A review of subgenus Bactrocera (Bactrocera) Macquart, 1835 (Diptera: Tephritidae: Dacinae)

73 RAFFLES BULLETIN OF ZOOLOGY 2025 A review of subgenus Bactrocera (Bactrocera) Macquart, 1835 (Diptera: Tephritidae: Dacinae) D. L. Hancock1* & R. A. I. Drew2 Abstract. The 439 described species in subgenus Bactrocera Macquart, 1835 of genus Bactrocera are referred to 90 species groups in 18 morphologically defined complexes. Semicallantra Drew, 1989 is placed as a new synonym of subgenus Bactrocera. Bactrocera (Bactrocera) wallacei Drew & Hancock, new species, is described from Maluku, Indonesia. Bactrocera clarkei Starkie, Strutt & Royer, 2022 and B. petila Drew, 1989 are transferred from subgenus Bactrocera to subgenera Neozeugodacus May, 1952a and Calodacus Hancock, 2015 respectively. Bactrocera incognita Doorenweerd & San Jose, 2024 and B. borneoensis Doorenweerd & San Jose, 2024 are regarded as new synonyms of B. occipitalis (Bezzi, 1919) and B. carambolae Drew & Hancock, 1994 respectively. Bactrocera irvingiae Drew & Hancock, 1994 is newly reported from Taiwan based on a previous misidentification as ‘Dacus parvulus’. Discussions are included on the history of research, biogeography, male lures, host plants and molecular evidence. Key words. Bactrocera, taxonomy, biogeography, morphology RAFFLES BULLETIN OF ZOOLOGY 73: 73–109 Date of publication: 28 February 2025 DOI: 10.26107/RBZ-2025-0007 http://zoobank.org/urn:lsid:zoobank.org:pub:FC07EE6B-DA0B-4FBC-8134-5E42B9ADC254 © National University of Singapore ISSN 2345-7600 (electronic) | ISSN 0217-2445 (print) INTRODUCTION The genus Bactrocera Macquart, 1835 is the most speciose in the family Tephritidae, with more than 700 described species (including those often included in its synonym Zeugodacus Hendel, 1927a) endemic to the Afrotropical, Oriental, Australian, and Oceanian Regions. It includes some of the world’s most damaging horticultural pests, several of which have become invasive beyond their normal distributions. Subgenus Bactrocera is the largest, with 439 described species endemic to the Oriental, Australian, and Oceanian Regions, three of which (B. invadens Drew, Tsuruta & White, 2005, B. latifrons (Hendel, 1915) and B. zonata (Saunders, 1842)) are invasive in Africa, with the first two also recorded from Italy but B. latifrons possibly not established (Gargiulo et al., 2021), two (B. dorsalis (Hendel, 1912) and B. latifrons) in Hawaii and one (B. carambolae Drew & Hancock, 1994) in NE South America. Comprehensive monographs covering the Bactrocera fauna of Southeast Asia, Australia, and the Pacific were published by Hardy (1973, 1974), Drew (1989), and Drew & Romig (2013, 2016, 2022), the latter publications resulting from extensive field collections from 1980 to 2010 across Southeast Asia and the Pacific and markedly increasing knowledge of the fauna. Seven subgenera were included in the Bactrocera group of subgenera by Hancock & Drew (2018), defined by the presence of a short posterior surstylus lobe and deeply concave (emarginate) abdominal sternite V in males (Drew, 1989). This is reduced here to six subgenera by the synonymy of Semicallantra Drew, 1989 with typical Bactrocera. Other included subgenera are Apodacus Perkins, 1939, defined by the presence of a yellow postsutural medial vitta on the scutum and narrow, transverse ceromata (shining spots) on abdominal tergite V in both sexes, Bulladacus Drew & Hancock, 1995, defined by the lack of ceromata and frequent presence of a ‘bulla’ on the male wing, Calodacus Hancock, 2015, defined by the lack of the pecten of cilia on abdominal tergite III in males, the monotypic Trypetidacus Drew, 1989 and Queenslandacus Drew, 1989, also defined by the lack of the pecten of cilia on abdominal tergite III in males, and typical Bactrocera, defined by the lack of a medial yellow vitta, presence of oval or rounded ceromata and presence of the abdominal pecten in males. [Note that figures of female B. (B.) daruensis Drew, 1989 and B. (B.) thistletoni Drew, 1989 (figs. 66 & 70) in Drew & Romig (2022) include the abdominal pecten in error]. Calodacus, Trypetidacus and Queenslandacus are possibly synonyms, the presence of B. (Trypetidacus) invisitata Drew, 1989 in methyl eugenol-baited traps, not repeated since its original description (Drew, 1989) and possibly accidental, mirrors a situation noted for B. (Tetradacus) minax (Enderlein, 1920) in Bhutan by Hancock & Drew (2018). Reviews of Calodacus, Bulladacus, and Apodacus were provided by Hancock (2015), Drew & Hancock (2016), and Hancock Taxonomy & Systematics Accepted by: Ang Yuchen 114 Yew Tree Court, Carlisle, Cumbria CA2 5QA, United Kingdom; Email: [email protected] (*corresponding author); ORCID iD: https://orcid.org/00000002-8478-7976 2Griffith University Centre for Planetary Health and Food Security, Queensland 4111, Australia; Email: [email protected]; ORCID iD: https://orcid.org/00000003-4973-5107 74 Hancock & Drew: Review of subgenus Bactrocera & Drew (2018) respectively, with additional species added by David et al. (2016) and Drew & Romig (2022). The 439 species included in subgenus Bactrocera are referred here to 90 species groups in 18 morphological complexes. Complexes (sensu Drew & Romig, 2013) are regarded as a group of morphologically similar species which might not be monophyletic [the quadrata complex, for example, is likely to be polyphyletic], while species groups (some monotypic) are proposed as actual or potential monophyletic assemblages of related species. HISTORY OF TAXONOMIC RESEARCH WITH REFERENCE TO SUBGENUS BACTROCERA Taxonomic research into the family Tephritidae has a long and valuable history dating back to the 18th century. With the expansion of European colonies across the tropics and subtropics of Africa, Southeast Asia, and the Pacific, several key biological expeditions were undertaken in the 19th century. Some of these expeditions focused on the collection of insects that included tephritid fruit flies. These specimens, including the resulting types of new species, were usually deposited in the national museums of the countries that either financed or undertook the expeditions. Consequently, many of the early type specimens are located in the Natural History Museum, London (NHML) and museums across Europe. A comprehensive list of these museums and a detailed history of taxonomic research in Tribe Dacini was published by Drew (1989) and Drew & Romig (2013). Taxonomic studies on the genus Bactrocera Macquart began with the description of the olive fruit fly, B. oleae (Gmelin, 1790), then under the generic name Musca Linnaeus, 1758. This was followed by the description of B. ferruginea (Fabricius, 1794) from India, also under Musca and, due to homonomy, now known as B. invadens [or, incorrectly, treated as a synonym of B. dorsalis] (see Drew & Hancock, 2022). Additional species were described by Fabricius: B. umbrosa and B. caudata in 1805, both under genus Dacus Fabricius, 1805; Macquart described B. longicornis, the type species of genus Bactrocera, in 1835; Saunders described B. zonata in 1842 under genus Dasyneura Saunders, 1842; Doleschall described B. maculigera in 1858; and Walker, based at the NHML, described a large number of species between 1859 and 1865 under Dacus, the specimens having been collected by Alfred Russel Wallace in his biological expedition to the Malay Archipelago (1854–1862). Further biological expeditions to the regions we now regard as Southeast Asia and Australasia resulted in Schiner describing B. frauenfeldi in 1868, Weyenbergh with B. ritsemai in 1869, de Meijere with B. apicalis, B. albistrigata and B. obscurata in 1911 and B. impunctata in 1914, all also under Dacus. Major Bactrocera pest species were described and/or revised during the 19th and 20th centuries. Major pest species of significance, listed in chronological order, are: B. tau (Walker, 1849), B. tryoni (Froggatt, 1897), B. psidii (Froggatt, 1899), B. cucurbitae (Coquillett, 1899), B. diversa (Coquillett, 1904), B. xanthodes (Broun, 1904), B. cucumis (French, 1907), B. curvipennis (Froggatt, 1909), B. kirki (Froggatt, 1910), B. passiflorae (Froggatt, 1910), B. facialis (Coquillett, 1910), B. melanotus (Coquillett, 1910), B. dorsalis (Hendel, 1912), B. latifrons (Hendel, 1915), B. correcta (Bezzi, 1916), B. occipitalis (Bezzi, 1919), B. minax (Enderlein, 1920), B. jarvisi (Tryon, 1927), B. musae (Tryon, 1927), B. neohumeralis (Hardy, 1951), B. kraussi (Hardy, 1951), B. caryeae (Kapoor, 1971), B. trivialis (Drew, 1971), B. decipiens (Drew, 1972), B. carambolae Drew & Hancock, 1994, B. kandiensis Drew & Hancock, 1994, B. papayae Drew & Hancock, 1994 and B. pyrifoliae Drew & Hancock, 1994, while B. invadens Drew, Tsuruta & White, 2005 and B. divenderi Maneesh, Hancock & Prabhakar, 2022 were added in the 21st century. Most of these belong in subgenus Bactrocera. MATERIAL AND METHODS The following review is based on detailed morphological descriptions and illustrations provided in Drew (1989), Drew & Hancock (1994, 1995), Drew et al. (1999, 2011), Drew & Romig (2001, 2013, 2022), Huxham et al. (2006), Leblanc et al. (2015a, 2015b, 2018, 2021), David et al. (2017), David & Ramani (2019), Doorenweerd et al. (2020), Maneesh et al. (2022, 2023), Korneyev et al. (2024) and Abhishek et al. (2024), supplemented by biogeographical data and information from other publications that are noted in the text, together with an overall familiarity with the fauna. A morphological approach, involving the visible expression of genes in all 439 recognised species, is considered to be more comprehensive and reliable than the limited molecular studies currently available that are based on genes of unknown significance and include no more that 10–20% of the fauna in even the most detailed of studies (e.g., San Jose et al., 2018; Dupuis et al., 2018; Starkie et al., 2022b); these studies also contain numerous anomalous pairings and placements that are contrary to morphological and