Male. Labelled: “Vietnam: Thừa Thiên-Huế Province, Bạch Mã National Park, 16.2297N, 107.8494E, 6–8-x-2015, M. San Jose and D. Rubinoff, FF485, zingerone lure. Molecular voucher ms6192”. Deposited in UHIM. Paratypes: 5 males. Same data as holotype. All specimens pinned and one specimen is molecular voucher ms6255. Three of the paratypes are deposited at UHIM, one at WFBM and one at VNMN.

Bactrocera ernesti is similar to other members of the subgenus Tetradacus in having an elongate oval abdomen with a petiolate base [oval in most Bactrocera] with separate terga [tightly joined in Dacus], and a slight concavity of sternum V and short surstylus lobe in the males. It is most similar to B. minax and B. brachycera, but differs from both in lacking a lateral yellow band connecting the postpronotal lobes to the notopleural suture, the absence of medial postsutural vitta, the anteriorly convergent lateral postsutural vittae, the lightly infuscate wing, and absence of distinct costal band, and the black bands on every abdominal segment.

B. ernesti sp. n. was referred to as Bactrocera species 73, represented by the holotype, in the seven-gene phylogeny presented in San Jose et al. (2018). Based on the sampling therein, its closest relative is B. (Tetradacus) tsuneonis (Miyake). The closest relative we could identify based on COI alone is the Australian species B. visenda, at a minimum intraspecific pairwise genetic distance of 13.52 % [14.89 % in COI5P and 12.27 % in COI3P] (Figure 3).

Figure 3. 

Maximum likelihood tree based on COI sequences of B. ernesti sp. n. and several of its genetically closest neighbors, which include Bactrocera, Zeugodacus and Dacus species. Bootstrap branch supports shown for intraspecific relationships. Abbreviations: HT holotype.

Head (Figure 4A). Vertical length 1.82 ± 0.04 (SE) (1.75–1.85) mm. Frons, of even width, length 1.16 ± 0.05 (1.10–1.26) times breadth; uniformly fulvous; anteromedial hump and frons covered by short red–brown hairs; orbital and frontal setae, when present, red–brown: orbital setae absent or at most very weak and frontal setae very weak and may be restricted to superior pair; lunule fuscous. Ocellar triangle black. Vertex fulvous. Face fulvous with elongate black spot in lower half of each antennal furrow, connected by a faint narrow fuscous band across mid height of face; length 0.69 ± 0.02 (0.65–0.70) mm. Genae fulvous, with fuscous sub–ocular spot; one large and numerous smaller red–brown setae present. Occiput fulvous with fuscous markings laterally on its lower half; occipital row with 9–13 dark setae and an irregular inner row of finer setae. Antennae with segments 1 (scape) and 2 (pedicel) fulvous, segment 3 (first flagellomere) fulvous with pale fuscous on outer surface; a strong red–brown dorsal seta on segment 2; arista black (fulvous basally); length of segments: 0.19 ± 0.01 (0.18–0.20) mm; 0.26 ± 0.02 (0.23–0.28) mm; 0.76 ± 0.04 (0.70–0.83) mm.

Figure 4. 

Bactrocera (Tetradacus) ernesti. A head B head and scutum C abdomen D wing E lateral view.

