, F2154, G. Walker et al, 2♀; Nausori, Naduruloulou R.S., 19.ii.1992, Garcinia mangostana, Bactrocera passiflorae, F2185, G. Walker et al, 5♂, 11♀; Saheed, Lautoka, 25.ii.1992, Psidium guajava, Bactrocera passiflorae, F2270, G. Walker et al, 3♂, 1♀; Nadi, Legalega R.S., 26.ii.1992, Psidium guajava, Bactrocera passiflorae, F2289, G. Walker et al, 1♂, 2♀; Nadi, Legalega R.S., 26.ii.1992, Psidium guajava, F2293, G. Walker et al, 1♀; Sigatoka, Sigatoka R.S., 26.ii.1992, Psidium guajava, Bactrocera passiflorae, F2305, G. Walker et al, 2♂, 1♀; Sigatoka, Sigatoka R.S., 26.ii.1992, Psidium guajava, Bactrocera passiflorae, F2310, G. Walker et al, 7♂, 10♀; Nausori, Naduruloulou R.S., 4.iii.1992, Psidium guajava, F2345, G. Walker et al, 1♂; Nausori, Naduruloulou R.S., 4.iii.1992, Psidium guajava, Bactrocera passiflorae, F2346, G. Walker et al, 1♂; Nausori, Naduruloulou R.S., 18.iii.1992, Psidium guajava, Bactrocera passiflorae, F2371, G. Walker et al, 1♂, 2♀; Nadi, Legalega R.S.,

25.i.1991, Psidium guajava, Bactrocera passiflorae, F241, G. Walker et al, 3♂, 5♀; Sigatoka Valley, 1st Deo, 1.iv.1992, Citrus reticulata, Bactrocera passiflorae, F2435, G. Walker et al, 1♂; 34km Nagali on Monosavu Rd, 6.iv.1992, Psidium guajava, Bactrocera passiflorae, F2498, A. Allwood et al, 2♀; Mana Is., 9.iv.1992, Terminalia catappa, Bactrocera passiflorae, F2568, G. Walker et al, 1♀; Nausori, Naduruloulou R.S., 22.iv.1992, Fortunella japonica, Bactrocera passiflorae, F2611, Ema Tora et al, 2♂; Nausori, Naduruloulou R.S., 22.iv.1992, Citrus maxima, Bactrocera passiflorae, F2616, Ema Tora et al, 1♂; Nausori, Naduruloulou R.S., 22.vi.1992, Psidium guajava, Bactrocera passiflorae, F2617, Ema Tora et al, 1♂; Nausori, Naduruloulou R.S., 22.vi.1992, Psidium guajava, Bactrocera passiflorae, F2618, Ema Tora et al, 3♂, 1♀; Nausori, Toga Village, 21.iv.1992, Averrhoa carambola, Bactrocera passiflorae, F2624, Ema Tora et al, 1♂, 2♀; Waimara R.S., 21.iv.1992, Citrus paradisi, Bactrocera passiflorae, F2631, G. Walker et al, 3♀; Rakiraki, A. Prasad, 21.iv.1992, Artocarpus altilis, Bactrocera xanthodes, F2665, G. Walker et al, 2♂, 5♀; 6km W of Tavika, 21.iv.1992, Inocarpus fagiferus, Bactrocera passiflorae, F2669, G. Walker et al, 7♂, 12♀; Ba, Pineapple grower, 22.iv.1992, Citrus sinensis, Bactrocera passiflorae, F2674, G. Walker et al, 4♂, 6♀; Ba, Pineapple grower, 21.vi.1992, Citrus paradisi, Bactrocera passiflorae, F2677, G. Walker et al, 2♂, 3♀; Saheed, Lautoka, 22.iv.1992, Fortunella, japonica, Bactrocera passiflorae, F2689, G. Walker et al, 1♂, 7♀; Saheed, Lautoka, 22.iv.1992, Annona, muricata, Bactrocera passiflorae, F2690, G. Walker et al, 2♂; Saheed, Lautoka, 22.iv.1992, Citrus sinensis, Bactrocera passiflorae, F2691, G. Walker et al, 3♂, 3♀; Orchard, Sigatoka R.S., 23.iv.1992, Citrus paradisi, Bactrocera passiflorae, F2713, G. Walker et al, 1♂, 4♀; Orchard, Sigatoka R.S., 23.iv.1992, Psidium guajava, Bactrocera passiflorae, F2715, G. Walker et al, 2♂; Nausori, Naduruloulou R.S., 4.v.1992, Fortunella japonica, Bactrocera passiflorae, F2746, G. Walker et al, 1♂, 2♀; Nausori, Naduruloulou R.S., 4.v.1992, Citrus maxima, Bactrocera passiflorae, F2747, G. Walker et al, 5♂, 3♀; Sigatoka R.S., 7.v.1992, Psidium guajava, Bactrocera passiflorae, F2751, G. Walker et al, 2♀; Nausori, Naduruloulou R.S., 21.v.1992, Fortunella japonica, Bactrocera passiflorae, F2781, G. Walker et al, 3♀; Nausori, Naduruloulou R.S., 11.vi.1992, Citrus reticulata, Bactrocera passiflorae, F2824, G. Walker et al, 3♀; Waimara, Rakiraki, 26.v.1992, Citrus sinensis, Bactrocera passiflorae, F2842, G. Walker et al, 2♂; Ba, Pineapple grower, 27.v.1992, Citrus sinensis, Bactrocera passiflorae, F2851, G. Walker et al, 1♂; Ba, Pineapple grower, 27.v.1992, Citrus reticulata, Bactrocera passiflorae, F2854, G. Walker et al, 2♂, 1♀; Saheed, Lautoka, 27.v.1992, Citrus paradisi, Bactrocera passiflorae, F2868, G. Walker et al, 1♂, 1♀; Colo-i-Suva Forest Park, 11.vi.1992,

Amaroria soulameiodes, Bactrocera passiflorae, F2876, G. Walker et al, 9♂, 2♀; Sigatoka, Sigatoka R.S. Orchard, 28.v.1992, Citrus reticulata, Bactrocera passiflorae, F2881, G. Walker et al, 1♂; Anil Kumar, Sigatoka Valley, 28.v.1992, Citrus sinensis, Bactrocera passiflorae, F2893, G. Walker et al, 1♀; Nausori, Naduruloulou R.S., 11.vi.1992, Psidium guajava, Bactrocera passiflorae, F2908, G. Walker et al, 1♂, 1♀; Pacific Harbour, Lomaloma Beach, 14.vi.1992, Bactrocera passiflorae, F2934, G. Walker et al, 8♀; Rakiraki Hotel, 28.vi.1992, Citrus paradisi, Bactrocera passiflorae, F2967, A. Allwood, 3♀; Ba, Pineapple grower, 29.vi.1992, Citrus sinensis, Bactrocera passiflorae, F2996, A. Allwood, 1♀; Saheed, Lautoka, 29.vi.1992, Fortunella japonica, Bactrocera passiflorae, F3001, A. Allwood, 2♂, 3♀; Saheed, Lautoka, 29.vi.1992, Citrus paradisi, Bactrocera passiflorae, F3004, A. Allwood, 2♂, 2♀; Nadi, Legalega R.S., 30.vi.1992, Syzygium jambos, Bactrocera passiflorae, F3009, A. Allwood, 1♀; Sigatoka R.S. Orchard, 30.vi.1992, Citrus reticulata, Bactrocera passiflorae, F3036, A. Allwood, 2♀; Nausori, Naduruloulou R.S., 9.vii.1992, Citrus maxima, Bactrocera passiflorae, F3088, G. Walker et al, 1♀; Colo-i-Suva Forest Park, 9.vii.1992, Amaroria soulameiodes, Bactrocera passiflorae, F3089, G. Walker et al, 12♂, 12♀; Nausori, Naduruloulou R.S., 23.vii.1992, Artocarpus altilis, Bactrocera xanthodes, F3143, G. Walker et al, 1♂, 2♀; Nausori, Naduruloulou R.S., 23.vii.1992, Citrus maxima, Bactrocera passiflorae, F3150, G. Walker et al, 1♂, 3♀; Saheed, Lautoka, 5.viii.1992, Citrus reticulata, Bactrocera passiflorae, F3228, G. Walker et al, 1♀; Nausori, Naduruloulou R.S., 3.ix.1992, Annona muricata, Bactrocera passiflorae, F3283, G. Walker et al, 1♂; Vanua Levu, Seaqaqa R.S., 11.ix.1992, Fortunella japonica, Bactrocera passiflorae, F3304, A. Allwood et al, 1♂, 5♀; Malevu, Reef Resort, 25.x.1992, Ochrosia oppositifolia, Sp., nov. 2, F3401, G. Walker, 8♂, 6♀. Tonga: Tongatapu. Fasi, 5.vi.2000, Terminalia catappa, SPC, (SPC), 1♂, 1♀; Tongatapu, Dateline Hotel, 21.iii.1991, Terminalia catappa, Bactrocera facialis & Bactrocera kirki, T0129, Pulotu, (QDPIF), 3♀; Tongatapu, Maufanga, 7.v.1991, Terminalia catappa, Bactrocera facialis & Bactrocera kirki, T0212, Pontiano, (QDPIF), 2♂, 7♀; Tongatapu, Dateline Hotel, 20.v.1991, Terminalia catappa, Bactrocera facialis & Bactrocera kirki, T0270, A. Allwood, (QDPIF), 16♂, 28♀; Tongatapu, Fasi, 21.xi.1991, Syzygium malaccense, Bactrocera facialis, T0877, Pontiano, (QDPIF), 5♂, 1♀; Tongatapu, Fasi, 21.xi.1991, Calophyllum inophyllum, Bactrocera facialis, T0894, Pontiano, (QDPIF), 4♂, 3♀; Tongatapu, Fasi, 21.xi.1991, Eugenia uniflora, Bactrocera facialis & Bactrocera kirki, T0895, Pontiano, (QDPIF), 2♂, 6♀; Tongatapu, Fasi, 27.xii.1992, Eugenia uniflora, T1079, Pontiano, (QDPIF), 1♂, 4♀; Tongatapu, Kolofo'ou, 28.i.1992, Terminalia catappa, Bactrocera facialis &

Bactrocera kirki, T1195, Sultoni, (QDPIF), 3♂, 7♀; Tongatapu, Kolofo'ou, 18.iii.1992, Terminalia catappa, Bactrocera facialis & Bactrocera kirki, T1274, Sultoni, (QDPIF), 5♂, 13♀; Tongatapu, Fasi, 23.viii.1992, Calophyllum, inophyllum, Bactrocera facialis, T1846, Pulotu, (QDPIF), 1♀; Tongatapu, Q. Salote Coll., 23.vii.1992, Phaleria disperma, Bactrocera facialis, T1851, Pulotu, (QDPIF), 15♀; Tongatapu, Q. Salote Coll., 23.vii.1992, Phaleria disperma, Bactrocera facialis, T1853, Pulotu, (QDPIF), 4♂, 8♀; Tongatapu, Maufanga, 23.ix.1992, Phaleria disperma, Bactrocera facialis, T1999, Una & Tali, (QDPIF), 4♂, 8♀; Haveloutu, 8.x.1992, Manilkara achrea, Bactrocera distincta, T2021, Una, (QDPIF), 1♂; Tongatapu, Fasi, 21.xi.1992, Phaleria disperma, Bactrocera facialis, T2090, Tali, (QDPIF), 3♂, 2♀; Tongatapu, Fasi, 21.xi.1992, Phaleria disperma, T2091, Tali, (QDPIF), 2♂, 4♀; Tongatapu, Basilica Kolofo'ou, 21.x.1992, Phaleria disperma, Bactrocera facialis, T2095, Tali, (QDPIF), 2♂, 6♀; Tongatapu, Q. Salote Coll., 21.x.1992, Phaleria disperma, Bactrocera facialis, T2096, Tali, (QDPIF), 4♂, 14♀; Tongatapu, Maufanga, 15.xii.1992, Syzygium jambos, Bactrocera facialis & Bactrocera kirki, T2121, Tali & Sultoni, (QDPIF), 9♂, 11♀; Tongatapu, Fasi, 31.xii.1992, Terminalia catappa, Bactrocera facialis & Bactrocera kirki, T2148, Sultoni, (QDPIF), 15♂, 9♀. Western Samoa: Nu'u Station, 15.v.2000, Terminalia catappa, SPC, (SPC), 2♀; Upolu, Tuanaimato, 1.x.1991, Bactrocera distincta, WS0221, B. Enosa et al, (QDPIF), 1♂, 4♀; Savoli, Aopo, 22.x.1991, Artocarpus altilis, Bactrocera xanthodes, WS0401, B. Enosa et al, (QDPIF), 1♂; Savoli, Asau, 22.x.1991, Syzygium malaccense, Bactrocera kirki, WS0421, B. Enosa et al, (QDPIF), 2♂, 1♀; Upolu, Alafua, 22.xi.1991, Syzygium jambos, Bactrocera kirki, WS0472, B. Enosa et al, (QDPIF), 1♂, 6♀; Upolu, Apia, 22.xi.1991, Syzygium aquem, Bactrocera kirki, WS0474, B. Enosa et al, (QDPIF), 4♂, 8♀; Savoli, Asau, 5.xii.1991, Syzygium malaccense, Bactrocera kirki, WS0538, B. Enosa et al, (QDPIF), 1♀; Savoli, Asau, 22.x.1991, Syzygium malaccense, Bactrocera kirki, WSO420, B. Enosa et al,

(QDPIF), 4♂, 1♀. Host Associations B. aberrans (Hardy) (1), B. barringtoniae (1,4), B. cacuminata (1,2,4), B. distincta (Malloch) (1), B. dorsalis (2,3,4) , B. endiandrae (Perkins & May) (1), B. facialis (Coquillett) (1), B. jarvisi (1,4), B. kirki (Froggatt) (1), B. kraussi (1,4), B. latifrons (2,4), B. manskii (Perkins & May) (1), B. melanotus (Coquillett) (1), B. neohumeralis (1,4), B. passiflorae (1,2,4), B. tryoni (1), B. xanthodes (1), and C. capitata (2). Discussion Morphologically similar to F. illusorius in its dark colouration and sculpture of the notauli, however, it can be readily distinguished in having dense longitudinal

striations on tergum 2. Fopius arisanus (as Opius oophilus) was introduced to Australia from Hawaii, for the control of B. tryoni, in 1956-57 and again in 1958-59, with the first liberation not thought to have established (Waterhouse 1993). Fopius arisanus has also been introduced to Fiji (1951 and 1954). Although there is no record of its introduction there, F. arisanus has been recovered on Cook Is (Waterhouse 1993).

Figure 3.5. Fopius arisanus. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view.

Fopius deeralensis (Fullaway, 1950) Opius deeralensis (Fullaway 1950a): 65. Opius (Biosteres) deeralensis: (Fischer 1963b): 230. Biosteres deeralensis: (Fischer 1971a): 23. Biosteres (Chilotrichia) deeralensis: (Fischer 1978a): 386. Fopius deeralensis: (Waterhouse 1993): 41. Taxonomic History This species was originally described as Opius deeralensis by Fullaway (1950a). Fischer (1963b) placed this species in the subgenus Biosteres [Opius (Biosteres) deeralensis]. Fischer (1971a) reviewed the subgeneric ranking and elevated Biosteres to a generic position [Biosteres deeralensis]. After further review, Fischer (1978a) reallocated this species into the subgenus Chilotrichia, therefore naming the species Biosteres (Chilotrichia) deeralensis. Wharton (1987) revised the Indo-Pacific species of Biosteres and formed a new genus Fopius, to which this species belongs [Fopius deeralensis] (Waterhouse 1993). Synopsis (Figure 3.6a-f) Frons and vertex densely setose; frons densely punctate, vertex more sparsely and weakly punctate. Frons with polished, weakly elevated projection extending from the median ocellus at least half the distance to torulus. Ocellar triangle arranged to form an obtuse triangle with a slight depression anteriad the median ocellus. Occipital carina well developed; clypeus with anterior margin very obtusely angulate, forming an indistinct median tooth. Notauli foveolate/crenulate to midpit; propodeum irregularly areolate with coarse rugose sculpture; median carina distinct on basal 0.3-0.4. Sternaulus broad, deep, crenulate. Wings infumate. Forewing r arising from or slightly distad midpoint of stigma, 2RS longer than 3RSa, (RS+M)a weakly sinuate; (RS+M)b varying from short to absent, m-cu arising basad or directly in line with 2RS. Hindwing RS absent; m-cu well developed and pigmented to wing margin. Petiole longitudinally striate, dorsal lateral carinae strongly elevated over basal two- thirds, converging posteriorly and extending to posterior margin. Metasomal tergum 2 sometimes smooth and laterally setose, frequently with weak longitudinal striae especially anteromedially. Ovipositor sheath equal to or longer than length of body including head. Whole body yellow/orange except hind tibiae and tarsi dark brown. Distribution Australia: Collected in north eastern Australia, from Stephen Island in the north to Brisbane in the south. Papua New Guinea: New Britain (but see discussion).