biogeographical evidence. Terminology follows White et al. (1999). SYSTEMATICS SYNONYMY OF SUBGENUS SEMICALLANTRA Semicallantra Drew, which includes seven species from Indonesia (Maluku) and New Guinea, is currently defined by the presence of elongate antennae and, when present, lateral postsutural yellow vittae that are narrower anteriorly than medially (Hancock & Drew, 2018). However, re-examination of the type-species of Bactrocera, B. longicornis Macquart, 1835 (which also has elongate antennae and anteriorly narrowed lateral postsutural yellow vittae) by Drew & Romig (2022), plus the description of B. malasaitiae Drew & Romig, 2022 from Papua New Guinea, show that neither character is a reliable indicator of subgeneric separation. With the removal of Semicallantra from the subgeneric key in Drew & Romig (2022), its component species run to either subgenus Bactrocera or, in the case of B. cerberae Drew & Romig, 2022, imperfectly to subgenus Calodacus; 75 RAFFLES BULLETIN OF ZOOLOGY 2025 it has a short cell bcu extension on the wing and lacks the male pecten on abdominal tergite III. Elongate antennae also occur in B. (Tetradacus) splendida (Perkins, 1938) and B. (Tetradacus) superba Drew & Romig, 2013 (transferred from Semicallantra by Hancock & Drew, 2018) and in the Dacus subgenera Callantra Walker, 1860 and Mellesis Bezzi, 1916 and this character is evidently homoplasious. Furthermore, the long-antennae species appear to belong in separate complexes and do not form a monophyletic entity. Hence, we do not recognise antennal length as a subgeneric character and regard Semicallantra Drew, 1989 as a new synonym of subgenus Bactrocera Macquart, 1835. Consequently, all species with short antennae currently included in subgenus Bactrocera are retained there, with no resurrection of subgenus Strumeta Walker, 1856 required. Bactrocera (Bactrocera) wallacei Drew & Hancock, new species (Fig. 1) Bactrocera (Bactrocera) epicharis: Drew & Romig, 2013: 79–80; 2016: 151; not Hardy, 1970: 119. Misidentification. Type material. Holotype: male, INDONESIA: North Moluccas [North Maluku] – Maluku, North Maluku, West Halmahera, Goal, 29.v.2007, coll. Raais Abdullah, attracted to cue lure in forest, deposited in Queensland Museum (QMIC), Brisbane. Paratypes: 2 males, 5.vi.2007, same label data as holotype; 1 male (3.vi.2007), 1 male (10.vi.2007), North Maluku, Kodya Ternate, Foramadiahi, coll. La Ruti, attracted to cue lure in forest. Paratypes in Department of Agriculture and Fisheries Collection (QDPC), Brisbane. Diagnosis. A large species (wing 7.5 mm); face fulvous with a pair of medium-sized to large circular black spots; postpronotal lobes and notopleural calli yellow; scutum black; lateral postsutural yellow vittae present, elongate and parallel-sided; medial postsutural yellow vitta absent; no yellow spot anterior to notopleural suture; anepisternal (mesopleural) stripe reaching midway between anterior margin of notopleuron and anterior notopleural seta dorsally; scutullum yellow; wing with cells bc and c fuscous, microtrichia in anteroapical corner of cell c only; a broad dark fuscous costal band becoming paler between R2+3 and R4+5 and remaining of uniform width throughout; a broad dark fuscous anal streak; abdominal terga III–V entirely black. Description. Male. Head: Height 2.0 mm. Frons length 1.57 times breadth, fuscous with fulvous along lateral and ventral margins and dark fuscous on anteromedial hump; orbital setae black: 1 s.or., 2 i.or.; lunule dark fuscous. Ocellar triangle black. Vertex fuscous. Face fulvous with a pair of medium-sized to large circular black spots; length 0.58 mm. Genae red-brown, dark fuscous subocular spot present; black seta present. Occiput fuscous, red-brown along eye margins; occipital row with 5–8 strong black setae. Antennae with segments 1 and 2 red-brown, segment 3 red-brown with fuscous on apex and outer surface; length of segments: 0.24 mm; 0.44 mm; 1.06 mm. Thorax: Scutum black with dark fuscous below and behind lateral postsutural vittae, around notopleural suture, along lateral margins between postpronotal lobe and notopleuron, inside postpronotal lobe. Pleural areas entirely black. Yellow markings as follows: postpronotal lobes; notopleural calli; anepisternal stripe reaching midway between anterior margin of notopleuron and anterior notopleural seta dorsally, continuing to katepisternum as a small transverse spot, anterior margin slightly convex; anatergite (posterior apex black); anterior ⅔ katatergite (remainder black); two broad parallel-sided lateral postsutural vittae ending just behind ia. seta. Postnotum black. Scutellum yellow except for narrow black basal band. Setae (pairs): sc. 1; prsc. 1; ia. 1; p.sa. 1; a.sa. 1; mpl. 1; npl. 2; scp. 2. Legs: All femora entirely fulvous; fore and hind tibiae dark fuscous, mid tibiae fuscous; fore tarsi with basal segment fulvous and apical four segments red-brown, mid and hind tarsi with all segments entirely fulvous; mid tibiae each with an apical black spur. Wings: Length 7.5 mm; cells bc and c fuscous, microtrichia in anteroapical corner of cell c only; remainder of wings colourless except dark fuscous cell sc, dark fuscous costal band confluent with R4+5 but tending fuscous between R2+3 and R4+5 and remaining of uniform width throughout, a broad dark fuscous anal streak; a dense aggregation of microtrichia around A1+CuA2; supernumerary lobe of medium development. Abdomen: Oval; terga free; pecten present on tergum III. Tergum I and sterna I and II wider than long. Tergum I black Fig. 1. Habitus of male Bactrocera (Bactrocera) wallacei Drew & Hancock, new species. 76 Hancock & Drew: Review of subgenus Bactrocera with a narrow transverse red-brown band across posterior margin but not reaching lateral margins; tergum II black with a narrow transverse fulvous band across posterior margin but not reaching lateral margins; terga III–V entirely black. A pair of oval black shining spots on tergum V. All sterna dark fuscous to black. Posterior lobe of surstylus short, sternum V with a deep concavity on posterior margin. Female. No known record. Male attractant. Cue lure. Distribution. Known only from Halmahera and Ternate, North Maluku, Indonesia. Host plants. No known record. Etymology. This species is named after Alfred Russel Wallace in recognition of his pioneering field work in Wallacea and beyond. Remarks. Bactrocera (Bactrocera) wallacei was misidentified as B. epicharis (Hardy, 1970) by Drew & Romig (2013, 2016), who also overlooked the description and discussion of B. epicharis based on new material from the Solomon Islands (Drew & Romig, 2001). It is included here in the trivialis group of the nigella complex and most resembles B. atrabifasciata Drew & Romig, 2001 from the Solomon Islands, differing from it in the narrower anepisternal stripe and fuscous tibiae, and from both B. illusioscutellaris Drew & Romig, 2013 from Bhutan and Vietnam and B. trivialis (Drew, 1971) from New Guinea in the broad costal band and darker abdomen. EXCLUDED SPECIES Bactrocera clarkei Starkie, Strutt & Royer, 2022 was provisionally placed in subgenus Bactrocera by Starkie et al. (2022a) but sternite V is broad with a relatively shallow [moderate] posterior emargination [Starkie et al., 2022a: fig 4]; it is here transferred to subgenus Neozeugodacus May, 1952a, treated as a distinct subgenus in the Melanodacus group of subgenera by Hancock & Drew (2018). As noted in the original description, it closely resembles B. (N.) aurea (May, 1952a) in abdominal markings and in having the apical half of the scutellum brown but differs in the lack of basal scutellar setae and transverse wing band; it also has a narrower anepisternal stripe. Variation in the depth of the sternite V emargination in Neozeugodacus is similar to that seen in subgenus Notodacus Perkins, 1937, which also belongs in the Melanodacus group (Hancock & Drew, 2017b). Bactrocera petila Drew, 1989, known from a single male, was placed in subgenus Bactrocera by Hancock (2015) but differs from all other included species in lacking the male pecten on abdominal tergite III and is provisionally included here in subgenus Calodacus. The original record at cue lure (Drew, 1989) has not been repeated and, as in the case of B. (Trypetidacus) invisitata discussed above, is also likely to have been accidental. Bactrocera cerberae Drew & Romig, originally described in subgenus Semicallantra by Drew & Romig (2022), also lacks the male pecten on abdominal tergite III and is included here in subgenus Calodacus. INCLUDED SPECIES The placement of B. pseudocucurbitae White & Evenhuis, 1999 has been uncertain. Drew & Romig (2013) and Hancock & Drew (2017c) included it in subgenus Parasinodacus Drew & Romig, 2013 (Zeugodacus group of subgenera) but Doorenweerd et al. (2018) returned it to subgenus Bactrocera based on its molecular affiliation with other species in that subgenus. In their original description, White & Evenhuis (1999) noted a V-shaped emargination to sternite V and lack of an elongate posterior surstylus lobe [long and stout in Drew & Romig (2013)]; the scutal and abdominal patterns are typical of several Bactrocera (Bactrocera) species (e.g., B. indecora (Drew, 1971) and B. vulgaris (Drew, 1971) in the indecora complex) and therefore we confirm its placement here. FURTHER NOTES ON THE DORSALIS COMPLEX Of the 18 recognised complexes, the dorsalis complex is the second largest and has attracted the most interest due to the economic implications relating to some major pest species, with B. citima (Hardy, 1973), B. ellenriederae Korneyev, Leblanc, Hauser, General & Gaimari, 2024, B. parafroggatti Drew & Romig, 2001 and B. youngi Korneyev, Leblanc, Hauser, General & Gaimari, 2024 here added to the 79 species included by Drew & Hancock (2022). Over the past decade, a part of the dorsalis complex has been left in a state of confusion that began with the erroneous synonymy of B. invadens and B. papayae with B. dorsalis by