THE AUSTRALIAN HANDBOOK FOR THE IDENTIFICATION OF Version 3.1

Species: facialis : Bactrocera aquilonis Species: Bactrocera cacuminata Species: Bactrocera frauenfeldi

Species: Bactrocera musae Species: Bactrocera tryoni Species: Species: Dacus aequalis

Species: Zeugodacus choristus Species: Bactrocera opiliae Species: Bactrocera rufofuscula Species: Bactrocera pallida

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Plant Health Australia (2018). The Australian Handbook for the Identification of Fruit Flies. Version 3.1. Plant Health Australia. Canberra, ACT. Contents Introduction 2 Bactrocera rufofuscula 90 How to use this handbook 4 Bactrocera trilineola 92 Bactrocera trivialis 94 Morphological identification 8 Bactrocera tryoni 96 Glossary 10 Bactrocera tsuneonis 98

Variation in appearance with Bactrocera tuberculata 100 preservation technique 15 Bactrocera umbrosa 102 Bactrocera xanthodes 104 Molecular identification 16 Bactrocera zonata 106 Groupings of fruit fly by ZEUGODACUS morphological similarity 18 Zeugodacus atrisetosus 108 BACTROCERA Zeugodacus choristus 110 Bactrocera albistrigata 26 Zeugodacus cucumis 112 Bactrocera aquilonis 28 Zeugodacus cucurbitae 114 Bactrocera bancroftii 30 Zeugodacus decipiens 116 Bactrocera breviaculeus 32 Zeugodacus depressus 118 Bactrocera bryoniae 34 Zeugodacus tau 120 Bactrocera cacuminata 36 DACUS Bactrocera carambolae 38 Dacus aequalis 122 Bactrocera caryeae 40 Dacus axanus 124 Bactrocera correcta 42 Dacus longicornis 126 Bactrocera curvipennis 44 Dacus solomonensis 128 46 Bactrocera endiandrae 48 CERATITIS Bactrocera facialis 50 Ceratitis capitata 130 Bactrocera frauenfeldi 52 Ceratitis quilicii 132 Bactrocera jarvisi 54 Ceratitis rosa 134 Bactrocera kandiensis 56 DIRIOXA Bactrocera kirki 58 Dirioxa pornia 136 Bactrocera kraussi 60 ANASTREPHA Bactrocera laticaudus 62 Anastrepha fraterculus 138 Bactrocera latifrons 64 Anastrepha serpentina 140 Bactrocera melanotus 66 Bactrocera minax 68 Rhagoletis cerasi 142 Bactrocera musae 70 Rhagoletis cingulata 144 Bactrocera neohumeralis 72 Rhagoletis completa 146 Bactrocera obliqua 74 Rhagoletis fausta 148 Bactrocera occipitalis 76 Rhagoletis pomonella 150 Bactrocera oleae 78 Bactrocera opiliae 80 TOXOTRYPANA Bactrocera pallida 82 Toxotrypana curvicauda 152 Bactrocera passiflorae 84 DROSOPHILA Bactrocera psidii 86 Drosophila suzukii 154 Bactrocera pyrifoliae 88 Reference list 156 Introduction Fruit flies are one of the world’s most destructive horticultural pests and pose risks to most commercial fruit and vegetable crops. This has major implications for the sustainable production and market access of Australia’s multi-billion-dollar horticultural industry. Worldwide there are over 4,000 species of fruit flies in the family of which around 350 species are of economic importance. More than 300 species of fruit fly occur in Australia although only a few of these have been found to have any degree of economic impact, with Queensland fruit fly (Bactrocera tryoni) and Mediterranean fruit fly (Ceratitis capitata) being the species of primary economic concern. It is therefore important to distinguish between the species that pose a threat to production and domestic market access and those that do not, including exotic species. Australia is free from many species that impact horticultural production elsewhere. Neighbouring countries in south east Asia and the South Pacific are home to numerous species of fruit fly that pose an immediate incursion risk to Australia. Rapid diagnosis of these flies, should they arrive in Australia, is therefore a critical prerequisite to containing and eradicating the populations before they establish. The accurate identification of fruit flies is a key component of Australia’s biosecurity system. It underpins the domestic movement of fruit and vegetables, maintains international market access for Australian producers, and protects Australia’s borders from exotic pest incursion. This completely revised version of The Australian Handbook for the Identification of Fruit Flies (Version 3.1) (the handbook) has been compiled by diagnosticians for diagnosticians using some of the latest research outcomes and resources available. The handbook is a compilation of diagnostic information for 65 fruit fly species, most of which are exotic to Australia, including the non-tephritid fruit infesting Drosophila suzukii (spotted-wing Drosophila). The handbook is intended to facilitate rapid diagnosis of fruit fly species and be a comprehensive guide for Australian diagnosticians and field officers involved in maintaining, supporting and enhancing Australia’s biosecurity system.

2 ABOUT THIS HANDBOOK This publication is the product of the Plant Biosecurity Cooperative Research Centre (PBCRC) project Next Generation National Fruit Fly Diagnostics and Handbook PBCRC2147. The handbook draws together hundreds of new high definition fruit fly images as well as new molecular diagnostic tools generated from this research. A companion website Fruit Fly Identification Australia (fruitflyidentification.org.au) has also been developed. The website contains an expanded selection of high definition images as well as an updated description of the latest molecular techniques and protocols used for the identification of fruit flies. The website also includes a new multi-entry Lucid key for identifying the Dacini species as well as comprehensive printable data sheets and an interactive glossary. The handbook and website were developed to support Australia’s fruit fly researchers, academics, surveillance officers, diagnosticians and laboratory scientists with the hope that the sharing of information and experience, as well as the constructive communication that brings this knowledge base together, will continue to grow.

CONTRIBUTORS

Mark Schutze, Jacinta McMahon, Queensland University of Technology Matt Krosch, Francesca Strutt Department of Agriculture and Jane Royer Fisheries, Queensland Melanie Bottrill, Nicholas Woods Plant Health Australia Stephen Cameron Purdue University Department of Agriculture and Food, Bill Woods Western Australia Department of Economic Development, Mark Blacket Jobs, Transport and Resources, Victoria

ACKNOWLEDGEMENTS Marc De Meyer (Department of Biology, Royal Museum for Central Africa) Gary Steck (Florida Department of Agriculture and Consumer Services) Allen Norrbom (Agricultural Research Service, United States Department of Agriculture)

3 How to use this handbook The handbook is designed as a diagnostic aid for entomologists and biosecurity and plant health professionals. Each fruit fly species described is illustrated with a dorsal photograph annotated with important diagnostic characteristics. Additional photographs provide information about variation within species as well as head, leg and lateral views. Maps indicate published distribution information and notes on easily confused or similar species are provided to aid correct identification. Photographs were taken by project members at Queensland University of Technology. A classic representative specimen (matching the current written morphological description in the literature) from each species has been photographed from a variety of angles showing key diagnostic features. Images for each individual include a head view, full body dorsal view, full body lateral view, dorsal and ventral abdomen views and a dorsal scutum view. Slide mounted fore, mid and hind legs and left and right wings were also photographed. Intraspecific morphological variants were also photographed illustrating differences where they occur, principally associated with colour pattern variation. As mentioned under ‘things to be wary of’, some of the fruit fly species included in this handbook are highly variable. Every effort has been made to include as many images of this variation as possible, however, due to limited space it was not always possible to include all images. Please note that the full image set can be found on the companion website Fruit Fly Identification Australia (fruitflyidentification.org.au). The coloured banner down the side of the species pages refers to whether the species is present in Australia and whether it is a pest or non-pest species. The functional categories are: Red (EXOTIC/PEST) Orange (PRESENT/PEST) Green (PRESENT/NON-PEST) The word ‘pest’ in the above context refers to whether the species is listed on the High Priority Pest list of the National Fruit Fly Strategy (planthealthaustralia.com.au/ national-programs/fruit-fly). Annotations have been included on the species pages to assist users with the identification of these species.

4 Bactrocera caryeae 5 THINGS TO BE WARY OF Variation: Some fruit fly species are highly variable, either morphologically, molecularly, or both. This often blurs the distinction between intra- versus inter- specific variation, with variants of some species often looking like something entirely different to what they actually are. A classic example of this is the guava fruit fly, Bactrocera correcta, which displays a wide variation in colour pattern such that the extremes look like completely different flies. Problematically, species likeB. frauenfeldi can often be confused with their close relatives, such as B. albistrigata; the former of which occurs in Australia, the latter not; or members of the Oriental fruit fly (B. dorsalis) species complex that contains almost 100 species, many of which are highly similar in appearance. Host and geographic distribution: Knowing the locality, host or lure a fruit fly was collected from can provide invaluable information towards making a diagnosis, but must also be used with caution to avoid falsely identifying a specimen as pest or non-pest. Geographic information presented in this handbook has been drawn from published records or available public records, such as Centre for Agriculture and Bioscience International (CABI). Please note that: • the markings on the maps are indicative for the sovereign states and may not present actual distribution ranges within each country as indicated • some locations are listed under their geographic region which may differ from their political association (e.g., Réunion Island is a region of France but is geographically placed under ‘Africa’ in this Handbook due to its proximity to that continent). Species complexes: Groups of species that share similar morphological characters are sometimes grouped together as species complexes. Typically, these groups are also very closely related and hence can be especially difficult to discern. This is particularly the case for cryptic species, which are species that, while biologically distinct, may not be distinguished morphologically. A number of species complexes exist within Bactrocera and Zeugodacus, such as the dorsalis, tau, and frauenfeldi- complexes. Further, species complexes are found in other genera, such as the Anastrepha fraterculus complex (also referred to as a ‘species group’). Particular care must be taken when identifying species that belong to complexes, with integration of multiple datasets and additional expert opinions generally recommended before making a definitive diagnosis.

6 Limitations of this handbook: While pest species of high priority to Australia have been included in this handbook, along with species that may be readily confused with them, it is by no means an exhaustive treatment of all fruit fly species. With several thousand species known from the family (and over 900 species from Bactrocera, Dacus and Zeugodacus), it is impossible to include them all, especially those that are more widely represented in other parts of the world (e.g. Anastrepha, Ceratitis, Rhagoletis). Therefore, we direct users to other excellent resources that deal in greater details for these other groups. This includes but is not limited to websites, primary literature, and other diagnostic publications. These include: • Anastrepha and Toxotrypana: descriptions, illustrations, and interactive keys: delta-intkey.com/anatox/intro.htm • A set of multi-entry identification keys to African frugivorous flies (Diptera, Tephritidae): fruitflykeys.africamuseum.be/ • USDA Compendium of Fruit Fly Host Information: coffhi.cphst.org/ Images were taken with grey photographic card as background; hence grey colouration on wings where they are transparent/hyaline in life. For example, see Toxotrypana curvicauda.

OTHER RESOURCES • Drew, R. A., & Romig, M. C. (2013). Tropical Fruit Flies (Tephritidae ) of South- East Asia: Indomalaya to North-West Australasia. CABI. • Drew, R. A., & Romig, M. C. (2016). Keys to the tropical fruit flies (Tephritidae: Dacinae) of South-East Asia: Indomalaya to North-West Australasia. CABI. • Drew, R. A. (1989). The tropical fruit flies (Diptera: Tephritidae: Dacinae) of the Australasian and Oceanian regions. Memoirs of the Queensland Museum, 26. • LUCID key for identification of fruit flies in Australia.

7 Morphological identification Approximately 90% of dacine pest species can be identified accurately and quickly by microscopic examination of the adult. For these species there is no need for supporting evidence. The remaining 10% (mainly members of species complexes) can be identified with this same method but require expert examination and may require additional supporting evidence (e.g., molecular data or host association records). Methods here are updated according to Drew & Romig (2013). Only morphological diagnostic procedures and information for adult fruit flies are contained in this document. Aside from molecular techniques, larval diagnosis has been excluded from this protocol. For routine morphological identification: • Collect flies in dry traps. • When clearing traps collect samples into a tissue. Put tissue in a small box with collection details on the outside. Samples in tissue can also be collected into a vial though this is less preferred particularly in the tropics as samples can ‘sweat’ causing specimens to deteriorate. If collecting into vial, a label written in pencil or permanent pen (not biro as it runs) should be put inside as writing on the outside can rub off. • Store samples in freezer until ready for identification. • Sort dry specimens in a petri dish under a dissecting stereo microscope. • If keeping specimens after identification store in freezer to prevent deterioration. • Do not store specimens in ethanol, alcohol or propylene glycol unless being kept for DNA analysis. They leach colours and patterns necessary for morphological identification. An option is to remove three legs from one side for molecular analysis, and keep the rest of the specimen on pinned material. For suspect specimens requiring further identification: • Store the specimen in a small vial with tissue to protect it until ready to pin or ship. Add a pencil or permanent ink label (not biro as it runs) detailing collection location, collection date, collection method, collector, tentative identification and identifier. • As manipulating loose specimens with forceps tends to damage them, suspect specimens should ideally be pinned to keep them as intact as possible. If the specimen is a suspect exotic and needs to be shipped to a specialist quickly, they can be sent unpinned in a tube as above. For pinning: • Put specimen in relaxing chamber with thymol (to prevent mould growth) for 6–12 hours. • Using a micropin, pin through the right hand side of the scutum. Mount micropinned specimen on a pith stage on a pin. • Add label to pin. • Store pinned specimens in reference collection conditions (21°C and 50% relative humidity).

8 Key features (see glossary on following page) used for the morphological diagnosis of adult fruit flies include: • Overall colour and colour patterning • Wing patterns • Presence and shape of thoracic vittae (yellow bands) • Presence or absence of various setae. Use the morphological diagnostic key and descriptions contained in the data sheets to identify the species of fruit fly under a microscope. If identification cannot be made and/or the specimen is suspected to be of quarantine concern, it should be referred to either a state or federal biosecurity agency. If the specimen is identified as an exotic fruit fly, the appropriate national authority should be notified as required under the Emergency Plant Pest Response Deed.

PRESERVATION OF LARVAE FOR MORPHOLOGICAL IDENTIFICATION Adult flies are generally always needed to make an accurate species identification; however under some circumstances larvae will need to be preserved for morphological examination to at least identify them to family or genus level. Under these circumstances, larvae are preserved by first placing them in hot (approximately 65°C) water for 2–4 minutes, after which they are removed and allowed to cool to room temperature. Once at room temperature, immerse larvae in 50% ethanol for 15- 30 minutes then transfer to 70% ethanol for storage ensuring all relevant collection information is included with the sample. Further details on subsequent procedures for slide mounting is provided in ISPM-27 DP9 (Diagnostic protocols for regulated pests: Genus Anastrepha Schiner).

Fruit fly specimens being prepared for pinning

9 Glossary

FEMALE DORSAL frons

postprontal lobe head a. npl. seta scutum lateral vitta p. sa. seta a. sa. seta

thorax prsc. setae ia. setae sc. setae scutellum

terga I+II

tergum III abdomen

tergum IV ovipositor (female) tergum V WING

costal cell R costal band basal costal cell r-m 2+3

R4+5

M

dm-cu cup cell

anal streak 10 MALE LATERAL

lateral vitta scutellum scutellar setae postprontal lobe

mesopleural stripe

a. npl. setae femur tibia notopleural calli

tarsi

HEAD

face

facial spots

11 Glossary HEAD face – an inverted smooth V-shaped area on the anterior surface between the antennae and ventral margin of the head (from which mouthparts protrude) facial spots – dark spots on the face in the antennal furrows and above the mouthparts facial mask – dark mask-like markings on face frons – upper-frontal area of the head immediately above the antennae and between the eyes antenna – located anteriorly on the head between the face and frons

THORAX scutum – dorsal surface of thorax excluding the scutellum mesonotum – dorsal surface of the thorax – in older texts used instead of scutum postpronotal lobes = ppn. lobes – rectangular protuberances on the anterolateral corners of the scutum (= humeral calli) ppn. lobes – postpronotal lobes humeral calli – in older texts used instead of postpronotal lobes notopleural calli – triangular protuberances on the notopleura, between the ppn. lobe and wing base scutellum – truncated triangle on the posterior edge of the scutum basal band – a band of dark colour on the base of the scutellum (where it joins the scutum) mesopleural stripe – yellow band on lateral surface of the thorax vitta – yellow stripe on the scutum (vittae plural) lateral vittae = lateral post-sutural vittae – yellow bands on lateral margin of scutum medial vitta – yellow, usually teardrop-shaped, band on the midline of the scutum mesonotal suture – an impressed line on the mesonotum running from the notopleural callus towards the centre of the scutum setae – longer bristle like hairs (= bristles) (seta singular) bristle – older term for seta haltere – modified hind wings, used for maintaining equilibrium during flight

12 THORACIC SETAE sc. = scutellar setae – 2 or 4 setae on the posterior dorsal edge of the scutellum a. sa. = anterior supra-alar setae, or abbreviated to sa. (supra-alar setae) – setae on the mid-dorsal edge of the smooth semicircular flat cavity above the wing – presence or absence is a taxonomic character ia. = intra-alar setae on the posterolateral corner of the mesonotum – used to measure length of the vittae prsc. = prescutellar setae – setae on the posterior of the scutum anterior to the scutellum – presence or absence is a taxonomic character a. npl. = anterior notopleural setae – setae between the notopleuron and ppn. lobes. – used to measure the width of the mesopleural stripe p. sa. = posterior supra-alar setae – setae on the posterior-dorsal edge of the smooth semicircular flat cavity above the wing – in older texts used to measure length of the lateral vittae

WING costal cells – cells on the basal anterior part of the wing – presence of microtrichia or tint are taxonomic characters bc – basal costal cell closest to the thorax (= first costal cell) c – costal cell, second costal cell from the thorax (= second costal cell) microtrichia – minute short hairs in the costal cells – presence, absence or amount is a taxonomic character

R2+3 – wing vein ending at the anterior apical third of the wing – often used to measure width of the costal band

R4+5 – wing vein below R2+3 ending at wing apex

M – wing vein below R4+5 ending at wing apex CuA – wing vein below M ending at posterior apical third of the wing r-m – cross vein between R4+5 and M – presence of infuscation is a taxonomic character dm-cu – cross vein between M and CuA ending near posterior edge of wing – presence of infuscation is a taxonomic character cup – cell with fingerlike extension running from near base of wing to posterior wing edge costal band – tinted band on anterior edge of wing – width and sometimes darkness are taxonomic characters

13 Glossary anal streak – band of colour outside cup cell – width is a taxonomic character infuscation – tint, usually referring to r-m or dm-cu cross veins

ABDOMEN terga – dorsal abdominal segments (there are five). Plural of tergum. Patterning on the posterior three (terga III-V) are often taxonomic characters tergum I – basal abdominal segment closest to the thorax tergum II – second from basal abdominal segment tergum III – middle abdominal segment which has the pecten on the lateral posterior edge tergum IV – second from posterior abdominal segment tergum V – posterior abdominal segment pecten – comb of hairs on posterolateral edge of tergum III, have the appearance of eyelashes ceromata – oval shining spots on tergum V – dark ceromata can be a taxonomic character

LEGS femur – basal leg segment basally joined to the coxa/e – colour patterns can be taxonomic (femora plural) tibia – apical leg segment between the femur and tarsi – colour patterns can be taxonomic (tibiae plural)

OTHER fuscous – dark red-brown fulvous – yellow brown

14 Variation in appearance with preservation technique

How adult fruit flies are captured, preserved, and stored can drastically affect their colour. Colour is frequently used as a defining characteristic of species, therefore knowing the impact of preservation techniques on the colour of a specimen is critical. The following methods of preservation are most frequently used: • propylene glycol (a non-toxic/non-flammable liquid often used in field collecting where DNA preservation is needed) • dry/pinned: how many adult flies are trapped and preserved, and considered the best method for retaining colour • ethanol: flies may be collected into or stored in ethanol, especially when needed for DNA work; however, ethanol will leach colour over time. As shown below for one species, the Queensland fruit flyBactrocera tryoni, the way each of these methods has impacted the colour is evident. Note especially the loss of colour in the propylene glycol specimen, and that while colour is retained in the ethanol specimen it will fade over time. Visual loss of the thoracic vittae and other yellow/pale characters is particularly affected.