Type Locality Deeral, Australia Material examined Australia: Freshwater, Queensland, i.1962, ex Valencia orange, G.J. Snowball, (ANIC), 2 sex undetermined; Innisfail, Queensland, 11.v.1960, ex yellow guava, T. G. Campbell, (ANIC), 1♀; East Palmerston, Queensland, 11.vi.1961, ex guava, R.G. Lukins, (ANIC), 3♀; Cairns, Queensland, 6.viii.1951, Endiandra discolor, Strumeta endiandrae, A.W.S. May, (ANIC), 1♂; Brisbane, Queensland, iii.1981, Psidium guajava, Dacus tryoni, G. Fitt, (ANIC), 1♂; Innisfail, Queensland, 11.v.1960, ex yellow guava, T.G. Campbell, (ANIC), 1♂; Maryborough, Queensland, 15.iv.1959, ex Guava, (ANIC), 1♂; Malanda Falls, Queensland, 25.xi.1980, Aglaia ferruginea, Dacus aglaiae, G. Fitt, (ANIC), 1♂, 1♀; Atherton Tableland, Queensland, vi.1949, N.L.H. Krauss, (ANIC), 1♂, 1♀; Cairns, Queensland, 7.viii.1951, Endiandra discolor, Strumeta endiandrae, A.W.S. May, (ANIC), 1♂, 2♀; Cairns, Queensland, ii.1962, Nauclea orientalis, G.J. Snowball, (ANIC), 1♂, 3♀, 1, sex undetermined; Freshwater, Queensland, 30.vii.1959, on comquat, T. G. Campbell, (ANIC), 2♂, 1♀; Ingham, Queensland, 21.vi.1959, ex Guava, (ANIC), 2♂, 1♀; Daintree, North Queensland, 25.vii.1959, T. G. Campbell, (ANIC), 5♂; Cairns, Queensland, .ii.1962, Nauclea orientalis, G.J. Snowball, (ASCU), 1♀; Deeral, near, .vi.1949, Acmena macrocarpa, N.L.H. Krauss, (ASCU), 1♂; North Queensland, Babinda Boulders at RFBB4, 7.v.1996, Fagraea cambagei, Bactrocera fagraea & Bactrocera musae, QDPI BB142, (QDPIF), 2♀; North Queensland, Cape Tribulation at RFCT 2, 20.iii.1996, Bactrocera kraussi & Bactrocera manskii, QDPI CT271, (QDPIF), 1♂; North Queensland, Goldsborough Valley at RFGV3, 23.i.1996, Barringtonia calyptrata, Bactrocera barringtoniae, QDPI GV13, (QDPIF), 4♀; North Queensland, Goldsborough Valley at RFGV7, 23.i.1996, Aglaia sapindina, Bactrocera aglaiae & Bactrocera tigrina, QDPI GV35, (QDPIF), 2♀; North Queensland, Goldsborough Valley at RFGV10, 23.i.1996, Aglaia sapindina, Bactrocera aglaiae, QDPI GV49, (QDPIF), 1♀; North Queensland, Mulgrave River at RF MR3, 23.i.1996, Barringtonia calyptrata, Bactrocera barringtoniae, QDPI MR10, (QDPIF), 3♂, 3♀; North Queensland, Mulgrave River at RFMR5, 23.i.1996, Barringtonia calyptrata, Bactrocera barringtoniae, QDPI MR21, (QDPIF), 1♂, 3♀; North Queensland, Old Palmerston Hwy, 23.xi.1995, Prunus persica, Bactrocera tryoni, QDPI NQ141, (QDPIF), 1♂, 1♀; North Queensland, Bot. Gardens, Cairns, 19.xii.1995, Barringtonia calyptrata, Bactrocera barringtoniae, QDPI NQ244, (QDPIF), 3♂, 2♀; North Queensland, Bloomfield R, 24.11.1996, Nauclea orientalis, Bactrocera pallida, QDPI NQ531, (QDPIF), 3♂, 6♀; North Queensland, Edmonton, 20.iii.1996, Carica papaya,

Bactrocera neohumeralis & Bactrocera tryoni, QDPI NQ662, (QDPIF), 1♂; North Queensland, Clifton Beach, 20.iii.1996, Nauclea orientalis, Bactrocera pallida & Bactrocera tryoni, QDPI NQ669, (QDPIF), 1♂. Stephen Island (Australia): Stephen Island, 23.ix.1993, Barringtonia racemosa, Bactrocera barringtoniae, SV015, E. Hamacek & J. Stephen, (QDPIF), 7♂, 10♀; Stephen Island, 30.xi.1993, Barringtonia racemosa, Bactrocera barringtoniae, SV063, J. Stephen, (QDPIF), 3♀; Stephen Island, 15.iii.1994, Barringtonia racemosa, Bactrocera barringtoniae, SV092, E. Hamacek & J. Stephen, (QDPIF), 2♂, 1♀; Stephen Island, Village, 24.iii.1995, Terminalia catappa, Bactrocera frauenfeldi, SV109, B. Sabine et al, (QDPIF), 2♂, 7♀; Stephen Island, 18.x.1993, Barringtonia racemosa, Bactrocera barringtoniae, SV23, B. Sabine & J. Stephen, (QDPIF), 5♂, 15♀. Host Associations B. aglaiae (Hardy) (1), B. barringtoniae (1,2), B. endiandrae (1,2), B. fagraea (Tryon) (2), B. frauenfeldi (1,2,3), B. jarvisi (2), B. kraussi (2), B. laticaudus (Hardy) (2), B. musae (Tryon) (2,3), B. pallida (1,2), and B. tryoni (1,2,3). Discussion Morphologically similar to F. schlingeri in colouration and notauli sculpture, however, F. deeralensis has a distinctly angulate clypeus and in many cases has weak striae on metasomal tergum 2. Fopius deeralensis was recorded from Australia (Queensland) and Papua New Guinea (New Britain Island) by (Wharton & Gilstrap 1983), however, this publication predates the description of F. schlingeri by Wharton (1999). No PNG material examined by us fit the description of F. deeralensis and its presence in that country must now be reassessed.

Figure 3.6. Fopius deeralensis. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view.

Fopius ferrari Carmichael & Wharton Etymology: Named after Dr Paul Ferrar, a retired Crop Sciences II Programme Manager from the Australian Centre for International Agricultural Research. Paul's long-term support for regional fruit fly research lead directly to several projects which helped generate the parasitoid collections upon which this paper is based. Synopsis (Figure 3.7a-f) Frons and vertex densely covered with white, decumbent setae; densely but finely punctate. Frons with polished, elevated projection extending from median ocellus almost to torulus; ocellar triangle otherwise margined by an impressed line. Occipital carina well developed, in lateral view extending dorsally nearly level with top of eye. Clypeus smooth, polished, very sparsely covered with long, erect setae, ventral margin obtusely angulate, forming a distinct median tooth; clypeus bulging along midline in profile. Notauli deep and foveolate-crenulate to midpit; midpit extending narrowly to posterior margin of scutum; propodeum exceptionally densely setose and rugose, the sculpture largely without obvious pattern except for median carina on basal 0.3; postpectral carina present. Sternaulus broad, crenulate/rugose throughout. Wings hyaline to weakly infumate. Forewing r arising slightly distad midpoint of stigma, 2RS longer than 3RSa, (RS+M)a weakly sinuate; (RS+M)b absent, m-cu arising directly in line with 2RS. Hindwing RS absent; m-cu developed and pigmented, but not reaching wing margin. Petiole weakly bicarinate longitudinally, otherwise polished, setose and weakly striate. Metasomal tergum 2 smooth (occasionally weakly striate) and densely setose; tergum 3 densely setose at least laterally. Ovipositor sheath almost twice length of metasoma; ovipositor not narrowed apically. Entire body dark brown to black. Fore and mid legs brown, hind legs dark brown to black. Diagnosis This species is similar to the Philippines species F. skinneri (Fullaway) in coloration, but differs from F. skinneri and darker specimens of F. arisanus by the absence of striae on metasomal tergum 2. The pattern of metasomal color and sculpture thus closely resembles that of F. illusorius. The clypeus, however, differs from that of F. illusorius and is similar to that of F. deeralensis. Fopius deeralensis is variable in T2 sculpture but is readily differentiated from the new species because of its yellow to orange coloration. Distribution Papua New Guinea: Collected from low altitude regions of the main island of New Guinea.

Types Holotype ♀; 3 labels, Label 1: Morobe Province, Lae: Bundun Conf Centre, 1.vi.2000, Label 2: Bred from: Persea americana Lauraceae N1516, Label 3: In association with [Bactrocera] frauenfeldi (QM). Paratypes 1♀ (ANIC): 3 labels, Label 1: PNG Morobe Province, Lae, Wampit village area, 16.ii.2000, Label 2: Bred from Musa ?, Musaceae, N 1263, Label 3: In association with [Bactrocera] musae. 1♀ (ANIC): Label 1: PNG Morobe Province Lae, Bukawa Tikeling 2 forest 27.v.1999 Label 2: Bred from ? N557 Label 3: In association with [Bactrocera] redunca; 3♂(ANIC): 3 labels, Label 1: PNG Morobe Province, Lae, Wampit village area, 16.ii.2000, Label 2: Bred from Musa ?, Musaceae, N 1263, Label 3: In association with [Bactrocera] musae. 2♀, 2♂ (QM): 3 labels, Label 1: PNG Morobe Province, Lae: FRI forest botanical garden, 16.v.2000, Label 2: Bred from: ? N 1462, Label 3: In association with [Bactrocera] frauenfeldi and [Bactrocera] trivialis. 3♀ (QM): 3 labels, Label 1: PNG Morobe Province, Lae, Gabensis Village, 6.iv.2000, Label 2: Bred from: Psidium guajava, Myrtaceae, N1384, Label 3: In association with [Bactrocera] frauenfeldi. 2♀, 1♂ (QM), 1♀ (ANIC): 3 labels, Label 1: PNG Morobe Province, Lae, Omsis forest, 11.vii.2000, Label 2: Bred from: ?, N1562, Label 3 In association with [Bactrocera] vulagaris [sic]. 1♀ (QM): 2 labels, Label 1: PNG Central Province, Bereina Station, Joe Aisa residence, 17.v.1999, Label 2: Bred from Averrhoa carambola Oxalidaceae, L 2552. Other Material examined Papua New Guinea: Madang Province, Baitabag, 16.viii.2000, ex Neisosperma oppositifolia, M 220, 10 sex undetermined (specimens damaged); Morobe Province, Lae: Bukawa, Tikeling 2 forest, 27.v.1999, in association with Bactrocera redunca, N 557, 1; Morobe Province, Lae: Omsis forest, 1.vi.2000, in association with Bactrocera trivialis?, N 1513(A), 1♂; Morobe Province, Lae, Gabensis Village, 6.iv.2000, ex Psidium guajava, Myrtaceae, in association with [Bactrocera] frauenfeldi N1384, 1♂, 1 sex undetermined (specimen damaged). Discussion Although this species appears at first glance to be just a somewhat darker, more densely setose variety of F. illusorius, the ovipositor and clypeus are sufficiently different to warrant description as a separate species. The tip of the ovipositor is not narrowed as it is in F. illusorius and F. arisanus. This species was formally described as a new species by Carmichael et al (2005)

Figure 3.7. Fopius ferrari. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view.

Fopius illusorius (Fischer, 1971) Biosteres illusorius (Fischer 1971b): 487. Biosteres (Chilotrichia) illusorius: (Fischer 1978a): 386. Taxonomic History This species was originally described as Biosteres illusorius by Fischer (1971). Fischer (1978) reallocated this species into the subgenus Chilotrichia, therefore naming the species Biosteres (Chilotrichia) illusorius. This species was transferred to Fopius by Carmichael et al (2005) Synopsis (Figure 3.8a-f) Frons and vertex densely setose; frons rugose-punctate, vertex densely punctate. Ocellar triangle arranged to form an obtuse triangle with a slight depression anteriad the median ocellus. Occipital carina well developed; clypeus in profile bulging very slightly dorsomedially, ventral margin slightly thickened medially, without distinct medial projection. Notauli deep, foveolate/crenulate to midpit; propodeum rugose, the sculpture largely without obvious pattern except for median carina on basal 0.3; postpectral carina present. Sternaulus deep, broad, crenulate. Wings infumate. Forewing r arising slightly distad midpoint of stigma, 2RS longer than 3RSa, (RS+M)a weakly sinuate; (RS+M)b usually present, m-cu arising distad or directly in line with 2RS. Hindwing RS virtually absent basally, represented only by a faint crease near the wing margin; m-cu well developed and at least weakly pigmented to wing margin. Petiole longitudinally striate, dorsal lateral carinae elevated over basal half and extending to posterior margin. Metasomal tergum 2 usually smooth, polished, and setose; occasionally with a few weak striae antero-medially. Ovipositor sheath equal in length to mesosoma plus metasoma; ovipositor tip narrower apically, with weak subapical constriction. Head orange-brown; mesosoma orange brown, with propodeum and usually metathorax dark brown to black; metasoma dark brown to black. Fore and mid legs yellow, except mid coxa dark brown dorsally; hind legs dark brown to black with apical tarsomeres lighter brown. Females tend to be slightly darker than males. Distribution Australia: Nambour. Papua New Guinea: Collected from low altitude regions of the main island of New Guinea, the Gazelle Peninsular of East New Britain and Duke of York Island. Solomon Islands: Visale/Guadalcanal and Ysabel. Type Locality Original reference: Waris, Hollandia, Papua New Guinea. [Hollandia is actually a province of Indonesia, now officially known as ‘Irian Jaya’ or ‘West Papua’].

Material examined Australia: Queensland, Nambour, 2.ii.1993, Carambola, Bactrocera tryoni & Bactrocera neohumeralis, Q64, M.M. Elson-Harris, (QDPIF), 2♂, 1♀. Papua New Guinea: E. New Britain, Kerevat: L.A.E.S., 12-Dec-97, MYRTACEAE, Syzygium, Bactrocera frauenfeldi & Bactrocera obliqua, I 118-122, (GU), 3♂; E. New Britain, Kerevat: L.A.E.S., Michael Wan house, 18-Dec-97, MYRTACEAE, Syzygium, Bactrocera frauenfeldi & Bactrocera obliqua, I 143-148, (GU), 7♀; E. New Britain, Sonoma: S.D.A. school, mechanics workshop, 22-Dec-97, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, I 162-163, (GU), 1♂, 1♀; E. New Britain, Kerevat: L.A.E.S., small arboretum orchard, 6-Mar-98, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, I 366-367, (GU), 4♂, 7♀; E. New Britain, Kerevat: L.A.E.S., Guava block 408, 28-May-98, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, I 563-578, (GU), 8♂, 4♀; E. New Britain, Kerevat: L.A.E.S., Cashew block 307, 28-Oct-98, ANACARDIACEAE, Anacardium occidentale, Bactrocera frauenfeldi, I 759-762, (GU), 1♀; E. New Britain, Kerevat: L.A.E.S., Guava block 408, 2-Feb-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, I 839- 844, (GU), 7♂, 7♀; Central, Lobulogo village Hiritano Highway, 23-Feb-99, OXALIDACEAE, Averrhoa carambola, Bactrocera frauenfeldi, L 1301-1400, (GU), 3♂, 2♀, 5 sex undetermined; Central, Laloki block A.Mehuze garden, 19-May-99, MUSACEAE, Musa x paradisiaca, Bactrocera musae, L 2937-2966, (GU), 1♂, 1♀; Central, NARI Laloki, 14-Jan-99, COMBRETACEAE, Terminalia catappa, Bactrocera frauenfeldi, L 501-600, (GU), 1♀, 2 sex undetermined; Morobe, Lae: Wampit village area, 16-Feb-00, MUSACEAE, Musa, Bactrocera musae, N 1263, (GU), 3♀; Morobe, Lae: Gabensis Village area, 28-Mar-00, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 1369, (GU), 5♀; Morobe, Lae: Omsis forest, 06- Apr-00, GNETACEAE, Gnetum gnemon, Bactrocera (Bulladacus) n.sp., N 1383, (GU), 1♂; Morobe, Lae: Gabensis Village area, 06-Apr-00, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 1384, (GU), 2♂, 12♀; Morobe, Lae: Gabensis Village area, 27-Apr-00, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 1404, (GU), 2♂, 9♀; Morobe, Lae:11-mile, Suambu plantation, 20-Jun-00, Bactrocera frauenfeldi & Bactrocera trivialis, N 1541, (GU), 1♂, 3♀; Morobe, Lae: Nadzarb airport area, 21-Aug-00, CARICACEAE, Carica papaya, Bactrocera frauenfeldi & Bactrocera musae, N 1594, (GU), 21♀; Morobe, Lae: University of Technology, 22-Apr-99, MYRTACEAE, Syzygium, Bactrocera frauenfeldi, N 359, (GU), 1♀; Morobe, Lae: Gabensis village area, 27-Apr-99, MYRTACEAE, Psidium

guajava, Bactrocera frauenfeldi, N 391, (GU), 1♂; Morobe, Lae: Gabensis village area, 27-Apr-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera trivialis, N 392, (GU), 1♀; Morobe, Lae: Gabensis village area, 27-Apr- 99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera trivialis,, N 395, (GU), 1♀; Morobe, Lae: 11-mile Suambu farm, 03-May-99, EUPHORBIACEAE, Baccaurea, Bactrocera frauenfeldi & Bactrocera trivialis, N 441, (GU), 2♀; Morobe, Lae: 11-mile Suambu farm, 03-May-99, MYRTACEAE, Syzygium, Bactrocera frauenfeldi, N 447, (GU), 1 sex undetermined; Morobe, Lae: Markham farm, Mararumi, 28-Jul-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 652, (GU), 5♀; Morobe, Lae: Markham farm, Mararumi, 25-Aug-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera trivialis, N 789, (GU), 1♀; Morobe, Lae: Markham farm, Mararumi, 25-Aug-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera trivialis, N 795, (GU), 2♂, 2♀; Morobe, Lae: Gabensis village area, 20-Sep-99, CARICACEAE, Carica, papaya, Bactrocera frauenfeldi, N 936, (GU), 2♂, 3♀; Morobe, Lae: Gabensis village area, 05-Oct-99, MUSACEAE, Musa, x paradisiaca, Bactrocera, N 981, (GU), 1♀; E. New Britain, Kerevat: L.A.E.S., Guava block 408, 4-Dec-98, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 2338-2438, (GU), 5♂, 10♀; E. New Britain, Kerevat C.I.S. prison, 2-Feb-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua,, R 2591-2640, (GU), 9♂, 10♀; E. New Britain, Kerevat C.I.S. prison, 10-Feb-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 2641-2690, (GU), 4♂, 8♀; E. New Britain, Kerevat C.I.S. prison, 24-Feb-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 2752-2801, (GU), 6♂, 9♀, 1 sex undetermined; E. New Britain, Kerevat C.I.S. prison, 17-Mar-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 2852-2901, (GU), 4♂, 11♀; E. New Britain, Kerevat C.I.S. prison, 26-May-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, R 3102-3151, (GU), 7♂, 4♀; E. New Britain, Kerevat C.I.S. prison, 9-Jun-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, R 3152- 3201, (GU), 22♂, 22♀; E. New Britain, Kerevat C.I.S. prison, 23-Jun-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 3202-3251, (GU), 7♂, 21♀; E. New Britain, Kerevat C.I.S. prison, 28-Jul-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 3302-3349, (GU), 3♂, 15♀, 1 sex undetermined; E. New Britain, Kerevat C.I.S. prison, 28-Jul-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 3556-3605, (GU), 6♂, 14♀, 2 sex undetermined; E. New Britain, Kerevat: L.A.E.S., Orchard nr. Country Club, 30-Aug-99, OXALIDACEAE,

Averrhoa, carambola, Bactrocera frauenfeldi, R 3606-3725, (GU), 2♂; E. New Britain, Kerevat: L.A.E.S., Guava block 408, 1-Sep-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 3726-3775, (GU), 2♂, 42♀, 10 sex undetermined; E. New Britain, Kerevat C.I.S. prison, 8-Sep-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 3776-3825, (GU), 9♂, 35♀, 15 sex undetermined; E. New Britain, Kerevat: L.A.E.S., Geoff Wiles house, 29-Jan-98, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 877-940, (GU), 18♂, 15♀; E. New Britain, Kerevat: L.A.E.S., F. Hela house (trap P202), 29-Jan-98, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 941-1015, (GU), 9♂, 9♀; E. New Britain, Kerevat C.I.S. prison, 3-Mar-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R2802-2851, (GU), 6♂, 3♀; E. New Britain, Kambain Village, 30-Nov-99, Taguak (DOY language), Bactrocera frauenfeldi, I 1300, (GU), 1♂, 2♀; E. New Britain, Kerevat: L.A.E.S., small arboretum orchard, 26-Nov-97, ANACARDIACEAE, Mangifera indica, Bactrocera frauenfeldi, I 85-87, (GU), 1♂, 19♀; E. New Britain, Duke of York Is: Kibil Station, 5-Jun-99, Utun (DOY language), I 975, (GU), 1♂; Central, Lobulogo village Hiritano Highway, 16- Feb-99, OXALIDACEAE, Averrhoa carambola, Bactrocera frauenfeldi, L 1201-1300, (GU), 1♀, 2 sex undetermined; Central, Lobulogo village Hiritano Highway, 30-Mar- 99, OXALIDACEAE, Averrhoa carambola, Bactrocera frauenfeldi, L 2001-2100, (GU), 4 sex undetermined; Central, Lobulogo village, Hiritano Highway, 19-Jan-99, OXALIDACEAE, Averrhoa carambola, Bactrocera frauenfeldi, L 601-700, (GU), 2♀, 12 sex undetermined; Central, Lobulogo village Hiritano Highway, 19-Jan-99, OXALIDACEAE, Averrhoa carambola, Bactrocera frauenfeldi, L 701-800, (GU), 6♂, 21♀, 2 sex undetermined;Central, Launakalana Estate Magi highway, 22-Sep-98, ANACARDIACEAE, Anacardium occidentale, L1-100, (GU), 1♀; Morobe, Lae: Bubia Agri Station, 18-Nov-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 1146, (GU), 1♀; Morobe, Lae: Bubia Agri Station, 18-Nov-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 1148, (GU), 4♀; Morobe, Lae: Omsis forest, 22-Feb-00, GNETACEAE, Gnetum gnemon, B.(Bulladacus) n.sp., N 1281, (GU), 2♀; Morobe, Lae:11-mile, Suambu plantation, 08-Mar-00, GUTTIFERAE, Garcinia mangostana, Bactrocera frauenfeldi, N 1318 (A), (GU), 1♀; Morobe, Lae: Bukawa, Tikeling 2 forest, 09-May-00, unidentified, N 1426, (GU), 1♀; Morobe, Lae: Labu-Butu area, 27-Apr-99, MORACEAE, Ficus, N 399, (GU), 2♀; Morobe, Lae: Erab Agri Station, 22-Jul-99, OXALIDACEAE, Averrhoa carambola, N 644, (GU), 2♂, 3♀; Morobe, Lae: Markham farm, Mararumi, 28-Jul-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 655, (GU), 4♀; Morobe, Lae: Markham farm, Mararumi, 28-