Schutze et al. (2015a, b), which has been followed by numerous publications that mainly focused on molecular analyses of specimens, generally without a detailed understanding of Bactrocera taxonomy. This limited approach has led to increased confusion in the diagnosis of species, particularly as most studies have resulted in conflicting results and incongruence between morphological and molecular data. Bactrocera youngi is a possible synonym of B. fulvifemur Drew & Hancock, 1994. Both were described from Luzon, Philippines (Drew & Hancock, 1994; Korneyev et al., 2024) and the presence of red-brown submedial scutal vittae covered with silvery pubescence in some specimens was recorded in the description of B. fulvifemur by Drew & Hancock (1994). This latter species was not mentioned by Korneyev et al. (2024) and we can find no reliable characters that separate the two taxa as currently known. However, further comparison between the black and red-striped forms is needed to confirm synonymy. A second species described from Luzon by Korneyev et al. (2024), B. ellenriederae, is very similar to B. lateritaenia Drew & Hancock, 1994 and 77 RAFFLES BULLETIN OF ZOOLOGY 2025 possibly synonymous, but the latter species is not yet known from the Philippines so we provisionally accept them as separate species pending further study. Bactrocera papayae and B. invadens were erroneously synonymised with B. dorsalis by Schutze et al. (2015a, b) and subsequently reestablished as valid species by Drew & Romig (2016). In his 1973 monograph, Hardy perceptively recognised B. papayae as a species distinct from B. dorsalis, although he incorrectly diagnosed it as the Philippine species B. pedestris (Bezzi, 1913) (see Hardy, 1973: p. 50). The specific status of both B. papayae and B. invadens was further confirmed by Drew & Hancock (2022) and Drew & Romig (2022). Contrary to Doorenweerd et al. (2024), Drew & Romig (2022) did not state that the dark foretibial stripe in B. papayae ‘quickly fades in deceased specimens’ but that it is more distinct in fresh specimens; it merely becomes harder to discern in dried material but is still evident, as in a specimen from Bangkok, Thailand examined by DLH (in NAQS Collection, Cairns, Queensland, Australia). Despite the study of Drew & Hancock (2022), confusion between this species, B. dorsalis, and B. invadens persists. The shorter and subovate glans of B. dorsalis is evident in Taiwanese specimens misidentified as ‘Dacus pedestris’ by Tseng et al. (1992) and is very similar to that illustrated by Drew & Hancock (2022) of Chinese dorsalis [cf. B. invadens in Maneesh et al. (2022)]. The ‘Dacus dorsalis’ male illustrated by Tseng et al. (1992) also differs from true B. dorsalis in its narrower anepisternal stripe, slight apical expansion of the costal band, short cell bcu extension, abdominal pattern and internal structure of the glans and appears to belong to a separate species. Part of the belief that B. dorsalis, B. papayae, and B. invadens are conspecific stems from methyl eugenol response studies by Hee et al. (2015a), using the IAEA Saraburi (Thailand) colony misidentified as ‘dorsalis’ instead of papayae and originating within the known distribution of the latter species (Drew & Hancock, 2022). A study that used true B. dorsalis from Taiwan (Wee et al., 2002) produced conflicting results that were dismissed as regional variation by Hee et al. (2015a), but indicated that B. dorsalis is twice as responsive to methyl eugenol as B. papayae, and that these two species are, respectively, 17 and 9 times more responsive than B. carambolae. The similarity in lure response between the Saraburi colony and B. papayae supports its actual identity as the latter species, with comparison in Hee et al. (2015a) thus being made between papayae and papayae. The response of B. invadens is similar to that of B. papayae (see Hee et al., 2015a) but morphological differences are significant (Drew & Hancock, 2022). Two ‘cryptic’ species in the dorsalis complex, B. incognita Doorenweerd & San Jose, 2024 and B. borneoensis Doorenweerd & San Jose, 2024, with holotypes from Cambodia and Sabah, respectively, were described recently by Doorenweerd et al. (2024) solely on molecular evidence with the bodies of the entire type series destroyed for DNA extraction, leaving only photographs of the holotypes while they were in ethanol and legs, wings, and terminalia mounted on slides. In the absence of supporting evidence such as female morphology, host plant preference or lure difference, we are unable to support specific status for these two taxa and regard them as genetic variants and new synonyms of B. occipitalis and B. carambolae respectively. These two species have distinctive abdominal patterns identical to those in the ‘cryptic’ variants, males of all four taxa are attracted to methyl eugenol and, although synonymies represent no more than taxonomic opinion based on available evidence, validation of currently inseparable taxa requires more convincing information. Genetic variation is extensive in at least some dorsalis complex species (e.g., B. carambolae: Aketarawong et al., 2015; Drosopoulou et al., 2019) and the gene fragments analysed are either known to be unreliable (COI) (e.g., Doorenweerd et al., 2024) or their reliability is currently unknown (RAD-seq and HiMAP). The holotype of B. incognita has a dark fore femoral spot and, despite the statement in Doorenweerd et al. (2024) that a dark fore femoral spot is never present in B. raiensis Drew & Hancock, 1994, such a spot was recorded in some specimens of both it and B. occipitalis by Drew & Hancock (1994). However, B. raiensis has a narrower costal band and more extensive lateral dark markings on the abdomen. The slightly narrower costal band in ‘incognita’ is also seen in some Philippine specimens of B. occipitalis and the darker scutum regarded as morphological variation, the extent of which in the incognita type series can no longer be determined. The distribution of B. occipitalis therefore ranges from southern Thailand, Cambodia, and Vietnam to West Malaysia, Borneo and the Philippines. Bactrocera occipitalis was recorded previously from Brunei, Sabah, and Kalimantan by Drew & Romig (2013). Bactrocera carambolae is common in the Danum Valley, Sabah [Drew & Hancock (1994) recorded 254 males], the type locality of B. borneoensis, and the two taxa cannot be separated morphologically. The subapical fore femoral spot is generally present in females but often absent in males; the type series of the latter is comprised solely of males and the statement that the fore femoral spot is always absent in borneoensis (in Doorenweerd et al., 2024) is based on too few specimens to be conclusive. The genetic similarity between incognita and borneoensis is likely due to introgression, as both taxa occur sympatrically with B. carambolae outside the Philippines and with each other in Borneo. It is considered inadvisable to describe species solely on molecular evidence where individual specimens cannot be identified without their destruction, leaving them unavailable for checking by taxonomists experienced in the particular fauna and thereby failing the results reproducibility test. Taiwanese specimens of ‘Dacus parvulus’ illustrated in Tseng et al. (1992) are regarded here as misidentifications of B. irvingiae Drew & Hancock, 1994 and run to it in the key of Drew & Hancock (1994); neither has been recorded from male lures. Taiwan is thus a new country record for B. irvingiae. 78 Hancock & Drew: Review of subgenus Bactrocera COMPLEXES AND SPECIES GROUPS A long-standing convention in taxonomy where a large and diverse fauna is being researched is to group species in complexes based on morphological similarities. While a complex is not a formal category and not necessarily monophyletic, experienced researchers in taxonomic studies of large faunas often place species within such complexes. This process assists in building diagnostic keys to species and, in part, understanding evolutionary and biogeographical relationships. Definition and scope of the term ‘complex’ varies among authors, with that used here (sensu Drew, 1989; Drew & Romig, 2013) used widely in the Dacinae. Over the past decade, many papers have been published by authors inexperienced in tephritid taxonomy and the processes and practice of systematic research. An example is that by Catullo et al. (2019), who appear to have misunderstood the concept of species complexes and stated that B. endiandrae (Perkins & May, 1949) should not be placed in the dorsalis complex based on molecular data. Based on morphology and lure response, B. endiandrae is best placed in the dorsalis group, close to B. parafroggatti Drew & Romig, 2001 [referred to the dorsalis complex by Doorenweerd et al., 2024] and nowhere near either B. aeroginosa (Drew & Hancock, 1981 [in Drew et al., 1981]) or B. umbrosa (Fabricius, 1805) as indicated by Catullo et al. (2019), B. lampabilis (Drew, 1971) as placed by Starkie et al. (2022b), or B. (Calodacus) calophylli (Perkins & May, 1949) as placed by San Jose et al. (2018), all on molecular grounds. Krosch et al. (2012) placed it in an unresolved clade that also included the dorsalis complex, B. murrayi (Perkins, 1939) and B. (C.) calophylli, while Dupuis et al. (2018) also included it in an unresolved clade. The following definitions of complexes and groups may be used as a ‘Key by elimination,’ reading through each complex in turn until a satisfactory placement is made and then using the group definitions for a more detailed placement. Monotypic groups reflect diversity within the complexes and might be combined with other groups when better known. Some species or groups do not fit readily in these complexes but are included to facilitate identification. longicornis complex: often largely black species with a very broad transverse wing band enclosing both R-M and DM-Cu crossveins, recurved towards wing base along vein Cu1 or