Thorax (Figure 4B, E). Scutum red–brown except a broad light fuscous lanceolate pattern on its posterior third, anteriorly prolonged into three very narrow lines reaching anterior margin, narrow elongate dark fuscous bands adjacent to inner margins of lateral postsutural vittae, broad lateral dark fuscous markings behind postpronotal lobes and two short and narrow parallel dark fuscous bands between postpronotal lobes. Pleural areas dark fuscous except red–brown below postpronotal lobes, anterior half of anepisternum and posterior portion of katepisternum. Yellow markings as follows: postpronotal lobes; notopleura; medium sized and parallel-sided mesopleural (anepisternal) stripe, reaching anterior margin of notopleuron, continuing to katepisternum as a transverse spot, anterior margin straight; entire katatergite except red–brown narrowly along posterior margin; lower quarter to half of anatergite (remainder dark fuscous and red-brown on posterior margin of lower quarter to half); two moderately broad parallel sided lateral postsutural vittae ending shortly before intra-alar setae and curved slightly inwards along notopleural suture. Postnotum medially red–brown and laterally black. Scutellum yellow except for narrow dark fuscous basal band. Setae: 2 scutellar; 1 intra-alar; 1 posterior supra-alar; 1 mesopleural; 4 notopleural; 4 or 6 scapular (often a second pair of median scapular, just behind each bristle); anterior supra-alar and prescutellar bristles absent; all setae well developed and red–brown.

Legs (Figure 4E). Fore coxae yellow with outer face dark fuscous; fore trochanters and mid coxae and trochanters yellow; hind coxae and trochanters dark fuscous. Femora yellow with broadly fuscous outer and inner surfaces. Fore and mid tibiae fulvous with dark fuscous on outer face of fore tibia and around base of mid tibia; hind tibiae dark fuscous. Tarsi fulvous with dark fuscous fore tarsomeres 2–5 and ventral face of fore basitarsus. Mid-tibiae each with an apical black spur.

Wings (Figure 4D). Length 7.56 ± 0.21 (7.22–7.78) mm; basal costal (bc) cell infuscate and costal (c) cells mostly colorless except at apex; microtrichia along costal margin of cell bc and along costal margin and outer corner of cell c; remainder of wings colorless except fuscous subcostal cell, and lightly infuscate membrane between R₁ and R₄₊₅; supernumerary lobe weakly developed.

Abdomen (Fig. 4C, E). Elongate oval and petiolate; terga free; pecten present on tergum III; posterior lobe of surstylus short; abdominal sternum V with a slight concavity on posterior margin. Tergum I as long as wide and tergum II and sterna I and II longer than wide. Tergum I medially fuscous with a faint narrow fuscous medial longitudinal band and laterally black. Tergum II yellow with a large inverted V-shaped medial marking. Tergum III black with large apical triangular yellow marking. Tergum IV black with apical fulvous band with a medial expansion; tergum V fulvous with a narrow basal black band expanded laterally to half the tergum length. Abdominal sterna dark, except yellow sternum II.

This species is named after Ernest James Harris (1928–2018), in honor of his long career working as a fruit fly ecologist for the USDA (1962–2006). Some of Dr. Harris’s important contributions include the field implementation of the first eradication program against invasive fruit flies in the Mariana Islands, the establishment of Mediterranean fruit fly suppression programs in North Africa and Chile and studies on its ecology and SIT in Hawaii, as a prelude to the initiation of the ongoing SIT program to prevent its establishment in California, and the development of mass rearing techniques of the fruit fly parasitoid Fopius arisanus (Sonan). He published over 120 scientific papers and was honored with distinctions by the State of Arkansas Black Hall of Fame (1999), the NAACP Hawaii Chapter (2012), the Alpha Phi Alpha Fraternity National award (2013), the US Congressional Gold Medal (2016), the USDA-ARS Hall of Fame (2017), and as CTAHR Outstanding Alumnus (2017). His emergence as African American from a modest cotton farming family in Arkansas to an internationally respected prominent scientist, through hard work and his love for his research, makes Ernie a true role model for the senior author of this paper. Biographic sketches of Dr. Harris were published by Riddick et al. (2015) and Leblanc and Vargas (2018, in press).

Bactrocera ernesti keys to couplet 2, page 314, in Drew and Romig (2016), where it can be differentiated from B. minax and B. brachycera based on the characters mentioned in the differential diagnosis.