B. tryoni - propylene glycol B. tryoni - dry pinned B. tryoni - ethanol

15 Molecular identification Molecular data, primarily generated via direct DNA sequencing of diagnostic markers, may be necessary to support inconclusive morphological identification. There are several circumstances for which this might be necessary. Firstly, for immature life stages, only third instar larvae (and sometimes pupae) of some species are identifiable via visual examination (e.g. White & Elson-Harris 1992). Secondly, there are numerous examples of ‘problematic’ species pairs and groups that are closely related and/or the adults are near morphologically identical. Thirdly, significant intraspecific variation in key morphological characters (e.g. scutum colour pattern), occurs in many widely distributed species. For all such situations, molecular data may improve species diagnosis. Diagnosticians should ideally obtain DNA from specimens using relatively non- destructive techniques, to ensure that a voucher specimen is available for future morphological re-examination (Floyd et al. 2010). For fruit fly adults up to three legs from the same side of the body can be destructively used, thereby retaining other diagnostically informative features of the specimen (such as wings, thorax and abdomen). For larvae, anterior and posterior sections can be sectioned off and retained, preserving the morphologically valuable mouthparts and spiracles. Alternatively, even less destructive methods – including Proteinase K digestion of internal tissues that retains the entire cuticle – can also be very effective (e.g. Gilbert et al. 2007). This handbook focuses only on DNA barcoding as the molecular tool of choice for diagnosing tephritid fruit flies. DNA barcoding of an unknown specimen involves obtaining a DNA sequence of a specific region of the genome (most commonly the mitochondrial cytochrome c oxidase subunit 1 gene [COI]), and then comparing it with a database of curated sequences from positively identified reference specimens. The molecular loci we refer to in this handbook include COI, and the nuclear genes EIF3L, RPA2, DDOSTs2, and POP4. In-depth protocols are not provided here, but are accessible on the companion website (fruitflyidentification.org.au). We do, however, provide a list of available diagnostic molecular markers and the species pairs/groups for which they are each most appropriately applied (Table 1).

16 Table 1. Genetic loci that best separate easily confused species

SPECIES PAIR MOLECULAR DIAGNOSTIC LOCUS COI POP4 EIF3L RPA2 DDOSTS2 B. dorsalis / B. carambolae B. dorsalis / B. kandiensis B. dorsalis / B. musae inconclusive B. dorsalis / B. endiandrae B. dorsalis / B. cacuminata inconclusive B. dorsalis / B. occipitalis N/A inconclusive B. dorsalis / B. opiliae N/A B. dorsalis / B. latifrons B. carambolae / B. kandiensis B. carambolae / B. opiliae N/A B. carambolae / B. musae B. carambolae / B. occipitalis inconclusive B. musae / B. opiliae N/A inconclusive B. musae / B. latifrons B. musae / B. endiandrae B. opiliae / B. endiandrae N/A B. musae / B. bancroftii inconclusive B. zonata / B. correcta / B. pallida / B. jarvisi B. kraussi / B. tryoni B. tryoni complex B. trivialis / B. rufofuscula N/A N/A N/A N/A B. trivialis / B. breviaculeus N/A N/A N/A N/A B. rufofuscula / inconclusive N/A N/A inconclusive B. breviaculeus B. decurtans / B. pallida N/A N/A B. passiflorae / B. facialis N/A N/A N/A B. frauenfeldi / B. albistrigata Z. atrisetosus / Z. cucumis N/A N/A N/A N/A Z. choristus / Z. cucurbitae Z. tau / Z. cucurbitae Z. depressus / Z. tau N/A

17 Groupings of fruit fly by morphological similarity DACINAE - DACINI - TWO VITTAE - DORSALIS COMPLEX

B. carambolae, page 38 B. caryeae, page 40 B. dorsalis, page 46

B. kandiensis, page 56 B. occipitalis, page 76 B. pyrifoliae, page 88

B. cacuminata, page 36 B. endiandrae, page 48 B. musae, page 70

KEY: Red (EXOTIC/PEST) Orange (PRESENT/PEST) Green (PRESENT/NON-PEST)

B. opiliae, page 80 18 TWO VITTAE - OTHERS WITH BLACK SCUTUM

B. curvipennis, page 44 B. bryoniae, page 34 B. latifrons, page 64

B. bancroftii, page 30

TWO VITTAE - INCOMPLETE COSTAL BAND

B. correcta, page 42 B. zonata, page 106 B. tuberculata, page 100

TWO VITTAE - BROWN FLIES - TRYONI COMPLEX

B. tryoni, page 96 B. neohumeralis, page 72 B. aquilonis, page 28

19 TWO VITTAE - BROWN FLIES CONFUSED WITH TRIVIALIS OR TRYONI (+ TRIVIALIS)

B. kraussi, page 60 B. trivialis, page 94 B. rufofuscula, page 90

B. breviaculeus, page 32

TWO VITTAE - GOLDEN FLIES

B. jarvisi, page 54 B. pallida, page 82

20 THREE VITTAE

Z. atrisetosus, page 108 Z. cucumis, page 112 Z. cucurbitae, page 114

Z. choristus, page 110 Z. decipiens, page 116 Z. tau, page 120

Z. depressus, page 118 B. minax, page 68 B. tsuneonis, page 98

B. xanthodes, page 104

21 ZERO VITTAE, OR SMALL FLIES BLACK SCUTUM

B. melanotus, page 66 B. passiflorae, page 84 B. oleae, page 78 no wing pattern no wing pattern no wing pattern

B. facialis, page 50 Vittae: 2 WING BAND

B. albistrigata, page 26 B. frauenfeldi, page 52 B. trilineola, page 92 WING PATTERN

B. umbrosa, page 102 22 WING INFUSCATION

B. kirki, page 58 B. obliqua, page 74 B. psidii, page 86

B. laticaudus, page 62

ZERO VITTAE - DACUS

D. longicornis, page 126 D. solomonensis, page 128 D. axanus, page 124

D. aequalis, page 122

23 CERATITIDINI

C. capitata, page 130 C. quilicii, page 132 C. rosa, page 134

TRYPETINAE

A. fraterculus, page 138 A. serpentina, page 140 R. cerasi, page 142

R. cingulata, page 144 R. completa, page 146 R. fausta, page 148

R. pomonella, page 150 T. curvicauda, page 152

24 PHYTALMIINAE

D. pornia, page 136

DROSOPHILIDAE

D. suzukii, page 154

25 Bactrocera albistrigata (de Meijere) WHITE STRIPED FRUIT FLY

postpronotal lobes yellow (anteromedial corners dark)

scutum glossy black

lateral vittae tapered, transverse band ending at ia. setae across wing

pale costal band

broad black basal broad anal band on scutellum streak

terga III-V with a narrow to medium large oval width medial line and facial spots dark lateral markings

1 mm

2 mm Lateral thorax Head Abdominal variation

2 mm

Scutum variation Fore, mid, hind legs 26 27

. EXOTIC ). and and LURE Cue lure. Cue Syzygium B. frauenfeldi HOST RANGE B. albistrigata SIMILAR SPECIES which is present in which is present OTHER COMMENTS OTHER can occasionally have can occasionally can be observed at EIF3L. at can be observed B. frauenfeldi, orange-brown postpronotal lobes. postpronotal orange-brown B. frauenfeldi B. frauenfeldi has a lesser pest status than the introduced than the introduced has a lesser pest status It is most similar to It B. caledoniensis, B. frauenfeldi, B. parafrauenfeldi B. albistrigata ASIA: Malaysia, Peninsular Thailand, Islands, Andaman and Indonesia Singapore East Malaysia, ISLANDS: OCEANIA/PACIFIC Island Christmas north Queensland. It is separated from this species by its yellow this species by from is separated north Queensland. It This species belongs to the frauenfeldi complex which includes complex the frauenfeldi to species belongs This B. trilineola. complex. Some resolution between especially especially between Some resolution complex. postpronotal lobes, although although lobes, postpronotal Anacardiaceae, Clusiaceae, Moraceae, Myrtaceae and Sapotaceae. and Sapotaceae. Myrtaceae Moraceae, Clusiaceae, Anacardiaceae, Polyphagous, recorded from 12 hosts from seven families including families seven 12 hosts from from recorded Polyphagous, Current molecular markers do not adequately separate species in this separate adequately do not molecular markers Current

Commercial hosts include mango, guava, jackfruit and several cultivated cultivated jackfruit and several guava, hosts include mango, Commercial having only been recorded from 12 hosts (compared to 109 for 109 for to 12 hosts (compared from been recorded only having SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION B. frauenfeldi Bactrocera aquilonis (May) NORTHERN TERRITORY FRUIT FLY

scutum yellow-brown

lateral vittae tapered, ending at costal cells tinted, with ia. setae microtrichia in both

narrow costal band broad anal streak

terga III-V pale orange-brown, may have a pale medial line and lateral bands

2 mm

Abdominal variation Lateral thorax

1 mm

2 mm Head Fore, mid, hind legs 28 PRESENT 29

). and LURE species Cue lure. Cue HOST RANGE B. tryoni and is difficult and is difficult to SIMILAR SPECIES B. neohumeralis , B. tryoni Terminalia ferdinandiana forms. No existing molecular No forms. B. tryoni belongs to the to belongs from other species in this complex. other from goat plum ( goat B. tryoni hybrid. Before this period it was recorded this period it was recorded Before hybrid. . It is most similar to is most similar to . It B. aquilonis B. tryoni Bactrocera aquilonis B. melas complex, which includes complex, distinguish from paler distinguish from OCEANIA: and the Australia Western of northern areas Australia: Territory Northern markers separate separate markers Polyphagous. Host records from the Northern Territory after 1984 are are 1984 after Territory the Northern from records Host Polyphagous. from 17 hosts from seven families including including Curcurbitaceae, including including Curcurbitaceae, families seven 17 hosts from from

Myrtaceae, Rosaceae, Rutaceae and Sapotaceae. Its major host is billy- major host is Its and Sapotaceae. Rutaceae Rosaceae, Myrtaceae, suspected to be of a be of to suspected SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera bancroftii (Tryon)

lateral vittae parallel black scutum sided, enclosing ia. setae

black band very narrow on apex of wide anal costal band scutellum streak

abdomen orange- brown and usually broad, may have a faint medial line on terga IV and V

2 mm

Abdominal variation Lateral thorax

legs entirely pale

2 mm 1 mm Fore, mid, hind legs Head 30 31

PRESENT . 2+3, LURE B. musae HOST RANGE and SIMILAR SPECIES It can be separated by can be separated It (Moraceae) and mulberry. (Moraceae) B. bancroftii Weakly attracted to methyl eugenol. eugenol. methyl to attracted Weakly The abdomen is also usually quite broad. quite abdomen is also usually The Bactrocera musae. Oligophagous on Moraceae: Indian cockspur Indian on Moraceae: Oligophagous adequately distinguish adequately Maclura cochinchinensis having a very narrow costal band confluent with R band confluent costal narrow a very having Several molecular markers, including COI, EIF3L and POP4, COI, including molecular markers, Several a wide anal streak and a band on the apex of the scutellum. and a band on the apex of a wide anal streak OCEANIA: of areas coastal and Islands Strait Torres Australia: Islands Guinea; Solomon New Papua Queensland; ASIA: Timor West Indonesia: This species is similar to species is similar to This

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera breviaculeus (Hardy)

lateral vittae bluntly tapered, ending at ia. setae red-brown costal cells clear or scutum with extremely pale tint

narrow costal band terga III-V with an broad anal usually confluent with

indistinct narrow streak R2+3 (can overlap) medial line and lateral bands

2 mm

Lateral thorax

1 mm 2 mm Head Fore, mid, hind legs 32 33

. PRESENT . . 2+3 2+3 and but LURE Cue lure. Cue B. trivialis B. rufofuscula HOST RANGE SIMILAR SPECIES (Harvey’s buttonwood). buttonwood). (Harvey’s but can be separated by but can be separated Glochidion ferdinandii is similar to the exotic is similar to B. rufofuscula It can be difficult to separate from this species. to separate can be difficult It Glochidion harveyanum and without the costal band distinctly overlapping R overlapping band distinctly and without the costal Bactrocera breviaculeus Oligophagous on Euphorbiaceae: on Euphorbiaceae: Oligophagous It is similar to the native is similar to It can be separated by having a brown scutum (instead of black), of scutum (instead a brown having by can be separated OCEANIA: Queenslandeastern and Islands Strait Torres Australia: Guinea New Papua Rockhampton; south to without the lateral bands on terga III broadly joining at the top, the top, joining at III broadly terga bands on without the lateral All molecular markers suggest close similarity to similarity to close suggest All molecular markers cells, and normally not having the costal band overlapping R band overlapping the costal having not and normally cells, III joining at the top, occasionally having a faint tint in the costal tint a faint having occasionally the top, III joining at being a paler medium brown, without the lateral bands on terga bands on terga without the lateral being a paler medium brown,

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera bryoniae (Tryon)

usually a larger species

cells bc and c tinted, microtrichia covering black scutum outer half of cell c only

broad costal band to R terga III-V broad anal 4+5 with a distinct streak wraparound T pattern

2 mm

Abdominal variation Lateral thorax

1 mm 2 mm Scutum variation Head Fore, mid, hind legs 34 PRESENT 35

. 4+5 from from LURE costal costal band. HOST RANGE B. bryoniae SIMILAR SPECIES except it has a distinct except complex species in having a species in having complex B. trivialis formate) than cue lure (Royer 2015). 2015). (Royer than cue lure formate) B. dorsalis black scutum and T on the abdomen, except it is black scutum and T on the abdomen, except including Cucurbitaceae, Loganiaceae, Musaceae, Musaceae, Loganiaceae, including Cucurbitaceae, wraparound T pattern on terga III-V and a very broad broad III-V and a very on terga T pattern wraparound Cue lure group. In north Queensland this species was In group. lure Cue the abovementioned morphologically similar species. morphologically the abovementioned cue-responsive and has a very broad costal band to R to band costal broad and has a very cue-responsive Polyphagous recorded from nine hosts from five families five nine hosts from from recorded Polyphagous significantly more attracted to Melolure (raspberry ketone (raspberry Melolure to attracted more significantly All molecular markers consistently separate separate consistently All molecular markers Superficially similar to similar Superficially OCEANIA: Territory, Northern Australia, northern Western Australia: New Papua south as Sydney; as far Australia eastern Guinea ASIA: (Papua) Indonesia Passifloraceae and Solanaceae. Main commercial host is chilli. commercial Main and Solanaceae. Passifloraceae In cue traps it is most similar to it is most similar to cue traps In

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera cacuminata (Hering) WILD TOBACCO FRUIT FLY, SOLANUM FRUIT FLY

red-brown scutum with lateral vittae a lanceolate tapered, ending at pattern ia. setae

costal band expands at wing apex terga III-V narrow anal around R with a distinct streak 4+5 wraparound T

2 mm

Abdominal Scutum variation Lateral thorax variation

1 mm 2 mm Head Head variation Fore, mid, hind legs 36 PRESENT 37

from from LURE B. opiliae (Solanaceae). HOST RANGE (tobacco tree) tree) (tobacco Methyl eugenol. Methyl SIMILAR SPECIES B. cacuminata , but differs in having , but differs , except it has an apically , except . setae. It also invariably has also invariably It . setae. ia B. opiliae lanceolate pattern on the scutum. pattern lanceolate B. carambolae Solanum mauritianum or B. dorsalis the abovementioned morphologically similar species. morphologically the abovementioned Monophagous on Monophagous a dark lanceolate pattern on the scutum, whereas on the scutum, whereas pattern a dark lanceolate OCEANIA: to Queensland York, Cape from Australia eastern Australia: district, Victoria the east Gippsland lanceolate pattern to W shape. It can also appear superficially W shape. It to pattern lanceolate can range from a mostly black scutum, to black centrally, to a to black centrally, to black scutum, a mostly from can range and tapering vittae ending at the and tapering vittae ending at All molecular markers consistently separate separate consistently All molecular markers Similar to the native non-pest the native to Similar similar to some similar to expanding costal band, a distinct wraparound T on the abdomen band, a distinct wraparound expanding costal a wider anal streak, a distinct wraparound T on the abdomen and T on the abdomen a distinct wraparound a wider anal streak,

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera carambolae Drew and Hancock CARAMBOLA FRUIT FLY

black scutum, may have small brown patches around mesonotal costal band slightly suture and inside overlapping R2+3 and postpronotal lobes expanding around

R4+5, may appear broad parallel sided lateral fish-hook like vittae enclosing ia. setae

narrow anal terga III-V with a streak wraparound T pattern

rectangular shaped band on anterolateral corners large facial tergum IV spots

1 mm 2 mm

Head Lateral thorax Abdominal variation

all tibia usually dark, sometimes oval preapical spot on outer surface of fore femora

2 mm

Scutum variation Fore, mid, hind legs 38 39

EXOTIC . but and LURE B. musae B. dorsalis B. opiliae B. opiliae HOST RANGE , Methyl eugenol. Methyl SIMILAR SPECIES B. musae from from is similar to the Australian the Australian is similar to , sometimes having a subapical spot on the spot a subapical having , sometimes B. carambolae 4+5 but differs in having a broader costal band overlappingcostal band a broader in having but differs rectangular bands on tergum IV. It is similar to the exotic the exotic is similar to It IV. bands on tergum rectangular include carambola, mango, sapodilla, jackfruit and guava. sapodilla, jackfruit and mango, include carambola, tergum IV and dark tibia. It is similar to the Australian the Australian to is similar IV and dark tibia. It tergum ASIA: East Malaysia, Malaysia, Peninsular Vietnam, Thailand, Islands Andaman Indonesia, Singapore, SOUTH AMERICA: Brazil Guiana, Guyana, French Suriname, but differs in having longer more parallel sided lateral vittae, an sided lateral parallel longer more in having but differs Bactrocera carambolae expanding costal band, a more defined T on the abdomen and band, a more expanding costal differs in having an expanding costal band, rectangular bands on costal band, an expanding in having differs fore femora, and having rectangular bands present on tergum IV. IV. on tergum bands present rectangular and having femora, fore B. dorsalis

Annonaceae, Clusiaceae, Lauraceae, Moraceae, Myrtaceae, Oxalidaceae, Myrtaceae, Moraceae, Lauraceae, Clusiaceae, Annonaceae, All molecular markers separate separate All molecular markers Rhamnaceae, Rutaceae, Sapotaceae, and Solanaceae. Major commercial hosts Major commercial and Solanaceae. Sapotaceae, Rutaceae, Rhamnaceae, and expanding apically around R around and expanding apically SIMILAR SPECIES, SPECIES, Polyphagous. Recorded from 75 hosts from 26 families including Anacardiaceae, including Anacardiaceae, 26 families 75 hosts from from Recorded Polyphagous. 2+3 HOST & LURE & HOST DISTRIBUTION R Bactrocera caryeae (Kapoor)

pale postpronotal lobes with dark anteromedial corners very narrow lateral vittae, parallel black scutum sided or tapering, narrow ending at or costal band before ia. setae confluent

with R2+3

scutellum yellow with a broad black basal band negligible anal streak terga III-V with a wraparound T pattern

large dark preapical spots on all femora

fore and hind tibiae dark 2 mm

Lateral thorax

1 mm large elongate facial spots

2 mm

Head Fore, mid, hind legs 40 41

EXOTIC but LURE HOST RANGE Methyl eugenol. Methyl B. endiandrae SIMILAR SPECIES defined T) on the abdomen. is similar to the Australian the Australian is similar to , but differs in having a wraparound T (instead of T (instead a a wraparound in having , but differs Commercial hosts include citrus, mango and guava. hosts include citrus, Commercial Anacardiaceae, Myrtaceae, Rutaceae and Sapotaceae. Rutaceae Myrtaceae, Anacardiaceae, and dark apical spots on all femora. It is similar to the exotic the exotic is similar to It on all femora. and dark apical spots ASIA: Southern India B. kandiensis lobes, very narrow vittae, very narrow anal streak, dark fore tibia dark fore anal streak, narrow vittae, very narrow very lobes, differs in having dark anteromedial corners on the postpronotal on the postpronotal corners dark anteromedial in having differs Bactrocera caryeae Polyphagous. Recorded from eight hosts from six families including six families hosts from eight from Recorded Polyphagous.