Jul-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 675, (GU), 1♂; Morobe, Lae: Markham farm, Mararumi, 28-Jul-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 677, (GU), 1♂, 1♀; Morobe, Lae: Markham farm, Mararumi, 28-Jul-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 678, (GU), 7♂, 5♀; Morobe, Lae: Markham farm, Mararumi, 28-Jul-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 679, (GU), 8♂, 14♀; Morobe, Lae: Markham farm, Mararumi, 28-Jul-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 682, (GU), 1♀; Morobe, Lae: Markham farm, Mararumi, 28-Jul-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 690, (GU), 1♂; Morobe, Lae: Markham farm, Mararumi, 25-Aug-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 787 , (GU), 6♂, 4♀, 1 sex undetermined; Morobe, Lae: Markham farm, Mararumi, 25-Aug-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 796, (GU), 7♀; Morobe, Lae: Markham farm, Mararumi, 25-Aug-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 801, (GU), 3♂, 1♀; Morobe, Lae: Markham farm, Mararumi, 6-Sep-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 863, (GU), 1♂, 1♀; Morobe, Lae: Markham farm, Mararumi, 6-Sep-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 864, (GU), 1♀; Morobe, Lae: Markham farm, Mararumi, 10-Sep-99, CARICACEAE, Carica papaya, frauenfeldi, N 888, (GU), 7♂, 5♀; Morobe, Lae: Mumeng (Zenag), 17-Sep-99, CARICACEAE, Carica papaya, N 915, (GU), 1♂; Morobe, Lae: Bulolo Forestry College, 20-Sep-99, MYRTACEAE, Psidium guajava, frauenfeldi, N 916, (GU), 2♂; Morobe, Lae: Gabensis village area, 05-Oct-99, MUSACEAE, Musa x paradisiaca, N 975, (GU), 1 sex undetermined; Morobe, Lae: Gabensis village area, 05-Oct-99, MUSACEAE, Musa x paradisiaca, N 978, (GU), 1♂, 1♀; E. New Britain, Kerevat: L.A.E.S., Guava block 408, 25-Jun-98, MYRTACEAE, Psidium guajava, R 1792-1794, (GU), 2♂, 3♀; E. New Britain, Kerevat: L.A.E.S., Cashew block 307, 12-Nov-98, ANACARDIACEAE, Anacardium occidentale, frauenfeldi, R 2113-2207, (GU), 2♂, 2♀; E. New Britain, Vudal College, 21-Dec-98, ANACARDIACEAE, Mangifera indica, R 2582, (GU), 7♀; E. New Britain, Kerevat C.I.S. prison, 14-Jul-99, MYRTACEAE, Psidium guajava, frauenfeldi, R 3252-3301, (GU), 14♂, 40♀, 2 sex undetermined; E. New Britain, Kerevat: L.A.E.S., Cashew block 307, 24-Aug-99, ANACARDIACEAE, Anacardium, occidentale, R 3506, (GU), 1♀; E. New Britain, Kerevat C.I.S. prison, 17-Sep-99, MYRTACEAE, Psidium guajava, frauenfeldi, R 3826-4144, (GU), 1♀; E. New Britain, Kerevat C.I.S. prison, 20/10/1999 to 2/2/00, MYRTACEAE, Psidium guajava, frauenfeldi, R 4145-4544, (GU), 2♂, 16♀, 4 sex undetermined. Solomon Islands: Visale, N.W., Guadalcanal, 29.v.1996, Terminalia catappa, Bactrocera frauenfeldi & Bactrocera tigrina, SI 1617, (QDPIF), 1♂, 1♀; Ysabel,

Buala Village, 13.viii.1996, Syzygium malaccense, Bactrocera frauenfeldi, Japhet Kaoga, (QDPIF), 1♀ Host Associations B. frauenfeldi (1), B. musae (1) & B.obliqua (1). Discussion This species was transferred to the genus Fopius Wharton by Carmichael et al. (2005) on the basis of mandibular morphology, wing venation and sculpture of the mesosoma. Additionally it has the long ovipositor typical of most Fopius species. This species is distinct from nearly all Biosteres species due to the absence of a strong basal tooth or lobe, a feature characteristic of all other Biosteres species except B. blandus (Wharton, 1997). Unlike the type species of Chilotrichia, the hindwing RS is absent basally and only represented by a faint crease near the apex of the wing. The propleuron has the strongly developed oblique ridge typical of Fopius and the postpectal carina is also well developed. As noted above, F. illusorius is similar in colouration to F. arisanus, but the second metasomal tergum is smooth or nearly so. Fopius illusorius was previously known only from the original description (Fischer 1971b) based on two males. The previously undescribed female has an ovipositor that is morphologically similar to that of F. arisanus and thus F. illusorius may oviposit in the eggs of its host as does F. arisanus. The hosts of F. illusorius were previously unknown as the males from the type series were collected with a sweep net.

Figure 3.8. Fopius illusorius. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view.

Fopius schlingeri Wharton, 1999 Fopius schlingeri (Wharton 1999): 58. Taxonomic History This species was originally described as Fopius schlingeri by Wharton (1999). No further revisions have taken place. Synopsis (Figure 3.9a-f) Frons and vertex densely setose; frons densely punctate, vertex more sparsely and weakly punctate. Frons with polished, weakly elevated projection extending from median ocellus at least half the distance to torulus. Ocellar triangle arranged to form an obtuse triangle with a slight depression anteriad the median ocellus. Occipital carina well developed; clypeus with rounded ventral margin. Notauli foveolate/crenulate to midpit; propodeum densely rugose, the sculpture without obvious pattern except for median carina basally. Sternaulus broad, deep, crenulate. Wings weakly infumate. Forewing r arising slightly distad midpoint of stigma, 2RS slightly longer than 3RSa, (RS+M)a sinuate; (RS+M)b present, m-cu arising basad 2RS. Hindwing RS virtually absent, represented only by a faint crease near wing margin; m-cu well developed and pigmented to wing margin. Petiole longitudinally, somewhat irregularly striate, dorsal lateral carinae well developed over basal two- thirds, weaker posteriorly. Metasomal tergum 2 smooth and setose laterally. Ovipositor sheath equal to or longer than length of body including head. Ovipositor tip strongly narrowed subapically. Whole body yellow/orange except hind tarsi dark brown. Distribution Australia: Collected along the eastern coast of Australia, from Cape Tribulation in the north, to Sydney in the south. Papua New Guinea: Collected from low altitude regions of the main island of New Guinea and the Gazelle Peninsula of East New Britain. Type Locality Mt Glorious, Australia. Material examined Australia: Condong, N.S.W., 6.iv.1959, T. G. Campbell, (ANIC), 2♀; Sydney, N.S.W., 28.vi.1954, G.J. Snowball, (ANIC), 5♀; Sydney, N.S.W., 2.vii.1954, G.J. Snowball, (ANIC), 1♂; Sydney, N.S.W., 29.i.1954, G.J. Snowball, (ANIC), 1♂; Dorrigo, N.S.W., xi.1980, Planchonella australis, Dacus halfordiae, G. Fitt, (ANIC), 1♂, 1♀; Coffs Harbour, N.S.W., 12.v.1954, ex loquat, (ANIC), 1♂, 2♀; Mullet Creek, Queensland, 25.iv.1959, ex cherry guava, (ANIC), 2♂; Deeral, near, .vi.1949,

Acmena macrocarpa, N.L.H. Krauss, (ASCU), 2♀; Queensland, Nambour, 14.i.1993, Wax Jambu, Bactrocera neohumeralis & Bactrocera tryoni, Q26, M.M. Elson-Harris, (QDPIF), 3♂; Queensland, Nambour, ii.1993, Feijoa, Bactrocera neohumeralis & Bactrocera tryoni, Q67, M.M. Elson-Harris, (QDPIF), 4♂, 5♀; Queensland, Nambour, 18.iii.1993, Carambola, Q99, M.M. Elson-Harris, (QDPIF), 1♀; North Queensland, Babinda Boulders at RFBB5, 20.v.1996, Acmena sp, Bactrocera kraussi, QDPI BB159, (QDPIF), 1♀; North Queensland, Cape Tribulation at RFCT40, 19.ix.1996, Niemeyera prunifera, Bactrocera kraussi, QDPI CT1234, (QDPIF), 2♂, 10♀; North Queensland, Cape Tribulation at RFCT1, 3.iv.1996, Acmena sp, Bactrocera kraussi, QDPI CT373, (QDPIF), 1♀; North Queensland, Cape Tribulation at RFCT1, 2.vi.1996, Acmena sp, Bactrocera kraussi, QDPI CT487, (QDPIF), 1♀; North Queensland, Cape Tribulation at RFCT20, 29.iv.1996, Acmena graveoleus, Bactrocera kraussi, QDPI CT643, (QDPIF), 1♂, 5♀; North Queensland, Cape Tribulation at RFCT9, 28.v.1996, Acmena sp, Bactrocera kraussi, QDPI CT890, (QDPIF), 2♀; North Queensland, Cape Tribulation at RFCT1, 28.v.1996, Acmena sp, Bactrocera kraussi, QDPI CT917, (QDPIF), 2♀; North Queensland, Cape Tribulation at RFCT 40, 15.viii.1996, Acmena graveoleus, Bactrocera kraussi, QDPI CT993, (QDPIF), 1♀; North Queensland, Josephine Falls at RFJF8, 4.ii.1996, Acmena graveolens, Bactrocera kraussi, QDPI JF29, (QDPIF), 3♀; North Queensland, Mulgrave River, 14.ii.1996, Terminalia seriocarpa, Bactrocera neohumeralis, QDPI MR2, (QDPIF), 2♂, 2♀; North Queensland, Redlynch Valley, 5.xii.1995, Chionanthus ramiflorus, Bactrocera nigra & Euphranta linocierae, QDPI NQ81, (QDPIF), 1♂. Papua New Guinea: E. New Britain, Kerevat: L.A.E.S., Cashew block 307, 28-Oct- 98, ANACARDIACEAE, Anacardium occidentale, Bactrocera frauenfeldi, I 759-762, (GU), 1♂, 3♀; Central, Laloki block A.Mehuze garden, 19-May-99, MUSACEAE, Musa x paradisiaca, Bactrocera musae, L 2937-2966, (GU), 1♀; Madang, Baitabag, 4-Jul-01, RUBIACEAE, Versteegia cauliflora, Bactrocera B. tinomiscii, M 1625, (GU), 2♂, 19♀; Madang, Baitabag, 11-Oct-00, LOGANIACEAE, Neuburgia corynocarpa, Bactrocera, Euphranta marginata, M 425, (GU), 2♀; Morobe, Lae: Omsis forest, 07-Feb-00, COMBRETACEAE, Terminalia, Bactrocera penefurva, N 1259, (GU), 2♀; Morobe, Lae: Omsis forest, 16-Feb-00, COMBRETACEAE, Terminalia, Bactrocera frauenfeldi & Bactrocera penefurva, N 1273, (GU), 4♀; Morobe, Lae: Bukawa, Tikeling 2 forest, 04-Apr-00, Bactrocera unidentified, N 1377, (GU), 1♀; Morobe, Lae: Gabensis Village area, 06-Apr-00, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 1384, (GU), 5♂, 5♀; Morobe, Lae: Gabensis Village area, 27-Apr-00, MYRTACEAE, Psidium guajava, Bactrocera

frauenfeldi, N 1404, (GU), 4♂, 3♀; Morobe, Lae: FRI forest/ botanical gard, 16-May- 00, Bactrocera frauenfeldi & Bactrocera trivialis, N 1462, (GU), 4♂, 1♀; Morobe, Lae: Omsis forest, 11-Jul-00, Bactrocera vulgaris (to check), N 1562, (GU), 7♀; Morobe, Lae: Gabensis village area, 27-Apr-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 391, (GU), 1♀; E. New Britain, Kerevat C.I.S. prison, 28- Jul-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 3302-3349, (GU), 1♀; E. New Britain, Kerevat: L.A.E.S., Geoff Wiles house, 29-Jan-98, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 877-940, (GU), 2♀; Madang, Baitabag, 19-Jul-01, sp. 127, B. tinomiscii, M 1731, (GU), 3♂, 10♀; Morobe, Lae: Markham, Nasuapum vill, 06-Apr- 00, GNETACEAE, Gnetum gnemon, B.(Bulladacus) , n.sp., N 1391, (GU), 1♀; Morobe, Lae: Gabensis village area, 27-Apr-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 389, (GU), 1♂, 2♀. Host Associations B. frauenfeldi (1), B. halfordiae (1,5), B. kraussi (1), B. musae (1), B. neohumeralis (1,5), B. penefurva (1), B. peninsularis (Drew & Hardy) (5), B. rufofuscula (Drew & Hancock) (5), B. tinomiscii Drew (1), B. tryoni (5), and B. (Bulladacus) sp. (1). Discussion Morphologically similar to F. deeralensis in colouration and in sculpture of the frons, vertex, and notauli. However, F. schlingeri has a rounded clypeus, without a distinct medial projection. Originally described as occurring along the eastern coast of Australia (Wharton 1999), the native range is now extended to include Papua New Guinea.

Figure 3.9. Fopius schlingeri. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view.

Fopius cf. vandenboschi Synopsis (Figure 3.10a-f) Frons and vertex densely setose and punctate. Frons with polished, weakly depressed, laterally carinate projection extending from median ocellus at least half the distance to torulus; frons otherwise weakly punctate, the punctures anteriorad ocellar field dense, with spacing between punctures slightly less than diameter of punctures; ocellar triangle margined by an impressed line. Occipital carina well developed. Clypeus in profile bulging dorsomedially; ventral margin of clypeus slightly thickened medially, and sufficiently convex to completely conceal labrum when mandibles closed. Notauli deep and foveolate/crenulate to midpit; propodeum rugose, the sculpture largely without obvious pattern. Sternaulus broad, deep, crenulate/rugose throughout. Wings weakly infumate. Forewing r arising slightly distad midpoint of stigma, second submarginal cell short, 2RS slightly longer than 3RSa, (RS+M)a sinuate; (RS+M)b short to absent, m-cu arising basad or directly in line with 2RS. Hind wing RS virtually absent; m-cu well developed and at least weakly pigmented to wing margin. Petiole longitudinally striate, dorsal lateral carina weakly developed to posterior margin. Metasomal tergum 2 striate. Ovipositor sheath about equal in length to body including head. Apex of ovipositor parallel sided with very weak dorsal node. Head and mesosoma yellow/orange, metasoma yellow orange with terga 2+3 and often 4 black, sternites pale yellow/brown to almost white; petiole varying from orange to dark brown. Distribution Papua New Guinea Kerevat. Material examined Papua New Guinea: E. New Britain, Kerevat: L.A.E.S., Michael Wan house, 18-Dec- 97, MYRTACEAE, Syzygium, Bactrocera frauenfeldi & Bactrocera obliqua, I 143- 148, (GU), 2♀; E. New Britain, Kerevat C.I.S. prison, 2-Feb-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 2591-2640, (GU), 1♂, 3♀; E. New Britain, Kerevat C.I.S. prison, 24-Feb-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 2752-2801, (GU), 3♂, 9♀; E. New Britain, Kerevat C.I.S. prison, 17-Mar-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 2852-2901, (GU), 3♂, 4♀; E. New Britain, Kerevat C.I.S. prison, 9-Jun-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, R 3152-3201, (GU) 1♀; E. New Britain, Kerevat: L.A.E.S., Orchard nr. Country Club, 30-Aug-99, OXALIDACEAE, Averrhoa carambola, Bactrocera frauenfeldi, R 3606-3725, (GU), 1♂, 2♀; E. New Britain, Kerevat C.I.S. prison, 3-

Mar-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R2802-2851, (GU), 1♂, 4♀; PNG , E. New Britain, Kerevat C.I.S. prison, 28-Apr-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, R 3002-3051, (GU), 1♂. Host Associations B. frauenfeldi (1) & B. obliqua (1). Discussion Fopius vandenboschi (Fullaway) appears to consist of a number of colour morphs and potentially cryptic species in the Indo-Pacific region. This colour form is potentially distinct from the true F. vandenboschi, having developed in isolation on the island of New Britain. Slight differences in clypeal morphology and punctation of the vertex and frons provide support for considering this distinctive colour morph as a separate species, but further studies are needed. The second metasomal tergum is more extensively striate than shown in Figure 3.10f both in typical F. vandenboschi and in the New Britain material.

Figure 3.10. Fopius cf. vandenboschi. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view.

Opius froggatti (Fullaway, 1950) Opius froggatti (Fullaway 1950a): 67. Opius (Opius) froggatti: (Fischer 1963b): 203. Opius (Utetes) froggatti: (Fischer 1987): 186. Opius froggatti: (Wharton 1997a): 29. Taxonomic History Opius froggatti was described by Fullaway (1950). In 1963b, Fischer placed this species in the subgenus Opius [Opius (Opius) froggatti]. The subgeneric position was again reviewed by Fischer (1987) and placed in the subgenus Utetes. [Opius (Utetes) froggatti]. In 1997, Wharton removed the subgeneric classification and replaced the species within the original genus Opius, therefore renaming the species Opius froggatti. Synopsis (Figure 3.11a-c) Frons and vertex polished and sparsely setose. Ocelli arranged in an equilateral triangle margined by a depression. Occipital carina well developed. Clypeus in profile bulging dorsomedially; labrum visible in frontal view; gap between mandible and ventral margin of clypeus large and distinct. Notauli beginning as deep grooves, converging and becoming shallower posteriorly, meeting in a deep circular midpit. Propodeum reasonably smooth, usually with a complete medial longitudinal carina. Sternaulus shallow, minutely crenulate throughout. Wings weakly infumate. Forewing r arising slightly basad midpoint of stigma, 2RS notably shorter than 3RSa, (RS+M)a weakly sinuate; (RS+M)b absent, m-cu arising distinctly distad 2RS. Hind wing RS and m-cu absent. Petiole with dorsal lateral carinae elevated, diverging posteriorly, but extending to posterior margin. Metasomal tergum 2 smooth and polished. Ovipositor sheath about two-thirds length of metasoma. Whole body yellow orange in colouration. Distribution Australia Deeral. Type Locality Deeral, Australia Material examined Australia: Deeral, near, vi.1949, Planchonella, N.L.H. Krauss, (ASCU), 1♀; near Deeral, .vii.1949, Planchonella, N.L.H. Krauss, (ASCU), 1♂, 2 sex undetermined. Host Associations B. kraussi (2), B. laticaudus (2), and B. psidii (2,3).