Male. Labelled: “Vietnam: Thừa Thiên–Huế Province, Bạch Mã National Park, 16.2098N, 107.8632E, 6-12-x-2015, M. San Jose and D. Rubinoff, FF494, zingerone trap.” ms6195 Deposited in UHIM. Paratypes: 3 males. Vietnam, Lâm Đồng Province, Cát Tiên National Park, 16–18-x-2015, at the following sites, identified by their geographical coordinates: 11.4485N, 107.4416E, (1 pinned, molecular voucher ms6105), 11.4447N, 107.4372E, (2 pinned, one molecular voucher ms6239). All specimens collected by Michael San Jose and Dan Rubinoff in zingerone baited traps. One paratype deposited at UHIM, one at WFBM and one at VNMN.

Bactrocera connecta shares morphological features common to all members of Asiadacus (absence of prescutellar and anterior supra-alar setae, scutellum with one pair of setae, and males with long posterior lobe on surstylus, slight concavity of sternum V and pecten of cilia present on the abdominal tergum III). It is distinguished from other members of Asiadacus by the elongate facial spots along antennal furrows and the small to extensive fuscous transverse band on face (Fig. 6A, B), the broad, apically expanded costal band (Figure 6H, I), and the absence of a medial longitudinal band and presence of extensive lateral black markings on the abdomen (Figure 6D–G).

Bactrocera connecta was referred to as Bactrocera species 68, sister to B. (Apodacus) visenda (Hardy), in the seven-gene phylogeny presented in San Jose et al. (2018). The nearest neighbor we could identify based on COI sequence data is Zeugodacus melanofacies, at 15.34 % minimum pairwise intraspecific genetic distance [16.18 % in COI5P and 14.47 % in COI3P] (Figure 5).

Figure 5. 

Maximum likelihood tree based on COI sequences of B. connecta sp. n. and several of its genetically closest neighbors, which include both Bactrocera and Zeugodacus species. Bootstrap branch supports shown for intraspecific relationships. HT = holotype.

Head (Figure 6A, B). Vertical length 1.64 ± 0.23 (SE) (1.45–1.88) mm. Frons, of even width, length 1.13 ± 0.07 (1.07–1.22) times as long as broad; fulvous with fuscous around orbital setae and on anteromedial hump; latter covered by short red–brown hairs; orbital setae black: one pair of superior and two pairs of inferior fronto-orbital setae present; lunule yellow. Ocellar triangle black. Vertex fuscous. Face fulvous with elongate black spots in each antennal furrow and a fuscous band across lower half of face varying from a small medial spot (Figure 6A) to an entire band across face (Figure 6B); length 0.63 ± 0.09 (0.53–0.75) mm. Genae fulvous, with dark fuscous sub-ocular spot; black seta present. Occiput light to dark fuscous (fulvous laterally in one specimen) and yellow along eye margins; occipital row with 9–11 dark setae. Antennae with segment 1 (scape) fulvous, segments 2 (pedicel) fulvous and fuscous on outer surface, and segment 3 (first flagellomere) fuscous and narrowly fulvous on inner surface; strong red–brown dorsal seta on segment 2; arista black (fulvous basally); length of segments: 0.23 ± 0.02 (0.20–0.25) mm; 0.34 ± 0.02 (0.33–0.38) mm; 0.76 ± 0.07 (0.70–0.83) mm.

Figure 6. 

Bactrocera (Asiadacus) connecta. A, B head C head and scutum D, E, F, G abdomen variation H, I wing.

Thorax (Figs 6C, 7). Scutum black except sometimes with red–brown as limited markings behind postpronotal lobes and anterior to lateral postsutural vittae and area below postsutural vittae. Pleural areas black except usually red–brown anterior portion of anepisternum. Yellow markings as follows: postpronotal lobes and notopleura (notopleural callus), usually connected by a lateral longitudinal band; medium sized mesopleural (anepisternal) stripe, reaching midway between anterior margin of notopleura and anterior notopleural seta dorsally, continuing to katepisternum as a transverse spot, anterior margin convex; anatergite (posterior apex black); anterior 75 % of katatergite (remainder black); moderately broad medial postsutural vitta beginning at level of or slightly anterior of prescutellar setae and ending at or slightly beyond the level of notopleural suture; two moderately broad parallel sided lateral postsutural vittae ending at or shortly before intra-alar setae and curved inwards along notopleural suture. Postnotum black with apical median red–brown spot. Scutellum yellow except for narrow black basal band. Setae (number of pairs): 1 scutellar; prescutellar absent; 1 intra-alar; 1 posterior supra-alar; anterior supra-alar absent; 1 mesopleural; 2 notopleural; 4 scapular; all setae well developed and red–brown.