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera correcta (Bezzi) GUAVA FRUIT FLY

black scutum, sometimes brown laterally and posteriorly

broad parallel both costal cells sided lateral vittae entirely devoid enclosing ia. setae of microtrichia

T on abdomen oval spot across apex of anal streak wing at R confined to 4+5 narrow black cell cup lateral margins on terga IV and V

2 mm

Abdominal variation Lateral thorax

Scutum variation

transverse elongate facial spots sometimes meeting in centre 1 mm 2 mm Head Fore, mid, hind legs 42 43

EXOTIC ). from from LURE Syzygium. B. zonata but differs but differs that is more is more that black scutum. HOST RANGE B. correcta SIMILAR SPECIES similar to similar to B. dorsalis B. correcta but has a predominantly but has a predominantly sapodilla and edible B. zonata is similar to the exotic the exotic is similar to attractive than methyl eugenol (Wee et al. 2017). et eugenol (Wee than methyl attractive All molecular markers consistently separate separate consistently All molecular markers microtrichia. It is similar to is similar to It microtrichia. in having transverse facial spots, an elongate apical wing spot apical wing an elongate spots, facial transverse in having instead of a continuous costal band and costal cells devoid of devoid cells band and costal costal a continuous of instead ASIA: Bhutan, Lanka, Sri Bangladesh, India, Nepal, Pakistan, Malaysia, Peninsular southern China, Thailand, Myanmar, Vietnam Bactrocera correcta the abovementioned morphologically similar species (but most morphologically the abovementioned

Polyphagous, including the families Anacardiaceae, Myrtaceae, Rhamnaceae, Myrtaceae, Anacardiaceae, including the families Polyphagous, Beta-caryophyllene is a species-specific attractant for attractant is a species-specific Beta-caryophyllene SIMILAR SPECIES, SPECIES, HOST & LURE & HOST and Sapotaceae. Commercial hosts include mango, guava, carambola, mandarin, carambola, guava, hosts include mango, Commercial and Sapotaceae. DISTRIBUTION Bactrocera curvipennis (Froggatt)

small species

black scutum cells bc and c slightly tinted lateral vittae taper to with microtrichia end before ia. setae in both

infuscation on r-m crossvein

wing with a broad fuscous costal very broad band to R anal streak 4+5

transverse band along anterior and lateral margin of tergum III, dark anterolateral corners of terga IV and V

2 mm

Lateral thorax

very small facial spots which may be present or absent 1 mm 2 mm Head Fore, mid, hind legs 44 45

EXOTIC LURE B. tryoni HOST RANGE SIMILAR SPECIES (Royer et al. 2018). et (Royer but has infuscation but has infuscation OTHER COMMENTS COMMENTS OTHER and is generally smaller. and is generally is almost absent, but where but where is almost absent, B. bryoniae is still an important pest species. pest is still an important B. curvipennis B. curvipennis This species is similar to species is similar to This was a dominant pest species in until the until Caledonia in New pest species was a dominant in the 1960s. Host fruit surveys show that where where that show fruit surveys in the 1960s. Host on the r-m vein, doesn’t have a medial line on the abdomen a medial have doesn’t vein, on the r-m peach, coffee, white sapote, citrus, sweet pepper and tomato. tomato. pepper and citrus, sweet sapote, white peach, coffee, OCEANIA: only) Island (Aneityum Vanuatu Caledonia, New Polyphagous. Recorded from 41 hosts from 20 families including 20 families 41 hosts from from Recorded Polyphagous. B. tryoni has low populations populations has low

B. tryoni Commercial hosts include mango, guava, custard apple, , carambola, apple, papaya, custard guava, hosts include mango, Commercial Anacardiaceae, Annonaceae, Caricaceae, Moraceae, Myrtaceae, Oxalidaceae, Myrtaceae, Moraceae, Caricaceae, Annonaceae, Anacardiaceae, Passifloraceae, Rhamnaceae, Rosaceae, Rubiaceae, Rutaceae and Solanaceae. Rutaceae Rubiaceae, Rosaceae, Rhamnaceae, Passifloraceae, is represented by dense populations populations dense by is represented SIMILAR SPECIES, SPECIES, Bactrocera curvipennis Isoeugenol (significantly more attractive than its previous known lure, cue lure) cue lure) lure, known than its previous attractive more (significantly Isoeugenol HOST & LURE & HOST introduction of of introduction DISTRIBUTION Bactrocera dorsalis (Hendel) ORIENTAL FRUIT FLY

scutum black to red-brown narrow costal band that dips in

at end of R2+3 broad parallel sided lateral vittae enclosing ia. setae

terga III-V with a wraparound T pattern, very narrow may be minimal or also anal streak have dark anterolateral corners of terga IV and V

2 mm

Scutum variation Abdominal variation Lateral thorax

1 mm large facial spots

fore and hind

tibiae dark 2 mm

Head Fore, mid, hind legs 46 47

.

EXOTIC ). ia. LURE B. musae

B. carambolae from from . RPA2 does not . RPA2 HOST RANGE Methyl eugenol. Methyl SIMILAR SPECIES B. dorsalis but differs in having broad in having but differs B. endiandrae . setae, a narrow costal band that costal a narrow . setae, and ia separate separate B. endiandrae but differs in having a narrow costal band a narrow in having but differs stage. The majority of edible fruit is susceptible. majority of stage. The and triangular shaped anterolateral dark corners and triangular shaped anterolateral B. carambolae 2+3 broadly wraps round the lateral edges of terga III-V. terga edges of lateral the round wraps broadly , a very narrow anal streak, and in not having a T that having and in not anal streak, narrow , a very but differs in having a very narrow anal streak, narrow anal streak, very narrow a in having but differs 2+3 from from , having broad parallel sided lateral vittae enclosing the vittae enclosing sided lateral parallel broad , having B. carambolae 2+3 B. musae Widespread, predominantly in sub-saharan predominantly Widespread, AFRICA: countries African Bhutan, Bangladesh, Nepal, Lanka, Sri India, ASIA: Pakistan, Vietnam, Thailand, Kong, Hong Taiwan, China, Myanmar, Indonesia Philippines, Singapore, Malaysia, Laos, Cambodia, Guinea, Palau, New Papua Island, Christmas OCEANIA: Tahiti Hawaii, Similar to the Australian the Australian to Similar B. dorsalis dips in at R dips in at Similar to to Similar confluent with R confluent setae, and in general having a more distinct T shape on the abdomen. distinct T shape on the a more having and in general setae, parallel sided vittae enclosing the sided vittae enclosing parallel on abdominal terga IV and V (if present) (rectangular in (rectangular (if present) IV and V on abdominal terga

Polyphagous on over 300 hosts and can oviposit in some fruit in the hard green green in some fruit in the hard and can oviposit 300 hosts on over Polyphagous SIMILAR SPECIES, SPECIES, DISTRIBUTION HOST & LURE & HOST costal band that dips in at R dips in at band that costal Similar to the Australian the Australian to Similar All markers separate separate All markers Bactrocera endiandrae (Perkins and May)

black scutum narrow costal band

that overlaps R2+3 lateral vittae taper to end before the ia. setae

terga III-V with a wraparound broad anal T pattern streak

2 mm

Abdominal variation Lateral thorax

1 mm 2 mm Head Fore, mid, hind legs 48 49

. PRESENT , 2+3 spp. from from LURE B. musae, terga III-V. terga B. musae

and HOST RANGE Endiandra Methyl eugenol. Methyl SIMILAR SPECIES B. endiandrae (see comments under (see comments B. dorsalis B. carambolae and

B. dorsalis Similar to to Similar All molecular markers adequately separate separate adequately All molecular markers northeast NSW AUSTRALIA: to south Australia eastern and Islands Strait Torres setae, and a wraparound T pattern with broad lateral bands on bands lateral with broad T pattern and a wraparound setae, those species for differentiating characters). Similar to Similar characters). differentiating those species for Primarily oligophagous on Lauraceae, particularly particularly oligophagous on Lauraceae, Primarily ia. but differs in having a narrower costal band that just overlaps R overlaps just costal band that a narrower in having but differs a wide anal streak, vittae tapering to a point and ending before the and ending before a point vittae tapering to a wide anal streak,

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera facialis (Coquillett)

small species black scutum, rarely partly red-brown

scutellum with narrow short narrow to very lateral vittae to deep basal band a.sa. setae, narrow that reaches across present or absent costal band scutellum

anal streak absent terga III-V with a or narrow and faint thick medial line

broad lateral bands mesopleural stripe or dark anterolateral reaching almost to corners of terga III-V postpronotal lobes

2 mm

Scutum variation Abdominal Lateral thorax variation

facial spots absent

1 mm 2 mm Head Fore, mid, hind legs 50 EXOTIC 51 LURE Cue Cue lure. B. frauenfeldi HOST RANGE SIMILAR SPECIES a band across the wing. a band across can appear superficially similar to similar can appear superficially

Polyphagous. Recorded on 64 hosts from 30 families on 64 hosts from Recorded Polyphagous. Myrtaceae, Passifloraceae, Rosaceae, Rubiaceae, Rutaceae, Rubiaceae, Rosaceae, Passifloraceae, Myrtaceae, Leguminosae, Malvaceae, Meliaceae, Moraceae, Musaceae, Musaceae, Moraceae, Meliaceae, Malvaceae, Leguminosae, Sapotaceae and Solanaceae. Main commercial hosts include Main commercial and Solanaceae. Sapotaceae mango, soursop, avocado, , guava, Brazilian cherry, cherry, Brazilian guava, breadfruit, avocado, soursop, mango, OCEANIA: Tonga including Anacardiaceae, Annonaceae, Caricaceae, Lauraceae, Lauraceae, Caricaceae, Annonaceae, including Anacardiaceae, Bactrocera facialis which is established in north Queensland, but differs in not having in not but differs which is established in north Queensland,

passionfruit, granadilla, citrus, sapodilla and star apple (Leblanc et al. 2012). et citrus, sapodilla and star apple (Leblanc granadilla, passionfruit, SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera frauenfeldi (Schiner) MANGO FRUIT FLY

postpronotal lobes black, occasionally red-brown

scutum glossy black

narrow pale lateral vittae transverse band costal band short or long across wing

broad anal streak

abdominal terga III-V scutellum yellow orange-brown with with a black triangle a broad medial and of variable size two broad lateral longitudinal black bands

2 mm 2 mm

Humeral calli variation Abdominal variation Lateral thorax

1 mm 2 mm Scutum variation Head Fore, mid, hind legs 52 53

PRESENT LURE B. kirki

HOST RANGE SIMILAR SPECIES but has lateral vittae but has lateral OTHER COMMENTS COMMENTS OTHER

can be observed at EIF3L. at can be observed , star apple, sapodilla and abiu. B. trilineola but has dark ppn. lobes, a medialbut has dark ppn. lobes, B. frauenfeldi Syzygium but has dark ppn. lobes (sometimes red- (sometimes but has dark ppn. lobes B. psidii and for this species in north Queensland (Royer 2015). 2015). this species in north Queensland (Royer for

B. albistrigata B. albistrigata brown ppn. lobes). Similar to to Similar lobes). ppn. brown OCEANIA: Guinea New Papua Papua, West Australia: Torres Strait and north Queensland south to south Queensland north and Strait Torres Australia: Marshall Kiribati, Micronesia, of States Federated Townsville; Moluccas, Island, Solomon Nauru, Palau, Islands, a distinct band on the wing. Current molecular markers do not molecular markers a distinct band on the wing. Current on the wing. Similar to to on the wing. Similar but has dark ppn. lobes, lateral vittae present and a distinct band vittae present lateral but has dark ppn. lobes, Similar to to Similar present and facial spots instead of a facial mask. Similar to to mask. Similar a facial of instead spots and facial present line and lateral bands on terga III-V, legs with dark patterning and legs with dark patterning III-V, bands on terga line and lateral was detected on the northernmost part of mainland Queensland in 1974 mainland Queensland in 1974 on the northernmost part of was detected adequately separate species in this complex. Some resolution between Some resolution species in this complex. separate adequately

Caricaceae, Moraceae, Musaceae, Myrtaceae, Oxalidaceae, Passifloraceae, Passifloraceae, Oxalidaceae, Myrtaceae, Musaceae, Moraceae, Caricaceae, Townsville, and this seems to be the natural southernmost limit for this species. this species. southernmost limit for be the natural and this seems to Townsville, SIMILAR SPECIES, SPECIES, Polyphagous. Recorded on 109 hosts from 37 families including Anacardiaceae, including Anacardiaceae, 37 families on 109 hosts from Recorded Polyphagous. citrus, carambola, guava, papaya, edible papaya, guava, citrus, carambola, HOST & LURE & HOST DISTRIBUTION B. frauenfeldi Rutaceae, Sapotaceae and Solanaceae. Commercial hosts include mango, banana,mango, hosts include Commercial and Solanaceae. Sapotaceae Rutaceae, no containment measures. While it occurs in high numbers in Cairns it is rarely found in found it is rarely in Cairns in high numbers it occurs While measures. no containment Cue group. Melolure (raspberry ketone formate) was found to be a stronger attractant be a stronger to was found formate) ketone (raspberry Melolure group. Cue and had spread to Townsville by 1997. It has not spread further since despite there being there despite further since spread has not 1997. It by Townsville to and had spread Bactrocera jarvisi (Tryon) JARVIS’ FRUIT FLY

scutum gold to red-brown

no sa. setae lateral vittae parallel sided enclosing ia. setae costal band expands at wing apex

T on terga III-V joining band between ppn. lobe and notoplueron

2 mm

Post notum variation Abdominal variation Lateral thorax

1 mm 2 mm Head Fore, mid, hind legs 54 55

PRESENT is LURE Syzygium. B. zonata HOST RANGE edible SIMILAR SPECIES but differs in having in having a but differs from all other species. all other from methyl eugenol-responsive. eugenol-responsive. methyl B. jarvisi B. zonata is similar to is similar to

also zingerone and cue-responsive whereas whereas and cue-responsive also zingerone joining band between the ppn. lobe and notopleural calli, a and notopleural the ppn. lobe joining band between Bactrocera jarvisi OCEANIA: Australia: Broome in Western Australia, Northern Territory, Territory, Northern Australia, in Western Broome Australia: Sydney south to South Wales New Queensland, anal streak present and a more defined T on the abdomen. It is defined T on the abdomen. and a more present anal streak continuous costal band (instead of an isolated apical oval spot), apical oval an isolated of band (instead costal continuous Strongly attracted to zingerone (Fay 2012) and weakly to cue lure. cue lure. to 2012) and weakly (Fay zingerone to attracted Strongly

Solanaceae. Commercial hosts include mango, guava, papaya, persimmon, papaya, guava, hosts include mango, Commercial Solanaceae. Punicaceae, Rosaceae, Rubiaceae, Rutaceae, Sapindaceae, Sapotaceae and Sapotaceae Sapindaceae, Rutaceae, Rubiaceae, Rosaceae, Punicaceae, All tested molecular markers clearly separate separate clearly molecular markers All tested Annonaceae, Arecaceae, Caricaceae, Clusiaceae, Curcurbitaceae, Lauraceae, Lauraceae, Curcurbitaceae, Clusiaceae, Caricaceae, Arecaceae, Annonaceae, Malpighiaceae, Moraceae, Musaceae, Myrtaceae, Oxalidaceae, Passifloraceae, Passifloraceae, Oxalidaceae, Myrtaceae, Musaceae, Moraceae, Malpighiaceae, soursop, avocado, banana, pomegranate, apple, peach, pear, coffee, citrus and coffee, apple, peach, pear, banana, pomegranate, avocado, soursop, SIMILAR SPECIES, SPECIES, Polyphagous. Recorded from 84 hosts from 29 families including Anacardiaceae, including Anacardiaceae, 29 families 84 hosts from from Recorded Polyphagous. HOST & LURE & HOST DISTRIBUTION Bactrocera kandiensis Drew and Hancock

ppn. lobes yellow with red-brown anteromedial corners black scutum, sometimes with brown patches around mesonotal suture and inside ppn. lobes very narrow parallel sided lateral vittae ending at ia. setae

narrow costal band confluent with R scutellum with 2+3 very narrow a broad black anal streak basal band

T on terga III-V

2 mm

Abdominal variation Scutum variation Lateral thorax

very large femora with facial spots dark apical markings

fore and mid

1 mm tibiae dark

Head Fore, mid, hind legs. Image: Luc Leblanc 56 57

EXOTIC . LURE ; data but has Garcinia scutellum. scutellum. B. caryeae HOST RANGE B. dorsalis Methyl eugenol. Methyl but has narrower but has narrower SIMILAR SPECIES B. dorsalis from from lacking for lacking for Oxalidaceae and Rutaceae. Oxalidaceae ). Similar to to ). Similar B. endiandrae B. kandiensis but differs in having a defined T on the in having but differs B. caryeae vittae, a very narrow anal streak, dark markings anal streak, narrow vittae, a very B. caryeae