Discussion Opius froggatti is morphologically similar to species in the genus Utetes (venation, readily visible labrum, distinct median pit on mesoscutum), but can be easily distinguished by the absence of a basal carina on the hind tibia. Opius froggatti is similar to Utetes perkinsi in having weakly developed notauli, but in addition to the presence of a tibial carina, the latter has a dark head.

Figure 3.11. Opius froggatti. a, head, anterior view; b, mesonotum, dorsal view; c, wing.

Psyttalia fijiensis (Fullaway, 1936) Opius fijiensis (Fullaway 1936): 179. Austroopius fijiensis: (Fischer 1963a): 177. Psyttalia (Austroopius) fijiensis: (Wharton 1987): 64. Psyttalia fijiensis: (Waterhouse 1993): 42. Taxonomic History Opius fijiensis was originally described by Fullaway (1936). Fischer (1963a) revised the generic status and placed the species with the genus Austroopius [Austroopius fijiensis]. Wharton (1987) described Psyttalia as a genus, which included this species, [Psyttalia fijiensis]. Synopsis See P. novaguineensis Distribution Australia: Collected in north eastern Australia, from Yam Island in the north, south to Brisbane; Fiji: Suva and surrounding area, Viti Levu. Solomon Islands: Bougainville Island. Tonga: Tongatapu and ‘Eua Islands. Type Locality Noainee, Fiji Islands Material examined Australia: Bingil Bay, Queensland, 1.v.1958, T. G. Campbell, (ANIC), 1 sex undetermined; Maryborough, Queensland, 15.iv.1959, ex guava, (ANIC), 2 sex undetermined; Mossman Gorge, N.Qld, 23.ii.1980, Castanospermum australei, Rioxa confusa, G. Fitt, (ANIC), 1♀; Wongabel Forest, Atherton, N.Qld, 17.i.1982, Castanospora alphandi, Dacus krausii, G. Fitt, (ANIC), 1♀; Mossman Gorge, N.Qld, 14.i.1982, Garcinia warrenii, Dacus visenda, G. Fitt, (ANIC), 1♀; Mullet Creek, Queensland, 9.iii.1960, ex yellow guava, T.G. Campbell, (ANIC), 1♀; Nambour, Queensland, 14.iv.1959, reared ex guava, (ANIC), 4♀; Malanda Falls, Atherton, N. Queensland, 25.ii.1980, Aglaia ferruginea, Dacus aglaiae, G. Fitt, (ANIC), 1♂; Wongabel Forest, Atherton, N.Qld, 13.i.1982, Castanospora alphandi, Dacus krausii, G. Fitt, (ANIC), 1♂; Brisbane, Queensland, .3.1981, guava, Dacus tryoni, G. Fitt, (ANIC), 1♂; Mossman Gorge, N.Qld, 14.i.1982, Garcinia warrenii, Dacus visenda, G. Fitt, (ANIC), 1♂; Condong, N.S.W, 6.iv.1959, ex guava, (ANIC), 1♂; Cooktown, Queensland, 1 km SE Mt Cook, 13.x.1980, collected at light, J.C. Cardale, (ANIC), 1♂; Ayr, Queensland, 10.iv.1953, Nauclea orientalis, Strumeta pallida, A.W.S. May, (ANIC), 1♂, 1 sex undetermined; Cape Tribulation, Queensland, 3.ix.1981, Endiandra microneura, Dacus krausii, G. Fitt, (ANIC), 1♂,

1♀; Mackay, Queensland, 20.iii.1950, Psidium guajava, Strumeta spp, A.W.S. May, (ANIC), 1♂, 1♀; Burrum River, Queensland, 8.iii.1960, ex yellow guava, T.G. Campbell, (ANIC), 1♂, 1♀; Maryborough, Queensland, 7.iii.1960, ex yellow guava, T.G. Campbell, (ANIC), 1♂, 1♀; N.L.H. Krauss, (ANIC), 1♂, 1♀; Mullet Creek, Queensland, 20.iv.1959, ex guava, (ANIC), 1♂, 1♀, 1 sex undetermined; Innisfail, Queensland, 11.v.1960, ex yellow guava, T. G. Campbell, (ANIC), 1♂, 2♀; Mullet Creek, Queensland, 9.iii.1960, ex cherry guava, T.G. Campbell, (ANIC), 1♂, 2♀; Cordelia, Queensland, 28.iv.1960, ex yellow guava, T.G. Campbell, (ANIC), 2♂; Maryborough, Queensland, 6.iii.1960, ex yellow guava, T.G. Campbell, (ANIC), 2♂; Freshwater, Queensland, .1.1963, G.I. Snowball, (ANIC), 2♂, 1♀; Mackay, Queensland, 10.iv.1960, ex yellow guava, T.G. Campbell, (ANIC), 2♂, 1♀; Atherton Tableland, Queensland, .xi.1949, Litsea leefeana, Dacus sp, NSW AG, N.L.H. Krauss, (ASCU), 1♀; Grafton, N.S.W., 16.iii.1934, ex avocado, Chaetodacus tryoni, NSW AG, (ASCU), 1♀; Grafton, N.S.W., 16.iii.1934, ex guava, Chaetodacus tryoni, NSW AG, (ASCU), 2♀; Cairns, Queensland, .xi.1949, Dacus sp., NSW AG, N.L.H. Krauss, (ASCU), 1♂; Atherton Tableland, Queensland, .xi.1949, Eugenia spp., Dacus sp., NSW AG, N.L.H. Krauss, (ASCU), 1♂, 1♀; Muller Creek, Queensland, 4.iii.1960, ex yellow guava, NSW AG, T.G. Campbell, (ASCU), 2♂, 2♀; Queensland, Nambour, 14.i.1993, K Wax Jambu, Q27, M.M. Elson-Harris, (QDPIF), 1♂; Queensland, Nambour, 17.ii.1993, Feijoa, Q80, M.M. Elson-Harris, (QDPIF), 2♂; Queensland, Downey Creek, 15.xi.1982, Pulogium galactoxyla, Bactrocera endiandrae, M.M. Elson-Harris, (QDPIF), 1♀; Queensland, Rockhampton, i.1995, ?, Planchonia careya, B. (Afrodacus) jarvisi, P. Leach, (QDPIF), 1♂. Fiji: Nausori, Naduruloulou R. S., 6.i.1992, Citrus maxima, Bactrocera passiflorae, F1926, A. Allwood et al, (QDPIF), 1♀; Ocean Pacific Club, 25km W of Suva, 23.i.1992, F1984P, G. Walker et al, (QDPIF), 1♂, 1♀; Suva, Colo-i-Suva Forest Park, 9.vii.1992, Amaroria soulameiodes, Bactrocera passiflorae, F3089, G. Walker et al, (QDPIF), 1♂, 3♀; Suva, Colo-i-Suva Forest Park, 18.vii.1991, FI042P, Ema Tora et al, (QDPIF), 6♂. Solomon Islands: Bougainville Island, Konga Village (Buin), 6.ii - 21.iii.1961, W.W. Brandt, (ANIC), 1♂. Tonga: Tongatapu, Nukunuku, 4.iii.1992, Psidium guajava, Bactrocera facialis & Bactrocera kirki, T1231, Pulotu, (QDPIF), 2♀; Tongatapu, Folaha, 5.iii.1992, Psidium guajava, Bactrocera facialis & Bactrocera kirki, T1243, Pulotu, (QDPIF), 2♂; Tongatapu, Vaini, 5.iii.1992, Terminalia catappa, Bactrocera facialis & Bactrocera kirki, T1250, Pulotu, (QDPIF), 2♂, 1♀; Lakufa'anga, 12.viii.1992, Ochrosia oppositifolia, Bactrocera facialis, T1949, Pontiano, (QDPIF), 2♂, 1♀; Eua,

Lakufa'anga, 12.viii.1992, Ochrosia oppositifolia, Bactrocera facialis, T1957, Pontiano, (QDPIF), 1♀; Tongatapu, Vaini, 15.xii.1992, Artocarpus altilis, Bactrocera facialis & Bactrocera xanthodes, T2112, Tali & Sultoni, (QDPIF), 2♂, 2♀; Tongatapu, Vaini, 15.xii.1992, Artocarpus altilis, Bactrocera facialis & Bactrocera xanthodes, T2114, Tali & Sultoni, (QDPIF), 4♂, 2♀; Tongatapu, Good Samaritan Beach, 15.xii.1992, Ochrosia oppositifolia, Bactrocera facialis, T2115, Tali & Sultoni, (QDPIF), 1♀; Tongatapu, Good Samaritan Beach, 15.xii.1992, Ochrosia oppositifolia, Bactrocera facialis, T2117, Tali & Sultoni, (QDPIF), 7♂, 12♀; Tongatapu, Good Samaritan Beach, 23.xii.1992, Ochrosia oppositifolia, Bactrocera facialis, T2132, Una & Tali, (QDPIF), 2♀; Tongatapu, Kanokopulu, 23.xii.1992, Ochrosia oppositifolia, Bactrocera facialis, T2133, Una & Tali, (QDPIF), 2♂, 4♀; Tongatapu, Taiafo'ou, 23.xii.1992, Cerbera manghas, T2137, Una & Tali, (QDPIF), 1♂; Tongatapu, Masilamea, 28.i.1993, Psidium guajava, Bactrocera facialis, T2193, Pulotu, (QDPIF), 2♂, 2♀; Tongatapu, Masilamea, 11.ii.1993, Psidium guajava, Bactrocera facialis & Bactrocera kirki, T2204, Tevita & 'Una, (QDPIF), 10♂, 3♀; Tongatapu, Good Samaritan Beach, 11.ii.1993, Cerbera manghas, Bactrocera facialis, T2205, Tevita & 'Una, (QDPIF), 2♂, 5♀; Eua, 12.v.1993, Psidium guajava, Bactrocera facialis & Bactrocera kirki, T2252, Pulotu, (QDPIF), 10♂, 8♀; Tongatapu Dateline Hotel, 10.vi.1991, Terminalia catappa, Bactrocera facialis & Bactrocera kirki, TO134, Pontiano, (QDPIF), 2♂; Eua, Eua Forestry Dept., 29.v.1991, Terminalia catappa, Bactrocera facialis & Bactrocera kirki, TO379, Sione & Pulotu, (QDPIF), 10♂, 19♀; Eua, Fuai, 'Eua, 29.v.1991, Psidium guajava, Bactrocera kirki, TO381, Sione & Pulotu, (QDPIF), 6♂, 8♀. Yam Island (Australia): Yam Island, 29.vi.1993, Moraceae, Bactrocera bancrofti, 2158B, J.F. Grimshaw, (QDPIF), 1♂, 2♀. Host Associations B. aglaiae (1), B. bancrofti (Tryon) (1), B. barringtoniae (2), B. curvipennis (2), B. endiandrae (1), B. facialis (1), B. fagraea (2), B. frauenfeldi (2,3), B. jarvisi (1,2), B. kirki (1), B. kraussi (1,2), B. laticaudus (2), B. musae (2,3), B. pallida (1,2), B. passiflorae (1,2,3), B. psidii (2), B. tryoni (1,2,3), B. visenda (Hardy) (1), and B. xanthodes (1,2,3). Discussion According to Fischer (1963a), P. fijiensis should be readily separated from P. novaguineensis on the basis of a broad, infuscated stripe running medially across the forewing. However, in almost all specimens examined from Australia (including some of those previously determined as P. fijiensis by Fischer) this stripe was not

readily apparent. In the specimens examined, the forewing pattern varied from distinctly infumate on the basal half to completely hyaline. The variation appeared to be continuous rather than representing two or more discrete patterns. In the absence of this distinctive wing pattern, we were unable to separate P. fijiensis from P. novaguineensis. The distribution and host records listed above are based solely on museum specimens of P. fijiensis previously determined as that species. All previously undetermined material is listed below under P. novaguineensis for reasons discussed under that species.

Psyttalia muesebecki (Fischer, 1963) Austroopius muesebecki Fischer, 1963a: 182. Psyttalia muesebecki: (Wharton 1987): 63. Taxonomic History Austroopius muesebecki was originally described by Fischer (1963a). Wharton (1987) reviewed the generic ranking, described Psyttalia as a genus, which included this species [Psyttalia muesebecki]. Synopsis Frons and vertex polished and sparsely setose. Clypeus in profile slightly bulged medially; labrum visible in frontal view; gap between mandible and ventral margin of clypeus distinct. Notauli deep at anterior margin, ending before level of tegula and absent posteriorly. Midpit absent; propodeum smooth with median longitudinal carina absent. Sternaulus impressed, short, with just a trace of sculpture. Wing with second submarginal and discal cells infumate. Forewing 2RS shorter than 3RSa; (RS+M)b absent, 2RS in line with m-cu; 2RS thickened medially, junction of 2RS and m-cu thickened. Hindwing RS and m-cu absent. Petiole with dorsal lateral carinae elevated on basal 0.3, diverging and less well developed posteriorly. Metasomal tergum 2 smooth and polished. Ovipositor sheath longer than metasoma. Entire body yellow/orange in colouration. Distribution New Caledonia: Noumea. Type Locality Noumea, New Caledonia. Material examined None. Host Associations Bactrocera psidii (7). Discussion Psyttalia muesebecki can be readily distinguished from other species of Psyttalia by the absence of a propodeal carina. No material was examined in this study.

Psyttalia novaguineensis (Szépligeti, 1900) Austroopius novaguineensis (Szépligeti 1900): 64. Austroopius novoguineensis: (Fischer 1963a): 184. Psyttalia novaguineensis: (Wharton 1987): 63. Taxonomic History Austroopius novaguineensis was originally described by Szépligeti (1900). Fischer (1963a) described this species with an incorrect spelling of the species name, Austroopius novoguineensis. Wharton (1987) reviewed the generic ranking, named Psyttalia as a genus, which included this species, thus renaming this species Psyttalia novaguineensis. Synopsis (Figure 3.12a-f) Frons and vertex polished and sparsely setose. Posterior ocelli margined laterally with deep anteriorly diverging grooves. Occipital carina well developed; clypeus in profile slightly bulged medially; labrum visible in frontal view; gap between mandible and ventral margin of clypeus distinct. Notauli deep at anterior margin, ending before level of tegula and absent posteriorly. Midpit absent; propodeum smooth with a median longitudinal carina. Sternaulus shallow, finely crenulate. Wing colouration variable (see discussion). Forewing r arising slightly basad midpoint of stigma, 2RS shorter than 3RSa; (RS+M)b absent, 2RS in line with m-cu; 2RS thickened medially, junction of 2RS and m-cu thickened. Hindwing RS and m-cu absent. Petiole with dorsal lateral carinae elevated, diverging posteriorly, but extending to posterior margin. Metasomal tergum 2 smooth and polished. Ovipositor sheath 1.5 times longer than metasoma. Entire body yellow/orange in colouration. Distribution Australia: Far north eastern Queensland. Papua New Guinea: Collected from low altitude regions of the main island of New Guinea, the Gazelle Peninsular of East New Britain and Duke of York Island. Type Locality Freidrich-Wilhelmshafen (now Madang), Papua New Guinea. Material examined Australia: North Queensland, Cape Kimberley at RFCK14, 25.iii.1996, Aglaia sapindina, Bactrocera aglaiae, QDPI CK269, (QDPIF), 1♀; North Queensland, Cape Kimberley, 5.viii.1996, Terminalia arenicola, Bactrocera neohumeralis, QDPI CK661, (QDPIF), 1♂, 1♀; North Queensland, Cape Kimberley, 6.viii.1996, Calophyllum inophyllum, Bactrocera calophylli, QDPI CK674, (QDPIF), 1♂, 5♀; North Queensland, Cape Kimberley, 3.viii.1996, Acmena graveoleus, Bactrocera kraussi,

QDPI CK675B, (QDPIF), 2♀; North Queensland, Cape Kimberley, 7.viii.1996, Planchonella, obovoidea, QDPI CK680, (QDPIF), 1♀; North Queensland, Cape Kimberley, 7.viii.1996, Syzygium sp., Bactrocera kraussi & Bactrocera neohumeralis, QDPI CK682, (QDPIF), 2♂, 5♀; North Queensland, Cape Kimberley at RF CK 3, 16.x.1996, Garcinia warrenii, Bactrocera visenda, QDPI CK872, (QDPIF), 2♂, 7♀; North Queensland, Cape Tribulation, 21.viii.1996, Fagraea cambagei, Bactrocera Fagraea QDPI CT1019, (QDPIF), 3♂, 6♀; North Queensland, Cape Tribulation, 17.x.1996, Ficus septica, Clusiosoma, semifuscum, QDPI CT1390, (QDPIF), 4♂, 6♀; North Queensland, Cape Tribulation at RFCT 30, 17.x.1996, Endiandra microneura, QDPI CT1397, (QDPIF), 2♀; North Queensland, Cape Tribulation at RF CT11, 24.x.1996, Endiandra microneura, QDPI CT1437, (QDPIF), 1♂, 2♀; North Queensland, Cape Tribulation at RFCT8, 5.iii.1996, Aglaia sapindina, Bactrocera aglaiae, QDPI CT15, (QDPIF), 1♀; North Queensland, Cape Tribulation at RFCT1, 2.vi.1996, Acmena sp, Bactrocera kraussi, QDPI CT487, (QDPIF), 2♀; North Queensland, Cape Tribulation at RFCT6, 14.v.1996, Fagraea cambagei, Bactrocera fagraea QDPI CT790, (QDPIF), 3♀; North Queensland, Cape Tribulation at RFCT21, 27.v.1996, Calophyllum inophyllum, Bactrocera calophylli, QDPI CT838, (QDPIF), 2♂, 2♀; North Queensland, Cape Tribulation, 7.viii.1996, Endiandra microneura, Bactrocera endiandrae, QDPI CT957, (QDPIF), 3♀; North Queensland, Cape Tribulation at RFCT 44, 15.viii.1996, Endiandra microneura, Bactrocera endiandrae, QDPI CT973, (QDPIF), 2♀; North Queensland, Cape Tribulation at RFCT 40, 15.viii.1996, Acmena graveoleus, Bactrocera kraussi, QDPI CT993, (QDPIF), 2♀; North Queensland, Goldborough Valley at RFGV1, 16.ii.1996, Syzygium tierneyanum, Bactrocera kraussi, QDPI GV2, (QDPIF), 1♂; North Queensland, Goldsborough Valley at RFGV7, 23.i.1996, Aglaia sapindina, Bactrocera aglaiae & Bactrocera tigrina, QDPI GV35, (QDPIF), 1♂, 2♀; North Queensland, Goldsborough Valley at RFGV8, 23.i.1996, Aglaia sapindina, Bactrocera aglaiae, QDPI GV41, (QDPIF), 2♀; North Queensland, Goldsborough Valley at RFGV10, 23.i.1996, Aglaia sapindina, Bactrocera aglaiae, QDPI GV49, (QDPIF), 1♂, 2♀; North Queensland, Kuranda at RFK19, 7.ii.1996, Garcinia warrenii, Bactrocera visenda, QDPI K152, (QDPIF), 1♀; North Queensland, Kuranda at RF K19, 7.ii.1996, Garcinia warrenii, Bactrocera visenda, QDPI K152, (QDPIF), 1♂, 2♀; North Queensland, Kuranda at RFK19, 25.i.1996, Garcinia warrenii, Bactrocera visenda, QDPI K85, (QDPIF), 2♀; North Queensland, Lake Morris at RFLM1, 5.xi.1996, Ficus opposita, QDPI LM1396, (QDPIF), 1♂, 1♀; North Queensland, Mount Lewis at RFLM3, 18.iv.1996, Aglaia sapindina, Bactrocera aglaiae, QDPI ML713, (QDPIF), 2♀; North Queensland, Mulgrave River at RFMR5,