alongside it in cell cu1 and usually expanded basal to R-M crossvein in cell dm and beyond line of DM-Cu crossvein in cells r4+5 and m; if wing almost entirely fuscous in apical 2/3 then with a longitudinal hyaline band in anterobasal quarter of cell dm, or without transverse bands or an isolated longitudinal hyaline band in cell dm but fuscous over all or most of wing or at least apically beyond line of R-M crossvein [17 species: Sulawesi and Maluku to New Caledonia and NE Australia]. ampla group: prescutellar acrostichal setae present; scutellum with basal black band narrow and not expanded; wing with transverse discal band not united with broad apical area in cell r4+5 and without a broad transverse basal band across BM-Cu crossvein; all femora fulvous; abdomen with a broad black medial vitta and lateral margins on tergites II–V; males respond to cue lure. Species: ampla (Drew, 1971). amplexiseta group: wing without transverse bands or an isolated longitudinal hyaline band in cell dm but fuscous over all or most of wing or at least apically beyond line of R-M crossvein; postpronotal lobes entirely yellow or apically fuscous; body and leg markings variable; costal cells bc and c pale to dark fuscous with microtrichia in anteroapical corner of cell c only; males respond to methyl eugenol. Species: amplexiseta (May, 1962a); ebenea (Drew, 1971); fumica Drew & Romig, 2022; fuscalata Drew, 1989; fuscoptera Drew & Romig, 2013. biarcuata group: prescutellar acrostichal setae present; scutellum with basal black band broadly convex or triangular but not crossing scutellum; wing with transverse discal band united with preapical band in cell r4+5 or preapical band absent; at least mid and hind femora apically fuscous; abdomen mostly black, with at most a pair of small posterior pale patches on tergite II and without a broad black medial vitta and lateral margins on tergites III–V; males respond to methyl eugenol. Species: biarcuata (Walker, 1865); malasaitiae Drew & Romig, 2022; retrorsa Drew, 1989. curvifer group: prescutellar acrostichal setae often absent; scutellum with basal black band narrow and not expanded; wing with transverse basal band across BM-Cu crossvein broad in cell dm; all femora fulvous; abdomen black or with a broad black medial vitta and lateral margins on tergites III–V and with a broad or medially intersected posterior pale band on tergite II; males respond to methyl eugenol. Species: confluens (Drew, 1971); curvifer (Walker, 1864); pepisalae (Froggatt, 1910); seguyi (Hering, 1939); speculifer (Walker, 1865); ternatiae Drew & Romig, 2013. longicornis group: prescutellar acrostichal setae present; scutellum with a broad medial vitta that reaches apex of scutellum; wing with transverse discal band not united with preapical band in cell r4+5; all femora apically fuscous; abdomen mostly black, with at most a pair of small posterior pale patches on tergite II and without a broad black medial vitta and lateral margins on tergites III–V; males respond to cue lure. Species: denigrata (Drew, 1971); longicornis Macquart, 1835. alyxiae complex: often largely black species with the transverse band enclosing R-M and DM-Cu crossveins either entire or separated in cell dm, not expanded basal to R-M crossvein in cell dm and beyond line of DM-Cu crossvein in cell r4+5 and recurved towards wing base along vein Cu1 or alongside it in cell cu1; or with the band uniformly broad and with a distinct preapical band; or with 2 separate bands over R-M and DM-Cu crossveins that are either united posteriorly or separated in cell dm into a Uor V-shaped band [16 species: Sulawesi to New Caledonia, Mariana Islands and NE Australia, with B. umbrosa also widespread in SE Asia. 79 RAFFLES BULLETIN OF ZOOLOGY 2025 alyxiae group: scutellum yellow with a narrow basal black band; transverse wing band broad and enclosing both R-M and DM-Cu crossveins and recurved towards wing base alongside vein Cu1 in cell cu1; all femora fulvous; abdomen entirely pale or black with tergite II broadly pale posteriorly and a broad pale medial stripe on tergites III–V; males respond to cue lure. Species: alyxiae (May, 1952b); repanda Drew, 1989. halmaherae group: scutellum yellow with a narrow basal black band; wing with 2 transverse bands over R-M and DM-Cu crossveins separated in cell dm and recurved towards wing base along vein Cu1; all femora diffusely darkened apically; all tibiae fuscous; abdomen with a narrow black T-shaped pattern on tergites III–V; males respond to cue lure. Species: halmaherae Drew & Romig, 2013. ochrosiae group: scutellum yellow with a broadly oval black basal band; anepisternal yellow stripe horizontal and reaching postpronotal lobe; wing with transverse band over R-M and DM-Cu crossveins broad and aligned with pterostigma, with a distinct but short preapical band and with a broad basal band united with anal stripe; costal band interrupted between medial and preapical bands; all femora fulvous; abdominal tergites III–V orange-brown with broad black posterior bands; males respond to cue lure. Species: ochrosiae (Malloch, 1942). This species is known only from Guam and the Northern Mariana Islands, a record from Hawaii being very doubtful (Leblanc, 2022). reclinata group: scutellum with either a broad black basal triangular patch or a medial stripe that reaches apex of scutellum; transverse wing band or bands enclosing both R-M and DM-Cu crossveins recurved or not towards wing base along vein Cu1 and either entire and of uniform width or separated in cell dm and basal band short or absent, not reaching anal stripe; fore and hind femora with apical half black, mid femur black; abdomen black with at most a pair of small pale posterior patches on tergite II; males respond to methyl eugenol. Species: ismayi Drew, 1989; lampabilis (Drew, 1971); reclinata Drew, 1989. recurrens group: scutellum yellow with a narrow basal black band; wing with 2 transverse bands over R-M and DM-Cu crossveins united posteriorly into a Uor V-shaped band; preapical band absent, reduced to a spot, or connected with band over DM-Cu crossvein; all femora fulvous; abdomen entirely pale, with a narrow black medial vitta on tergites II–V or III–V or with tergite II broadly pale posteriorly and tergites III–V with a black T-shaped pattern and broad black lateral margins; males respond to cue lure. Species: absidata Drew, 1989; anfracta Drew, 1989; manskii (Perkins & May, 1949); nigrescentis (Drew, 1971); recurrens (Hering, 1941a); redunca (Drew, 1971); resima (Drew, 1971). umbrosa group: scutellum yellow with a narrow basal black band; wing with transverse band over R-M and DM-Cu crossveins broad and aligned with apex of pterostigma, with a distinct preapical band and with or without a broad basal band united with anal stripe; all femora fulvous; abdomen with tergites III–V with variable fuscous medial and/or lateral bands; males respond to cue lure or methyl eugenol. Species: bifasciata (Hardy, 1982); umbrosa (Fabricius, 1805). distincta complex: often largely black species with costal band distinct and a transverse wing band enclosing both R-M and DM-Cu crossveins not expanded beyond line of DM-Cu crossvein in cell r4+5 and sometimes absent over anterior part of R-M crossvein; preapical band absent; scutellum with a broad black medial band or mostly yellow; femora usually entirely pale or narrowly darkened apically [40 species: Maluku to New Caledonia, Fiji, and Australia]. angustifasciata group: scutellum with a broad black medial vitta; scutum black; males respond to cue lure. Species: angustifasciata Drew, 1989; hollingsworthi Drew & Romig, 2001; raunsepnaensis Drew & Romig, 2022; rounaensis Drew & Romig, 2022; unilineata Drew, 1989. distincta group: scutellum yellow with at most an apical fuscous band and a narrowly concave black basal band; males respond to cue lure, zingerone or response unknown. Species: allodistincta Leblanc & Doorenweerd, 2021 [in Leblanc et al., 2021]; anomala (Drew, 1971); atriliniellata Drew, 1989; avittata Drew & Romig, 2013; curreyi Drew, 1989; decumana (Drew, 1972); distincta (Malloch, 1931); fergussoniensis Drew, 1989; furvilineata Drew, 1989; fuscohumeralis White & Evenhuis, 1999; latilineata Drew, 1989; monostriata Drew & Romig, 2022; morobiensis Drew, 1989; neofulvicauda Drew & Romig, 2013; oblineata Drew, 1989; penephaea Drew & Romig, 2013; pisinna Drew, 1989; propedistincta Drew, 1989; pseudodistincta Drew, 1989; pulchra Tryon, 1927; rhabdota Drew, 1989; tikelingiae Drew & Romig, 2022; torresiae Huxham & Hancock, 2006 [in Huxham et al., 2006]; tortuosa White & Evenhuis, 1999; truncata Drew & Romig, 2013; tsatsiai Leblanc & Doorenweerd, 2021 [in Leblanc et al., 2021]; unifasciata (Malloch, 1939); unitaeniola Drew & Romig, 2001. fulvicauda group: as for distincta group but males respond to methyl eugenol. Species: fulvicauda (Perkins, 1939); inconspicua Drew & Romig, 2013; maculigera Doleschall, 1858; manusiae Drew & Romig, 2022; obliquivenosa Drew & Romig, 2001; paranigrita Drew & Romig, 2013; unistriata Drew, 1989. frauenfeldi complex: largely black species with costal band very faint or absent beyond pterostigma, preapical band absent and transverse wing band distinct or faint and posteriorly diffuse and enclosing both R-M and DM-Cu crossveins; R-M crossvein distinctly oblique and less than its own length from DM-Cu crossvein; scutellum basally broadly black or with a broad black medial vitta [8 species: New Guinea to New Caledonia and Australia, except B. albistrigata, which is widespread from Andaman Islands and southern Thailand to Maluku and Timor]. frauenfeldi group: abdomen with a broad black medial vitta on tergites II–V and broad black lateral margins on tergites II–V or III–V (sometimes coalesced with medial 80 Hancock & Drew: Review of subgenus Bactrocera vitta); males respond to cue lure. Species: albistrigata (de Meijere, 1911); frauenfeldi (Schiner, 1868); parafrauenfeldi Drew, 1989; trilineola Drew, 1989. Proposed synonymy of