Similar to to Similar on the apices of the femora and a broad basal band on the basal and a broad the femora of on the apices abdomen (instead of wraparound) and in having blunt ended blunt and in having wraparound) of abdomen (instead ASIA: Lanka Sri the Australian rainforest species rainforest the Australian narrower vittae, dark markings on the apices of the femora and a the femora of vittae, dark markings on the apices narrower broad basal band on the scutellum. Similar to some specimens of some specimens to Similar basal band on the scutellum. broad vittae (sometimes taper in taper vittae (sometimes

and Allwood et al. (1999) only record this species from mango and mango this species from record only al. (1999) et and Allwood Polyphagous. The pest status of this species in unclear. Drew & Romig (2013) & Romig Drew species in unclear. this of pest status The Polyphagous. SIMILAR SPECIES, SPECIES, spp. However, Tsuruta et al. (1997) record it from 21 hosts from 13 plant families 13 plant 21 hosts from it from al. (1997) record et Tsuruta spp. However, All tested molecular markers clearly separate separate clearly molecular markers All tested HOST & LURE & HOST DISTRIBUTION including Anacardiaceae, Caricaceae, Clusiaceae, Lauraceae, Moraceae, Myrtaceae, Myrtaceae, Moraceae, Lauraceae, Clusiaceae, Caricaceae, including Anacardiaceae, Bactrocera kirki (Froggatt)

postpronotal lobes yellow (anteromedial margins black) tinge around r-m and dm-cu crossveins scutum glossy black

narrow pale lateral vittae absent costal band

scutellum glossy black with extreme lateral margins yellow

all terga black except for two longitudinal orange-brown bands over terga II-V

2 mm

Lateral thorax

large facial spots

1 mm 2 mm Head Fore, mid, hind legs 58 59

EXOTIC LURE Cue lure. Cue , carambola, HOST RANGE SIMILAR SPECIES but has facial spotsbut has facial Syzygium complex but differs in not but differs complex B. trilineola (instead of a mask), yellow ppn. lobes. ppn. lobes. a mask), yellow of (instead B. frauenfeldi but lacks vittae, has a wide black band on the but lacks vittae, has a wide black band on

B. psidii but lacks vittae, has yellow ppn. lobes, very wide band very ppn. lobes, but lacks vittae, has yellow on the scutellum and very thick medial line on abdomen; and very on the scutellum the r-m and dm-cu cross veins), and additionally: similar to and additionally: veins), and dm-cu cross the r-m but lacks vittae, has a very wide band on the scutellum and wide band on the scutellum but lacks vittae, has a very Similar to to Similar OCEANIA: Western Samoa, American Samoa, Fiji Islands (Rotuma), Islands Fiji Samoa, American Samoa, Western Tahiti , and Futuna, Wallis Tonga, Similar to members of the of members to Similar having a distinct transverse band on the wing, (only infuscation on infuscation on the wing, (only band a distinct transverse having scutellum and has two yellow longitudinal bands on the abdomen. longitudinal yellow and has two scutellum B. frauenfeldi passionfruit, noni, citrus, abiu, sweet pepper, tomato and eggplant. tomato pepper, noni, citrus, abiu, sweet passionfruit, B. albistrigata

very thick medial line on abdomen, similar to abdomen, similar to thick medial line on very Annonaceae, Caricaceae, Lauraceae, Myrtaceae, Oxalidaceae, Passifloraceae, Passifloraceae, Oxalidaceae, Myrtaceae, Lauraceae, Caricaceae, Annonaceae, custard apple, soursop, papaya, avocado, guava, edible guava, avocado, papaya, soursop, apple, custard Polyphagous. Recorded on 42 hosts from 19 families including Anacardiaceae, including Anacardiaceae, 19 families on 42 hosts from Recorded Polyphagous. SIMILAR SPECIES, SPECIES, similar to similar to Rosaceae, Rutaceae and Solanaceae. Commercial hosts include , mango, hosts include cashew, Commercial and Solanaceae. Rutaceae Rosaceae, HOST & LURE & HOST DISTRIBUTION Bactrocera kraussi (Hardy)

red-brown scutum

may have narrow slightly slightly tinted costal band tapering vittae costal cells overlapping to ia. setae and wing R2+3

broad basal band on scutellum broad anal streak

terga III and IV with narrow lateral spots (most mesopleural defined on terga V) stripe

1 mm

2 mm

Head Lateral thorax dark apices of femora

2 mm Abdominal variation Fore, mid, hind legs 60 PRESENT 61 LURE

(Royer 2015). (Royer HOST RANGE SIMILAR SPECIES but differs in having tint in having but differs B. tryoni is similar to is similar to on the scutellum and dark apices of the femora. of and dark apices on the scutellum

Torres Strait Islands and northeast Queensland northeast and Islands Strait Torres Rosaceae, Rubiaceae, Rutaceae, Sapindaceae, Sapotaceae Sapotaceae Sapindaceae, Rutaceae, Rubiaceae, Rosaceae, and Solanaceae. Commercial hosts include mango, banana, hosts include mango, Commercial and Solanaceae. Bactrocera kraussi OCEANIA: : Australia south to Townsville south to longer less tapered vittae, lateral spots on the abdomen (instead on spots vittae, lateral less tapered longer only in both costal cells (instead of tint and microtrichia in both), and microtrichia tint of (instead cells costal in both only of a wraparound T), narrow mesopleural stripe, broad basal band stripe, broad mesopleural T), narrow a wraparound of Polyphagous. Recorded on 106 hosts from 31 families including 31 families on 106 hosts from Recorded Polyphagous. grumichama, guava, feijoa, carambola, peach, citrus and tamarind. carambola, feijoa, grumichama, guava, Anacardiaceae, Musaceae, Myrtaceae, Oxalidaceae, Passifloraceae, Passifloraceae, Oxalidaceae, Myrtaceae, Musaceae, Anacardiaceae,

Isoeugenol (significantly more attractive than its previous known lure, cue lure) cue lure) lure, known than its previous attractive more (significantly Isoeugenol SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera laticaudus (Hardy)

scutum dark red-brown

lateral vittae taper infuscation on ending at or before r-m cross vein ia. setae

broad anal narrow costal streak band terga III-V dark brown to black

femora with dark 2 mm apices

Scutum variation Lateral thorax

1 mm

2 mm Abdominal Head Fore, mid, hind legs 62 variation 63

PRESENT ). LURE setae. ia. HOST RANGE Methyl eugenol. Methyl SIMILAR SPECIES Planchonella obovoidea ( responsive (instead of cue lure). of (instead responsive in having infuscation on the r-m infuscation in having B. psidii

is similar to Oligophagous on Sapotaceae. Major host is black ash on Sapotaceae. Oligophagous It differs in having a brown scutum, a yellow scutellum and no scutellum yellow scutum, a a brown in having differs It ustralia: Torres Strait Islands and northeast Queensland and northeast Islands Strait Torres ustralia: OCEANIA: A Townsville south to infuscation on the dm-cu cross vein and being methyl vein on the dm-cu cross infuscation eugenol- B. laticaudus vein, a dark abdomen and lateral vittae tapering to end at vittae tapering to and lateral a dark abdomen vein,

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera latifrons (Hendel) SOLANUM FRUIT FLY

black scutum

costal band lateral postsutural narrow, expanding vittae parallel around apex of R4+5 sided enclosing ia. setae

narrow anal streak all terga entirely orange-brown

mesopleural stripe extending to anterior npl. seta

apical spots

2 mm on femora

Lateral thorax

large oval facial spots

1 mm 2 mm Head Fore, mid, hind legs 64 65

EXOTIC . LURE B. musae but differs HOST RANGE from from complex species. complex SIMILAR SPECIES but differs in havingbut differs but differs in having but differs B. carambolae B. latifrons B. dorsalis B. musae B. dorsalis and contradict this (Mziray et al. (2010)). et this (Mziray contradict eggplant, winter melon and ivy gourd melon winter eggplant, in having no markings on the abdomen. in having is similar to is similar to

Bactrocera latifrons an expanding costal band, no markings on the abdomen, andan expanding costal an expanding costal band, longer parallel sided vittae and apical parallel band, longer an expanding costal ASIA: Southern Myanmar, Bangladesh, Lanka, Sri India, Pakistan, Vietnam, Laos, Thailand, Taiwan, (Okinawa), China, Japan Brunei Singapore, Malaysia, Peninsular OCEANIA: Burundi Kenya, Tanzania, Hawaii AFRICA: spots on the femora. It is similar to is similar to It on the femora. spots All molecular markers clearly distinguish clearly All molecular markers apical spots on the femora. It is similar to is similar to It on the femora. apical spots

other species, it is more attractive than protein bait; however some records some records bait; however than protein attractive species, it is more other 59 hosts from 14 families including Myrtaceae, Oxalidaceae, Passifloraceae, Passifloraceae, Oxalidaceae, including Myrtaceae, 14 families 59 hosts from Peck 2001). While it’s not as attractive as cue lure and methyl eugenol are to eugenol are and methyl as cue lure as attractive not it’s 2001). While Peck Rhamnaceae and Rutaceae. Main commercial hosts include , chilli, hosts include commercial Main and Rutaceae. Rhamnaceae Alpha-ionol and cade oil is the best attractant for this species (McQuate and this species (McQuate for Alpha-ionol and cade oil is the best attractant Polyphagous. Primarily a pest of Solanaceae and Cucurbitaceae. Recorded on Recorded and Cucurbitaceae. Solanaceae a pest of Primarily Polyphagous. SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera melanotus (Coquillett)

postpronotal lobes yellow

black scutum

pale infuscation black notopleura on r-m and dm- cu crossveins vittae absent

scutellum narrow pale black costal band

all terga black

2 mm legs mostly dark

Lateral thorax

1 mm 2 mm

facial spots absent or small and pale

Head Fore, mid, hind legs 66 EXOTIC 67 LURE Cue lure. Cue HOST RANGE in having a black in having SIMILAR SPECIES B. passiflorae is similar to is similar to

pale postpronotal lobes, a black scutellum and dark legs. a black scutellum lobes, pale postpronotal scutum without vittae and black abdomen, but differs in having but differs scutum without vittae and black abdomen, Polyphagous. Recorded on 31 hosts from 18 families including 18 families on 31 hosts from Recorded Polyphagous. OCEANIA: Cook Island Cook Bactrocera melanotus Anacardiaceae, Annonaceae, Caricaceae, Lauraceae, Moraceae, Moraceae, Lauraceae, Caricaceae, Annonaceae, Anacardiaceae, cherry, guava, rose apple, carambola, citrus, sapodilla and tomato. citrus, sapodilla and tomato. apple, carambola, rose guava, cherry,

Myrtaceae, Oxalidaceae, Rubiaceae, Rutaceae, Sapotaceae and Solanaceae. Sapotaceae Rutaceae, Rubiaceae, Oxalidaceae, Myrtaceae, SIMILAR SPECIES, SPECIES, Commercial hosts include mango, papaya, avocado, breadfruit, jackfruit, Brazilian breadfruit, avocado, papaya, hosts include mango, Commercial HOST & LURE & HOST DISTRIBUTION Bactrocera minax (Enderlein) CHINESE CITRUS FRUIT FLY

large Dacus-like species

lateral and medial vittae cells bc and c tinted

a. sa. setae absent

broad costal band

overlapping R4+5 with darker spot terga III-V with at apex a T pattern and dark anterolateral corners on terga IV and V

2 mm yellow band from postpronotal lobe to near notopleuron

Lateral thorax

very elongated facial spots

1 mm 2 mm Head Fore, mid, hind legs 68 EXOTIC 69 LURE HOST RANGE in being a larger SIMILAR SPECIES OTHER COMMENTS COMMENTS OTHER setae, having the costal having setae, tangerine and cumquat. tangerine and cumquat. a. sa. B. tsuneonis lateral bands on terga IV and V. bands on terga lateral Weak response to methyl eugenol. methyl to response Weak with a dark spot at the apex, and having the apex, at with a dark spot is similar to 4+5 is univoltine, having only one generation per year. one generation only having is univoltine, Oligophagous. Recorded on eight hosts in Rutaceae. hosts in Rutaceae. on eight Recorded Oligophagous. like fly with lateral and medial vittae, a band between the and medial vittae, a band between with lateral fly like

Bactrocera minax Dacus ASIA: China southern India, northern Nepal, Bhutan, Commercial hosts include orange, lemon, , mandarin, lemon, pomelo, hosts include orange, Commercial the abdomen. It differs in lacking the differs the abdomen. It band well overlapping R overlapping band well postpronotal lobe and notopleuron, a wide costal band, and a T on a wide costal and notopleuron, lobe postpronotal Bactrocera minax

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera musae (Tryon) BANANA FRUIT FLY

black scutum, can be partly brown

lateral vittae costal band dark slightly taper to and distinctly

touch or enclose overlapping R2+3 ia. setae

narrow anal streak terga III-V varies from having T shape to being uniformly orange-brown

2 mm

Abdominal variation Scutum variation Lateral thorax

1 mm

2 mm Head Fore, mid, hind legs 70 71

. . PRESENT ; ia , a but 2+3 LURE

B. dorsalis B. bancroftii B. opiliae HOST RANGE and Methyl eugenol. Methyl and SIMILAR SPECIES B. opiliae has a distinct T). Similar to has a distinct T). Similar , a slightly wider anal streak, , a slightly 2+3 from from B. endiandrae B. opilae from from B. musae papaya and guava are occasional hosts. hosts. occasional are and guava papaya but differs in having a wider costal band,costal a wider in having but differs B. musae , but differs in not having an apical band on the having in not , but differs setae, and usually less patterning on the abdomen. less patterning and usually setae,

Torres Strait Islands and northeast Queensland northeast and Islands Strait Torres B. endiandrae B. bancroftii to but differs in having a wider costal band that overlaps at R overlaps band that costal a wider in having but differs OCEANIA: Australia: associated Guinea and New Papua Townsville, south to Islands, the Solomon and Archipelago islands, Bismarck West Papua West Similar to to Similar T or wraparound T on the abdomen. Very similar to similar to T on the abdomen. Very T or wraparound Similar to several other methyl other several to Similar eugenol-responsive species. Similar narrower anal streak, longer less tapered vittae and less of a distinct vittae and less of less tapered longer anal streak, narrower scutellum and having a wider costal band and narrower anal streak. band and narrower a wider costal and having scutellum B. dorsalis

however RPA2 does not separate separate does not RPA2 however Polyphagous. Recorded on 16 hosts from nine families, although these are these are although nine families, on 16 hosts from Recorded Polyphagous. All molecular markers separate separate All molecular markers Musaceae, Caricaceae and Myrtaceae. Primary economic host is banana, but host economic Primary and Myrtaceae. Caricaceae Musaceae, and usually less patterning on the abdomen ( less patterning and usually differs in having a wider costal band that overlaps R overlaps band that costal a wider in having differs SIMILAR SPECIES, SPECIES, mostly single records or specimens that may be dubious. Confirmed families are are families be dubious. Confirmed may or specimens that single records mostly the exotic the exotic HOST & LURE & HOST DISTRIBUTION slightly wider anal streak, slightly tapering and shorter vittae that usually end at the end at usually that vittae tapering and shorter slightly wider anal streak, slightly Bactrocera neohumeralis (Hardy) LESSER QUEENSLAND FRUIT FLY

postpronotal lobes dark brown

narrow lateral postsutural costal band vittae taper scutum dark ending at or brown before ia. setae

costal cells terga III-V with a tinted with medial line and broad anal microtrichia in broad lateral bands streak both joining across terga III and IV

2 mm

Scutum variation Lateral thorax

1 mm

2 mm Abdominal variation Head Fore, mid, hind legs 72 73

PRESENT from LURE Cue lure. lure. Cue B. tryoni HOST RANGE , mate at dusk. at , mate B. neohumeralis. SIMILAR SPECIES OTHER COMMENTS COMMENTS OTHER B. tryoni sapodilla, abiu, capsicum and tomato. sapodilla, abiu, capsicum and tomato. but differs in having dark postpronotal lobes dark postpronotal in having but differs is unusual in mating in the middle of the day. the day. in the middle of is unusual in mating

B. tryoni, No current molecular markers separate separate markers molecular current No Torres Strait Islands, eastern Australia south to Australia eastern Islands, Strait Torres Most species, including the similar Most Similar to to Similar and being generally darker, with more dark marking on the legs. with more darker, and being generally OCEANIA: Australia: Guinea New Papua Harbour, Coffs Polyphagous. Recorded on over 160 hosts from 44 families including 44 families 160 hosts from on over Recorded Polyphagous. Bactrocera neohumeralis

Moraceae, Musaceae, Myrtaceae, Oxalidaceae, Passifloraceae, Rosaceae, Passifloraceae, Oxalidaceae, Myrtaceae, Musaceae, Moraceae, Anacardiaceae, Annonaceae, Caricaceae, Clusiaceae, Ebenaceae, Lauraceae, Ebenaceae, Clusiaceae, Caricaceae, Annonaceae, Anacardiaceae, SIMILAR SPECIES, SPECIES, Rubiaceae, Rutaceae, Sapotaceae, Solanaceae, and Vitaceae. Commercial hosts Commercial and Vitaceae. Solanaceae, Sapotaceae, Rutaceae, Rubiaceae, include mango, custard apple, rollinia, date palm, persimmon, mulberry, banana, mulberry, palm, persimmon, date apple, rollinia, custard include mango, HOST & LURE & HOST DISTRIBUTION carambola, passionfruit, loquat, apple, plum, peach, pear, citrus, coffee, star apple, star citrus, coffee, apple, plum, peach, pear, loquat, passionfruit, carambola, Bactrocera obliqua (Malloch)

r-m cross vein black scutum oblique with pale infuscation

lateral vittae very narrow short and tapering costal band

scutellum yellow with a broad medial medium longitudinal band anal streak

all abdominal terga black

femora with dark apical markings

2 mm Fore, mid, hind legs

2 mm

black face

1 mm

Head Lateral thorax 74 EXOTIC 75 LURE HOST RANGE

SIMILAR SPECIES and longer r-m cross-vein. r-m and longer (water apple, Malay apple). apple, Malay (water Syzygium , but differs in having a band across the a band across in having , but differs B. psidii Weakly attracted to isoeugenol (Royer et al. 2018). et (Royer isoeugenol to attracted Weakly

setae, dark apices of the femora, and a more obliqueand a more the femora, of dark apices setae, Similar to to Similar a. sa. the scutellum (instead of a triangle), shorter vittae that end level to end level vittae that a triangle), shorter of (instead scutellum OCEANIA: Papua New Guinea: Bismarck Archipelago and Archipelago Bismarck Guinea: New Papua Bougainville Island Bougainville

Edible hosts include guava and edible hosts include guava Edible Polyphagous. Recorded on six hosts from three families including Myrtaceae. Myrtaceae. including families three on six hosts from Recorded Polyphagous. SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera occipitalis (Bezzi)

black scutum, may have small red-brown patches peripherally

costal band broad parallel sided distinctly or subparallel lateral overlapping R2+3 vittae ending at or and expanding at enclosing ia. setae apex

terga III-V with a distinct narrow anal wraparound T, or dark streak anterolateral corners

very thick femora fulvous, large oval medial line tibia dark facial spots

1 mm

2 mm Leg variation Fore, mid, hind legs Head

2 mm

Abdominal variation Scutum variation Lateral thorax 76 77

EXOTIC . LURE HOST RANGE Methyl eugenol. Methyl SIMILAR SPECIES on terga IV and sometimes V) IV and sometimes on terga but differs in usually having subparallel having in usually but differs , a very thick medial line and defined wraparound and defined wraparound thick medial line , a very 2+3 B. carambolae

ASIA: Indonesian Brunei, Sabah, (Malaysian Borneo Philippines, Kalimantan) Similar to to Similar overlapping R overlapping T or dark anterolateral corners on terga III-V (instead of rectangles rectangles of III-V (instead on terga corners T or dark anterolateral vittae, no spot on the fore femora, a broader costal band distinctly costal a broader femora, on the fore vittae, no spot Polyphagous. Recorded from four hosts from four families including families four hosts from four from Recorded Polyphagous.