23.i.1996, Barringtonia, calyptrata, Bactrocera barringtoniae, QDPI MR21, (QDPIF), 1♂; North Queensland, Mulgrave River at RFMR8, 23.i.1996, Cryptocarya, sp, Bactrocera endiandrae, QDPI MR29A, (QDPIF), 4♀; North Queensland, Bot. Gardens, Cairns, 19.xii.1995, Barringtonia calyptrata, Bactrocera barringtoniae, QDPI NQ244, (QDPIF), 1♂; North Queensland, Lake Placid, 20.xii.1995, Endiandra longipedicellata, Bactrocera endiandrae, QDPI NQ259, (QDPIF), 2♂, 4♀. Papua New Guinea: E. New Britain, Rabaul Town Market, 30-Mar-00, MYRTACEAE, Syzygium, Bactrocera frauenfeldi & Bactrocera obliqua, I 1428, (GU), 1♀; E. New Britain, Kerevat: L.A.E.S., Kavieng compound, 19-Dec-97, MYRTACEAE, Syzygium malaccense, Bactrocera frauenfeldi & Bactrocera obliqua, I 156-158, (GU), 1♂; E. New Britain, Sonoma: S.D.A. school, mechanics workshop, 22-Dec-97, MYRTACEAE, Syzygium aqueum, Bactrocerafrauenfeldi, I 169-170, (GU), 1♂, 1♀; E. New Britain, Kerevat: L.A.E.S., M. Powell house, 15-Jul-98, MYRTACEAE, Syzygium malaccense, Bactrocera frauenfeldi & Bactrocera obliqua, I 724-725, (GU), 1♀; E. New Britain, Sonoma: S.D.A. school, mechanics workshop, 6-Nov-98, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, I 802-804, (GU), 3♂, 2♀, 1 sex undetermined; E. New Britain, Kerevat: L.A.E.S., near Labourer compound, 9-Feb-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, I 904-905, (GU), 1♀; Central, Laloki block A.Mehuze garden, 19-May-99, MUSACEAE, Musa x paradisiaca, Bactrocera musae, L 2937-2966, (GU), 1♂, 1♀; Madang, Baitabag, 4-Jul-01, RUBIACEAE, Versteegia cauliflora, Bactrocera tinomiscii, M 1625, (GU), 2♀; Morobe, Lae: Omsis forest, 07-Feb-00, COMBRETACEAE, Terminalia, Bactrocera penefurva, N 1259, (GU), 3♂, 1♀; Morobe, Lae: Wampit village area, 16-Feb-00, MUSACEAE, Musa, Bactrocera musae, N 1263, (GU), 1♀; Morobe, Lae: Omsis forest, 16-Feb-00, GNETACEAE, Gnetum gnemon, Bactrocera B.(Bulladacus) n.sp., N 1272, (GU), 1♂; Morobe, Lae: Omsis forest, 16-Feb-00, COMBRETACEAE, Terminalia, Bactrocera frauenfeldi & Bactrocera penefurva, N 1273, (GU), 5♂, 4♀; Morobe, Lae: Omsis forest, 06-Apr- 00, GNETACEAE, Gnetum gnemon, Bactrocera B.(Bulladacus) n.sp., N 1383, (GU), 1♂; Morobe, Lae: Gabensis Village area, 06-Apr-00, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 1384, (GU), 5♀; Morobe, Lae: Gabensis Village area, 27-Apr-00, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 1404, (GU), 2♀; Morobe, Lae: FRI forest/ botanical gard, 16-May-00, Bactrocera frauenfeldi & Bactrocera trivialis, N 1462, (GU), 1♂; Morobe, Lae: Omsis forest, 11- Jul-00, Bactrocera vulgaris (to check), N 1562, (GU), 3♂, 6♀; Morobe, Lae: Lae town, FRI office area, 13-Apr-99, COMBRETACEAE, Terminalia catappa, Bactrocera exima & Bactrocera frauenfeldi, N 325, (GU),1♀; Morobe, Lae: Gabensis

village area, 27-Apr-99, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi, N 391, (GU), 1♂; Morobe, Lae: 11-mile Suambu farm, 03-May-99, MYRTACEAE, Syzygium Bactrocera frauenfeldi & Bactrocera trivialis, N 449, (GU), 1♂, 1♀; Morobe, Lae: Markham farm, Mararumi, 25-Aug-99, MYRTACEAE, Psidium guajava, Bactrocera trivialis & frauenfeldi, N 795, (GU),1♀; Morobe, Lae: Gabensis village area, 05-Oct-99, MUSACEAE, Musa, x paradisiaca, Bactrocera N 981, (GU), 1♂, 1♀; E. New Britain, Kerevat C.I.S. prison, 6-Aug-98, MYRTACEAE, Psidium guajava, Bactrocera frauenfeldi & Bactrocera obliqua, R 2062, (GU), 2♂, 4♀, 1 sex undetermined; E. New Britain, Kerevat: L.A.E.S., Orchard nr. Country Club, 20-Oct- 99, MYRTACEAE, Syzygium malaccense, Bactrocera frauenfeldi, I 1159-1168, (GU), 1♀; E. New Britain, Kerevat: L.A.E.S., Kavieng cpd Godrey house, 7-Jul-98, MYRTACEAE, Syzygium malaccense, Bactrocera frauenfeldi, I 650, (GU), 1♂; E. New Britain, Duke of York Is: Butlivuan Comm. School, 4-Jun-99, Aila (DOY language), frauenfeldi, I 957, (GU), 1♂; Central, Lobulogo village Hiritano Highway, 5-Jun-99, OXALIDACEAE, Averrhoa carambola, frauenfeldi, L 3099-3142, (GU), 1♂, 2♀; Madang, Baitabag, 27-Jun-01, APOCYNACEAE, Cerbera manghas, Bactrocera sp.3, M 1592, (GU), 2♀; Madang, Baitabag, 4-Jul-01, APOCYNACEAE, Cerbera manghas, Bactrocera sp.3, M 1621 (B), (GU), 1♂, 3♀; Madang, Baitabag, 4-Oct-00, APOCYNACEAE, Neisosperma oppositifolia, Bactrocera sp 1., M 399, (GU), 6♂; Madang, Ohu, 13-Oct-00, APOCYNACEAE, Cerbera manghas, Bactrocera sp.3, M 430, (GU), 3♀; Madang, Ohu, 19-Oct-00, RUBIACEAE, Morinda bracteata, Euphranta perkinsi, M 446, (GU), 5♀; Morobe, Lae: FRI forest/ botanical gard, 31-May-00, calophylli, N 1502, (GU), 1♂; Morobe, Lae: Omsis forest, 30-Jul-00, Xanthophyll, Xanthophyllum papuana, musae complex, N 1567, (GU), 1♀; Morobe, Lae: Bubia Agr. Station, 27-Dec-98, GNETACEAE, Gnetum gnemon, B.(Bulladacus)n.sp., N 161, (GU), 1♂, 1♀, 1 sex undetermined; Morobe, Lae: Gabensis village area, 27-Apr-99, MYRTACEAE, Psidium, guajava, N 394, (GU), 1♀; Madang, Madang: Miss Consv area, 28-Jun-99, B. (Gymnodacus) n.sp., N 623, (GU), 1♀; Morobe, Lae: Gabensis village area, 05- Oct-99, MUSACEAE, Musa x paradisiaca, Bactrocera musae, N 972, (GU), 1♂; Western, Waidoro village (Western Prov.) 15-Oct-99, MUSACEAE, Musa x paradisiaca, musae, S 619, (GU), 1♀. Host Associations B. aglaiae (1), B. barringtoniae (1), B. calophylli (Perkins & May) (1), B. endiandrae (1), B. fagraea (1), B. frauenfeldi (1), B. kraussi (1), B. musae (1), B. neohumeralis (1), B. penefurva (1), B. tinomiscii (1), B. visenda (1), B. (Bulladacus) sp. (1), and B. (Gymnodacus) sp. (1).

Discussion Psyttalia novaguineensis is defined by (Fischer 1963a) as having a uniformly hyaline or yellowish wing. The wing of P. fijiensis, in contrast, was described by the same author as having a broad infuscated band across the middle of the forewing. An indistinct band matching this description was visible in only a very few previously determined specimens of that species. However, several of the undetermined specimens examined by us have infuscate mottling all across the basal half of the wing. The infuscation appears to be more distinct in the males but there is also variation across females. The colour pattern is different from the broad median band described for P. fijiensis. A careful examination of material from northeastern Queensland, including both reared and unreared material, indicates that at least five species of Psyttalia occur in this area, four of which have an “Austroopius” wing vein pattern (sensu Fischer 1963a: with a medially thickened 2RS). The most obvious difference among these species is in the length of the ovipositor. The reared material falls into three general categories: 1) ovipositor sheath about as long as mesosoma; 2) ovipositor sheath nearly twice as long as mesosoma; and 3) ovipositor sheath more than twice length of mesosoma. Previously undetermined material in which the ovipositor sheath is nearly twice as long as the mesosoma is tentatively placed under the name P. novaguineensis, since they most closely fit the redescription by Fischer (1963a). Note, however, that Szépligeti (1900) originally described the ovipositor of this species as very short. Further study is required in order to correctly place these specimens.

Figure 3.12. Psyttalia novaguineensis. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view; e, propodeum, dorsal view; f, metasoma, dorsal view.

Utetes cf. albimanus Synopsis (Figure 3.13) Frons and vertex densely setose; and densely but finely punctate. Frons with median pit adjacent anterior ocellus, posterior ocelli margined laterally with weak, anteriorly diverging grooves. Occipital carina well developed. Clypeus in profile weakly protruding, overhanging base of labrum; clypeus in frontal view large and hemispherical, ventral margin thin, truncate to weakly convex; labrum visible in frontal view; gap between mandible and ventral margin of clypeus distinct but not large. Antennae 41 segmented. Notauli deep from anterior margin to level of tegula and absent or nearly so posteriorly. Midpit long, narrow, deep. Propodeum rugose throughout. Sternaulus broad, deep, crenulate throughout. Wings infumate. Forewing r arising from midpoint of stigma, 2RS slightly shorter than 3RSa, (RS+M)a weakly sinuate; (RS+M)b absent, m-cu entering second submarginal cell. Hind wing RS and m-cu absent. Basal carina on posterior side of hind tibia well developed (Fig.17). Petiole with dorsal lateral carinae strongly elevated over basal half, much weaker posteriorly, but extending to posterior margin. Metasomal tergum 2 smooth and polished. Head and mesosoma dark brown to black; metasoma dorsally black. Coxae and trochanters brown to dark brown, fore femur and tibia yellow brown, mid and hind femur and tibia brown to dark brown, tarsi 1-4 white, 5th tarsus brown. Distribution Papua New Guinea: Lae. Material examined: Papua New Guinea: Morobe, Lae: Omsis forest, 16-Feb-00, GNETACEAE, Gnetum gnemon, Bactrocera (Bulladacus) n.sp., N 1272, (GU), 1♂. Discussion Information based on a single male. The tibial carina and a small, but distinct, gap between clypeus and mandibles clearly place this species in the genus Utetes Foerster as delimited by (Wharton 1997a). The holotype of Utetes albimanus (Szépligeti), in the Hungarian Natural History Museum was examined by R.A.Wharton. and appears to be identical, but some slight uncertainty must remain because the holotype is damaged and some features therefore cannot be compared. The second submarginal cell is somewhat shorter than in many species of Utetes, but the general pattern of wing venation is otherwise typical of Utetes.

Figure 3.13. Utetes cf. albimanus. Habitus.

Utetes perkinsi (Fullaway, 1950) Opius perkinsi (Fullaway 1950a): 66. Opius (Opius) perkinsi: (Fischer 1963b): 210. Opius (Utetes) perkinsi: (Fischer 1987): 223. Utetes perkinsi: (Wharton 1997a): 30. Taxonomic History This species was originally described as Opius perkinsi by Fullaway (1950). Fischer (1963b) reviewed the subgeneric position and placed the species within the subgenus Opius [Opius (Opius) perkinsi]. In 1987, after further examination Fischer revised the placement and designated the species to fit within subgenus Utetes [Opius (Utetes) perkinsi]. Utetes was subsequently elevated to generic rank (Wharton 1997a) thus the species is named Utetes perkinsi. Synopsis (Figure 3.14a-d) Frons and vertex polished and very sparsely setose and punctate. Frons with median pit adjacent anterior ocellus, posterior ocelli margined laterally with weak, anteriorly diverging grooves. Occipital carina well developed. Clypeus in profile weakly protruding, overhanging base of labrum; clypeus in frontal view large and hemispherical, ventral margin thin, truncate to weakly convex; labrum visible in frontal view; gap between mandible and ventral margin of clypeus distinct but not large. Notauli deep from anterior margin to level of tegula, continuing very weakly posteriorly. Midpit long, narrow, deep; propodeum weakly rugulose to rugose, usually with irregular transverse carina. Sternaulus broad, deep, weakly crenulate throughout. Wings infumate. Forewing r arising basad midpoint of stigma, 2RS notably shorter than 3RSa, (RS+M)a sinuate; (RS+M)b absent, m-cu entering second submarginal cell. Hind wing RS and m-cu absent. Basal carina on the posterior side of the hind tibia well developed (Figure 3.15). Petiole with dorsal lateral carinae strongly elevated over basal two-thirds, much weaker posteriorly, but extending to posterior margin. Metasomal tergum 2 smooth and polished. Head dark red-brown to black; mesosoma yellow-orange; apical metasomal terga black. Fore and mid legs yellow orange, except tarsi 5 brown to dark brown. Hind coxa and trochanter brown, femur, tibiae and tarsi dark brown. Distribution Australia: Far north eastern Queensland. Type Locality Deeral, Australia

Material examined Australia: Deeral, 1944, N.L.H. Krauss, (ASCU), 1♂, 2♀; North Queensland, Babinda Boulders at RFBB5, 20.v.1996, Acmena sp, Bactrocera kraussi, QDPI BB159, (QDPIF), 1♂; North Queensland, Cape Kimberley, 7.viii.1996, Syzygium sp., Bactrocera kraussi & Bactrocera neohumeralis, QDPI CK682, (QDPIF), 2♀; North Queensland, Mulgrave R, 14.ii.1996, Bactrocera neohumeralis, QDPI MR8, (QDPIF) 3♀. Host Associations B. cacuminata (2), B. jarvisi (2), B. kraussi (1,2), B. laticaudus (2), B. neohumeralis (1), and B. tryoni (2,3). Discussion Utetes perkinsi is similar to U. cf. albimanus in having a basal carina on the posterior side of the hind tibia. These species can be separated from each other on the basis of colouration and notauli development.

100

Figure 3.14. Utetes perkinsi. a, head, anterior view; b, head, lateral view; c, mesosoma, lateral view; d, mesonotum, dorsal view.

Figure 3.15. Hind femur and tibia showing tibial carina (arrow) typical of Utetes.

101

Chapter 4 Identification and Diagnostics

Introduction Taxonomy, by definition, is the science of naming and classifying organisms, while Identification is the process of ascertaining the taxonomic identity of an organism (Cranston et al. 1991). In Chapter 3 I present the major taxonomic component of my thesis, but also include, by way of a traditional paper based dichotomous key, one identification component. In this chapter, I explore the identification process further and also move into diagnostics - the field of distinguishing or differentiating a species, genus, etc from others which are similar. Diagnostics packages are often more than just a key; they can contain a variety of supporting information in which the identification method forms just a small part (Stevenson et al. 2001).

Identification Keys A key is a tool that is used to identify specific objects or specimens. Keys are a series of choices or questions that guide the user to the correct name of a specimen (Anonymous 2004; Walter & Winterton 2007). Most keys fit into one of two types, ‘dichotomous’ or ‘multi-access’. As the name suggests, dichotomous keys, which are those traditionally used in taxonomy, give the user two choices in each step (or couplet). Ideally a perfect couplet gives the user a “yes/no” pair of questions and divides unidentified taxa equally into either the “yes” or “no” groups, or alternatively separates some uniquely grouped set of taxa (eg a genus or subgenus) from all other taxa. The identification process continues through successive couplets until each group, becoming progressively smaller as the key progresses, resolves itself to the identification of one or two taxa (Stevenson et al. 2001). Unfortunately taxonomic character states are often not clear cut, meaning that couplets can commonly become complex and rarely perfectly separate taxa or groups of taxa.

Multi-access [= matrix keys], in contrast, also use a process of elimination, but in this case the user is presented with an unstructured question set, each question of which asks the user to describe features of the specimen (Stevenson et al. 2001). The questions can be answered in any order and multiple features can be selected in any question. By selecting those features relevant to the specimen to be identified, the taxa that do not match the selected choices are immediately rejected. The process continues until only one or a short list of names remains, identifying the specimen or at least providing a short list of possible identities (Anonymous 2004).

102

Multi-access keys have been demonstrated to have many advantages over traditional dichotomous keys (White & Sandlant 1998). Reduced chances of error, a reduction in the number of questions required to arrive at a solution, and the elimination of the ‘unanswerable couplet’ problem are three.

Dichotomous keys are traditionally printed; however, these can be readily converted to be internet based and somewhat more interactive with the use of hyperlinks and basic web programming (eg Carmichael et al. 2008). Multi-access keys are almost invariably computer or web-based and, when first starting to appear commonly in the 1990s, were often incorrectly labelled as “computer-based” keys. There are a number of computer programs available that can be used to build multi-access keys (Dallwitz 2007a), including Lucid (Anonymous 2004), DELTA (DEscription Language for TAxonomy) (Dallwitz 2007b) and INTKey (Dallwitz 2008) (Walter & Winterton 2007). The most widely used is the Lucid software package and Lucid keys are available both online (www.lucidcentral.com) and via CD-Rom publications (e.g. Walter & Proctor 2001; Lawson et al. 2003; Seeman & Beard 2007; Malipatil & Ridland 2008). One of the problems with multi-access keys is that not all computer systems will support or enable access to their use; however, this issue is being overcome with new compatibility requirements.

The processes by which dichotomous and multi-access key structures are developed have some similarities. Initially, all keys start with a list of taxa to be identified and a series of character states upon which the taxa will be separated. To create a dichotomous key, the initial couplet is designed to divide off either a single specific taxa group (eg a subgenus) or cut the potential taxa into two groups of approximately equal size. The first couplet in my dichotomous key in Chapter 3, for example, involves the presence or absence of a basal carina on the hind tibia. The genus Utetes is characterised by the presence of the hind tibial basal carina and, thus, this genus is split off at the very beginning of the key. Only species in the genus Utetes need then be considered in couplet 2, allowing the character state question in that couplet to be specific to that small group. Conversely, character state questions asked in couplets deeper within the key need have no relevance to Utetes species, as these species have already been resolved.

In contrast, when developing a multi-access key, every taxa must be scored for each character state, as questions can be answered in any order and the key works by considering all [non-discarded] taxa simultaneously. The structure of a multi-

103

access key does not allow the key developer to direct questions at a smaller or specific group, as in Utetes example above. While quicker in identifying taxa for the end user, the development and scoring of a multi-access key is more difficult than scoring a dichotomous key as every character state for every taxon needs to be examined and entered into the key. For these reasons, it is generally not possible to take a dichotomous key and turn it into a multi-access key, because a dichotomous key will not contain all the information a multi-access key will require (Stevenson et al. 2001).