albistrigata with frauenfeldi by Doorenweerd et al. (2023a) was considered to be based on insufficient evidence by Drew & Hancock (2022) and was not supported by the molecular studies of Yong et al. (2024). obliqua group: abdomen black with or without paler areas posterolaterally on tergite II; males respond to cue lure, zingerone or possibly isoeugenol. Species: caledoniensis Drew, 1989; obliqua (Malloch, 1931); vargasi Leblanc & Doorenweerd, 2021 [in Leblanc et al., 2021]; yayamiae Drew & Romig, 2022. atramentata complex: often black or mostly black species; wing with costal band very narrow and linear or pale and indistinct; preapical band absent; transverse band narrow (strigata) or absent, with or without a pale fuscous tint over R-M, DM-Cu or both R-M and DM-Cu crossveins; scutum with postsutural lateral yellow vittae often very narrow, narrowing anteriorly or absent; notopleural calli yellow or orange-brown to black; if scutum largely red-brown or orange-brown then postpronotal lobes brown or notopleural calli orange-brown to fuscous; scutellum black, with a broad black or red-brown medial stripe reaching apex, with a triangular black basal band not reaching apical margin, with a broad red-brown apical band or mostly yellow; legs often with femora and/or tibiae mostly black [18 species: Eastern Australia and Papua New Guinea to Cook, Marquesas, Henderson, Austral and Society Islands, and the Tuamotu Archipelago]. atra group: notopleural calli black; scutellum black or with lateral yellow margins and a broad medial black stripe reaching apical margin; abdomen black or with pale submedial patches apically on tergites II–IV and basally on tergites IV–V; males respond to cue lure or response unknown. Species: atra (Malloch, 1938); carbonaria (Hendel, 1927b); melanotus (Coquillett, 1910); perfusca (Aubertin, 1929); setinervis (Malloch, 1938). atramentata group: notopleural calli yellow; scutellum with a broad medial black stripe reaching apical margin or with a broadly triangular black basal band; if costal band pale and indistinct then R-M or R-M and DM-Cu crossveins with a pale fuscous tint; abdomen black with paler apical or subapical areas on tergite V at most; males respond to cue lure. Species: atramentata (Hering, 1941c); hypomelaina Drew, 1989; morula Drew, 1989; psidii (Froggatt, 1899). kirki group: notopleural calli yellow; scutellum with lateral yellow margins and a very broad medial black stripe reaching apical margin; costal band pale; transverse wing band reduced to a pale tint over R-M and DM-Cu crossveins or absent; all femora pale; abdomen with a broad black medial vitta and broad black lateral margins on tergites I–V or II–V; males respond to cue lure. Species: enochra (Drew, 1972); kirki (Froggatt, 1910); trifaria (Drew, 1971). luteola group: scutum mostly shining orange-brown without lateral yellow vittae; notopleural calli shining orange-brown; scutellum yellow with a broad orange-brown basal band; costal cells hyaline; costal band narrow; crossveins R-M and DM-Cu with a faint infuscation; anal streak faint or absent; facial spots absent; all femora and tibiae fulvous; abdomen entirely shining orange-brown with small fuscous patches anterolaterally and medially on tergite III and medially on tergite IV; males show no response to known lures. Species: luteola (Malloch, 1931). picea group: scutum black with elongate lateral yellow vittae; notopleural calli yellow; scutellum with a broad medial black stripe reaching apical margin; costal band pale and indistinct and R-M and DM-Cu crossveins without a pale fuscous tint; mid and hind femora black; abdomen black with small paler areas posteriorly on tergite II; males respond to methyl eugenol. Species picea (Drew, 1972). strigata group: scutum red-brown or orange-brown with short or elongate lateral yellow vittae; notopleural calli red-brown or fuscous; costal band narrow; costal cells hyaline or fulvous; wing with or without a transverse band across R-M, DM-Cu or both R-M and DM-Cu crossveins; abdomen orange-brown with or without a narrow black medial vitta on abdominal tergites III–V or V; scutellum yellow, broadly red-brown apically or red-brown with a pair of large basolateral yellow spots; facial spots small or absent; female aculeus with 1–3 pairs of subapical lobes; males show no response to known lures. Species: brunnea (Perkins & May, 1949); hispidula (May, 1957); phaleriae (May, 1955); strigata (Perkins, 1934). These four species are restricted to Eastern Australia, with a reported response of B. phaleriae to isoeugenol based on a single specimen (Starkie et al., 2022a) that was possibly accidental in the trap. laticaudus complex: scutum dark fuscous to black; notopleural calli yellow; wing without transverse or preapical bands but with a distinct infuscation over R-M crossvein that sometimes extends weakly into cell dm; costal band distinct and extending across vein R2+3 and often reaching vein R4+5; costal cells bc and c hyaline to pale fuscous and with microtrichia confined to anteroapical corner or apical half of cell c; scutellum yellow with a narrow black basal band [6 species: Maluku to Solomon Islands and Australia]. laticaudus group: scutum dark fuscous to black; abdomen mostly black with a pair of large, pale posterior patches or broadly pale on tergite II; all femora apically dark fuscous to black; males respond to methyl eugenol. Species: laticaudus (Hardy, 1950); melanogaster Drew, 1989; neonigrita Drew, 1989; quasineonigrita Drew & Romig, 2013. phaea group: scutum largely black; abdomen largely pale with a dark T-shaped pattern over tergites III–V; all femora apically pale fulvous; males respond to cue lure. Species: kunvawaensis Drew & Romig, 2022; phaea (Drew, 1971). 81 RAFFLES BULLETIN OF ZOOLOGY 2025 tryoni complex: wing without transverse or preapical bands, at most with a pale or distinct infuscation over R-M crossvein; costal band distinct and narrow or broad, often reaching or almost reaching vein R4+5 throughout its length; costal cells bc and c with at least a fulvous tint and at least most of cell c (and often part of cell bc) covered with microtrichia; hind tibiae often fuscous; abdominal tergite I wider than long and broadest posteriorly except in B. incompta [39 species: Malaysia to New Caledonia and Australia]. assita group: costal band broad, crossing vein R2+3 and often reaching or almost reaching vein R4+5 throughout its length; costal cells bc and c pale to dark fuscous; femora pale or at least apically fuscous; hind tibiae fuscous or pale; males respond to cue lure. Species: apicopicta Drew & Romig, 2013; assita Drew, 1989; careofascia Drew & Romig, 2013; fuscolobata Drew & Romig, 2013; fuscoformosa Drew & Romig, 2013; gabensiae Drew & Romig, 2022; labubulu Drew & Romig, 2022; ochracea Drew, 1989; paraochracea Drew & Romig, 2022; pusilla (Hardy, 1983); ustulata Drew, 1989. curvipennis group: costal band broad, reaching vein R4+5 throughout its length; crossveun R-M infuscated; costal cells bc and c pale fuscous; femora pale; hind tibiae pale; males respond to cue lure, isoeugenol and dihydroeugenol. Species: curvipennis (Froggatt, 1909). incompta group: scutum largely dull black (fuscous posteriorly) without lateral yellow vittae; postpronotal lobes and notopleural calli entirely dark fuscous; supra-alar setae absent; antennae with segment I elongate; abdomen elongate with tergite I longer than wide and almost parallel-sided, not widest posteriorly; tergites II–V red-brown without a dark T-shaped pattern and with darker lateral margins on tergite III; costal band narrow and apically linear; costal cells pale fuscous with dense microtrichia over all of cell c; all femora red-brown; all tibiae fuscous at least basally; males respond to cue lure. One species: incompta Drew & Romig, 2013. This species is distinct and provisionally included in the tryoni complex pending further study. mediorufula group: costal band narrow or broadened apically; costal cells bc and c fuscous; scutum black with a broad rufous medial vitta ending before prescutellar acrostichal setae; femora fulvous or with a fuscous apical spot; fore, at least base of mid and hind tibiae fuscous; males respond to methyl eugenol. Species: mediorufula Drew & Romig, 2013; tapahensis Drew & Romig, 2013. notatagena group: costal band narrow or broad, often reaching or almost reaching vein R4+5 throughout its length; costal cells bc and c fulvous to fuscous; femora pale, apically dark or with fore, mid and apical third to half of hind femora fuscous; male response to lures uncertain or unknown; isolated reports of B. daruensis at methyl eugenol (Huxham & Hancock, 2002) and B. mutabilis at isoeugenol (Starkie et al., 2022a) appear to be accidental. Species: buloloensis Drew, 1989; caliginosa (Hardy, 1970); commina Drew, 1989; daruensis Drew, 1989; humilis (Drew & Hancock, 1981) [in Drew et al., 1981]; kelaena Drew, 1989; laensis Drew & Romig, 2022; mendosa (May, 1957); mutabilis (May, 1952a); notatagena (May, 1952b); pectoralis (Walker, 1859); pometiae Drew & Romig, 2022; popondettiensis Drew, 1989; trivirgulata Drew & Romig, 2022. romigae group: costal band broad, reaching or almost reaching vein R4+5 throughout its length; costal cells bc and c pale to dark fuscous; fore and at least part of mid and hind femora fuscous; males respond to methyl eugenol or dihydroeugenol and isoeugenol. Species: nigrovittata Drew, 1989; quadrisetosa (Bezzi, 1928) [= varipes Drew, 1989]; romigae (Drew & Hancock, 1981) [in Drew et al., 1981]. speewahensis group: costal band narrow; costal cells bc and c fuscous; all femora fulvous; scutellum broadly redbrown posteriorly; males respond to zingerone. Species: speewahensis Fay & Hancock, 2006 [in Huxham et al., 2006]. sylvania group: costal band broad, extending well beyond vein R4+5 over most of its length; costal cells bc and c fuscous (cell bc paler); fore, mid and apical third of hind femora fuscous; males respond to methyl eugenol or