Anacardiaceae, Cucurbitaceae, Myrtaceae and Rutaceae. Commercial hosts Commercial Rutaceae. and Myrtaceae Cucurbitaceae, Anacardiaceae, SIMILAR SPECIES, SPECIES, include mango, calabash, guava and calamondin (a cumquat mandarin hybrid). mandarin hybrid). and calamondin (a cumquat calabash, guava include mango, HOST & LURE & HOST DISTRIBUTION Bactrocera oleae (Rossi) OLIVE FRUIT FLY

small fly

black scutum costal band absent, small lateral and spot around

medial vittae apex of R4+5 absent

anal streak terga II-IV with dark absent anterolateral corners

1 mm

Scutum variation Lateral thorax

1 mm 2 mm Abdominal variation Head Fore, mid, hind legs 78 79

EXOTIC . LURE

in having Oleaceae). HOST RANGE apex of the wing apex of SIMILAR SPECIES B. passiflorae Olea europea: and in having a black scutum and spot in having B. melanotus is quite distinct in lacking vittae and having a distinct in lacking vittae and having is quite

B. correcta the latter has been found to be more attractive. attractive. be more to has been found the latter and Monophagous on olives ( on olives Monophagous

spot at the apex of the wing. It is superficially similar to similar superficially is It the wing. the apex of at spot

Bactrocera oleae EUROPE: Albania, Croatia, Cyprus, France, Italy, Malta, Montenegro, Montenegro, Malta, Italy, France, Cyprus, Croatia, Albania, Switzerland Spain, Slovenia, Serbia, Portugal, AFRICA: Algeria, Angola, Egypt, Eritrea, Ethiopia, Kenya, Libya, Kenya, Ethiopia, Eritrea, Egypt, Algeria, Angola, South, Seychelles, Island Réunion Morocco, Mauritius, Tunisia Sudan, Africa, ASIA: Saudi Pakistan, Lebanon, Jordan, Israel, Iran, India, Georgia, Turkey Syria, Arabia, THE AMERICAS: Mexico USA (California), abdomen with lateral colouring on terga II-IV and a spot at the at II-IV and a spot on terga colouring abdomen with lateral B. tuberculata is superficially similar to similar is superficially at the wing apex instead of a costal band, but lacks lateral vittae. It lateral band, but lacks a costal of wing apex instead the at a black scutum and lacking lateral vittae, but has an orange brown brown vittae, but has an orange a black scutum and lacking lateral

Olive fruit fly lure or protein bait in McPhail traps attracts both sexes. In the USA sexes. both attracts McPhail traps bait in or protein lure fruit fly Olive SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera opiliae (Drew and Hardy) FALSE ORIENTAL FRUIT FLY

scutum dull black or red-brown with irregular longitudinal black bands narrow costal band confluent with or slightly

lateral vittae parallel overlapping R2+3 or subparallel, touch or enclose ia. setae

narrow anal T shape on terga III-V, streak may be reduced to medial line

2 mm

Abdominal variation Scutum variation Lateral thorax

medium sized facial spots

fore tibiae slightly tinted,

1 mm hind tibiae dark 2 mm Head Fore, mid, hind legs 80 PRESENT 81 LURE specimens but differs in but differs HOST RANGE B. cacuminata. Methyl eugenol. Methyl of the Northern of (Cucurbitaceae). 2 SIMILAR SPECIES All tested molecular All tested from from OTHER COMMENTS COMMENTS OTHER B. opiliae B. musae 2+3. (Opiliaceae) as its host. (Opiliaceae) (if at all). These two species two all). These (if at B. opiliae 2+3 in having a red-brown scutum a red-brown in having . This caused alarm and intensive intensive caused alarm and . This Mukia maderaspatana is most similar to is most similar to Opilia acmentata but has a slightly wider anal streak, and a wider anal streak, but has a slightly B. dorsalis (Opiliaceae), although there are two records records two are there although (Opiliaceae), from from B. cacuminata

B. dorsalis markers clearly distinguish clearly markers Bactrocera opiliae band only slightly overlapping R overlapping slightly band only can be very difficult to distinguish. Some difficult can be very can be similar to can be similar to Opilia amentacea OCEANIA: Australia: northern Western Australia, Northern Territory and Territory Northern Australia, northern Western Australia: Queensland and north central north-west with a lanceolate pattern but differs in not having an expanding having in not but differs pattern with a lanceolate costal band and having a straight T on the abdomen (instead of (instead T on the abdomen a straight band and having costal usually having a more defined T on the abdomen and the costalT on the abdomen and the defined a more having usually

was collected in methyl eugenol traps in Melville Island in the Northern Island in Melville eugenol traps in methyl was collected Territory and Western Australia. Ecological, physiological and genetic studies and genetic physiological Ecological, Australia. and Western Territory field surveys showed the fly was distributed over 358 000km fly was distributed the showed field surveys SIMILAR SPECIES, SPECIES, showed it to be a distinct species with it to showed wraparound T). It is similar to is similar to T). It wraparound B. opiliae Monophagous on Monophagous HOST & LURE & HOST DISTRIBUTION slightly wider costal band that doesn’t dip in at the end of R the end of dip in at doesn’t band that wider costal slightly Territory in 1975 and was thought to be to in 1975 and was thought Territory Bactrocera pallida (Perkins and May)

overall golden fly

lateral vittae tapered costal band but square ended to overlaps R end at ia. setae 2+3

narrow anal streak terga III-V with T or medial line ventral thorax black around legs

1 mm

Lateral thorax

1 mm

2 mm Head Fore, mid, hind legs 82 83

PRESENT ). LURE B. zonata Rubiaceae). HOST RANGE Methyl eugenol. Methyl SIMILAR SPECIES (absent in (absent from all similar species. from Nauclea orientalis: B. pallida but differs in having a complete costal band complete a in having but differs

B. zonata streak (instead of no anal streak), and microtrichia in cell c in cell and microtrichia no anal streak), of (instead streak The COI barcode separates separates barcode COI The OCEANIA: Australia: Northern Territory, Torres Strait Islands and Islands Strait Torres Territory, Northern Australia: Guinea New Papua Brisbane, south to Queensland (instead of an oval spot on the apex of the wing), a narrow anal the wing), a narrow on the apex of spot an oval of (instead Similar to to Similar Monophagous on Leichhardt tree ( tree on Leichhardt Monophagous

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera passiflorae (Froggatt) FIJIAN FRUIT FLY

small species

black scutum postpronotal lobes black

vittae absent

scutellum yellow narrow anal streak

all terga black, sometimes tergum V with paler posterior margin or with a medial line 1 mm

Abdominal variation Lateral thorax

1 mm facial spots absent

pale legs

2 mm Head Fore, mid, hind legs 84 85

EXOTIC LURE from the from Cue lure. Cue HOST RANGE SIMILAR SPECIES B. melanotus is similar to is similar to scutellum and black postpronotal lobes. and black postpronotal scutellum

Bactrocera passiflorae Polyphagous. Recorded from 49 hosts in 28 families including 49 hosts in 28 families from Recorded Polyphagous. OCEANIA: Fiji Islands, Niue, Wallis Niue, Islands, Fiji in being all black but differs in having pale legs and in having in being all black but differs Islands Cook

guava, rose apple, Malay apple, cocoa, coffee, citrus, star apple and chilli. citrus, coffee, apple, cocoa, apple, Malay rose guava, mango, papaya, avocado, passionfruit, Tahitian chestnut, breadfruit, jackfruit, breadfruit, chestnut, Tahitian passionfruit, avocado, papaya, mango, Malvaceae, Moraceae, Myrtaceae and Solanaceae. Commercial hosts include Commercial and Solanaceae. Myrtaceae Moraceae, Malvaceae, SIMILAR SPECIES, SPECIES, Anacardiaceae, Caricaceae, Rutaceae, Passifloraceae, Lauraceae, Leguminosae, Lauraceae, Passifloraceae, Rutaceae, Caricaceae, Anacardiaceae, HOST & LURE & HOST DISTRIBUTION Bactrocera psidii (Froggatt) SOUTH SEA GUAVA FRUIT FLY

postpronotal lobes black yellow except scutum anteromedial corner black very narrow very pale costal lateral vittae band taper ending at psa. setae

scutellum with a large black triangle very narrow anal streak infuscation on r-m and dm-cu crossveins

all terga black,

tergum V may be 1 mm pale centrally with a medial line

Lateral thorax

small facial spots present or absent

1 mm 2 mm Head Fore, mid, hind legs 86 EXOTIC 87 LURE Cue lure. Cue HOST RANGE SIMILAR SPECIES granadilla and plum. granadilla in possessing a black setae, face fulvous with or fulvous face setae, a. sa. B. obliqua is similar to

than dm-cu crossvein, and being cue lure responsive. responsive. and being cue lure than dm-cu crossvein, abdomen and thorax, infuscation on the r-m and dm-cu on the r-m infuscation abdomen and thorax, longer vittae that end at the the end at vittae that longer Bactrocera psidii a black triangle on the scutellum (instead of a band across it), band across a of (instead a black triangle on the scutellum OCEANIA: New Caledonia New crossveins and a dark pattern on the scutellum. It differs in having differs It on the scutellum. and a dark pattern crossveins without small facial spots, legs entirely pale, r-m cross vein shorter vein cross pale, r-m legs entirely spots, without small facial

persimmon, acerola, mulberry, guava, rose apple, Malay apple, carambola, apple, Malay rose guava, mulberry, acerola, persimmon, Vitaceae. Commercial hosts include mango, soursop, custard apple, papaya, custard soursop, hosts include mango, Commercial Vitaceae. Myrtaceae, Oxalidaceae, Passifloraceae, Punicaceae, Rosaceae, Rutaceae andRutaceae Rosaceae, Punicaceae, Passifloraceae, Oxalidaceae, Myrtaceae, Polyphagous. Recorded on 31 hosts from 16 families including Anacardiaceae, including Anacardiaceae, 16 families on 31 hosts from Recorded Polyphagous. SIMILAR SPECIES, SPECIES, Annonaceae, Caricaceae, Ebenaceae, Euphorbiaceae, Malpighiaceae, Moraceae, Moraceae, Malpighiaceae, Euphorbiaceae, Ebenaceae, Caricaceae, Annonaceae, HOST & LURE & HOST DISTRIBUTION Bactrocera pyrifoliae Drew and Hancock

black scutum, may be partly brown narrow costal band confluent

with R2+3, dips in lateral vittae at end of R taper to end 2+3 before ia. setae

very narrow anal streak

terga III and IV each with its own ‘T’ pattern femora dark which continues down apically terga V, terga III-V may also have a wraparound tibiae dark T or straight T

2 mm Abdominal variation Fore, mid, hind legs

2 mm

large facial spots

1 mm Head Lateral thorax 88 89

EXOTIC in LURE the abdomen. HOST RANGE , all tibiae dark, and nashi pear. B. endiandrae 2+3 SIMILAR SPECIES OTHER COMMENTS COMMENTS OTHER is similar to the Australian the Australian is similar to Found only at elevations of 1,000–1,500m. of elevations at only Found

Possible weak response to cue lure (Drew and Romig 2013). and Romig (Drew cue lure to response weak Possible ASIA: Vietnam northern Thailand, Northern Bactrocera pyrifoliae having a black scutum, tapered vittae and wraparound T on the vittae and wraparound a black scutum, tapered having dark apices of the femora and sometimes having a double T on having and sometimes the femora of dark apices streak, a costal band that dips in at the end of R the end of dips in at band that a costal streak, abdomen. It differs in having large facial spots, a very narrow anal very narrow a spots, facial large in having differs abdomen. It

and Rosaceae. Commercial hosts include guava, sour cherry, plum, peach sour cherry, hosts include guava, Commercial and Rosaceae. SIMILAR SPECIES, SPECIES, Polyphagous. Recorded from seven hosts from five families including Myrtaceae families including five hosts from seven from Recorded Polyphagous. HOST & LURE & HOST DISTRIBUTION Bactrocera rufofuscula (Drew and Hancock)

red-brown scutum lateral vittae taper slightly ending at ia. setae narrow costal band slightly overlapping R costal cells 2+3 colourless

broad anal terga III-V with indistinct streak wraparound T that joins across the top of tergum III

1 mm

Lateral thorax

1 mm

2 mm Head Fore, mid, hind legs 90 91

.

PRESENT ). LURE and the Cue lure. Cue B. trivialis 2+3 HOST RANGE B. breviaculeus SIMILAR SPECIES except it is dark red-brown it is dark red-brown except all of tergum III in tergum all of except it has a dark brown scutum it has a dark brown except B. breviaculeus Recorded from seven hosts in four families. hosts in four seven from Recorded B. trivialis lateral bands join across the top of tergum III. tergum of the top bands join across lateral

Polyphagous on native rainforest (no economic hosts). fruits (no economic rainforest on native Polyphagous (instead of red-brown), the costal band overlaps R band overlaps costal the red-brown), of (instead (instead of black), and the lateral bands on the abdomen join bands black), and the lateral of (instead OCEANIA: Australia: Torres Strait Islands, Cape York Peninsula, north Peninsula, York Cape Islands, Strait Torres Australia: Mackay south to Queensland Similar to the native the native to Similar All molecular markers suggest close similarity to similarity to close suggest All molecular markers Similar to the exotic the exotic to Similar narrowly across the top of tergum III (instead of broadly covering covering broadly of III (instead tergum of the top across narrowly

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera trilineola (Drew) VANUATU FRUIT FLY

dark postpronotal lobes very narrow black scutum pale costal band

transverse band across wing scutellum black with yellow lateral margins broad anal streak

terga black except for two broad longitudinal yellow bands on terga II-V either side of a broad medial line

2 mm

Scutum variation Lateral thorax

face entirely black

1 mm 2 mm Head Fore, mid, hind legs 92 EXOTIC 93 LURE Cue lure. Cue HOST RANGE SIMILAR SPECIES complex. It is similar to It complex. frauenfeldi but differs in lacking vittae, having but differs in having dark postpronotal lobes. dark postpronotal in having but differs in having a distinct band on the in having but differs wing, dark postpronotal lobes and a black face. and a lobes wing, dark postpronotal B. albistrigata B. albistrigata, B. kirki

and differs from from differs OCEANIA: Vanuatu It is similar to is similar to It a black face, having a broad black band on the scutellum, it also on the scutellum, black band a broad having a black face, Similar to other species in the other to Similar B. frauenfeldi Annonaceae, Caricaceae, Lauraceae, Moraceae, Musaceae, Myrtaceae, Myrtaceae, Musaceae, Moraceae, Lauraceae, Caricaceae, Annonaceae,

Oxalidaceae, Rutaceae and Sapindaceae. Commercial hosts include mango, hosts include Commercial and Sapindaceae. Rutaceae Oxalidaceae, Polyphagous. Recorded on 31 hosts from 17 families including Anacardiaceae, including Anacardiaceae, 17 families on 31 hosts from Recorded Polyphagous. SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION soursop, avocado, breadfruit, guava, rose apple, Malay apple, carambola and citrus. apple, carambola apple, Malay rose guava, breadfruit, avocado, soursop, Bactrocera trivialis (Drew)

black scutum

lateral vittae subparallel, touch or costal band

enclose ia. setae overlaps R2+3

terga III-V dark with a wide anal streak medial pale area from posterior margin of tergum III to tergum V, may have indistinct medial line

1 mm

Lateral thorax

1 mm 2 mm Head Fore, mid, hind legs 94 EXOTIC 95 LURE Cue lure. Cue HOST RANGE SIMILAR SPECIES but differs in having but differs OTHER COMMENTS COMMENTS OTHER fruit fly incursions for 20 years. for 20 incursions fruit fly B. breviaculeus and lateral bands broadly joining on tergum III. joining on tergum bands broadly lateral

B. rufofuscula a black scutum (instead of dark brown or red-brown) and the or red-brown) dark brown of a black scutum (instead OCEANIA: Papua New Guinea and West Papua West and Guinea New Papua Polyphagous. Recorded on 17 hosts from 10 families including 10 families on 17 hosts from Recorded Polyphagous. Similar to to Similar

Commercial hosts include mango, guava, carambola, peach, citrus and chilli. carambola, guava, hosts include mango, Commercial most years. These are promptly detected and eradicated under the Exotic Fruit Fruit the Exotic under and eradicated detected promptly are These most years. Anacardiaceae, Myrtaceae, Oxalidaceae, Rosaceae, Rutaceae, and Solanaceae. Rutaceae, Rosaceae, Oxalidaceae, Myrtaceae, Anacardiaceae, SIMILAR SPECIES, SPECIES, Fly in Torres Strait Response Plan, jointly run by the federal and state agriculture agriculture and state the federal run by Plan, jointly Response Strait Torres in Fly Seasonal incursions into the Torres Strait Islands from Papua New Guinea occur New Papua from Islands Strait the Torres into Seasonal incursions HOST & LURE & HOST DISTRIBUTION departments. This response program has been successfully eradicating exotic pest exotic eradicating has been successfully program response This departments. Bactrocera tryoni (Froggatt) QUEENSLAND FRUIT FLY

postpronotal lobes yellow scutum red-brown costal band

cells bc and c tinted with overlaps R2+3 microtrichia in both and usually tinted

lateral vittae taper ending broad anal before ia. setae streak

terga III-V generally red- brown with an indistinct wraparound T

1 mm

Abdominal variation Lateral thorax

1 mm 2 mm Head Fore, mid, hind legs 96 97 . PRESENT ;