To illustrate the scoring differences between dichotomous and multi-access keys, I present below a simple dichotomous key to four fruits; Orange, Grapefruit, Banana and Strawberry.

Dichotomous Fruit Identification Key: 1a. Large (greater than 8cm in diameter) sized fruit ...... 2 1b. Small (less than 8cm in diameter) sized fruit ...... Strawberry

2a. Fruit is predominantly yellow in colour ...... 3 2b. Fruit is predominantly orange in colour ...... Orange

3a. Fruit is round in shape ...... Grapefruit 3b. Fruit is elongate...... Banana

Converted to a matrix format, the scoring matrix for this dichotomous key is presented in Table 4.1. To convert this dichotomous key into a full multi-access key additional scores and additional character states are required. Strawberry, for example, is identified in the first couplet of dichotomous key and so is not scored for character states dealing with fruit colour or shape. These states need to be scored for a multi-access key (Table 4.2).

104

Table 4.1 Character state matrix required for creating the dichotomous fruit identification key above. “1” indicates that the character state was scored positively for the particular fruit type. Blank cells are not negative scores, but illustrate where a character state did not need to be scored for the key to be developed.

Strawberry Orange Grapefruit Banana Fruit size: Large 1 1 1

Fruit size: Small 1

Colour: Orange 1 Colour: Yellow 1 1

Shape: Round 1

Shape: Elongate 1

Table 4.2 Character state matrix required for creating a multi-access fruit identification key. The presence or absence of a character state is indicated by the following code: 0 absence; 1 presence; 2 possibly misinterpreted.

Strawberry Orange Grapefruit Banana Fruit size: Large 0 1 1 1

Fruit size: Small 1 0 0 0

Colour: Orange 0 1 0 0 Colour: Yellow 0 0 1 1

Colour: Red 1 0 0 0

Shape: Round 2 1 1 0

Shape: Elongate 0 0 0 1

Shape: Heart-shaped 1 0 0 0

Both keys are used for identifying the same taxon set and use similar character states. As can be seen, however, the dichotomous matrix has many cells remaining empty while the multi-access matrix is complete. If some cells remain empty in a multi-access matrix, species can be erroneously discarded as the key interprets an empty cell as being negative for the character state. From a developmental point of

105

view, far greater effort is required to develop a multi-access key, however, experience has shown that usability and functionality of the resulting key is greater, particularly for non-experts.

Diagnostic Tools The actual definition and requirements of what constitutes a diagnostic tool is an often debated point (Cranston et al. 1991; Bisby et al. 2002; Anonymous 2004; Dallwitz 2007a; Walter & Winterton 2007). Diagnostic tools may or may not contain identification keys as well as information on an organism’s biology, ecology and behaviour, morphological descriptions, photographs, illustrations and geographical range information (Walter & Winterton 2007).

A spectrum of packages that can be described as diagnostic tools, are available online or on CD. At one end are the broad user based packages, such as PaDIL (www.padil.gov.au); an image rich identification resource that contains biological, taxonomic and photographic descriptions of over 2500 taxa. At the other end are what is often known as lucid style packages, such as Dorsalis (Lawson et al. 2003), a specialist identification package to a small complex of pest fruit fly species. These lucid-style packages have at their core a multi-access identification key developed using Lucid software, on to which supporting information, tutorials and fact sheets are often attached. The packages that do not contain identification keys often have other search-driven ways of identifying the taxon in question. It would be almost impossible to develop and maintain an identification key to very large and/or diverse taxon datasets. For example, if PaDIL was an identification key, each time a new taxon was to be entered the identification key would need to be re-developed. This in itself is not a problem for a multi-access key development package when the taxon list is related (eg within an order, family, or genus), but in the PaDIL example the database contains a taxonomically diverse range of organisms, (eg insects, plant pathogens and marine pests) and maintaining a functional matrix would be very difficult, if not impossible.

Structure of Chapter

To help alleviate some of the problems with identification of the parasitoids in the Pacific region, an interactive diagnostics package (“OpiineID”) was developed, the centre of which is a Lucid based multi-access key. The diagnostics package is

106

computer based, so there is no pictorial (SEM photographs, full colour photographs, line drawings and fully rendered illustrations) limit, unlike printed keys, which are often limited by cost and print space. The identification key is only one small component of OpiineID and the key is supported by fact sheets with morphological descriptions, host associations, geographical information and images.

OpiineID provides a platform enabling users to perform identifications and access both general (host association and geographic distribution) and taxonomic information (morphological descriptions and synonymies). This makes this package a ‘one stop shop’ for taxonomy and identification of tephritid fruit fly parasitoids in the Pacific region. It is intended that OpiineID be a functional tool for fruit fly workers of the region and will aid biological control and IPM decisions by allowing rapid diagnostics. In addition to OpiineID, diagnostic images were also placed on the PaDIL web-site as an example of an alternative diagnostic tool.

Materials and Methods

OpiineID Multi-access key development The Lucid package was chosen to develop a multi-access key to the opiine braconids in my research, as it is commonly used in both entomological and taxonomic applications. Within Lucid, there are six possible scores to describe presence or absence of a character state, but only four of these were necessary within the OpiineID matrix and these are: Absent - the state does not occur in the entity. Common - the state occurs commonly (or always) in the entity. Rare - the state occurs rarely in the entity. Uncertain - it is not known whether the state occurs in the entity or not

The multi-access matrix developed for the key contains 460 character scores (Table 4.3).

Additional resources within OpiineID In addition to the development of the identification tool, supporting information was required to make OpiineID a true diagnostic tool, rather than simply an identification key. Morphological tutorials were developed requiring 60 line illustrations,

107

demonstrating each morphological region or term. A comprehensive fact sheet was completed for each of the 14 wasp species, including full colour and scanning electron photographs of each body region from various angles and full rendered illustrations. A tutorial page was created for each character state within the key; these were constructed from full colour or SEM photographs as appropriate. Each of these pages were programmed for access within the OpiineID homepage and from within the Lucid key component.

108

Table 4.3. Character state matrix required for creating the multi-entry or matrix key to opiine fruit fly parasitoids of the South Pacific as presented in OpiineID. The presence or absence of a character state is indicated by the following code: 0 absent; 1 present; 2 rarely present; 3 uncertain.

LUCID MATRIX Diachasmimorpha fullawayi fullawayi Diachasmimorpha hageni Diachasmimorpha kraussii Diachasmimorpha longicaudata Diachasmimorpha tryoni Diachasmimorpha arisanus Fopius cf. vandenboschii Fopius deeralensis Fopius ferrari Fopius Fopius illusorius schlingeri Fopius vandenboschii Fopius Opius froggatti concolor Psyttalia Psyttalia fijiensis Psyttalia novaguineensis Psyttalia incisi* Psyttalia muesebecki Utetes cf. albimanus Utetes perkinsi HEAD Labrum:Labrum exposed, not concealed by clypeus 0 0 0000 0 00000111111 0 0 Labrum:Labrum weakly exposed, visible in frontal view 0 0 0000 0 00000000000 1 1 Labrum:Labrum concealed by clypeus 1 1 1111 1 11111000000 0 0 THORAX Notauli:Crenulate / sculptured 000001 1 11111000000 0 0 Notauli:Smooth 111110 0 00000011111 1 1 Midpit of Mesoscutum:Absent 000000 0 00000011111 0 0 Midpit of Mesoscutum:Present 111111 1 11111100000 1 1 LEGS Hind Tibia - Basal Carina:Present 000000 0 00000000000 1 1 Hind Tibia - Basal Carina:Absent 111111 1 11111111111 0 0 ABDOMEN Metasomal tergum II:Smooth and polished 3 1 1010 0 11110111131 1 1 Metasomal tergum II:Longitudinally striate 3 0 0101 1 12001000030 0 0

109

Petiole:Weakly longitudinally striate 3 0 1000 0 11010000000 0 0 Petiole:Longitudinally striate 3 0 0101 1 00101000000 0 0 Petiole:Smooth 310010 0 00000111111 1 1 COLOURATION Head Colouration:Dark brown to black 000000 0 01000000000 1 1 Head Colouration:Yellow-orange 111110 1 10211111111 0 0 Head Colouration:Orange-Brown 000001 0 00100000000 0 0 Mesosomal Colouration:Dark Brown to black 000001 0 01100000000 1 0 Mesosomal Colouration:Orange-Brown 0 0 0001 0 00100000000 0 1 Mesosomal Colouration:Yellow-Orange 111110 1 10011111111 0 1 Metasomal Colouration:Dark brown to black 010011 0 01100000000 1 0 Metasomal Colouration:Predominantly yellow- orange with areas of dark brown to black 000010 1 00001000000 1 1 Metasomal Colouration:Yellow-Orange 101100 1 10010111111 0 0

110

PaDIL (Pest and Diseases Image Library – www.padil.gov.au) Each species contained within the OpiineID package has also been uploaded onto the PaDIL website. All morphological colour photographs and scanning electron micrographs have been uploaded via the content editors upload portal. Reference list, host records, distribution data and brief morphological descriptions of the species are also included. I have not been directly involved in the programming of the PaDIL website, and take no credit for the functionality of the website. However, I have been heavily involved in the development and population of the site and have produced a users guide to the site (Appendix 3).

111

Results

OpiineID The OpiineID key is available on the accompanying CD on the back page of this thesis.

Installation of Lucid software From the CD, double click on the install.html icon. If required, click on the top yellow menu bar to ‘Allow Blocked Content’. If required, install Java Virtual Machine by clicking on the link on the left hand side of the page. Otherwise, click ‘Install Lucid 3 Player’. Underneath the ‘Recommended Installation for your platform title, click ‘Download Installer for windows (your operating system will automatically be detected). Run the file when asked, this occurs twice. Follow the prompts until you reach the ‘Done’ button. Close the window.

Instructions for OpiineID From the CD, open the OpiineID folder. Double click the index.html file to open. If required, click on the top yellow menu bar to ‘Allow Blocked content’. To open the key, click ‘Identify’. You will be asked to verify that the content is safe and then once again, click on the top yellow bar to ‘Allow Blocked content’.

A tour of OpiineID OpiineID opens to display the home page with a menu bar down the left hand side of the screen. Separate web pages are available for each species, character state, morphological help (ie tutorials), and a bibliography. The menu bar is available on each page within the website; to navigate to another page, click any item in the menu bar. Each of the items in the menu bar is described in detail below.

Identify Click on ‘Identify’ to enter the Lucid key. The opening page of OpiineID is split into 4 panels. The top left hand panel displays the Features available or character states of your specimen. Clicking on the + beside each feature title, will display the couplet choices available. To make a selection, check the box beside the matching feature. Multiple features can be selected and features can be selected in any order. The top right hand panel (Entities Remaining) displays a list of potential species. After selecting a matching feature, the list of potential species may be reduced. The lower left hand panel (Features Chosen) displays a list of your selections from the features panel. The

112

lower right hand panel (Entities Discarded) displays species which do not match your selection. Continue describing your specimen to the key by selecting features in the features available panel, until one or a few taxa remain. You can select multiple states of any character and you can describe characters in any order. To view a larger version of the image, click on the thumbnail, this will open the image in a new window. To view the factsheet about each species or feature, click on the factsheet icon ( ), this will open the fact sheet in a new window. For a more detailed description of the fact sheets, see ‘species pages’ or ‘character pages’ below.

Species Pages The ‘Species Page’ menu has a full list of all species with fact sheets found within OpiineID. By clicking on a species name you can view the fact sheet for that species. The species fact sheet presents information on synonymies, a morphological synopsis, type locality, material examined and host information. At the top of the page is a link to images, clicking on this link will take you to an interactive page of images, scrolling your mouse across different parts of the stylised wasp displays scanning electron microscope images and full colour photographs of that particular part of the wasp species. The species fact sheets are available from either the navigation menu or from within the identification key itself.

113

Table 4.4. An example species page within OpiineID: Fopius ferrari Carmichael and Wharton. Fopius ferrari Carmichael & Wharton

- Taxonomic Information - Images

114

115

116

Etymology Named after Dr Paul Ferrar, recently retired as Crop Sciences II Programme Manager at the Australian Centre for International Agricultural Research. Paul's long-term support for regional fruit fly research lead directly to several projects which helped generate the parasitoid collections upon which this paper is based.

Synopsis Frons and vertex densely covered with white, decumbent setae; densely but finely punctate. Frons with polished, elevated projection extending from median ocellus almost to torulus; ocellar triangle otherwise margined by an impressed line. Occipital carina well developed, in lateral view extending dorsally nearly level with top of eye. Clypeus smooth, polished, very sparsely covered with long, erect setae, ventral margin obtusely angulate, forming a distinct median tooth; clypeus bulging along midline in profile. Notauli deep and foveolate-crenulate to midpit; midpit extending narrowly to posterior margin of scutum; propodeum exceptionally densely setose and rugose, the sculpture largely without obvious pattern except for median carina on basal 0.3; postpectral carina present. Sternaulus broad, crenulate/rugose throughout. Wings hyaline to weakly infumate. Forewing r arising slightly distad midpoint of stigma, 2RS longer than 3RSa, (RS+M)a weakly sinuate; (RS+M)b absent, m-cu arising directly in line with 2RS. Hindwing RS absent; m-cu developed and pigmented, but not reaching wing margin. Petiole weakly bicarinate longitudinally, otherwise polished, setose and weakly striate. Metasomal tergum 2 smooth (occasionally weakly striate) and densely setose; tergum 3 densely setose

117

at least laterally. Ovipositor sheath almost twice length of metasoma; ovipositor not narrowed apically. Entire body dark brown to black. Fore and mid legs brown, hind legs dark brown to black.

Diagnosis This species is similar to the Philippines species F. skinneri (Fullaway) in coloration, but differs from F. skinneri and darker specimens of F. arisanus by the absence of striae on metasomal tergum 2. The pattern of metasomal color and sculpture thus closely resembles that of F. illusorius. The clypeus, however, differs from that of F. illusorius and is similar to that of F. deeralensis . Fopius deeralensis is variable in T2 sculpture but is readily differentiated from the new species because of its yellow to orange coloration.

Distribution Papua New Guinea Collected from low altitude regions of the main island of New Guinea.

Types

Holotype ; 3 labels, Label 1: Morobe Province , Lae: Bundun Conf Centre, 1.vi.2000, Label 2: Bred from: Persea americana Lauraceae N1516, Label 3: In association with [Bactrocera] frauenfeldi (QM). Paratypes;

1 (ANIC): 3 labels, Label 1: PNG Morobe Province, Lae, Wampit village area, 16.ii.2000, Label 2: Bred from Musa?, Musaceae, N 1263, Label 3: In association with [ Bactrocera ] musae.

1 (ANIC): Label 1: PNG Morobe Province Lae, Bukawa Tikeling 2 forest 27.v.1999 Label 2: Bred from ? N557 Label 3: In association with [Bactrocera] redunca;

3 (ANIC): 3 labels, Label 1: PNG Morobe Province. Lae, Wampit village area, 16.ii.2000, Label 2: Bred from Musa?, Musaceae, N 1263, Label 3: In association with [Bactrocera] musae.

2 , 2 (QM): 3 labels, Label 1: PNG Morobe Province, Lae: FRI forest botanical garden, 16.v.2000, Label 2: Bred from: ? N 1462, Label 3: In association with [Bactrocera] frauenfeldi and [Bactrocera] trivialis.

3 (QM): 3 labels, Label 1: PNG Morobe Province, Lae, Gabensis Village,

118

6.iv.2000, Label 2: Bred from: Psidium guajava, Myrtaceae, N1384, Label 3: In association with [Bactrocera] frauenfeldi.

2 , 1 (QM), 1 (ANIC): 3 labels, Label 1: PNG Morobe Province, Lae, Omsis forest, 11.vii.2000, Label 2: Bred from: ?, N1562, Label 3 In association with [Bactrocera] vulagaris [sic].

1 (QM): 2 labels, Label 1: PNG Central Province , Bereina Station, Joe Aisa residence, 17.v.1999, Label 2: Bred from Averrhoa carambola Oxalidaceae, L 2552.

Other Material Examined PAPUA NEW GUINEA: Madang Province, Baitabag, 16.viii.2000, ex Neisosperma oppositifolia, M 220, 10 sex undetermined (specimens damaged); Morobe Province, Lae: Bukawa, Tikeling 2 forest, 27.v.1999, in association with Bactrocera redunca, N 557, 1; Morobe Province, Lae: Omsis forest, 1.vi.2000, in association with Bactrocera trivialis?, N 1513(A), 1; Morobe Province, Lae, Gabensis Village, 6.iv.2000, ex Psidium guajava, Myrtaceae, in association with [Bactrocera] frauenfeldi N1384, 1 , 1 sex undetermined (specimen damaged).

Discussion Although this species appears at first glance to be just a somewhat darker, more densely setose variety of F. illusorius, the ovipositor and clypeus are sufficiently different to warrant description as a separate species. The tip of the ovipositor is not narrowed as it is in F. illusorius and F. arisanus.

119

Character Pages The ‘Character Pages’ access a full list of character states used within the key. Clicking on each character state accesses an image and description of the character state of interest. Some of the images within these pages are interactive and may be viewed by scrolling a mouse over the image to see an arrow pointing out the exact feature. The character page fact sheets are available from either the navigation menu or from within the identification key itself.

Figure 4.1. An example of a character page with OpiineID.

Morphology The ‘Morphology’ link takes the user to a comprehensive morphological tutorial library. Clicking on a body region will lead to a tutorial for that particular area. Scrolling a mouse over the terminology will highlight the body part defined by each term. Table Table 4.5 is a static version of one of these pages for the face.

120

Table 4.5. Static version of morphological tutorial of the face.

Static Image

Upon mousing over the words, the image displayed changes to:

121

122

Bibliography The bibliography link takes the user to a comprehensive bibliography of all taxonomic references pertaining to the opiine species that OpiineID covers. An alphabetical quick-link bar at the top of the page allows rapid access to papers based on first letter of the primary author’s surname.

PaDIL The relevant Opiine pages of PaDIL have been included on the accompanying CD. These pages are also available on the PaDIL website (www.padil.gov.au) and can be found via the Target Species menu or by entering ‘Braconidae’ into the search bar. Figure 4.2 is a screen capture of the species menu. Figure 4.3 is a comparative image table of three of the species; Fopius deeralensis, Fopius schlingeri and Fopius arisanus. Figure 4.2. Screen capture of the species menu of the Braconidae within the PaDIL website.

123

Figure 4.3. A comparative image table illustrating three morphological characters of three Fopius species; Fopius arisanus Fopius deeralensis and Fopius schlingeri.

Discussion This chapter (including the electronic components) presents two “soft” (i.e. computer based) formats for the delivery of the traditional taxonomic information generated in Chapter 3. Neither is better than the other, they are simply alternative delivery styles. Lucid based multi-access keys can supply more supporting information than PaDIL is designed to do and can “walk” a user through an identification, while PaDIL is better designed to handle large and diverse taxa data sets.

PaDIL is a stand alone, custom written data-base with a single manager, and as such has strong internal quality controls. Multi-access keys, in contrast, can now be made by anyone with access to a computer and one of several commercially available software packages (Walter & Winterton 2007). Given this, it is not surprising that publically available multi-access keys are extremely variable in their functionality and quality. There are some very good and very poor examples of

124

multi-access keys. The common flaws in the less successful keys include some or all of the following; lack of good quality illustrative material; absence of informative fact sheets; and reliance on technical jargon.