response unknown. Species: grandifasciata White & Evenhuis, 1999; sylvania Drew & Romig, 2022. tryoni group: costal band narrow, often crossing vein R2+3 but not reaching or almost reaching vein R4+5 except at apex; costal cells bc and c fulvous to fuscous; all femora pale; hind tibiae fuscous; males respond to cue lure. Species: aquilonis (May, 1965); melas (Perkins & May, 1949) [hybrid?]; neohumeralis (Hardy, 1951); tryoni (Froggatt, 1897). nigrotibialis complex: wing without transverse or preapical bands or a distinct infuscation over R-M crossvein; costal band distinct and narrow to broad, often faintly reaching but not crossing vein R4+5 except at apex; costal cells bc and c hyaline or with a pale fulvous to fuscous tint and not covered with microtrichia; face with a pair of dark spots or black; scutum black without paler areas and with postsutural lateral yellow vittae often very narrow, narrowing anteriorly or absent; scutellum with a narrow or broad black basal band not reaching apex; legs with femora and tibiae mostly or at least partly black; abdomen black with at most pale areas posteriorly on tergites I–II and dorsocentrally-posteriorly on tergites III–V; tergite I not broadly pale medially [30 species: Pakistan and India to Japan, Solomon Islands, and northern Australia]. aquila group: costal band broad but not crossing vein R4+5 except at apex; scutellum yellow with a narrow black basal band; antennae often elongate; all femora largely or entirely black; abdomen black with at most small paler areas posteriorly on tergites II or V; males respond to cue lure or response unknown. Species: aquila (Drew, 1989); epicharis (Hardy, 1970); exspoliata (Hering, 1941b); memnonia (Drew, 1989); nigricula (Drew, 1989); toxopeusi (Hering, 1953). diospyri group: costal band narrow; scutum with lateral postsutural yellow vittae short and triangular or absent; 88 Hancock & Drew: Review of subgenus Bactrocera yellow with anterior black or fuscous area; scutellum with a broad black apical band or patch; abdominal tergites III–V with a black T-shaped pattern and broad lateral margins; costal band narrow and either fulvous in cell r1 or with an isolated apical spot; costal cells hyaline with microtrichia reduced to a narrow anteromedial stripe in cell c; anal stripe narrow; all femora fulvous or mid and hind femora narrowly black at apex; hind tibiae fuscous; response to male lures unknown. Species: hyalina (Shiraki, 1933); venefica (Hering, 1938). versicolor group: scutum red-brown with fuscous markings; costal band very narrow and linear; scutellum with or without a fuscous apical patch; femora fulvous or with pale fuscous subapical spots on fore and mid femora and around extreme apex of hind femora; hind tibiae fuscous or all tibiae fulvous with fuscous bases; abdominal tergites III–V with a black T-shaped pattern and very broad lateral fuscous bands; ceromata dark red-brown to black; aculeus needle-like; males respond to methyl eugenol. Species: aethriobasis (Hardy, 1973); versicolor (Bezzi, 1916). zonata group: scutum red-brown with pale fuscous markings or black; costal band interrupted in cell r2+3 and with an isolated apical spot; scutellum without a fuscous apical patch; femora fulvous; abdominal tergites III–V black, mostly black with a black medial vitta on tergites IV–V, or pale with a narrow T-shaped pattern often reduced to anterolateral streaks on tergite III and medially on tergites IV–V or V; ceromata pale or dark red-brown to black; aculeus needlelike or apically trilobed; males respond to methyl eugenol. Species: affinis (Hardy, 1954); correcta (Bezzi, 1916); penecorrecta Drew, 2002; tuberculata (Bezzi, 1916); zonata (Saunders, 1842). LIST OF SPECIES The 439 species, their complex and group placements, male lures and distributions are listed in Table 1. Invasive, vagrant, and presumed misidentified records are excluded from the distributions. Invasive populations within the Asia-Pacific Region (i.e., excluding those noted in the Introduction) include B. carambolae in Bangladesh, B. tryoni in New Caledonia, French Polynesia and Pitcairn Island [eradicated from Easter Island], B. dorsalis in Tahiti, B. latifrons in Timor-Leste, and B. papayae in Palau and New Guinea (Drew, 1989; Bellis et al., 2017; Drew & Romig, 2022; Hoskins et al., 2023). Both B. latifrons and B. parvula were recorded as invasive in Japan’s Ryukyu Islands by Hisaoka et al. (2024): B. parvula (= ‘Haplotype A’) on Yonaguni Island and B. latifrons (= ‘Haplotype B’) throughout. Vagrants in Australia’s Torres Strait islands were noted by Huxham & Hancock (2002) and Hancock (2013). Presumed misidentifications include the Sri Lankan B. hantanae from Sulawesi (Doorenweerd et al., 2020). India’s Andaman Islands and Australia’s Christmas Island are included in Zone B and the Mariana Islands in Zone D. Most species can be identified by illustrations and keys in Drew (1989), Drew & Romig (2001, 2013, 2016, 2022), Leblanc (2022), Korneyev et al. (2024) and Abhishek et al. (2024). Note, however, that illustrations of ‘B. dorsalis’ in Korneyev et al. (2024) are of B. invadens. BIOGEOGRAPHY The biogeography of the Dacini was discussed in detail by Drew & Hancock (1999) and Drew (2004) and, in Papua New Guinea, by Clarke et al. (2004). The endemic habitat of genus Bactrocera is the tropical and subtropical rainforest ecosystem recognised as the Indomalayan rainforest flora. Because there is a close association between fruit fly species and their host plants, whereby their reproductive biology is dependent upon factors within the host plant environment, Drew & Hancock (1999) proposed that the fruit fly species and their host plants continued to co-evolve over the Tertiary and Quaternary Periods. Within subgenus Bactrocera, there have been high levels of speciation in the Asian-Pacific Region (Table 2). The largest number of known species is in Papua New Guinea and the Melanesian Archipelago (193), followed by Southeast Asia (102), Wallacea (84), Australia (60), the Indian subcontinent (49), and the South Pacific (24). This pattern matches the distribution of Indomalayan rainforest plant species, which is at a peak in Papua New Guinea and Southeast Asia and then markedly declines to Australia and the South Pacific. The two largest complexes within subgenus Bactrocera are the dorsalis complex (83 species) and the quadrata complex (86 species) (Tables 3 and 4). The distribution of species in these complexes demonstrates a concentration of the dorsalis-type melanic morphological characters in Southeast Asia and the quadrata-type pale characters in Papua New Guinea and the Pacific region. These character states overlap in Wallacea, which is a zone of integration between the two major faunas of Southeast Asia and the Pacific region. Wallacea is now recognised as a transitional zone with shared species from West and East as well as having experienced considerable independent speciation. The dorsalis complex has undergone prolific speciation in the continental area of Southeast Asia, with fewer species originating in the Philippines, Indonesia, and Borneo. It is important to recognise that the Philippines and Borneo possess several Bactrocera species in common that are not shared with other areas of Southeast Asia. This has probably resulted from these two land areas remaining connected for a longer period after isolation from the rest of Southeast Asia. A practical consequence of this has been the inability to separate, on morphological characters, the Bactrocera papayae populations that currently occur in the Philippines, Borneo, and Java. Minor differences between Philippine and other populations of B. papayae were later judged insufficient to support species status (Drew & Romig, 2013) and thus B. philippinensis Drew & Hancock, 1994 is presently regarded as a synonym of B. papayae. Extensive speciation in the quadrata complex has occurred in Papua New Guinea and northeastern Australia. These two land areas also show some species indicative of periods of both isolation and unification. 89 RAFFLES BULLETIN OF ZOOLOGY 2025 Table 1. List of species in subgenus Bactrocera, with their complex and group placements, known lures and biogeographic zones in the Indo-Australian region (A = Indian Subcontinent; B = Southeast Asia; C = Wallacea; D = New Guinea & Solomon Islands; E = Australia; F = South Pacific) [Zones after Hancock & Drew (2015)]. Species Complex Group Lure Zone abdofuscata quadrata abdofuscata Unknown D abdolonginqua indecora abdolonginqua ME D abdonigella quadrata abdofuscata Cue CD aberrans musae aberrans Isoeugenol E abscondita quadrata silvicola Cue DE absidata alyxiae recurrens Unknown D absoluta quadrata dispar Unknown C abundans quadrata silvicola Cue D adamantea quadrata adamantea Zingerone B aemula dorsalis pedestris Cue CD aeroginosa quadrata quadrata Cue, zingerone DE aethriobasis zonata versicolor ME AB affinibancroftii quadrata barringtoniae ME C affinidorsalis dorsalis pedestris Cue BC affinis zonata zonata ME A aithogaster quadrata barringtoniae Unknown D albistrigata frauenfeldi frauenfeldi Cue BC allodistincta distincta distincta Cue D allwoodi musae brevistriata Cue E alyxiae alyxiae alyxiae Cue, zingerone DE amarambalensis dorsalis caryeae ME A ampla longicornis ampla Cue D amplexiseta longicornis amplexiseta ME E andamanensis quadrata andamanensis Cue B anfracta alyxiae recurrens Cue D angustifasciata distincta angustifasciata Cue D anomala distincta distincta Cue F anthracina nigrotibialis nigrotibialis Cue D antigone quadrata quadrata Cue DE apicofuscans quadrata apicofuscans ME A apiconigroscutellata nigrotibialis nigrotibialis Cue A apicopicta tryoni assita Cue C aquila nigrotibialis aquila Cue D aquilonis tryoni tryoni Cue E arecae dorsalis arecae None B assita tryoni assita Cue D aterrima nigrotibialis nigrotibialis Cue D atra atramentata atra Cue F atrabifasciata nigella trivialis Cue D atramentata atramentata atramentata Cue, zingerone D atrifemur