LURE B. tryoni B. kraussi but has pale and HOST RANGE SIMILAR SPECIES OTHER COMMENTS COMMENTS OTHER to be very close to to close be very to B. neohumeralis B. breviaculeus but differs in having tinted costal tinted in having but differs from but differs in having microtrichia in both microtrichia in having but differs but is overall red-brown instead of golden of instead red-brown but is overall B. erubescentis B. tryoni complex species. COI barcodes suggest the suggest barcodes species. COI complex apple, sapodilla, capsicum, chilli and tomato. B. breviaculeus B. kraussi

B. tryoni B. aquilonis and has more of a wraparound T on the abdomen (instead a wraparound of and has more postpronotal lobes and is overall red-brown (instead of dark of (instead red-brown and is overall lobes postpronotal Australian native native Australian of minimal markings). Similar to to minimal markings). Similar of OCEANIA: Australia: Northern Territory, Queensland, South Wales New Queensland, Territory, Northern Australia: Island Pitcairn , French Caledonia, New and Victoria; Similar to to Similar cells with microtrichia in both and sharply tapering vittae that end tapering vittae that and sharply in both with microtrichia cells brown). Similar to to Similar brown). . setae. Similar to to Similar . setae. ia

lacking lateral spots on tergum V and lacking dark apices of the femora. femora. the of dark apices V and lacking on tergum spots lacking lateral in in 1970. It has become a dominant pest in both places. pest in both a dominant has become in 1970. It Polynesia in French Annonaceae, Caricaceae, Curcurbitaceae, Lauraceae, Moraceae, Musaceae, Musaceae, Moraceae, Lauraceae, Curcurbitaceae, Caricaceae, Annonaceae, was accidentally introduced into New Caledonia around 1969 and detected and detected 1969 around Caledonia New into introduced was accidentally All molecular markers distinguish All molecular markers SIMILAR SPECIES, SPECIES, costal cells, sharply tapered vittae, lacking a broad basal band on the scutellum, vittae, lacking a broad tapered sharply cells, costal before the before Cue lure. Melolure (raspberry ketone formate) was 1.5 times more attractive in north attractive was 1.5 times more formate) ketone (raspberry Melolure lure. Cue custard apple, papaya, carambola, passionfruit, loquat, apple, peach, coffee, star peach, coffee, apple, loquat, passionfruit, carambola, papaya, apple, custard Sapotaceae, Solanaceae, and Vitaceae. Major commercial hosts include mango, Major commercial Vitaceae. and Solanaceae, Sapotaceae, HOST & LURE & HOST DISTRIBUTION Myrtaceae, Oleaceae, Oxalidaceae, Passifloraceae, Rosaceae, Rubiaceae, Rutaceae, Rubiaceae, Rosaceae, Passifloraceae, Oxalidaceae, Oleaceae, Myrtaceae, however, no markers separate no markers however, Queensland (Royer 2015) but was slightly less attractive in Sydney (Dominiak et al. 2015). (Dominiak et in Sydney less attractive 2015) but was slightly Queensland (Royer Polyphagous. Recorded from over 200 hosts in 49 families including Anacardiaceae, including Anacardiaceae, families 200 hosts in 49 over from Recorded Polyphagous. B. tryoni Bactrocera tsuneonis (Miyake) CITRUS FRUIT FLY

large Dacus-like species broad costal band

confluent with 4+5R and overlapping towards wing apex cells bc and c tinted

lateral and medial vitta anal streak absent terga III-V with a T, sometimes with dark anterolateral corners tergum IV

2 mm

yellow band connecting postpronotal lobe and notopleuron

Lateral thorax

oval facial spots

1 mm 2 mm Head Fore, mid, hind legs 98 99

EXOTIC LURE Fortunella HOST RANGE and SIMILAR SPECIES No known record. known No Citrus in general body shape in general B. minax is similar to is similar to and lacking a dark apical spot on the costal band. on the costal and lacking a dark apical spot 4+5

with R . setae, having a narrower costal band that is mostly confluent confluent is mostly band that costal a narrower having . setae, between the postpronotal lobes and notopleuron, possessing and notopleuron, lobes the postpronotal between ASIA: Taiwan China, southern Japan, Bactrocera tsuneonis and colour patterns but differs in having a complete joining bandcomplete a in having but differs patterns and colour a.sa

Oligophagous. Recorded on nine hosts in Rutaceae, all on nine hosts in Rutaceae, Recorded Oligophagous. species. Commercial hosts include orange, mandarin, tangerine and cumquat. mandarin, tangerine orange, hosts include species. Commercial SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera tuberculata (Bezzi)

black scutum

lateral vittae costal band parallel sided reduced to apical enclosing ia. oval spot setae

scutellum yellow anal streak confined within cell cup abdomen black

2 mm

Lateral thorax

1 mm 2 mm Head Fore, mid, hind legs 100 EXOTIC 101 in in LURE B. oleae B. correcta HOST RANGE and Methyl eugenol. Methyl SIMILAR SPECIES in having lateral vittae lateral in having and a black abdomen. B. correcta B. oleae is similar to is similar to papaya, wax jambu, peach and sapodilla. papaya,

elongate spot and black scutum. It differs from from differs and black scutum. It spot elongate Bactrocera tuberculata ASIA: Vietnam, Thailand, China, southern Myanmar, Bhutan, Bangladesh having a black abdomen, and from a black abdomen, and from having possessing an incomplete costal band that is reduced to an apical to is reduced band that costal possessing an incomplete

Caricaceae, Myrtaceae and Sapotaceae. Commercial hosts include mango, Commercial and Sapotaceae. Myrtaceae Caricaceae, SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Polyphagous. Recorded on 11 hosts from eight families, including Anarcardiaceae, including Anarcardiaceae, families, eight on 11 hosts from Recorded Polyphagous. Bactrocera umbrosa (Fabricius) BREADFRUIT FRUIT FLY

black scutum

lateral vittae broad subparallel

three bands across wing

terga III-V vary from having no markings to having a short medial line to having a medial line and two broad lateral bands

1 mm

Abdominal variation Lateral thorax

1 mm

2 mm Scutum variation Head Fore, mid, hind legs 102 103

EXOTIC . LURE

B. confluens HOST RANGE Methyl eugenol. Methyl SIMILAR SPECIES and jackfruit and chempadek. jackfruit and chempadek. B. seguyi bears no close resemblance to other to resemblance no close bears

pest methyl eugenol-responsive species that have bands have species that eugenol-responsive pest methyl

superficially appear similar e.g. appear similar superficially Bactrocera umbrosa on the wings and similar pattern on the abdomen that may on the abdomen that on the wings and similar pattern species. It is easily recognised by the three broad transverse transverse broad the three by recognised is easily species. It bands across the wings. However, there are other PNG non- other are there the wings. However, bands across ASIA: Caledonia New Malaysia, southern Thailand, Philippines, Indonesia Philippines, Thailand, southern Malaysia, ISLANDS: OCEANIA/PACIFIC Vanuatu, Islands, Solomon Guinea, New Papua Palau,

Oligophagous. Recorded on four hosts in Moraceae including breadfruit, including breadfruit, hosts in Moraceae on four Recorded Oligophagous. SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Bactrocera xanthodes (Broun) PACIFIC FRUIT FLY

overall golden

long narrow medial vitta lateral vittae begins anterior to mesonotal suture

narrow costal scutellum band bilobed with lateral yellow margins joining band between postpronotal lobes and notopleuron

large yellow spot on pleural region in place of the normal mesopleural stripe

2 mm Lateral thorax

1 mm 2 mm Head Fore, mid, hind legs 104 105

EXOTIC and LURE HOST RANGE SIMILAR SPECIES methyl eugenol) (Royer et al. 2018). et eugenol) (Royer methyl B. neoxanthodes, B. paraxanthodes ( is unique in having a medial vitta, joining is unique in having

) undescribed species) which are not pests and have limited pests and have not ) undescribed species) which are apple, papaya, breadfruit, jackfruit, citrus, star apple and abiu. jackfruit, citrus, star apple breadfruit, apple, papaya, band between the postpronotal lobe and notopleuron and a and notopleuron lobe the postpronotal band between large bilobed scutellum. It is similar to the other species in the species in the other is similar to It scutellum. bilobed large N. Notodacus subgenus OCEANIA: Fiji, Tonga, Niue, Samoa, American Samoa, southern group southern Samoa, American Samoa, Niue, Tonga, Fiji, Polynesia French and Futuna, Wallis Islands, Cook of ( Bactrocera xanthodes distributions (Vanuatu, New Caledonia and Samoa respectively). Caledonia New distributions (Vanuatu, B.

Polyphagous. Recorded on 34 hosts in 20 families including Anacardiaceae, Anacardiaceae, including on 34 hosts in 20 families Recorded Polyphagous. Methyl-isoeugenol (significantly more attractive than its previous known lure, lure, known than its previous attractive more (significantly Methyl-isoeugenol SIMILAR SPECIES, SPECIES, Annonaceae, Caricaceae, Euphorbiaceae, Lauraceae, Moraceae, Passifloraceae, Passifloraceae, Moraceae, Lauraceae, Euphorbiaceae, Caricaceae, Annonaceae, HOST & LURE & HOST DISTRIBUTION Rutaceae, Sapotaceae and Solanaceae. Commercial hosts include mango, custard custard mango, hosts include Commercial and Solanaceae. Sapotaceae Rutaceae, Bactrocera zonata (Saunders) PEACH FRUIT FLY

red-brown scutum costal band reduced to oval spot at narrow parallel sided apex of wing lateral vittae touching or enclosing ia. setae

terga III-V with a T anal streak absent

1 mm

Lateral thorax

1 mm 2 mm Head Fore, mid, hind legs 106 107

EXOTIC . from from LURE

B. correcta notopleuron. . zonata HOST RANGE and Methyl eugenol. Methyl SIMILAR SPECIES but has an incomplete but has an incomplete B. tryoni but has an incomplete costal but has an incomplete B. pallida guava, rose apple, apple and peach. rose guava, B. jarvisi but has a red-brown scutum (instead of scutum (instead but has a red-brown

B. correcta

costal band reduced to an oval apical spot and anal streak and anal streak apical spot an oval to band reduced costal All tested molecular markers clearly separate B separate clearly molecular markers All tested Similar to to Similar band and no joining band between the postpronotal lobe and lobe the postpronotal band and no joining band between ASIA: Bangladesh, India, Lanka, Sri Nepal, Bhutan, Pakistan, Laos, Myanmar Vietnam, Thailand, AFRICA: Island Réunion Libya, Sudan, Egypt, Mauritius, MIDDLE EAST: Yemen Oman, Israel, Iraq, Iran, UAE, black or partly black). Similar to to Similar black). black or partly restricted to cup. Similar to to cup. Similar to restricted

Polyphagous. Recorded on 20 hosts from 15 families including Anacardiaceae, including Anacardiaceae, 15 families on 20 hosts from Recorded Polyphagous. Annonaceae, Caricaceae, Malpighiaceae, Malvaceae, Myrtaceae, Rosaceae and Rosaceae Myrtaceae, Malvaceae, Malpighiaceae, Caricaceae, Annonaceae, SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Rutaceae. Commercial hosts include mango, custard apple, papaya, acerola, okra, acerola, apple, papaya, custard hosts include mango, Commercial Rutaceae. Zeugodacus atrisetosus Perkins

scutum golden or orange brown with dark brown markings lateral vittae medial vitta begin anterior to mesonotal narrow costal suture band

prsc. and a. sa. setae present

narrow anal streak

no pecten on male

medial line on tergum V, sometimes terga III with darker lateral margins

1 mm 1 mm

Head Lateral thorax

108 109

EXOTIC . prsc. LURE Luffa in the . setae. a. sa Zeugodacus HOST RANGE and atrisetosa ) SIMILAR SPECIES OTHER COMMENTS OTHER alternative to fruit rearing. to alternative , but differs in having in having , but differs . For a full list of full list of a . For Austrodacus to generic level. Now known Now generic level. to ( Z. cucumis species see Doorenweerd et al. (2018). et species see Doorenweerd atrisetosa ) Bactrocera Zeugodacus group of subgenera (Hancock and Drew 2015) until and Drew (Hancock subgenera of group Austrodacus ( Cucurbitaceae and Solanaceae. Commercial hosts include Commercial and Solanaceae. Cucurbitaceae Previously known as known Previously OCEANIA: altitudes higher particularly mainland, Guinea New Papua watermelon, cucumber, pumpkin, bitter melon and edible melon pumpkin, bitter cucumber, watermelon, Zeugodacus No known record. Protein or orange ammonia traps are the only are traps ammonia or orange Protein record. known No Very similar to the Australian Australian the similar to Very Zeugodacus Polyphagous. Recorded on eight hosts from three families including families three hosts from on eight Recorded Polyphagous. as Virgilio et al. (2015) elevated al. (2015) elevated et Virgilio

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Zeugodacus choristus May

larger species

red-brown broad sharply scutum with tapered lateral costal band dark markings vittae that begin expands at anterior to wing apex mesonotal suture

medial vitta

infuscation on dm-cu crossvein terga III-V with distinct wraparound T, or T with lateral colouring on terga IV and V

2 mm

Abdominal variation Lateral thorax

1 mm 2 mm Head Fore, mid, hind legs 110 PRESENT 111

LURE Cue lure. Cue HOST RANGE in Cucurbitaceae. SIMILAR SPECIES (striped cucumber) but differs in being more in being more but differs Z. cucurbitae Diplocyclos palmatus Similar to the exotic to Similar Monophagous on Monophagous OCEANIA: Guinea New Papua and Queensland eastern Australia: semicircle) and not having a faint infuscation on the r-m vein. on the r-m infuscation a faint having and not semicircle) vittae, having the costal band expanding less apically (not into a into (not band expanding less apically the costal vittae, having red-brown (rather than golden), having wider medial and lateral wider medial and lateral than golden), having (rather red-brown

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Zeugodacus cucumis (French) CUCUMBER FRUIT FLY

scutum golden

medial vitta

lateral vittae narrow costal begin anterior to band mesonotal suture

prsc. and a. sa. narrow anal setae absent streak

medial line on tergum V, sometimes terga III-V no pecten dark laterally with on male a medial line

2 mm Abdominal variation Lateral thorax

1 mm 2 mm Head Fore, mid, hind legs 112 PRESENT 113

LURE setae. (2018). a. sa. and tomato. ZeugodacusZeugodacus HOST RANGE and SIMILAR SPECIES in the but differs in lacking but differs prsc. OTHER COMMENTS OTHER cucumis ) species see Doorenweerd et al. et species see Doorenweerd Z. atrisetosus Austrodacus ( Zeugodacus Polyphagous. Recorded from 40 hosts in 15 families from Recorded Polyphagous. Bactrocera Similar to the exotic the exotic to Similar OCEANIA: including Queensland Territory, Northern Australia: South Wales New northeast Islands, Strait Torres including the families Caricaceae, Curcurbitaceae, Myrtaceae, Myrtaceae, Curcurbitaceae, Caricaceae, including the families Passifloraceae, and Solanaceae, although many recorded hosts recorded many although and Solanaceae, Passifloraceae,

are single records and may require confirmation. Major commercial hosts commercial Major confirmation. require and may single records are Previously known as known Previously sexes than orange-ammonia or protein traps (Ceratraps) (Royer et al. 2014). et (Royer (Ceratraps) traps or protein than orange-ammonia sexes Cucumber volatile blend was found to generally be more attractive to both to attractive be more generally to blend was found volatile Cucumber to generic level. For a full list of a full list of For generic level. to No known attraction to male lures, although sporadically trapped at cue lure. cue lure. at trapped sporadically although male lures, to attraction known No SIMILAR SPECIES, SPECIES, group of subgenera (Drew 1989) until Virgilio et al. (2015) elevated al. (2015) elevated et Virgilio until 1989) (Drew subgenera of group HOST & LURE & HOST include papaya, cucumber, pumpkin, squash, zucchini, guada bean, passionfruit guada bean, pumpkin, squash, zucchini, cucumber, include papaya, DISTRIBUTION Zeugodacus cucurbitae (Coquillett) MELON FLY golden to red- brown scutum with or without darker markings narrow medial vitta costal band expanding into a lateral vittae semicircular spot begin anterior to at wing apex mesonotal suture

terga III-V with a T, infuscation on dm- dark anterolateral cu, pale infuscation corners of terga IV on r-m and V

1 mm

Scutum variation Abdominal variation Lateral thorax

1 mm 1 mm 2 mm Wing variation Head Fore, mid, hind legs 114 EXOTIC

ASIA: Widely distributed over Asia through the Indian subcontinent, Southeast Asia and southern China DISTRIBUTION AFRICA: Occurs in a number of sub-Saharan countries OCEANIA: Papua New Guinea, Mariana Islands, Solomon Islands, Nauru, Kiribati, Guam, Hawaii

SIMILAR SPECIES Similar to Z. choristus but differs in being more golden, having SIMILAR narrower medial and lateral vittae, having a more semicircular SPECIES, HOST & LURE spot at the wing apex, sometimes having a faint infuscation on the r-m cross vein and having a less defined narrower T on the abdomen. All tested molecular markers clearly separate Z. cucurbitae and Z. choristus. HOST RANGE Polyphagous. Primarily a pest of cucurbits but recorded on 44 hosts from 12 families including Cucurbitaceae, Fabaceae, Malvaceae, Myrtaceae and Solanaceae. Main commercial hosts include watermelon, rockmelon, cucumber, pumpkin, bitter melon, edible Luffa, ivy gourd, wax gourd, bean and tomato. LURE Cue group. Melolure (raspberry ketone formate) was found to be a stronger attractant than cue lure in Hawaii (Casana-Giner et al. 2003). OTHER COMMENTS Previously known as Bactrocera (Zeugodacus) cucurbitae until Virgilio et al. (2015) elevated Zeugodacus to generic level. Now known as Zeugodacus (Zeugodacus) cucurbitae. For a full list of Zeugodacus species see Doorenweerd et al. (2018).