Figure 7. 

Bactrocera (Asiadacus) connecta: lateral view.

Legs (Figure 7). Femora fulvous, except for dark fuscous ventral faces of basal three quarters of fore femur, apical three quarters of mid femur and apical two–fifths of hind femur, and yellow basal quarter of mid femur and basal three–fifths of hind femur; tibiae dark fuscous, with or without ventral faces narrowly fulvous; mid-tibiae each with an apical black spur; tarsi fulvous.

Wings (Figure 6H, I). Length 5.75 ± 0.67 (5.11–6.67) mm; basal costal (bc) and costal (c) cells fuscous and both covered with microtrichia; remainder of wings colorless except broad fuscous costal band nearly confluent with R₄₊₅ and broadly expanded apically; anal streak absent; dense aggregation of microtrichia around A₁ + CuA₂; supernumerary lobe of medium development.

Abdomen (Figs 6D–G, 7). Elongate oval; terga free; pecten present on tergum III; posterior lobe of surstylus long; abdominal sternum V with a slight concavity on posterior margin. Tergum I and sterna I and II wider than long. Tergum I black. Tergum II yellow with sub-basal or basal broad medial transverse black band that may be extended to cover entire basal half of tergum. Terga III–V black with fulvous areas as small markings on apex of tergum III and narrow median area on tergum IV (or restricted to small apical median marking), continued on base of tergum V and expanded around ceromata. A pair of dark fuscous ceromata (shining spots) on tergum V. Abdominal sterna black, except fulvous sternite II.

The species name is an adjective that refers to the longitudinal yellow band connecting the postpronotal lobes and notopleura.

We have tentatively assigned B. connecta to Asiadacus based on the following combination of characters: absence of prescutellar and anterior supra-alar setae, scutellum with one pair of setae, a long posterior lobe of the male surstylus, slight concavity of sternum V and pecten of cilia present on the abdomen (Drew 1989, Drew and Romig 2016). All known species of Asiadacus were transferred to Zeugodacus, based on the surstylus and sternum V concavity characters which made them part of the Zeugodacus group of subgenera (de Meyer et al. 2015). Only more recently was one species, B. (Asiadacus) apicalis, genetically confirmed to belong to Zeugodacus (San Jose et al. 2018). In that same molecular phylogeny, however, B. connecta is only distantly related to B. apicalis and is placed in the basal grade of subgenera in Bactrocera. Recently, Hancock and Drew (2017, 2018) transferred all but two species of Asiadacus to the subgenera Parasinodacus, Sinodacus and Zeugodacus, leaving in Asiadacus the two species with a large, oval apical wing spot that reaches but does not cross vein M. Because B. connecta falls within Bactrocera in the recently published phylogeny (San Jose et al. 2018), it cannot be assigned to any of these three subgenera. Moreover, in B. connecta the lateral postsutural vittae are not extended as yellow markings beyond the notopleural suture, a character shared by the Zeugodacus group of subgenera and genus Dacus, referred to as “small notopleural xanthines” by White (2006) and “lateral yellow triangles that reach the notopleural lobes” by Hancock and Drew (2018). Because B. connecta cannot be fitted in existing subgenera and to avoid creating a new subgenus, we arbitrarily assign this species to the subgenus Asiadacus sensu Drew and Romig (2016), until the subgeneric relationships within Bactrocera are more clearly elucidated. It cannot be keyed beyond couplet 4, page 48, in Drew and Romig (2016), where it can be singled out by the costal band on the wing overlapping R₂₊₃ and expanding apically (Figs 6H, I).