dorsalis dorsalis ME B atriliniellata distincta distincta Cue D atriscuta bryoniae froggatti ME D aurantiaca quadrata quadrata Cue DE avittata distincta distincta Cue C balagawii musae musae ME D bancroftii musae musae ME weak CE barringtoniae quadrata barringtoniae Isoeugenol E batemani nigella nigella ME E 90 Hancock & Drew: Review of subgenus Bactrocera Species Complex Group Lure Zone beckerae bryoniae bryoniae Cue C bellisi nigrotibialis nigrotibialis Cue C bhutaniae quadrata silvicola Cue AB biarcuata longicornis biarcuata ME D bidentata quadrata bidentata Isoeugenol E bifasciata alyxiae umbrosa Cue C bimaculata dorsalis pedestris Cue BC binhduongiae dorsalis dorsalis ME B bisianumu quadrata silvicola Cue D bitungiae dorsalis pedestris Cue C bivittata dorsalis dorsalis ME B blairiae quadrata commensurata ME B bogiae quadrata obfuscata Cue D breviaculeus quadrata quadrata Cue, zingerone DE brevistriata musae brevistriata Cue D bruneiae bryoniae froggatti ME B brunnea atramentata strigata None E brunneola quadrata silvicola Cue A bryoniae bryoniae bryoniae Cue, zingerone DE bubiae musae brevistriata Cue D bukaensis quadrata silvicola Cue D buloloensis tryoni notatagena None D caccabata nigrotibialis nigrotibialis Cue, zingerone D cacuminata dorsalis dorsalis ME E caledoniensis frauenfeldi obliqua Cue F caliginosa tryoni notatagena None D carambolae dorsalis dorsalis ME B carbonaria atramentata atra Cue D careofascia tryoni assita Cue C caryeae dorsalis caryeae ME A centraliae quadrata barringtoniae ME D ceylanica quadrata silvicola Cue A cibodasae dorsalis pedestris Cue B cinnamea quadrata silvicola Cue D circamusae musae brevistriata Cue D citima dorsalis pyrifoliae Cue B cognata dorsalis pedestris Unknown B collita dorsalis dorsalis ME B commensurata quadrata commensurata ME BC commina tryoni notatagena None D confluens longicornis curvifer ME D congener quadrata congener Cue D consectorata dorsalis pedestris Cue D contermina musae musae ME D contigua musae musae ME CD correcta zonata zonata ME AB costalis bryoniae bryoniae Cue B curreyi distincta distincta Cue D curtivitta quadrata silvicola Unknown B curvifer longicornis curvifer ME CD curvipennis tryoni curvipennis Isoeugenol, cue F curvosterna bryoniae bryoniae Cue C 91 RAFFLES BULLETIN OF ZOOLOGY 2025 Species Complex Group Lure Zone dapsiles dorsalis dorsalis ME D daruensis tryoni notatagena ME accidental? DE decumana distincta distincta Cue D decurtans quadrata bidentata ME DE denigrata longicornis longicornis Cue D diallagma diallagma diallagma ME D digressa quadrata digressa Cue, zingerone A diospyri nigrotibialis diospyri ME weak? E dispar quadrata dispar Unknown C distincta distincta distincta Cue F divenderi nigrotibialis nigrotibialis Cue A dongnaiae dorsalis pedestris Cue B dorsalis dorsalis dorsalis ME B dorsaloides dorsalis pedestris Cue B dyscrita quadrata quadrata Cue D dysoxyli nigrotibialis nigrotibialis Unknown D ebenea longicornis amplexiseta ME F ellenriederae dorsalis pedestris Cue B elongata musae elongata Cue C endiandrae dorsalis dorsalis ME DE enochra atramentata kirki Cue D epicharis nigrotibialis aquila Cue D erubescentis quadrata silvicola Cue DE ettinabhuja quadrata commensurata ME A eurycosta bryoniae bryoniae Cue B expandosa bryoniae bryoniae Cue D exspoliata nigrotibialis aquila Unknown D facialis passiflorae obscura Cue F fagraea quadrata quadrata Cue weak E fastigata diallagma laticosta Cue A fergussoniensis distincta distincta Unknown D fernandoi dorsalis pedestris Cue A finitima musae musae Unknown D flavipennis quadrata dispar Cue C flavoscutellata dorsalis pedestris Cue B flavosterna dorsalis pedestris Cue C floresiae dorsalis dorsalis ME B frauenfeldi frauenfeldi frauenfeldi Cue, zingerone CDE froggatti bryoniae froggatti ME D fuliginus dorsalis pedestris Cue DE fulvicauda distincta fulvicauda ME D fulvifemur dorsalis pedestris Cue B fumica longicornis amplexiseta ME D furcata zonata furcata Unknown A furfurosa quadrata silvicola Cue D furvescens furvescens furvescens Cue D furvilineata distincta distincta Cue D fuscalata longicornis amplexiseta ME D fuscitibia dorsalis pedestris Cue, zingerone BC fuscoformosa tryoni assita Cue C fuscohumeralis distincta distincta Unknown D fuscolobata tryoni assita Cue C 92 Hancock & Drew: Review of subgenus Bactrocera Species Complex Group Lure Zone fuscoptera longicornis amplexiseta ME C gabensiae tryoni assita Cue D geminosimulata bryoniae bryoniae Cue D gombokensis dorsalis pedestris Cue B grandifasciata tryoni sylvania Unknown D grandistylus passiflorae samoae None F halfordiae quadrata barringtoniae Isoeugenol E halmaherae alyxiae halmaherae Cue C hantanae dorsalis pedestris Cue A hispidula atramentata strigata None E hollimgsworthi distincta angustifasciata Cue D holtmanni dorsalis pedestris Cue B humilis tryoni notatagena None E hyalina zonata venefica None known B hypomelaina atramentata atramentata Cue D illusioscutellaris nigella trivialis Cue, zingerone AB impunctata musae impunctata ME BC incompta tryoni incompta Cue C inconspicua distincta fulvicauda ME C inconstans musae brevistriata Cue D indecora indecora indecora Cue D indonesiae dorsalis dorsalis ME B infulata dorsalis dorsalis ME C invadens dorsalis caryeae ME A involuta dorsalis pedestris Cue C irvingiae dorsalis dorsalis Unknown B ismayi alyxiae reclinata ME D jaceobancroftii quadrata barringtoniae ME B jarvisi quadrata jarvisi Zingerone, cue weak DE kaiauiae nigrotibialis nigrotibialis Cue D kalimantanae dorsalis pedestris Cue B kanchanaburi dorsalis dorsalis ME B kandiensis dorsalis caryeae ME A kauiae quadrata quadrata Cue D kelaena tryoni notatagena ME accidental? D keravatiae quadrata barringtoniae ME D kinabalu dorsalis pedestris Cue B kirki atramentata kirki Cue F kohkongiae quadrata silvicola Cue B kokodiae furvescens furvescens Cue D kraussi quadrata barringtoniae Isoeugenol E kunvawaensis laticaudus phaea Cue D kyrdemkulai quadrata commensurata ME A labubulu tryoni assita Cue D laensis tryoni notatagena None D laithieuiae dorsalis pedestris Cue B lampabilis alyxiae reclinata ME D lata nigrotibialis nigrotibialis Cue B lateritaenia dorsalis pedestris Cue B laticaudus laticaudus laticaudus ME E laticosta diallagma laticosta Cue D latifrons musae latifrons Latilure ABC 93 RAFFLES BULLETIN OF ZOOLOGY 2025 Species Complex Group Lure Zone latilineata distincta distincta Unknown D latilineola dorsalis dorsalis ME B latissima bryoniae bryoniae Cue D limbifera bryoniae bryoniae Cue ABC linduensis bryoniae bryoniae Cue C lineata nigrotibialis nigrotibialis Cue D lombokensis dorsalis pedestris Cue BC longicornis longicornis longicornis Cue D luteola atramentata luteola None F maculigera distincta fulvicauda ME C makilingensis dorsalis pedestris Cue B malasaitiae longicornis biarcuata ME D malaysiensis dorsalis pedestris Cue B mamaliae quadrata silvicola Cue C manskii alyxiae recurrens Cue E manusiae distincta fulvicauda ME D mayi quadrata mayi ME E mediorufula tryoni mediorufula ME B megaspilus quadrata quadrata Cue C melanogaster laticaudus laticaudus ME D melanothoracica musae aberrans ME DE melanotus atramentata atra Cue F melas tryoni tryoni Cue E melastomatos dorsalis melastomatos Cue B memnonia nigrotibialis aquila Cue D mendosa tryoni notatagena None E meraiensis bryoniae froggatti ME D merapiensis dorsalis pedestris Cue B mimulus bryoniae froggatti ME D minuscula dorsalis dorsalis ME BC minuta passiflorae passiflorae Cue DF moluccensis quadrata moluccensis Cue BCD monostriata distincta distincta Cue D morobiensis distincta distincta Cue D morula atramentata atramentata Cue D mucronis passiflorae obscura Cue F muiri dorsalis dorsalis Unknown B murrayi quadrata murrayi Methyl isoeugenol E musae musae musae ME DE mutabilis tryoni notatagena Isoeugenol? E nanoarcuata bryoniae bryoniae Cue C nationigrotibialis nigrotibialis diospyri ME C naucleae quadrata barringtoniae ME D neoabdonigella quadrata silvicola Cue D neoaeroginosa quadrata barringtoniae ME D neoarecae dorsalis caryeae ME A neocognata dorsalis pedestris Cue BC neofulvicauda distincta distincta Cue C neohumeralis tryoni tryoni Cue, zingerone DE neonigrita laticaudus laticaudus ME D neonigrotibialis nigrotibialis nigrotibialis Cue A neopropinqua dorsalis propinqua Cue B 94 Hancock & Drew: Review of subgenus Bactrocera Species Complex Group Lure Zone neoritsemai bryoniae bryoniae Cue C nigella nigella nigella ME D nigrescens musae musae ME D nigrescentis alyxiae recurrens Cue D nigricula nigrotibialis aquila Cue D nigrifacia diallagma laticosta Cue AB nigrifemorata nigrotibialis nigrotibialis Cue AB nigrita musae nigrita ME accidental? B nigrofemoralis nigrotibialis nigrotibialis Cue A nigroscutata furvescens opacovitta Unknown D nigrotibialis nigrotibialis nigrotibialis Cue BC nigrovittata tryoni romigae Methyl isoeugenol DE niogreta quadrata dispar Zingerone C notatagena tryoni notatagena None E nothaphoebe diallagma nothaphoebe Unknown B obfuscata quadrata obfuscata Cue D oblineata distincta distincta Cue D obliqua frauenfeldi obliqua Isoeugenol weak? D obliquivenosa distincta fulvicauda ME D obscura passiflorae obscura Cue F obscurata nigrotibialis nigrotibialis Unknown B obscurivitta quadrata obscurivitta Cue C occipitalis dorsalis dorsalis ME B ochracea tryoni assita Cue D ochroma dorsalis dorsalis ME BC ochromarginis quadrata ochromarginis ME D ochrosiae alyxiae ochrosiae Cue D ochroventer quadrata quadrata Cue? C ohuiae quadrata obfuscata Cue D opacovitta furvescens opacovitta ME C opiliae dorsalis dorsalis ME E osbeckiae dorsalis melastomatos Cue B pallescentis musae nigrita Unknown A pallida quadrata mayi ME E papayae dorsalis dorsalis ME B paraarecae dorsalis caryeae ME AB parabancroftii musae brevistriata Cue D parabarringtoniae quadrata barringtoniae ME E paradiospyri nigrotibialis diospyri ME A paraendiandrae bryoniae paraendiandrae ME D parafrauenfeldi frauenfeldi frauenfeldi Cue E parafroggatti dorsalis dorsalis ME D paralatissima bryoniae bryoniae