The Polyphagous Agromyzid Leafminer (Malipatil & Ridland 2008) Lucid key illustrates a multi-access key with weaknesses. This key has ample illustrations and photographs, but lacks informative fact sheets for each couplet step and does not assist those new to the field of Agromyzidae identification. The CD contains instruction pages detailing specimen collection and genitalia preparation, but stops short of assisting throughout the identification process. In contrast Mites in Soil (Walter & Proctor 2001), despite being developed and published in an older Lucid version with limited functionality, is easy to use, informative and has help tutorials throughout the identification process and factsheets on each species to aid in the confirmation of the identification.

Plankton.net (http://planktonnet.awi.de/#content) is a web based multimedia package that does not utilise the Lucid software, but rather relies of various search functions. Without an extensive prior knowledge, this package would be difficult to identify a species of interest. Each species page has excellent high magnification images and detailed three-dimensional mapping images, but no useful identification tools. TaroPest (Carmichael et al. 2008) is another non-Lucid based diagnostic package. Initially a Lucid based project to develop a taro (Colocasia esculenta) pest and diseases toolkit, the Lucid component was eventually abandoned due to the diversity of the organisms to be identified (viruses, snails, insects, fungal and bacterial diseases): the lucid platform was found unsuitable when dealing with such a variety of organisms and character states (pers obs.). The Lucid platform is ideally suited to identify (either morphologically or taxonomically) closely related organisms. Similar organisms allow the key structure to follow the matrix methodology, where commonalities among the organisms are scored for all organisms. In contrast, when dealing with morphologically diverse taxa, a couplet based approach (albeit web- based) was found to be more suitable as very few commonalities were present among all species.

The above discussion simply highlights the fact that there are no hard and fast rules when developing diagnostic packages, and also that simply developing a computer- based package is not a solution in itself. The diversity and number of taxa being covered, the end user group(s), available technology platforms, and resources

125

available to develop supporting information, will all influence the decision being made as to the potentially most fruitful approach.

126

Chapter 5

Taxonomy, Identification and Diagnostics

Introduction & thesis summary This thesis was initially prompted not through a fundamental interest in opiine braconids, but by a much more applied requirement; the need to identify parasitoids reared from fruit flies collected as part of pest management projects. Failure to meet what should have been a relatively simple task for an important pest group led me into this thesis topic. Initial study identified two key problems facing fruit fly workers: (i) a poor underlying taxonomic basis upon which to base parasitoid identifications and; (ii) a complex, scattered scientific literature largely interpretable only by braconid specialists. This highlighted the need for both taxonomic research and development of diagnostic tools.

In this thesis I have carried out a regional revision of opiine braconids (Chapter 3) that attack fruit flies, providing a sound taxonomic basis for 15 species, one of which was new. Built upon that taxonomic base, I have created user friendly tools for specialists and non-specialists alike to identify those species (Chapter 4). This is particularly important given that most workers interested in this group are not braconid experts, or indeed even likely to be taxonomists, but rather will be fruit fly researchers and managers who simply want to know the name of that “little yellow wasp”. By addressing the two needs, taxonomy and diagnostics, this thesis has achieved its aim of providing sound scientific knowledge, in a user friendly format, for the future development of sustainable fruit fly management.

Taxonomy, identification and diagnostics A key underlying theme of this thesis is the link between taxonomy, identification and diagnostics. As raised in the Introduction (Chapter 1), these are not identical activities, although the terms are often used interchangeably. In this major section of the Discussion, I examine the various ways of presenting taxonomic information and diagnostic tools.

Diagnostics is newly emerging field and this is evidenced by the fact that there are very few formal literature references available which discuss the nexus between taxonomy, identification and diagnostics. Nor in this constantly developing area do

127

there seem to be set rules about what constitutes a diagnostic toolkit versus an identification tool, versus a taxonomic revision which includes electronic based delivery. Given these limitations, much of the following discussion is unreferenced and is based upon my personal experience in the area, which extends significantly beyond the limits of this thesis. I have been working on identification/diagnostic projects since 2000, when I began developing a lucid key to the Bactrocera dorsalis complex of tropical fruit flies (Lawson et al. 2003)3, a particularly difficult taxonomic group. I then moved on to develop TaroPest (Carmichael et al. 2008a), which became more of a diagnostic package supporting the identification method. I have also been a core team member in the development of PaDIL (www.padil.gov.au) and the Plant Biosecurity Toolbox (www.padil.gov.au/pbt). Throughout these projects I have presented at many national (e.g. Carmichael et al. 2008b) and international (e.g. Lawson et al. 2002; Carmichael et al. 2009) conferences, meetings and workshops4. This has given me a great opportunity to develop a network of interested researchers in the field. The beauty about ‘diagnostics’ as a field, is the seemingly endless variety of applications and this promotes a diverse array of interests and discussions between colleagues. The following discussion is based on this nearly decade of conversations and debates with peers about the link between, and presentation of, taxonomy, identification and diagnostics. I believe that utilising electronic media for diagnostic processes is the path of the future. Technology is becoming faster, easier to use and cheaper to purchase. A prime example of this being the mobile phone; in 2007, 72% of Australians had mobile phones, compared with just 24% in 1996 (Linacre 2007). The portability and acceptance of such devices makes them prime assets for dissemination of diagnostic tools. In recent years, this technology has been utilised to a greater extent than ever before in this field. The following discussion considers where we have come from and my view of the future directions of taxonomy, identification and diagnostics.

Changes in Taxonomy Traditionally, taxonomists describe, name and curate specimens, as well as constructing keys, usually dichotomous keys, allowing users to identify their species correctly. Central to these efforts is the use of scientific names for species. Formal species designation allows the scientific community to communicate about

3 Lawson is my maiden name. 4 5th Pacific Plant Protection Organisation and 13th Regional Technical Meeting for Plant Protection, Suva, Fiji Islands, June 2006; TaroPest training workshops in Australia, Fiji and Samoa, October-November 2006; 16 PaDIL workshops throughout Australia and New Zealand, November – December, 2007; Crawford fund International Master Class in Plant Biosecurity, Kuala Lumpur, Malaysia, June 2008;

128

individuals that occur in nature. As in all scientific endeavours, standards are critical (Rumble 1999). Taxonomists clearly recognise the necessity of naming standards and three, slightly different but well established, international codes of nomenclature exist for animals, plants and bacteria (Lapage et al. 1992; International Commission of International Nomenclature 1999; Greuter et al. 2000; Knapp et al. 2004).

In taxonomy, however, new names emerge and names change. Species are clumped or split depending upon new information becoming available or the judgement of the authority involved. This flexibility is necessary, but detracts from the concept of a standard and makes communication more difficult for people, especially non-scientists. These communication difficulties, combined with the fact that most biologists work on a few closely related species at a time, has led to a perception that taxonomy is an insular and arbitrary science, open to trivialisation by the biological sciences community (Stevenson et al. 2001). Taxonomy has undergone a long and persistent decline since the 1950s, in terms of both the number of contributors and the number of papers contributed (Hopkins & Freckleton 2002). Recently, as taxonomy has moved into the bioinformatics age, the image of taxonomists relegated to old dusty basements of museums is being replaced by digital technology, imaging equipment and the World Wide Web. This new outlook has inspired greater effort by some of the taxonomic community to share expertise and information. In the 21st century, taxonomy is slowly creeping its way back into awareness within the scientific community. For some, with this new attitude, the actual definition of taxonomy has transformed and the term is now often used interchangeably with identification or diagnostics. Whether this is correct is a debatable point. However, the lines between the definitions are greying. Diagnostic tools are becoming easier to use, faster to develop and more widely available, thus fading the lines further as the current perceived function of taxonomy is purely for identification.

Taxonomy remains of paramount importance in biology. As one example, discovery and maintenance of knowledge about species and their distribution is at the core of biodiversity conservation. Biodiversity knowledge is, however, unfortunately limited, with less than 20% of the estimated species on earth having scientific names (Stevenson et al. 2001). This problem is worsened by the information and original articles often being difficult to access and only available in specialised technical publications with limited distribution, or recorded on card files and specimen labels in museum collections. Taxonomic knowledge is also needed for all other areas of

129

biology, including conservation biology, pest management, biosecurity, theoretical ecology, and so on (Wheeler et al. 2004).

Efforts to make changes in how taxonomy is done and outputs communicated are increasing, especially with regard to sharing collection information over the internet. There are many computer programs being developed to assist taxonomists in this ever-changing world. Web sites can now be developed by anyone with basic word- processing knowledge, rather than the exhaustively detailed html (HyperText Markup Language) code of only a few years ago (Wright et al. 1995). Many software companies offer easy to use web developing software with pre-packaged templates, allowing users to simply cut and paste information, and at the press of a button a web page is created. With these continuing advances in technology and its availability, field and web guides are being produced and are in greater demand than ever before; allowing members of the non-scientific community access to information and to gain understanding of various life forms and species. This is no more apparent than in the field of ornithology, which is a sport, an industry and an educational tool: “Birding encourages people to reconnect with nature and therefore value it” (Stevenson et al. 2001). A simple Google search using the terms “bird”, “identification” and “electronic” pulls up over 90,000 hits and even a quick view of the first few pages highlights the diversity of electronic tools available to ornithologists. This implementation of electronic media can and should be embraced by other disciplines. To date, entomology has been slow on the uptake, but is gradually gaining momentum with the recent development of a wide variety of electronic keys, field guides and interactive web resources (Walter & Proctor 2001; Seeman & Beard 2007; Malipatil & Ridland 2008).

Progression of Identification Interactive keys have evolved from systems in which expert diagnostic knowledge was initially presented in the form of unillustrated dichotomous keys and has now moved into user friendly visual based approaches (Walter & Winterton 2007). Regardless of the method of identification used, most identification systems are becoming system-centric rather than user-centric. By this I mean it is the system itself that is in charge of the identification, and the user is basically an input device, used to supply the system with observations.

One of the problems facing developers of interactive keys and electronic resources is that the interactive format is reasonably new in both technology and approach to

130

identification. Builders of such keys must learn new programming methods to develop interactive keys, and end-users must learn new and often complicated software, in order to perform an identification. This problem is lessening, as both the building and identification software is becoming more consistent in its appearance and use with commonly used programs, such as internet explorer, and pre- packaged templates become available.

As part of the electronic change, traditional printed dichotomous keys are being transferred to electronic format, being converted from printed matter to computer based hypertext linked keys (e.g. see Phoenix software at http://www.lucidcentral. com). Presenting such keys on the web in a digital format allows easy linkage to illustrative and descriptive information; however, this does not afford the flexibility that interactive keys permit. Hypertext dichotomous keys are still limited by their logic to a single path to enable identification. Dichotomous keys utilising hypertext links to define paths are not strictly interactive keys. An interactive identification key is an interactive computer program in which the user enters characteristics of the specimen. The program continuously analyses the user input and eliminates those taxa whose characteristics do not match those of the specimen. This process continues until only one taxon remains. ‘Non-interactive’ keys follow a structured pathway, answering questions in a pre-defined order.

Diagnostics in a digital age It is telling that few bird enthusiasts use a key to identify birds. This is largely because (i) bird field guides are based on pictures which are more intuitive to use, (ii) field guides are readily available for most locations and (iii) people have a good general knowledge of basic bird forms (duck, hawk, sparrow, and pigeon) (Stevenson et al. 2001). These are sound reasons which can and have been extrapolated into the entomological world. The PaDIL website does not contain keys, but allows searches via basic insect form (bees, butterflies, cockroaches) which are common knowledge among the broader audience. A comparison of different approaches taken by field guides and keys suggest field guides and illustrated media are easier for most people to use accurately than printed dichotomous keys (Morse et al. 1996; Stevenson et al. 2001)

In addition to their primary identification information, field guides (both printed and electronic) generally contain information such as general biology, illustrations labelled to teach morphological terminology and tips for accurate identification, as

131

well as usually being accompanied by a wealth of images. Often simple keys are provided which are usually based on illustrated characters (Walter & Winterton 2007). The amount and detail of information takes into account the audience and the target list involved. Electronic guides do have some significant advantages over printed field guides and these include the capacity to contain greater amounts of information and illustrative material and hyperlinks between similar species and definitions of scientific terms (Walter & Winterton 2007). Electronic field guides and identification keys are rapidly gaining a foot-hold because of the extra information they can contain and the ease of use provided by hyper-links. Electronic field guides are often very powerful tools except, unlike their paper based counterparts, that they cannot easily be taken in the field. This too, however, is rapidly changing with the development of PDA (personal digital assistants) and handheld computers which can support electronic guides. As such, paper-based field guides may disappear within a short time.

Electronic guides are evolving into much more than just tools for identification, they are becoming diagnostic packages in their own right. Often the identification tool is merely the backbone onto which other information is attached. Two global initiatives, perhaps “blue-sky” but technically quite achievable, illustrate the potential end-point of electronic data presentation and identification. IdentifyLife (www.identifylife.org) is an international collaborative project to help identify the world’s living organisms. At its core is a library of identification keys to various taxon groups. It is planned that gradually this library will expand to include all organisms, worldwide. The Encyclopaedia of Life (EOL) (http://www.eol.org/) is the second ambitious project, aimed at making all information about every species on earth available electronically. EOL is to be made up of a web page for each of the approximately 1.8 million currently known species. EOL is still in the developmental stages, so testing of their menu structures and key alternative navigation strategies is not currently possible. What fires my imagination are the possibilities if Identifylife and Encyclopaedia of Life could unite to link identification keys with informative web pages.

Communication and Collaboration Aside from identification keys and diagnostic packages, the internet and its vast number of communication tools have incredible application in the field of science. Blogs and wiki’s are the new craze, with many people, groups and associations having their own internet space (e.g. http://en.wordpress.com/tag/ask-an-

132

entomologist/). Blogs, abbreviated from web log, resemble online diaries. Blogs are usually maintained by an individual, with regular entries of commentary, descriptions of events, or other material such as graphics or video (Anonymous 2008a). In the scientific community, blogs are being utilised as newsletters or discussion boards for research and collaborative groups. Many landcare and environmental groups have blogs to communicate with members and interact with potentially interested participants or funding bodies. Blogs are freely available to access and construct, and are able to be edited from any computer anywhere. Wiki, coming from the Hawaiian word for fast, is a collection of web pages designed to enable anyone who accesses it to contribute or modify content, using a simplified programming language (Anonymous 2008b). Wikis are often used to create collaborative websites. A wiki enables documents to be written cooperatively and can function as a central repository for collaborative documents and communications. A defining characteristic of wiki technology is the ease with which pages can be created and updated. This simplicity, speed and functionality has been embraced by the scientific community in recent times. Wiki’s allow password protection to the sites, and enable collaboration between scientists who are often geographical separate. Rather than emailing documents to and fro, wikis offer a single repository and a single version of the document. These communication tools, wikis and blogs, can open the communication channels between collaborating scientists or enable scientists to communicate effectively and easily with the general public. The development of the Plant Biosecurity Toolbox was greatly aided through the creation of a project specific wiki which allowed team members instant access to the latest version of a document, regardless of by who, or where, it was being edited.

Each of these tools; identification keys, diagnostic packages, web development software, wiki’s and blogs, are just building blocks and communication conduits. Despite these technological changes, taxonomy still remains the foundation of this ever-changing field. I firmly believe all of these tools can and should be embraced as they aid collaboration, communication, education, learning and extension. Not one single methodology will work for all applications, but a combined approach can improve the delivery of science, while also raising the image of science. It needs to remembered, however, that without the underpinning science of taxonomy, technological advances will simply improve the delivery of previous knowledge, but will not lead to the creation of new knowledge.

133

Chapter 6 Appendices

Appendix 1 Carmichael AE, Wharton RA & Clarke AR. 2005. Opiine parasitoids (Hymenoptera: Braconidae) of tropical fruit flies (Diptera: Tephritidae) of the Australia and South Pacific region. Bulletin of Entomological Research 95, 545-569.

134

135

136

137

138

139

140

141

142

143

144

145

146

147

148

149

150

151

152

153

154

155

156

157

158

159

Appendix 2 UserGuide for Photomicroscopy equipment A comprehensive step-by-step user guide was developed for the use of Automontage and the microscopic equipment at QUT. It is presented here as this equipment was extensively used in this study.

160

161

162

163

164

165

166

167

168

169

170

171

172

173

174

175

176

177

178

179

180

181

182

183

184

185

186

187

188

189

Appendix 3 PaDIL UserGuide A step-by-step guide was developed to assist naïve users get the most from the PaDIL website (www.padil.gov.au). The guide was designed to lead the user through a series of steps to perform both simple and advanced searches. The guide takes a hands-on approach, containing example activities to gain user confidence.

190

191

192

193

194

195

196

197

198

199

200

201

202

203

204

205

206

207

208

209

210

211

Chapter 7 Bibliography

Anonymous. 2004 What are keys? http://www.lucidcentral.com/keys/ Accessed: 18.Nov.2008,

Anonymous. 2008a Blog. http://en.wikipedia.org/wiki/Blog Accessed: August 13, 2008.

Anonymous. 2008b Wiki. http://en.wikipedia.org/wiki/Wiki Accessed: August 13, 2008.

Ashmead WH. 1905. Additions to the recorded hymenopterous fauna of the Philippine Islands, with descriptions of new species. Proceedings of the United States National Museum 28, 957-971.

Belokobylskij SA, Wharton RA & La Salle J. 2004. Australian species of the genus Opius Wesmael (Hymenoptera: Braconidae) attacking leaf-mining Agromyzidae, with the description of a new species from South-east Asia. Australian Journal of Entomology 43, 138-147.

Bisby FA, Shimura J, Ruggiero M, Edwards J & Haeuser C. 2002. Taxonomy, at the click of a mouse. Nature 418, 367.

Brèthes J. 1924. Varios himenópteros de la América del Sud. Nuanquam. Otiosus 2, 6-16.

Cameron P. 1911. On a collection of parasitic Hymenoptera (chiefly bred) made by Mr. W.W. Froggatt, F.L.S., in New South Wales, with descriptions of new genera and species. Proceedings of the Linnaean Society of New South Wales 34, 333- 346.

Cantrell B, Chadwick B & Cahill A. 2001. Fruit Fly Fighters: Eradication of the Papaya Fruit Fly. CSIRO Publishing, Collingwood, Vic.

Capek M. 1970. A new classification of the Braconidae based on the cephalic structures of the final instar larva and biological evidence. Canadian Entomologist 102, 846-875.

Carmichael AE, Harding R, Jackson G, Kumar S, Lal SN, Masamdu R, Wright J & Clarke AR. 2008a. TaroPest: An illustrated guide to pests and diseases of taro in the South Pacific. Australian Centre for International Agricultural Research.

Carmichael AE, Kong G & Beasley D. 2009. Australia's EPP diagnostic database: The plant biosecurity toolbox. In: IUFRO International Forest Biosecurity Conference, Rotorua, New Zealand.

Carmichael AE, Wharton R & Clarke AR. 2008b. Opiine parasitoids (Hymenoptera: Braconidae) of tropical fruit flies (Diptera: Tephritidae) of the Australian and South Pacific region. In: Australian Entomological Society 39th Annual General Meeting and Scientific Conference, Orange, New South Wales, Australia.

212

Carmichael AE, Wharton RA & Clarke AR. 2005. Opiine parasitoids (Hymenoptera: Braconidae) of tropical fruit flies (Diptera: Tephritidae) of the Australia and South Pacific region. Bulletin of Entomological Research 95, 545-569.