Male. Labelled: “Vietnam: Thừa Thiên–Huế Province, Cát Tiên NP, 11.4480N, 107.3826E, 14-18-x-2015, M. San Jose and D. Rubinoff, FF540, zingerone lure, molecular voucher ms6176”. Deposited in UHIM. Paratypes: 48 males. Vietnam, Thừa Thiên-Huế Province, Bạch Mã National Park, 6–8-x-2015, at the following sites, identified by their geographical coordinates: 16.2279N, 107.8557E, (1 in ethanol, molecular voucher ms6191), 16.1943N, 107.8490E, (2 in ethanol, including molecular voucher ms6194), 16.2006N, 107.8481E, (1 pinned). Vietnam, Lâm Đồng Province, Cát Tiên National Park, 14–18-x-2015, at the following sites, identified by their geographical coordinates: 11.4920N, 107.3855E, (7 pinned), 11.4867N, 107.3834E, (1 in ethanol, molecular voucher ms6175), 11.4764N, 107.3817E, (2 in ethanol, including molecular voucher ms6249), 11.4715N, 107.3809E, (2 in ethanol), 11.4480N, 107.3826E, (1 in ethanol, molecular voucher ms6176), 11.4436N, 107.3925E, (8 in ethanol), 11.4412N, 107.4026E, (3 pinned), 11.4419N, 107.4080E, (1 in ethanol, molecular voucher ms6177), 11.4394N, 107.4241E, (5 pinned and 2 in ethanol, including molecular vouchers ms6094 and ms6095, in ethanol), 11.4531N, 107.3557E, (1 in ethanol), 11.4486N, 107.3584E, (1 in ethanol), 11.4564N, 107.3686E, (1 in ethanol), 11.4381N, 107.4267E, (1 in ethanol), 11.4398N, 107.4290E, (1 in ethanol), 11.4539N, 107.4430E, (4 pinned), 11.4485N, 107.4416E, (3 pinned); 11.4472N, 107.4392E, (1 pinned). All specimens collected by Michael San Jose and Daniel Rubinoff in zingerone baited traps. All paratypes are deposited at UHIM, except five at WFBM and three at VNMN.

Bactrocera clarifemur (Figure 9A–G) is genetically and morphologically closely related to B. pendleburyi (Perkins) (Figure 10A–E). Both share the defining characters of subgenus Parazeugodacus (two pairs of scutellar setae; male with lateral pecten on tergum III (though present or absent in different species of Parazeugodacus according to Hancock and Drew 2015), a slight concavity on posterior margin of abdominal sternum V, and posterior lobe of male surstylus short), as well as the absence of medial postsutural vitta and the entirely yellow scutellum. Bactrocera clarifemur differs from B. pendleburyi in that all femora are entirely fulvous (Figure 9G), whereas apices of all femora in B. pendleburyi are apically broadly dark (Figure 10E).

This species was referred to as Bactrocera sp 70 and is a close relative yet distinctly monophyletic sister to B. (Parazeugodacus) pendleburyi, in the San Jose et al. (2018) seven-gene phylogeny. Based on COI sequence data the two species are in two monophyletic clusters, with one exception. We found an aberrant COI haplotype from specimen (ms6095) that groups with B. pendleburyi in ML analysis, though at a minimum of 2.66 % pairwise genetic distance from any of the other specimens (Figure 8). Morphologically, specimen ms6095 fits with B. clarifemur (Figure 9A–G). We sequenced an additional nuclear gene to confirm the genetic distinction. Based on EF1a sequences, both species, with specimens from the same localities, were separated in monophyletic groups with a minimum pairwise distance of 1.35 %. A haplotype network of COI sequences further shows that specimen ms6095 is relatively distantly related to both and may eventually be found to represent a cryptic species. The nearest neighbor to both B. pendleburyi and B. clarifemur is B. abbreviata at 6.91 % minimum intraspecific pairwise distance in COI.