115 Zeugodacus decipiens (Drew) PUMPKIN FRUIT FLY

red-brown scutum, dark large species bands either side lateral vittae of medial vitta begin anterior to mesonotal suture broad costal broad medial vitta band

an irregular recurved pale broad anal fuscous marking streak across wing

terga III-V with a medial line

2 mm Lateral thorax

1 mm 2 mm Head Fore, mid, hind legs 116 EXOTIC 117

LURE in the species see Zeugodacus on the wings. much slower. HOST RANGE decipiens was elevated to was elevated (Cucurbitaceae). (Cucurbitaceae). ) SIMILAR SPECIES OTHER COMMENTS OTHER Zeugodacus Doorenweerd et al. (2018). et Doorenweerd Paradacus ( Zeugodacus ) but its rate of development is development of ) but its rate Cucurbita pepo Bactrocera . For a full list of full list of a . For larvae can co-exist in pumpkin with melon larvae can co-exist placed within pumpkin crops (Royer et al. 2017). et (Royer within pumpkin crops placed decipiens subgenus group until until subgenus group ) Zeugodacus cucurbitae Distinctive in having a medial vitta and recurved pattern a medial vitta and recurved in having Distinctive Previously known as known Previously Monophagous on pumpkin Monophagous ASIA: Britain) (New Guinea New Papua Paradacus ( genus level by Virgilio et al. (2015). Now known as known al. (2015). Now et Virgilio by genus level No known male attractant. Responds to Ceratrap protein traps protein Ceratrap to Responds male attractant. known No Zeugodacus fly larvae ( fly Zeugodacus decipiens

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Zeugodacus depressus (Shiraki)

red-brown scutum, dark bands either side of medial vitta costal band slightly medial vitta overlapping R2+3 expanding into lateral vittae apical spot begin anterior to mesonotal suture

pale tint around medial broad anal posterior area streak of wing margin terga III-V with a T, dark anterolateral corners of terga IV large species and V

2 mm Wing variation Lateral thorax

1 mm

2 mm Head Fore, mid, hind legs 118 EXOTIC 119 until LURE . For a full . For depressa HOST RANGE ) SIMILAR SPECIES depressus ) OTHER COMMENTS COMMENTS OTHER Zeugodacus No known male attractant. known No ( Zeugodacus ( except has pale tint around medial around has pale tint except tau

species see Doorenweerd et al. (2018). et species see Doorenweerd Bactrocera Zeugodacus was elevated to genus level by Virgilio et al. (2015). et Virgilio by genus level to was elevated Zeugodacus Zeugodacus include watermelon, cucumber, pumpkin and tomato. tomato. pumpkin and cucumber, include watermelon,

posterior area of wing margin, and all leg segments pale. leg segments and all wing margin, of area posterior Oligophagous. Recorded from nine hosts in two families nine hosts in two from Recorded Oligophagous. list of list of Previously known as known Previously including Cucurbitaceae and Solanaceae. Commercial hosts Commercial and Solanaceae. including Cucurbitaceae ASIA: Taiwan, Japan, Korea, China Korea, Japan, Taiwan, Similar to to Similar Now known as known Now Zeugodacus

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Zeugodacus tau (Walker)

lateral vittae begin anterior to mesonotal suture costal band

overlapping R2+3 and expanding medial vitta into apical spot

black scutum with large areas of red- broad anal brown centrally streak

terga III-V with a T pattern dark anterolateral dark apical corners of terga IV and V markings on femora

2 mm

Leg variation Fore, mid, hind legs

1 mm

1 mm

Head Scutum Abdominal Lateral thorax variation variation 120 121

EXOTIC LURE Cue lure. Cue complex species see Zeugodacus HOST RANGE Z. tau until until SIMILAR SPECIES OTHER COMMENTS COMMENTS OTHER tau ) bitter melon and guava. melon bitter Zeugodacus Doorenweerd et al. (2018). et Doorenweerd Luffa, Zeugodacus except is smaller, has dark tibia and is smaller, except ( contains several other similar species. similar species. other several contains . For a full list of full list of a . For tau ) Bactrocera Z. depressus

femora with dark markings apically. The The apically. with dark markings femora Zeugodacus cucumber, pumpkin, edible pumpkin, cucumber, Similar to to Similar ( ASIA: India, Sri Lanka, Bangladesh, Bhutan, Vietnam, southern Vietnam, Bhutan, Bangladesh, Lanka, Sri India, Philippines, Laos, Cambodia, Thailand, China, Taiwan, Brunei, East Malaysia, Singapore, Malaysia, Peninsular Indonesia Polyphagous, but primarily a pest of cucurbits. Recorded on 34 cucurbits. Recorded a pest of but primarily Polyphagous, Commercial hosts include wax gourd, watermelon, rockmelon, rockmelon, watermelon, hosts include wax gourd, Commercial hosts from nine families including Curcurbitaceae and Myrtaceae. and Myrtaceae. including Curcurbitaceae nine families hosts from was elevated to genus level by Virgilio et al. (2015). Now known as known al. (2015). Now et Virgilio by genus level to was elevated Previously known as known Previously

Zeugodacus SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Dacus aequalis (Coquillet)

yellow to brown large wasp-like postpronotal lobes narrow yellow triangle anterior to species mesonotal suture

lateral and medial postsutural vittae absent

costal band scutellum with confluent with M broad basal band broad anal or all brown streak

abdomen mostly red brown with a paler oval spot on centre of tergum V

2 mm Abdominal variations Scutum Lateral thorax variation 1 mm large elongate facial spots

all femora and tibiae dark 2 mm Head Fore, mid, hind legs 122 PRESENT 123 LURE Cue lure. Cue but differs in but differs in but differs HOST RANGE SIMILAR SPECIES (Asclepiadaceae). D. absonifacies D. solomenensis and Marsdenia rostrata and M). Similar to to and M). Similar D. longicornis 4+5 Monophagous on Monophagous Similar to to Similar between R between OCEANIA: Australian Wales, South New Queensland, Australia: Territory Capital having the hind femora all red-brown (instead of pale basally). pale basally). of (instead all red-brown the hind femora having having the costal band equal to the M vein (instead of midway of (instead the M vein band equal to the costal having

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Dacus axanus (Hering)

postpronotal large wasp-like lobes yellow narrow yellow species to brown triangle anterior to mesonotal suture

lateral and medial costal band

postsutural vittae between R4+5 absent and M

scutellum with indistinct broad basal band anal streak or all brown

abdomen mostly red brown with a paler oval spot on centre of tergum IV and V

2 mm

Lateral thorax

irregularly round facial spots

all femora and tibiae dark 1 mm 2 mm Head Fore, mid, hind legs 124 PRESENT 125

LURE Cue lure. Cue but clearly [guada bean]). HOST RANGE SIMILAR SPECIES Trichosanthes anguina but differs in having dark in having but differs D. longicornis, and , but differs in having an anal in having , but differs differentiated using COI barcodes. using differentiated having a less elongated abdomen. a less elongated having D. solomonensis streak, not having mid femora pale basally. pale basally. mid femora having not streak, D. longicornis Morphologically similar to similar to Morphologically Luffa acutangulata, L. cylindrica postpronal lobes, not having a protrusion on tergum V, and V, on tergum a protrusion having not lobes, postpronal ( Oligophagous. Recorded on three hosts from Cucurbitaceae Cucurbitaceae hosts from on three Recorded Oligophagous. OCEANIA: south toQueensland and Islands Strait Torres Australia: (Moluccas Guinea; Indonesia New Papua Gladstone; and Timor) Similar to the exotic the exotic to Similar Similar to the exotic to Similar

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Dacus longicornis (Wiedemann)

postpronotal lobes large wasp-like yellow to brown a narrow yellow species triangle anterior to mesonotal suture costal band lateral and medial between R4+5 postsutural yellow and M vittae absent

scutellum with anal streak very pale, broad basal band appears absent or all brown

abdomen strongly petiolate, pale centrally on terga IV and V

2 mm

Lateral thorax very small irregularly round facial spots

all femora and tibiae dark, except mid femora pale 1 mm on basal 1/4 2 mm Head Fore, mid, hind legs 126 EXOTIC 127

LURE Cue lure. Cue L. aegyptica , HOST RANGE SIMILAR SPECIES , but differs in having no in having , but differs but differs in not having having in not a but differs Trichosanthes cucumerina. the mid femora pale basally. pale basally. the mid femora Luffa acutangula and D. axanus D. solomonensis and narrower triangles anterior to the mesonotal suture. suture. the mesonotal to triangles anterior and narrower Oligophagous. Recorded on four hosts in Cucurbitaceae, hosts in Cucurbitaceae, on four Recorded Oligophagous. Similar to the Australian the Australian to Similar including the Asian vegetables including the Asian vegetables ASIA: Thailand, Myanmar, Laos, Brunei, Malaysia, Indonesia, Bangladesh Vietnam, Bhutan, India, China, Philippines, discernible anal streak, usually smaller facial spots and having spots smaller facial usually anal streak, discernible protrusion on tergum V and having a less elongate abdomen, elongate a less V and having on tergum protrusion Similar to the exotic the exotic to Similar

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Dacus solomonensis (Malloch)

postpronotal large wasp-like lobes yellow yellow triangle species to brown anterior to mesonotal suture costal band lateral and medial between R4+5 postsutural vittae and M absent

scutellum usually anal streak yellow, sometimes indistinct brown

abdomen very elongate with round facial spots, protrusion on can join centrally tergum V as a mask

2 mm 1 mm Abdominal variation Lateral thorax Head

2 mm Scutum variation Fore, mid, hind legs Abdomen lateral 128 EXOTIC 129 . LURE Luffa Cue lure. Cue HOST RANGE SIMILAR SPECIES except has a protrusion has a protrusion except anterior to the mesonotal suture. suture. the mesonotal to anterior D. axanus and including pumpkin, cucumber and edible D. longicornis Oligophagous. Recorded from five hosts in Cucurbitaceae, hosts in five from Recorded Oligophagous. on tergum V, a longer oval abdomen, and broader triangles abdomen, and broader oval a longer V, on tergum OCEANIA: Islands Solomon and (Bougainville) Guinea New Papua Similar to to Similar

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Ceratitis capitata (Wiedemann) MEDITERRANEAN FRUIT FLY

scutum yellowish with small to medium-sized numerous black areas in brightly coloured species a characteristic pattern

scutellum swollen, rounded above, shiny black with a thin sinuate yellow streak near base dorsally

wing relatively broad in small dark irregular- comparison with its length, shaped streaks cloudy yellow, with three brown within the cells in the bands on apical two-thirds, all proximal half of wing separated from each other

1 mm 2 mm

Head Lateral thorax

1 mm 2 mm

Wing Abdomen Fore, mid, hind legs 130 PRESENT 131 LURE HOST RANGE SIMILAR SPECIES Sapotaceae, Solanaceae and Vitaceae. and Solanaceae Sapotaceae, Easily separated from other members of the family. the family. of members other from separated Easily which appears to be more powerful than trimedlure. powerful be more to which appears Widespread Rosaceae, Rubiaceae, Rutaceae, Santalaceae, Sapindaceae, Santalaceae, Rutaceae, Rubiaceae, Rosaceae, Apocynaceae, Arecaceae, Cactaceae, Caricaceae, Clusiaceae, Caricaceae, Cactaceae, Arecaceae, Apocynaceae, AFRICA: Europe EUROPE: Southern Saudi Lebanon, Jordan, Israel, Iraq, Iran, MIDDLE EAST: Yemen Turkey, Syria, Arabia, Australia and Western Hawaii OCEANIA: in or restricted widespread THE AMERICAS: Variably Guatemala, Salvador, El Rica, (Costa America Central America and South Panama) Nicaragua, Honduras, Ecuador, Chile, Colombia, Bolivia, Brazil, (Argentina, Venezuela) Uruguay, Peru, Paraguay, Polyphagous including the families Anacardiaceae, Annonaceae, Anacardiaceae, including the families Polyphagous Combretaceae, Ebenaceae, Juglandaceae, Lauraceae, Lythraceae, Lythraceae, Lauraceae, Juglandaceae, Ebenaceae, Combretaceae, Trimedlure/capilure and terpinyl acetate. Also attracted to EGO lure lure EGO to Also attracted acetate. and terpinyl Trimedlure/capilure Malpighiaceae, Malvaceae, Muntingiceae, Myrtaceae, Passifloraceae, Passifloraceae, Myrtaceae, Muntingiceae, Malvaceae, Malpighiaceae,

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Ceratitis quilicii (De Meyer, Mwatawala & Virgilio) CAPE FRUIT FLY

scutum ground colour greyish to greyish-brown, sometimes with orange tinge

scutellum yellowish- white, apically with three separate black spots

tergites II and IV on posterior half with greyish microtrichosity; abdomen anterior margin sometimes ground colour narrowly brownish coloured, mainly yellow especially laterally

black coloration on tibia not reaching the ventral and dorsal margins throughout the full length

slender tibia, gradually tapering towards the base

2 mm 1 mm Fore, mid, hind legs Head

132 133

EXOTIC LURE HOST RANGE SIMILAR SPECIES in arrangement of in arrangement these are inseparable these are as a separate as a separate C. rosa powerful than trimedlure. powerful Rubiaceae and Sapotaceae. Rubiaceae C. rosa: C. quilicii setae on the male mid-tibial comb. comb. on the male mid-tibial setae , these distributions may require require may , these distributions Polyphagous including Myrtaceae, Rosaceae, Rosaceae, including Myrtaceae, Polyphagous C. rosa Morphologically similar to similar to Morphologically using COI barcodes. Differs from from Differs barcodes. using COI AFRICA: Swaziland, Africa, South Malawi, Kenya, Botswana, Island Réunion Zimbabwe, Tanzania, of distinction the recent Due to further resolution as more knowledge is gained of each of is gained knowledge as more further resolution range geographic species’ true species from species from Capilure. Also attracted to EGO lure which appears to be more be more to which appears lure EGO to Also attracted Capilure.

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Ceratitis rosa (Karsch) NATAL FRUIT FLY

scutellum marked black and yellow, with yellow lines or areas meeting margin

male mid-femora without stout ventral setae broader mid-tibia with black coloration reaching the ventral and dorsal margins of the tibia throughout

2 mm Fore, mid, hind legs

1 mm 2 mm

Head Lateral thorax 134 EXOTIC 135 LURE but this C. quilicii. C. capitata. HOST RANGE and from from C. quilicii, SIMILAR SPECIES as a separate as a separate and C. rosa C. capitata C. quilicii C. rosa more powerful than trimedlure. powerful more Similar to Similar , these distributions may require require may , these distributions C. rosa marker does distinguish marker Polyphagous including the families Anacardiaceae, Anacardiaceae, including the families Polyphagous Annonaceae, Apocynaceae, Caricaceae, Clusiaceae, Caricaceae, Apocynaceae, Annonaceae, Sapindaceae, Sapotaceae, Solanaceae, and Vitaceae. Solanaceae, Sapotaceae, Sapindaceae, Trimedlure. Also attracted to EGO lure which appears to be to which appears lure EGO to Also attracted Trimedlure. Oxalidaceae, Rhamnaceae, Rosaceae, Rubiaceae, Rutaceae, Rubiaceae, Rosaceae, Rhamnaceae, Oxalidaceae, COI barcodes do not separate separate do not barcodes COI AFRICA: Tanzania Africa, South Mozambique, Malawi, Kenya, of distinction the recent Due to further resolution as more knowledge is gained of each of is gained knowledge as more further resolution range geographic species’ true species from species from Combretaceae, Lauraceae, Malvaceae, Moraceae, Myrtaceae, Myrtaceae, Moraceae, Malvaceae, Lauraceae, Combretaceae,

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Dirioxa pornia (Walker) ISLAND FLY

thorax and scutellum golden

wing six scutellar setae pattern as per figure*

terga IV and V black, tergum III black laterally *note there are other trypetines with patterned wings and the pattern needs to closely match the figure

1 mm Abdominal variation Lateral thorax

1 mm 2 mm Head Fore, mid, hind legs 136 PRESENT 137 LURE Anastrepha HOST RANGE SIMILAR SPECIES detected at cue lure. at detected different wing patterns. wing patterns. different (Dacinae: Ceratitidini) have very have Ceratitidini) (Dacinae: Ceratitis and Annonaceae, Caricaceae, Curcurbitaceae, Lauraceae, Curcurbitaceae, Caricaceae, Annonaceae, Polyphagous, but on overripe, damaged or fallen fruit. damaged or fallen but on overripe, Polyphagous, carambola, peach, pear, citrus, capsicum and eggplant. citrus, capsicum peach, pear, carambola, Rhagoletis) wings, but has a distinct wing pattern and is the only one and is the only wings, but has a distinct wing pattern associated with edible fruit. Exotic ( Trypetinae with edible fruit. Exotic associated Passifloraceae, Rosaceae, Rubiaceae, Rutaceae, Sapotaceae Rutaceae, Rubiaceae, Rosaceae, Passifloraceae, and Solanaceae. Commercial hosts include mango, papaya, hosts include mango, Commercial and Solanaceae. and OCEANIA: York Cape Range, Iron from Australia Eastern Australia: to Introduced South Wales. New southern to Peninsula, Australia Western Perth, Attracted to protein lures and orange ammonia. Occasionally and orange lures protein to Attracted Recorded on 83 hosts in 27 families including Anacardiaceae, including Anacardiaceae, on 83 hosts in 27 families Recorded Lecythidaceae, Moraceae, Musaceae, Myrtaceae, Oxalidaceae, Oxalidaceae, Myrtaceae, Musaceae, Moraceae, Lecythidaceae, Similar to some other Australian Trypetinae in having patterned patterned in having Trypetinae Australian some other to Similar

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Anastrepha fraterculus (Wiedemann) SOUTH AMERICAN FRUIT FLY

scutum posteriorly V-band usually without brown not connected mesonotum markings or with anteriorly to with three pale only a single medial S-band; may postsutural brown spot on scuto- be connected vittae scutellar suture in some cryptic species

abdominal tergites V-band distal arm without brown complete markings

USDA 2 mm

Aculeus and aculeus tip, ventral. Image: USDA Lateral thorax

1 mm

frons without brown markings

2 mm Head Fore, mid, hind legs 138 EXOTIC 139 LURE revision. HOST RANGE Further, it is now Further, SIMILAR SPECIES A. distincta. ammonium acetate + putrecine. ammonium acetate and is difficult to diagnose and is very similar very similar to diagnose and is to is difficult Rubiaceae, Rutaceae, Sapotaceae and Vitaceae. Sapotaceae Rutaceae, Rubiaceae, Polyphagous including the families Actinidiaceae, including the families Polyphagous A. fraterculus Moraceae, Myrtaceae, Oleaceae, Oxalidaceae, Rosaceae, Rosaceae, Oxalidaceae, Oleaceae, Myrtaceae, Moraceae, A. obliqua, A. suspensa, Anacardiaceae, Annonaceae, Combretaceae, Ebenaceae, Combretaceae, Annonaceae, Anacardiaceae, Fabaceae, Juglandaceae, Lauraceae, Lythraceae, Malvaceae, Lythraceae, Lauraceae, Juglandaceae, Fabaceae, with decaying protein (or similar), or with multilure traps withtraps similar), or with multilure (or protein with decaying composing multiple cryptic species that await full taxonomic await species that cryptic multiple composing THE AMERICAS: and Trinidad Argentina, northern to Mexico Northern Island Galapagos Tobago, No known record, but can can be captured using McPhail traps using McPhail but can can be captured record, known No recognised through using integrative taxonomic approaches as approaches taxonomic using integrative through recognised

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Anastrepha serpentina (Wiedemann) SAPOTE FRUIT FLY; BLACK FRUIT FLY

scutum with very dark wing S-band slender mostly dark markings contrast and not brown base strongly with the connected to the colour strongly light hyaline areas V-band contrasting with of the wing lighter stripes

apical arm of V-band absent abdominal tergites mostly brown with a somewhat ‘T-shaped’ medial yellow or white area subscutellum and mediotergite are very dark, with a lighter mesopleuron brownish or yellowish partially brown spot or stripe dorsally