Chinajariyawong A, Clarke AR, Jirasurat M, Kritsaneepiboon S, Lahey HA, Vijaysegaran S & Walter GH. 2000. Survey of opiine parasitoids of fruit flies (Diptera: Tephritidae) in Thailand and Malaysia. The Raffles Bulletin of Zoology 48, 71-101.

Clausen CP, Clancy DW & Chock QC. 1965. Biological control of the Oriental Fruit Fly (Dacus dorsalis Hendel) and Other Fruit Flies in Hawaii. United States Department of Agriculture, Washington, D.C. pp. 1-100

Cranston PS, Gullan PJ & Taylor RW. 1991. Principles and Practice of Systematics. In: The Insects of Australia: A textbook for students and research workers. Melbourne University Press, Melbourne, Australia.

Dallwitz MJ. 2007a A Comparison of Interactive Identification Programs. http://delta- intkey.com/www/comparison.htm Accessed: 18.Nov.2008,

Dallwitz MJ. 2007b Overview of the DELTA System. http://delta- intkey.com/www/overview.htm Accessed: 18.Nov.2008,

Dallwitz MJ. 2008 DELTA Programs and Documentation. http://delta- intkey.com/www/programs.htm Accessed: 18.Nov.2008,

Duan JJ & Messing RH. 1997. Biological Control of fruit flies in Hawaii: Factors affecting non-target risk analysis. Agriculture and Human Values 14, 227-236.

Enderlein G. 1920. Zur Kenntnis aussereuropäischer Braconiden. . Archiv für Naturgeschichte 84a, 51-224.

Ferrar P. 1997. Fruit fly research and development in the South Pacific. In: Management of Fruit Flies in the Pacific. ACIAR Proceedings. No. 76. (eds AJ Allwood & RAI Drew) pp. 43-45. ACIAR, Canberra.

Fischer M. 1959. Die europaeischen Arten der Gattung Opius Wesm. (Hymenoptera, Braconidae) Teil V a. Mitteilungen der Muenchener Entomologischen Gesellschaft 49, 1-35.

Fischer M. 1963a. Das Genus Austroopius Szépligeti (Hymenoptera, Braconidae, Opiinae). Mitteilungen aus dem Zoologischen Museum in Berlin 39, 173-186.

Fischer M. 1963b. Die orientalischen und australischen Arten der Gattung Opius Wesmael (Hymenoptera, Braconidae). Acta Entomologica Musei Nationalis Pragae 35, 197-242.

Fischer M. 1967a. Redeskriptionen von einigen Opiinen (Hymenoptera, Braconidae). Zeitschrift der Arbeitsgemeinschaft Oesterreichischer Entomologen 19, 59-69.

Fischer M. 1967b. Zusammenfassung der neotropischen Opiinae mit Ausschluss der Gattung Opius Wesm. Beitraege zur Neotropischen fauna 5, 1-21.

Fischer M. 1971a. Hym. Braconidae. World Opiinae. In: Index of Entomophagous Insects pp. 189, Paris.

213

Fischer M. 1971b. Opiinae aus Neu-Guinea und von den Bismarck-Inseln (Hymenoptera: Braconidae). Pacific Insects 13, 487-512.

Fischer M. 1972. Hymenoptera: Braconidae (Opiinae I). Das Tierreich 91, 1-620.

Fischer M. 1977. Hymenoptera, Braconidae (Opiinae II - Amerika). Das Tierreich 96, 1-1001.

Fischer M. 1978a. Neue Opiinae (Hymenoptera, Braconidae) von der australischen Region, besonders aus Tasmanien. Polskie Pismo Entomologiczne 48, 371-412.

Fischer M. 1978b. Neue Opiinae (Hymenoptera, Braconidae) von der australischen Region, besonders aus Tasmanien. Polskie Pismo Entomologiczne 48, 371-412.

Fischer M. 1984. Deskriptionen und Redeskriptionen von Opiinae (Hymenoptera, Braconidae) aus der palaarktischen Region. Annalen des Naturhistorischen Museums in Wien Serie B Botanik und Zoologie. 1984; 86: 85 131.

Fischer M. 1986. Neue Bestimmungsschlussel fur palaarktische Opiinae, neue Subgenera, Redeskriptionen und eine neue Art (Hymenoptera, Braconidae). Annalen des Naturhistorischen Museums in Wien Serie B Botanik und Zoologie. 88/89, 607-662.

Fischer M. 1987. Hymenoptera: Opiinae III - athiopische, orientalische, australische und ozeanische Region. Das Tierreich 104, 1-734.

Fischer M. 1988a. Neues von der australischen Opiinen-Fauna (Hymenoptera, Braconidae, Opiinae). Stapfia. 1988: 239 272; No. 17.

Fischer M. 1988b. Neues von der australischen Opiinen-fauna (Hymenoptera, Braconidae, Opiinae). Stapfia 17, 239-272.

Förster A. 1862. Synopsis der Familien und Gattungen der Braconen. Naturhistorische Verein der Rheinlande Verhandelingen 19, 225-288.

Frank JH & Gillett JL. 2006 Glossary of Expressions in Biological Control. http://edis.ifas.ufl.edu/IN673 Accessed: 18 August, 2008.

Fullaway DT. 1926. A new species of fruit fly parasite from Formosa (Braconidae). Proceedings of the Hawaiian Entomological Society 6, 283-284.

Fullaway DT. 1936. Description of a new fruit fly parasite from Fiji. Proceedings of the Hawaiian Entomological Society 9, 179-180.

Fullaway DT. 1950a. Fruit fly parasites collected in Queensland by N.L.H. Krauss in 1949. Proceedings of the Hawaiian Entomological Society 14, 65-67.

Fullaway DT. 1950b. Notes on parasites of fruit flies. Proceedings of the Hawaiian Entomological Society 14, 27-28.

Fullaway DT. 1951. Review of the Indo-Australasian parasites of the fruit flies (Tephritidae). Proceedings of the Hawaiian Entomological Society 14, 243-250.

Fullaway DT. 1952. New species of Opius (Hymenoptera: Braconidae). Proceedings of the Hawaiian Entomological Society 14, 411-413.

214

Fullaway DT. 1953. New species and varieties of Opius (Hymenoptera: Braconidae). Proceedings of the Entomological Society of Washington 55, 308-314.

Gahan AB. 1915. A revision of the North American ichneumon-flies of the subfamilty Opiinae. Proceedings of the United States National Museum 49, 63-95.

Greuter W, McNeill J, Barrie FR, Burdet H-M, Demoulin V, Filgueiras TS, Nicolson DH, Silva PC, Skog JE, Trehane P, Turland NJ & Hawksworth DL. 2000. International Code of Botanical Nomenclature (St Louis Code). Regnum Vegetabile 138. Koeltz Scientific Books, Königstein.

Hopkins GW & Freckleton RP. 2002. Declines in the numbers of amateur and professional taxonomists: implications for conservation. Animal Conservation 5, 245- 249.

Infante F, Hanson P & Wharton R. 1995. Phytophagy in the genus Monitoriella (Hymenoptera: Braconidae) with description of new species. Annals of the Entomological Society of America 88, 406-415.

Keiser I. 1968. Residual effectiveness of foliar sprays against the Oriental fruit fly, melon fly adn Mediterranean fruit fly. Journal of Economic Entomology 61, 438-443.

Knapp S, Lamas G, Lughadha EN & Novarino G. 2004. Stability or stasis in the names of organisms: the evolving codes of nomenclature. Philosophical Transactions of the Royal Society B 359, 611-622.

Knipling EF. 1992. Principles of insect parasitism analysed from new perspectives. Practical implications for regulating insect populations by biological means. United States Department of Agriculture, Agriculture Research Service, Agriculture Handbook 693.

Lapage SP, Sneath PHA, Lessel EF, Skerman VBD, Seeliger HPR & Clark WA. 1992. International Code of Nomenclature of Bacteria, 1980 Revision. Washington.

Lawson AE, McGuire DJ, Yeates DK, Drew RAI & Clarke AR. 2003. Dorsalis: an interactive identification tool to fruit flies of the Bactrocera dorsalis complex. Griffith University, Brisbane, Australia.

Lawson AE, Yeates DK, McGuire DJ, Clarke AR & Drew RAI. 2002. Dorsalis - an interactive key to fruit flies of the Bactrocera (Bactrocera) dorsalis complex. In: 5th International Congress of Dipterology, pp. 132-133, Brisbane, Australia.

LeBlanc L & Allwood AJ. 1997. Mango fruit fly (Bactrocera frauenfeldi): Why so many in the Federated States of Micronesia? In: In Management of Fruit Flies in the Pacific (eds AJ Allwood & RAI Drew) pp. 125-130. Australian Centre for International Agricultural Research, Canberra.

Lever RJAW. 1938. Fruit flies and their control: biological and chemical. Agricultural Journal 9, 19-20.

Linacre S. 2007. Trends in household consumption. Australian Social Trends 4102.0, 1-12.

Macedo MV & Monteiro RF. 1989. Seed predation by a braconid wasp, Allorhogas sp. (Hymenoptera) Journal of the New York Entomological Society 97, 358-363.

215

Malipatil M & Ridland P. 2008. Polyphagous Agromyzid Leafminers: Identifying polyphagous agromyzid leafminers (Diptera: Agromyzidae) threatening Australian primary industries. The Department of Agriculture, Fisheries and Forestry.

Marshall TA. 1891. A monograph of British Braconidae. Part IV. Transactions of the Entomological Society of London, 7-61.

Matthews RW. 1974. Biology of Braconidae. Annual Review of Entomology 19, 15- 32.

Messing RH, Klungness LM, Purcell M & Wong T. 1993. Quality control parameters of mass-reared opiine parasitoids used in augmentative biological control of tephritid fruit flies in Hawaii. Biological Control 3, 140-147.

Morse DR, Tardivel GM & Spicer J. 1996. A Comparison of the Effectiveness of a Dichotomous Key and a Multiaccess Key to Woodlice. Accessed: from http://www.cs.kent.ac.uk/pubs/1996/44/content.ps.gz.

Naumann ID. 1991. Hymenoptera. In: The Insects of Australia: a textbook for students and research workers pp. 916-1000. CSIRO, Division of Entomology.

Nomenclature ICoZ. 1999. International Code of Zoological Nomenclature. The International Trust for Zoological Nomenclature.

Norrbom A. 2004 The Diptera Site. http://www.sel.barc.usda.gov/diptera/tephriti/tephriti.htm Accessed: 4 August, 2008,

Norrbom AL, Carroll LE, Thompson FC, White IM & Freidberg A. 1998. Systematic database of names. Myia 9, 65-251.

O'Connor BA. 1960. A decade of biological control work in Fiji. Agriculture Journal 30, 44-54.

Ovruski S, Aluja M, Sivinski J & Wharton RA. 2000. Hymenoptera parasitoids on fruit infesting Tephritidae (Diptera) in Latin America and the southern United States: Diversity, distribution, taxonomic status and their use in fruit fly biocontrol. Integrated Pest Management Reviews 5, 81-107.

Purcell M, Herr JC, Messing RH & Wong T. 1998. Interactions between augmentatively released Diachasmimorpha longicaudata (Hymenoptera: Braconidae) and a complex of opiine parasitoids in a commercial guava orchard. Biocontrol Science and Technology 8, 139-151.

Quimio GM & Walter GH. 2001. Host preference and host suitability in an egg-pupal fruit fly parasitoid, Fopius arisanus (Sonan) (Hym., Braconidae). J appl entomol. Berlin : P. Parey, c1986 . Apr 2001. v. 125 (3) p. 135 140.

Ramadan MM. 2004. Mass-rearing biology of Fopius vandenboschi (Hym., Braconidae). Journal of Applied Entomology 128, 226-232.

Rumble J. 1999. Standards for Scientific and Technical Data. Accessed: 14 August, 2007 from http://www.tdwg.org/standards-tdwg.htm.

Seeman OD & Beard JJ. 2007. Mites of Quarantine Importance. Queensland Museum, Queensland Government.

216

Sharkey MJ & Wahl DB. 1992. Cladistics of the Ichneumonoidea (Hymenoptera). Journal of Hymenoptera Research 1, 15-24.

Sharkey MJ & Wharton RA. 1997. Morphology and Terminology. In: Manual of the New World Genera of the Family Braconidae (Hymenoptera). (eds RA Wharton, PM Marsh & MJ Sharkey) pp. 19-37. The International Society of Hymenopterists, Washington, D.C.

Silvestri F. 1913. Viaggio in Africa per cercare parassiti di mosche dei frutti. Bollettino del Laboratorio di Zoologia General e Agraria, Portici 8, 1-164.

Silvestri F. 1914. Report of an expedition to Africa in search of the natural enemies of Fruit Flies (Trypaneidae) with descriptions, observations and biological notes. Division of Entomology, Board of Agriculture and Forestry, Territory of Hawaii. Bulletin no. 3, 1-176.

Silvestri F. 1916. Descrizione di alcuni Imenotteie Braconidi parasiti Ditteri Triponeidi nell'India. Bollettino del Laboratorio di Zoologia General e Agraria, Portici 11, 160- 169.

Snowball GJ & Lukins RG. 1964. Status of introduced parasites of Queensland Fruit Fly (Strumeta tryoni), 1960-62. Australian Journal of Agricultural Research 15, 586- 608.

Sonan J. 1932. Notes on some Braconidae and Ichneumonidae from Formosa, with descriptions of 18 new species. Transactions of the Natural History Society of Formosa 22, 66-87.

Steiner LF. 1952. Fruit fly control in Hawaii with poison-bait sprays containing protein hydrolysates. Journal of Economic Entomology 45, 838-843.

Stevenson RD, Haber WA & Morris. RA. 2001. Standing on the Shoulders of Taxonomists: User Communities and Electronic Field Guides in the Ecoinformatics Revolution. Accessed: 13 August 2007 from http://www.cs.umb.ed/Researches/Publications/PDFs/TR-01-03.pdf

Szépligeti G. 1900. Braconiden aus New-Guinea in der Sammlung des Ung. National-Museums. Termeszetrajzi Fuzetek 23, 49-65.

Tobias VI. 1967. A review of the classification, phylogeny and evolution of the family Braconidae. Entomological Review 46, 387-399.

Tobias VI. 1977. The genus Opius Wesm. (Hymenoptera, Braconidae) as parasites of fruit flies (Diptera, Tephritidae). Entomologicheskoe Obozrenie. 1977; 56(2): 420 430.

Tobias VI & Jakimavicius AB. 1986. [A guide to the insects of the European part of the USSR. Hymenoptera. Family Braconidae.]. Opredeliteli po Faune SSSR. 1986; 3(5): 7 231, 284 292, 296 299; No. 147.

Van Achterberg C & Maeto K. 1990. Two new and aberrant species of Braconidae (Hymenoptera) from Japan. Zoologische Mededelingen. 1990, 64: 1 6, 59 70; 12 ref.

Viereck HL. 1913. Descriptions of six new genera and twelve new species of ichneumon-flies. Proceedings of the United States National Museum 44, 639-648.

217

Walker F. 1860. Characters of some apparently undescribed Ceylon Insects. Annals and Magazine of Natural History 3, 304-311.

Walter DE & Proctor HC. 2001. Mites in Soil. An interactive key to mites and other soil microarthropods. CSIRO Publishing.

Walter DE & Winterton S. 2007. Keys and the Crisis in Taxonomy: Extinction or Reinvention? Annual Review of Entomology 52, 193-208.

Waterhouse DF. 1993. Pest Fruit flies in the Oceanic Pacific. In: Biological Control: Pacific Prospects - Supplement 2 pp. 4-47. Australian Centre for International Agricultural Research, Canberra.

Wesmael C. 1835. Monographie des Braconides de Belgique. Nouveaux memoires de l'academie royale des sciences et belles-lettres de Bruxelles 9, 1-252.

Wharton RA. 1980. Review of the Narctic Alysiini (Hymenoptera, Braconidae). With discussion of generic relationships with the tribe. University of California Publications in Entomology 88, 1-112.

Wharton RA. 1983. Variation in Opius hirtis Fischer and discussion of Desmiostoma Foerster (Hymenoptera: Braconidae). Proceedings of the Entomological Society of Washington 85, 327-330.

Wharton RA. 1987. Changes in nomenclature and classification of some opiine Braconidae (Hymenoptera). Proceedings of the Entomological Society of Washington 89, 61-73.

Wharton RA. 1988. Classification of the braconid subfamily Opiinae (Hymenoptera). The Canadian Entomologist 120, 333-360.

Wharton RA. 1993. Bionomics of the Braconidae. Annual Review of Entomology 38, 121-143.

Wharton RA. 1997a. Generic relationships of Opiine Braconidae (Hymenoptera) parasitic on fruit infesting Tephritidae (Diptera). Contributions of the American Entomological Institute 30, 3-53.

Wharton RA. 1997b. Introduction. Manual of the New World Genera of the Family Braconidae (Hymenoptera). In: Manual of the New World Genera of the Family Braconidae (Hymenoptera). (eds RA Wharton, PM Marsh & MJ Sharkey) pp. 1-18. The International Society of Hymenopterists, Washington, D.C.

Wharton RA. 1999. A review of the Old World genus Fopius Wharton (Hymenoptera: Braconidae: Opiinae), with description of two new species reared from fruit-infesting Tephritidae (Diptera). Journal of Hymenoptera Research 8, 48-64.

Wharton RA. 2008 Parasitoids of Fruit-Infesting Tephritidae. http://hymenoptera.tamu.edu/paroffit/?taxcpl=tax&taxcpl_id=7462. Accessed 27/7/2005. Accessed: 16th September, 2008.

Wharton RA & Gilstrap FE. 1983. Key to and status of opiine braconid (Hymenoptera) parasitoids used in biological control of Ceratitis and Dacus s.l. (Diptera: Tephritidae). Annals of the Entomological Society of America 76, 721-742.

218

Wharton RA & Marsh PM. 1978. New World Opiinae (Hymenoptera: Braconidae) parasitic on Tephritidae (Diptera). Journal of the Washington Academy of Sciences 68, 147-167.

Wharton RA, Trostle MK, Messing RH, Copeland RS, Kimani-Njogu SW, Lux S, Overholt WA, Mohamed S & Sivinski J. 2000. Parasitoids of medfly, Ceratitis capitata, and related tephritids in Kenyan coffee: a predominantly koinobiont assemblage. Bulletin of Entomological Research 90, 517-526.

Wharton RA & Van Achterberg C. 2000. Family Group names in Braconidae (Hymenoptera: Ichneumonoidea). Journal of Hymenoptera Research 9, 254-270.

Wheeler QD, Raven RH & Wilson EO. 2004. Taxonomy: impediment or expedient? Science 303, 285.

White IM & Elson-Harris MM. 1992. Fruit Flies of Economic Significance: their identification and bionomics. C.A.B. International in association with A.C.I.A.R., Wallingford.

White IM & Sandlant GR. 1998. Computerised insect identification: a comparison of differing approaches and problems. In: Information Technology, Plant Pathology and Biodiversity (eds P Bridge, P Jeffries, DR Morse & PR Scott) pp. 261-272. CAB International, Wallingford.

Whitfield JB. 1997. Subfamily Microgastrinae. In: Manual of the New World genera of the Family Braconidae (Hymenoptera) (eds R Wharton, PM Marsh & MJ Sharkey) pp. 333-364. The International Society of Hymenopterists, Washington, D.C.

Wright JF, Morse DR & Tradivel GM. 1995. An investigation into the use of hypertext as a user-interface to taxonomic keys. Computer Applications in the Biosciences 11, 19-27.

219

CD Envelope

Contents: OpiineID

220