Cue C paralimbifera bryoniae bryoniae Cue C paramusae bryoniae bryoniae Cue D paranigrita distincta fulvicauda ME C paraochracea tryoni assita Cue D paraosbeckiae quadrata silvicola Cue A paraverbascifoliae dorsalis caryeae ME A parvula musae latifrons None known B passiflorae passiflorae passiflorae Cue F patula quadrata quadrata Cue B 95 RAFFLES BULLETIN OF ZOOLOGY 2025 Species Complex Group Lure Zone pedestris dorsalis pedestris Cue B pectoralis tryoni notatagena Unknown C peneallwoodi musae musae ME? C penebeckerae musae musae ME? C penecognata dorsalis pedestris Cue BC penecorrecta zonata zonata ME A penecostalis bryoniae bryoniae Cue C peneobscura passiflorae obscura Cue F penephaea distincta distincta Cue C peninsularis quadrata quadrata Cue DE pepisalae longicornis curvifer ME D perfusca atramentata atra None known F perigrapha nigrotibialis nigrotibialis Cue, zingerone A perkinsi quadrata silvicola Cue E pernigra nigrotibialis pernigra Cue BC phaea laticaudus phaea Cue D phaleriae atramentata strigata None E picea atramentata picea ME D pictipennis musae pictipennis ME B pisinna distincta distincta Cue D pometiae tryoni notatagena None D popondettiensis tryoni notatagena None D prabhakari musae latifrons None known A prabhui nigrotibialis nigrotibialis Unknown A profunda dorsalis pedestris Cue A prolixa musae musae ME D propedistincta distincta distincta Cue D propinqua dorsalis propinqua Cue BC pruniae musae pruniae Unknown B pseudobeckerae quadrata selenophora Cue C pseudocucurbitae indecora indecora Cue BC pseudodistincta distincta distincta Cue CD pseudoversicolor quadrata commensurata ME A psidii atramentata atramentata Cue F pulchra distincta distincta None E pusilla tryoni assita Cue B pyrifoliae dorsalis pyrifoliae Cue weak? B quadrata quadrata quadrata Cue DE quadrusetosa tryoni romigae Isoeugenol DF quasienochra quadrata obfuscata Cue D quasiinfulata dorsalis pedestris Cue B quasineonigrita laticaudus laticaudus ME C quasipropinqua dorsalis propinqua Unknown B quasisilvicola quadrata silvicola Cue D raiensis dorsalis dorsalis ME B ramuensis musae brevistriata Cue D ranganathi dorsalis dorsalis ME B raunsepnaensis distincta angustifasciata Cue D reclinata alyxiae reclinata ME D recurrens alyxiae recurrens Cue CD redunca alyxiae recurrens Cue DF repanda alyxiae alyxiae Cue D 96 Hancock & Drew: Review of subgenus Bactrocera Species Complex Group Lure Zone resima alyxiae recurrens Cue BD retrorsa longicornis biarcuata ME D rhabdota distincta distincta Cue D ritsemai bryoniae bryoniae Cue BC robertsi musae brevistriata Cue D robiginosa quadrata barringtoniae Unknown E romigae tryoni romigae ME DE rounaensis distincta angustifasciata Cue D rubigina quadrata silvicola Cue, zingerone AB rufescens quadrata sylvicola Cue E rufivitta musae brevistriata Cue D rufofuscula quadrata silvicola Cue, zingerone DE russeola quadrata quadrata Cue E rutengiae diallagma diallagma ME C rutila quadrata moluccensis Cue D rutilana quadrata obfuscata Cue D samoae passiflorae samoae None F sapaensis dorsalis pedestris Cue B saramandiae furvescens furvescens Cue D sari quadrata silvicola Cue D seguyi longicornis curvifer ME D selenophora quadrata selenophora Cue A sembaliensis dorsalis pedestris Cue BC setinervis atramentata atra None known F silvicola quadrata silvicola Cue, zingerone E simulata bryoniae bryoniae Cue DF speculifer longicornis curvifer ME D speewahensis tryoni speewahensis Zingerone E strigata atramentata strigata None E sulawesiae dorsalis dorsalis ME C suliae quadrata commensurata ME C sumbawaensis dorsalis pedestris Cue BC sylvania tryoni sylvania ME D syzygii dorsalis pedestris Zingerone ABC tapahensis tryoni mediorufula ME B tenuifascia quadrata mayi ME E terminaliae nigrotibialis nigrotibialis Cue D ternatiae longicornis curvifer ME C thailandica dorsalis pyrifoliae Cue AB thistletoni nigrotibialis nigrotibialis Cue CD tikelingiae distincta distincta Cue D tinomiscii musae brevistriata Cue D torresiae distincta distincta Cue D tortuosa distincta distincta Unknown D toxopeusi nigrotibialis aquila Unknown D trifaria atramentata kirki Cue D trifasciata quadrata dispar Cue C trilineola frauenfeldi frauenfeldi Cue F trivialis nigella trivialis Cue, zingerone D trivirgulata tryoni notatagena None D truncata distincta distincta Cue C tryoni tryoni tryoni Cue, zingerone E 97 RAFFLES BULLETIN OF ZOOLOGY 2025 Species Complex Group Lure Zone tsatsiai distincta distincta Zingerone D tuberculata zonata zonata ME AB turneri quadrata silvicola Cue D umbrosa alyxiae umbrosa ME BCDF unifasciata distincta distincta Cue D unimacula dorsalis dorsalis ME B unilineata distincta angustifasciata Cue D unistriata distincta fulvicauda ME D unitaeniola distincta distincta Cue D usitata dorsalis pedestris Cue BC ustulata tryoni assita Cue D uvariae musae brevistriata Cue D vargasi frauenfeldi obliqua Zingerone D venefica zonata venefica None known B verbascifoliae dorsalis caryeae ME AB versicolor zonata versicolor ME A vishnu dorsalis pedestris Cue A vulgaris indecora indecora Cue D waaiae quadrata barringtoniae ME C waidoriae bryoniae paraendiandrae ME D wallacei nigella trivialis Cue C wuzhishana nigrotibialis diospyri ME ABC yayamiae frauenfeldi obliqua Cue D yorkensis quadrata ochromarginis M-isoeugenol, ME E youngi dorsalis pedestris Cue B zonata zonata zonata ME AB Table 2. List of species complexes in subgenus Bactrocera, with the total number of included species and the number of species occurring in each of the biogeographic zones in the Indo-Australian region (A = Indian Subcontinent; B = Southeast Asia; C = Wallacea; D = New Guinea & Solomon Islands; E = Australia; F = South Pacific). Complex Species A B C D E F longicornis 17 – – 3 13 1 1 alyxiae 16 – 2 3 13 2 1 distincta 40 – – 8 29 1 2 frauenfeldi 8 – 1 2 5 2 2 atramentata 18 – – – 7 4 7 laticaudus 6 – – 1 4 1 – tryoni 39 – 3 6 22 11 2 nigrotibialis 30 9 6 6 14 1 – passiflorae 8 – – – 1 – 8 diallagma 6 2 2 1 2 – – indecora 4 – 1 1 3 – – nigella 6 1 1 1 3 1 – bryoniae 25 1 5 10 11 1 1 musae 32 3 6 7 18 5 – dorsalis 83 13 58 18 6 4 – furvescens 5 – – 1 4 – – quadrata 86 12 11 16 38 26 – zonata 10 8 6 – – – – Totals 439 49 102 84 193 60 24 104 Hancock & Drew: Review of subgenus Bactrocera Further, the use of the COI gene by Schutze et al. (2015a, b) and Hee et al. (2015b) to argue conspecificity has been proven inadequate for accurate description of species (Zamani et al., 2022; Doorenweerd et al., 2024). Prior to their description by Drew & Hancock (1994), White & Elson-Harris (1992) referred to papayae as species B and philippinensis as species C and those two species were noted as having an aculeus length / wing cell dm ratio of 0.77–0.93; in comparison, dorsalis [at that time also including populations later described as B. invadens] and species A (= carambolae] only had an aculeus length / cell dm ratio of 0.57–0.75. In view of such data, coupled with morphological differences, the withdrawal of B. invadens and B. papayae from synonymy with B. dorsalis by Drew & Romig (2016) was justified and is further corroborated by Drew & Romig (2022) and Drew & Hancock (2022), especially in the light of a more recent understanding of the field host plant-based reproductive biology and data on the structure and function of the male genitalia. Drew & Romig (2013) had recognised B. philippinensis as a synonym of B. papayae after several years of intensive study of the dorsalis complex populations from the Philippines, across Borneo and into Java. Schutze et al. (2015a, b) then erroneously placed B. papayae, B. philippinensis and B. invadens as synonyms of B. dorsalis. Hee et al. (2015a, b) implied that this synonymy would significantly facilitate global trade, yet ten years later this has not eventuated. Indeed, the expanded host list has made matters worse (see Drew & Hancock, 2022). If the WTO, IPPC, and IAEA wish to aggregate several dorsalis-type species under one name for possible trade implications, then that is a separate matter entirely from the definition of species based on sound and consistent scientific principles. Due to the known or potential unreliability of COI and other gene fragments, an overreliance on molecular data and a single ‘best’ tree selected from a multitude of alternatives, in many cases molecular studies have created more problems than they have resolved. This is especially the case in the large and complex subgenus Bactrocera, where such studies not only fail to agree with each other but also are at odds with morphology, biology and biogeography. Three allopatric species of major agricultural and biosecurity concern, B. dorsalis, B. papayae, and B. invadens, allegedly were synonymised to facilitate international trade. However, such trade is better served by the recognition that these three species appear to be unable to establish in each other’s territories (suggested by limited introgression and maintenance of allopatry), rather than by a false synonymy that has caused nothing but confusion and continues to compromise many recent studies (e.g., Wang et al., 2024). There is no guarantee that laboratory-reared colonies of one species will interact successfully with the others in field-based SIT programs and any comparisons between B. papayae and the IAEA Saraburi colony of ‘B. dorsalis’ will have been between papayae and papayae. Morphology-based systematics, used judiciously, still provides the best evidence of a species’ identity and phylogenetic relationships. Proper taxonomic training is essential to this process. ACKNOWLEDGEMENTS We thank Michelle Baker for preparing Fig. 1 and Ian White and Allen Norrbom for constructive comments that improved the initial manuscript. LITERATURE CITED Abhishek V, David KJ & Pradeep S (2024) Two new species and a new record of Bactrocera Macquart (Diptera: Tephritidae: Dacinae: Dacini) from India with an updated key to species of subgenus Bactrocera. Zootaxa, 5486: 283–300. 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