1 mm

frons without brown markings except ocellar tubercle

2 mm Head Lateral thorax

140 EXOTIC 141

LURE HOST RANGE SIMILAR SPECIES using ammonia traps. any other major pest species. other any Myrtaceae, Rosaceae and Rutaceae. Rosaceae Myrtaceae, No known record, but can be captured but can be captured record, known No from Anacardiaceae, Annonaceae, Clusiaceae, Lauraceae, Clusiaceae, Annonaceae, Anacardiaceae, from THE AMERICAS: Honduras, Guatemala, Rica, Costa Belize, Mexico, and Tobago, Trinidad Panama, Antilles, Netherlands Guyana, Guiana, French Ecuador, Colombia, Brazil, Bolivia, Argentina, northern Venezuela, Suriname, Peru, Nicaragua Paraguay, This species is very distinctive and unlikely to be confused with be confused to and unlikely distinctive species is very This Polyphagous, predominantly on Sapotaceae, but also recorded but also recorded on Sapotaceae, predominantly Polyphagous,

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Rhagoletis cerasi (Linnaeus) EUROPEAN CHERRY FRUIT FLY

short narrow short broad transverse fuscous transverse fuscous band from anterior band towards base margin to vein R4+5 of wing

inverted L-shaped band at apex of wing, stem encloses dm-cu crossvein

broad transverse fuscous band across centre from anterior margin to hind margin and enclosing r-m crossvein

yellow scutellum

1 mm 1 mm 2 mm Scutum Head Lateral thorax

142 EXOTIC 143

LURE HOST RANGE and Rosaceae. unknown status. unknown SIMILAR SPECIES No known record. known No , some of which are of which are , some of Rhagoletis

is a recent invasive (2015) to North (2015) to invasive is a recent Polyphagous including the families Caprifoliaceae Caprifoliaceae including the families Polyphagous

Similar to other species of species of other to Similar hagoletis cerasi R Native to Europe/Asia; introduced to North America North to introduced Europe/Asia; to Native ASIA: Georgia, Iran, Kazakhstan, Kyrgyzstan, Tajikistan, Turkey, Turkey, Tajikistan, Kyrgyzstan, Kazakhstan, Iran, Georgia, Uzbekistan Turkmenistan, EUROPE: Czech Croatia, Bulgaria, Britain, Belgium, Austria, Germany, France, Finland, Denmark, Estonia, Republic, Lithuania, Moldova, Latvia, Italy, Hungary, Greece, Romania, Portugal, Poland, Norway, Netherlands, Switzerland, Sweden, Spain, Serbia, Slovakia, Russia, Ukraine THE AMERICAS: America (southern Ontario, Canada and New York York and New Canada America (southern Ontario, USA) State,

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Rhagoletis cingulata (Loew) CHERRY FRUIT FLY

dark fuscous spot

inverted ‘U’- across apex of R4+5; shaped dark may be connected fuscous pattern or disconnected to across apex main band

accessory pale transverse band absent bands on abdomen

a broad dark fuscous band from anterior margin to hind margin in centre of wing

1 mm

Wing

1 mm

1 mm

Head Lateral thorax 144 EXOTIC 145 ). LURE P. cerasus HOST RANGE SIMILAR SPECIES No known species. known No

. COI does not effectively effectively does not . COI does (Johannesen et al. 2013). et does (Johannesen , including sour cherries ( Prunus

Rhagoletis indifferens genus

Similar to to Similar Oligophagous, attacking a small number of host species in a small number of attacking Oligophagous, Native to North America. Distribution in the USA is in the Distribution America. North to Native several from species is absent but this widespread, true represent does not as presented the map states; USA. within the distributions geographic THE AMERICAS: northern Quebec), (Ontario, Canada America: North USA eastern Mexico, and central EUROPE: Austria, Belgium, Britain, Croatia, Czech Republic, Czech Croatia, Belgium, Britain, Austria, Poland, Netherlands, Italy, Hungary, Germany, France, Switzerland Slovenia, distinguish these taxa, but a microsatellite panel is available that panel is available distinguish these taxa, but a microsatellite

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Rhagoletis completa Cresson WALNUT HUSK FLY

thorax orange/ pale brown; abdomen partly four transverse wing dark brown bands present and connected

scutellum concolorous yellow

Lateral variation

1 mm 2 mm

Dorsal variation Head Lateral thorax 146 EXOTIC 147

LURE Juglans R. zoqui R. completa , and e.g., peaches). HOST RANGE SIMILAR SPECIES emitting ammonia. from other species. other from R. ramosae , COI separates separates COI R. completa R. suavis. R. completa and distinguish

, but not , but not No known record, but can be captured in traps but can be captured record, known No

R. berberis (Frey et al. 2013). Morphological characters can reliably reliably can characters al. 2013). Morphological et (Frey R. suavis and Established in southern Europe Established THE AMERICAS: Mexico. northern including USA and Central Southern this species is but the USA is widespread, in Distribution does not map as presented the states; several from absent within the USA distributions true geographic represent EUROPE: Similar to to Similar Polyphagous, including the families of Juglandaceae ( Juglandaceae of including the families Polyphagous, spp. are predominant hosts) and Rosaceae (sub-optimal hosts, (sub-optimal hosts) and Rosaceae predominant spp. are

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Rhagoletis fausta (Osten-Sacken) BLACK CHERRY FRUIT FLY

scutellum pale thorax and abdomen predominantly very broad black connection between the discal and subapical bands in cell dm with the presence of both an anterior and posterior apical band

posterior apical band arises from the subapical band in the vicinity of vein r-m, making an F-shaped pattern in the apical half of the wing

1 mm 2 mm Head Lateral thorax 148 EXOTIC 149

, LURE species. R. striatella HOST RANGE and SIMILAR SPECIES emitting ammonia. Rhagoletis suavis

No known record, but can be captured in traps but can be captured record, known No commercial hosts are sweet cherry and sour cherry). cherry and sour sweet hosts are commercial Oligophagous - affecting the Rosaceae family (majorfamily Rosaceae the - affecting Oligophagous but molecular markers (eg., 16S) can separate these taxa. 16S) can separate (eg., but molecular markers Morphologically similar to similar to Morphologically Morphological characters can be used to reliably identify this identify reliably can be used to characters Morphological North America – western and eastern North America. North and eastern – western America North but either is widespread, the USA and Canada in Distribution states several from, or is absent distributions, has restricted true represent does not map below The and provinces. USA and Canada within the distributions geographic THE AMERICAS:

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Rhagoletis pomonella (Walsh) APPLE MAGGOT FLY

wing pattern consists of a slightly oblique discal band to which the anterior overall black and posterior apical bands body colour are connected, forming a characteristic F-shaped pattern in the apical half of the wing

absence of the subapical band

1 mm

presence, in most specimens, of heavy black shading on the posterior surface of the fore femur

Lateral thorax

2 mm 1 mm Fore, mid, hind legs Head 150 EXOTIC 151 LURE complex. HOST RANGE SIMILAR SPECIES OTHER COMMENTS COMMENTS OTHER R. pomonella spp. are the main hosts. spp. are Crataegus COI does not separate these species. separate does not COI species of the species of It is a major pest of cultivated apples. It is apples. It cultivated is a major pest of It

R. mendax. family: apple and family: Rhagoletis. Difficult to distinguish from several other non-pest cryptic from several to distinguish Difficult Similar to to Similar Oligophagous – affects numerous species of the Rosaceae of the species numerous – affects Oligophagous THE AMERICAS: Canada, USA and Mexico. Distribution in the USA is wide- in the USA Distribution Mexico. USA and Canada, the states; several from is absent but this species spread, geographic true represent not does map as presented within the USA distributions No known record, but can be captured in traps emitting ammonia. in traps but can be captured record, known No It has been the subject of extensive taxonomic, ecological and pest and ecological taxonomic, extensive has been the subject of It

of the USA and is now widespread in that region (pers. comm. Drew 2010). Drew comm. (pers. region in that widespread the USA and is now of distributed over the central and north-eastern regions of mainland USA and of regions and north-eastern the central over distributed species within the genus SIMILAR SPECIES, SPECIES, management research in the USA and is considered the major economic pest the major economic in the USA and is considered research management extreme southern Canada. In 1979 it was introduced into the western coastline the western into 1979 it was introduced In southern Canada. extreme HOST & LURE & HOST DISTRIBUTION Toxotrypana curvicauda Gerstaecker PAPAYA FRUIT FLY

predominantly yellow marked with black

male fly resembles female, but without the ovipositor (body length: 11–13.5 mm; wing width: 8.5–11 mm)

very long, slender abdomen; females with a greatly elongated ovipositor

2 mm Female dorsal view Lateral thorax

1 mm 2 mm

Head Fore, mid, hind legs 152 153

EXOTIC ). LURE HOST RANGE SIMILAR SPECIES No known record. known No Zeugodacus, Dacus , online keys are available. are online keys Bactrocera may initially be confused with other be confused initially may (e.g. (e.g. Toxotrypana;

and are readily distinguished from other fruit fly genera fruit fly other distinguished from readily and are Anacardiaceae (mango), and Apocynaceae (milkweed). and Apocynaceae (mango), Anacardiaceae members of genus of members All members of this genus are exotic to Australia, however, however, Australia, to exotic are this genus of All members Toxotrypana curvicauda THE AMERICAS: America. in North and Texas Florida to Restricted in particularly the Caribbean, throughout Distributed Cuba Trinidad, Republic, the Dominican Rico, Puerto Central in Mexico, is also found It and the Bahamas. Honduras, Guatemala, Rica, Costa America (Belize, America and northern South Panama) Nicaragua, Venezuela) (Columbia, Polyphagous including the families Caricaceae (papaya: primary host), (papaya: Caricaceae including the families Polyphagous

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Drosophila suzukii (Matsumura) SPOTTED-WING DROSOPHILA (SWD)

wing pattern as per figure

thorax and scutellum golden uniform apricot coloured thorax free of stripes, spots or patterns, terga IV and V with unbroken dorsal black, tergum bands on its abdomen III black laterally

a single row of apical dark combs running across (rather than parallel with) each six scutellar of the basal two tarsal setae Lateral thorax - female. Image: Mark Blacket segments

0.5 mm

0.5 mm Scutellum Head Male comb. Image: Mark Blacket

2 mm males have a single apical wing spot

strongly sclerotized ovipositor with 30–36 robust teeth; six–seven times longer than the diameter of the spermatheca Lateral thorax - male. Image: Mark Blacket Ovipositor. Image: Mark Blacket 154 EXOTIC 155 LURE HOST RANGE SIMILAR SPECIES

. Several molecular. Several (see Murphy et al. 2015). et (see Murphy D. simulans and

Rhamnaceae, Rosaceae, Rutaceae and Vitaceae. Rutaceae Rosaceae, Rhamnaceae,

D. melanogaster Caprifoliaceae, Cornaceae, Ebenaceae, Elaeagnaceae, Ebenaceae, Cornaceae, Caprifoliaceae, Adults are known to be attracted to traps baited with a baited traps to be attracted to known are Adults

combination of wine, vinegar and soap (Landolt et al. 2012). et wine, vinegar and soap (Landolt of combination diagnostic tools are available that can distinguish this species can that available are diagnostic tools ASIA: Bangladesh, China, India, Iran, Japan, Korea, Myanmar, Myanmar, Korea, Japan, Iran, India, China, Bangladesh, Thailand Taiwan, Pakistan, THE AMERICAS: Brazil Argentina, USA, Mexico, Canada, EUROPE: Republic, Czech Croatia, Belgium, Bosnia-Herzegovina, Portugal, Poland, Netherlands, Italy, Germany, France, Spain, Serbia, Slovenia, Federation, Russian Switzerland, UK Switzerland, Polyphagous including the families Actinidiaceae, Adoxaceae, Adoxaceae, Actinidiaceae, including the families Polyphagous Similar to to Similar Ericaceae, Grossulariaceae, Moraceae, Myricaceae, Myrtaceae, Myrtaceae, Myricaceae, Moraceae, Grossulariaceae, Ericaceae,

SIMILAR SPECIES, SPECIES, HOST & LURE & HOST DISTRIBUTION Reference list Allwood, A.J., Chinajariyawong, A., Drew, R.A.I., Hamacek, E.L. and Hancock, D.L. (1999). Host plant records for fruit flies (Diptera: Tephritidae) in South East Asia.Raffles Bulletin of Zoology Supplement 7: 1-92. Casana-Giner V., Oliver J.E., Jang E.B. & Carvalho L.A. (2003). Syntheses and behavioral evaluations of fluorinated and silylated analogs of raspberry ketone as attractants for melon fly, Bactrocera cucurbitae (Coquillett).Journal of Entomological Science 38: 111–119. Dominiak, B.C., Campbell, A.J., Jang, E.B., Ramsey, A. and Fanson, B.G. (2015). Field evaluation of melolure, a formate analogue of cuelure, and reassessment of fruit fly species trapped in Sydney, New South Wales, Australia. Journal of Economic Entomology 108: 1176-1181. Doorenweerd, C., Leblanc, L., Norrbom, A. L., Jose, M. S., & Rubinoff, D. (2018). A global checklist of the 932 fruit fly species in the tribe Dacini (Diptera, Tephritidae).ZooKeys 730: 17–54. Drew, R. A., and Romig, M. C. (2013). Tropical Fruit Flies (Tephritidae Dacinae) of South- East Asia: Indomalaya to North-West Australasia. CABI. Drew, R. A. (1989). The tropical fruit flies (Diptera: Tephritidae: Dacinae) of the Australasian and Oceanian regions. Memoirs of the Queensland Museum 26. Fay, H.A.C. (2012). A highly effective and selective male lure for Bactrocera jarvisi (Tryon) (Diptera: Tephritidae). Australian Journal of Entomology 51: 189–197. Floyd, R., Lima, J., deWaard, J., Humble, L., and Hanner, R. (2010). Common goals: policy implications of DNA barcoding as a protocol for identification of pests. Biological Invasions 12: 2947-2954. Frey, J.E., Guillén, L., Frey, B., Samietz, J., Rull, J. and Aluja, M., (2013). Developing diagnostic SNP panels for the identification of true fruit flies (Diptera: Tephritidae) within the limits of COI-based species delimitation. BMC Evolutionary Biology, 13(1), p.106. Gilbert, M.T.P., Moore, W., Melchior, L., and Worobey, M. (2007). DNA extraction from dry museum beetles without conferring external morphological damage. PLoS ONE 2: e272. doi:10.1371/journal.pone.0000272 Hancock, D. L. and Drew, R. A. I., (2015). A review of the Indo-Australian subgenus’ Parazeugodacus’ Shiraki of ‘Bactrocera’ Macquart (Diptera: Tephritidae: Dacinae). The Australian Entomologist, 42(2): 91. Johannesen, J., Keyghobadi, N., Schuler, H., Stauffer, C., & Vogt, H. (2013). Invasion genetics of American cherry fruit fly in Europe and signals of hybridization with the European cherry fruit fly.Entomologia Experimentalis et Applicata, 147(1): 61-72. Landolt P. J., Adams T., Davis T. S., & Rogg H. (2012). Spotted Wing Drosophila, Drosophila suzukii (Diptera: Drosophilidae), Trapped with Combinations of Wines and Vinegars. Florida Entomologist 95: 326-332.

156 Leblanc L., Vueti E. T., Drew R. A. I. & Allwood A. J. (2012). Host plant records for fruit flies (Diptera: Tephritidae: Dacini) in the Pacific islands. Proceedings of the Hawaiian Society 44: 11-53. McQuate G. T. & Peck S. L. (2001). Enhancement of attraction of alpha ionol to male Bactrocera latifrons (Diptera: Tephritidae) by addition of a synergist cade oil. Journal of Economic Entomology 94: 39–46. Murphy K. A., Unruh T. R., Zhou L. M., Zalom F. G., Shearer P. W., Beers E. H., Walton V. M., Miller B., & Chiu J. C. (2015). Using comparative genomics to develop a molecular diagnostic for the identification of an emerging pest Drosophila suzukii.Bulletin of Entomological Research 105, 364-372. Mziray, H. A., Makundi, R. H., Mwatawala, M., Maerere, A., & De Meyer, M. (2010). Spatial and temporal abundance of the solanum fruit fly,Bactrocera latifrons (Hendel), in Morogoro, Tanzania. Crop Protection, 29(5), 454-461. Royer, J. E., Agovaua, S. Bokosou, J. Kurika, K., Mararuai, A., Mayer, D. G. and Niangu, B. (2018). Responses of fruit flies to new attractants in Papua New Guinea.Austral Entomology 57: 40-49. Royer J. E., DeFaveri S. & Lowe G. (2014). Cucumber volatile blend, a promising female- biased lure for Bactrocera cucumis (French 1907) (Diptera: Tephritidae: Dacinae), a pest fruit fly that does not respond to male attractants.Australian Journal of Entomology 53: 347–352 Royer, J. E., Mille, C., Cazares, S., Brinon, J. and Mayer, D. G. (2018). Isoeugenol, a more attractive male lure for the cuelure-responsive pest fruit fly Bactrocera curvipennis (Froggatt) (Tephritidae: Dacinae), and new records of species responding to zingerone in New Caledonia. In prep. Royer, J. E., Teakle, G. and Mayer, D. G. (2018). Methyl-isoeugenol, a significantly more attractive male lure for the methyl eugenol-responsive Pacific fruit fly,Bactrocera xanthodes (Broun) (Diptera: Tephritidae: Dacinae). In prep. Royer, J. E. (2015). Responses of fruit flies (Tephritidae: Dacinae) to novel male attractants in north Queensland Australia, and improved lures for some pest species. Austral Entomology 54: 411-426. Tsuruta, K., White, I.M., Bandara, H.M.J., Rajapakse, H., Sundaraperuma, S.A.H., Kahawatta, S.B.M.U.C., and Rajapakse, G.B.J.P. (1997). A preliminary notes on the host plants of fruit flies of the tribe Dacini (Diptera: Tephritidae) in Sri Lanka.Esakia 37: 149- 160. Virgilio, M., Jordaens, K., Verwimp, C., White, I. M., & De Meyer, M. (2015). Higher phylogeny of frugivorous flies (Diptera, Tephritidae, Dacini): Localised partition conflicts and a novel generic classification. Molecular phylogenetics and 85: 171-179. Wee, S. L., Chinvinijkul, S., Tan, K. H. and Nishida, R. (2018). A new and highly effective male lure for the guava fruit fly Bactrocera correcta. Journal of Pest Science 2: 691-698. White, I. M. and Elson-Harris, M. M. (1992). Fruit Flies of Economic Significance: Their Identification and Bionomics. CAB International. Oxon, UK. 601 p. 157 Notes

158

Species: Bactrocera facialis Species: Bactrocera aquilonis Species: Bactrocera cacuminata Species: Bactrocera frauenfeldi

Species: Bactrocera musae Species: Bactrocera tryoni Species: Ceratitis capitata Species: Dacus aequalis

Species: Zeugodacus choristus Species: Bactrocera opiliae Species: Bactrocera rufofuscula Species: Bactrocera pallida

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