Figure 8. 

Maximum likelihood trees and haplotype network based on COI (A) and EF1a (B) sequences of B. clarifemur sp. n. and several of its genetically closest neighbors. C displays a TCS haplotype network, with notches on the connections to indicate mutations, based on COI data of B. clarifemur and B. pendleburyi and shows that specimen ms6095 is relatively distantly related from all others. Bootstrap branch supports shown for intraspecific relationships. Abbreviation: HT holotype.

Figure 9. 

Bactrocera (Hemigymnodacus) clarifemur. A head B head and scutum C, D, E abdomen variation F wing G lateral view.

Figure 10. 

Bactrocera (Hemigymnodacus) pendleburyi. A head B head and scutum C abdomen D wing E lateral view.

Head (Figure 9A). Vertical length 1.40 ± 0.13 (SE) (1.20–1.55) mm. Frons, of even width, length 1.32 ± 0.06 (1.22–1.43) times as long as broad; fulvous with usually fuscous spot around orbital setae and on anteromedial hump; latter covered by short red–brown hairs; orbital setae dark fuscous: one pair of superior and two pairs of inferior fronto-orbital setae present; lunule yellow. Ocellar triangle black. Vertex fuscous. Face fulvous with medium sized oval black spots in each antennal furrow; length 0.47 ± 0.04 (0.40–0.53) mm. Genae fulvous, with fuscous sub-ocular spot; red–brown seta present. Occiput dark fuscous and yellow along eye margins; occipital row with 5–7 dark setae. Antennae with segments 1 (scape) and 2 (pedicel) fulvous and segment 3 (first flagellomere) fulvous with pale fuscous on outer surface; a strong red–brown dorsal seta on segment 2; arista black (fulvous basally); length of segments: 0.21 ± 0.02 (0.18–0.23) mm; 0.24 ± 0.01 (0.23–0.25) mm; 0.62 ± 0.05 (0.53–0.68) mm.

Thorax (Figure 9B, G). Scutum black and frequently red–brown below lateral postsutural vittae. Pleural areas black and usually narrowly red–brown on anterior corner and frequently posterior corner of proepisternum anterior to postpronotal lobes. Yellow markings as follows: postpronotal lobes; notopleura (notopleural callus); medium sized mesopleural (anepisternal) stripe, reaching anterior margin of notopleura dorsally, continuing to katepisternum as a transverse spot, anterior margin slightly convex; anatergite (posterior apex black); anterior 60 % of katatergite (remainder black); two moderately broad and short postsutural vittae, tapering posteriorly and ending a short distance behind anterior intra-alar setae. Postnotum black. Scutellum yellow except for narrow black basal band. Setae (number of pairs): 2 scutellar; 1 prescutellar; 1 intra-alar; 1 posterior supra-alar; 1 anterior supra-alar; 1 mesopleural; 1 notopleural; 4 scapular; all setae well developed and black.

Legs (Figure 9G). Femora entirely fulvous; fore and hind tibiae dark fuscous, mid tibia basally dark fuscous and apically becoming fulvous; mid-tibiae each with an apical black spur; tarsi fulvous.

Wings (Figure 9F). Length 4.68 ± 0.35 (4.22–5.00) mm; basal costal (bc) and costal (c) cells colorless; microtrichia in outer corner of cell c only; remainder of wings colorless except fuscous subcostal cell, very narrow light fuscous costal band confluent with R_2+3, not widened apically and ending just slightly past the extremity of R₄₊₅; anal streak absent; dense aggregation of microtrichia around A₁ + CuA₂; supernumerary lobe of medium development.

Abdomen (Figure 9C–E). Oval; terga free; pecten present on tergum III; posterior lobe of surstylus short; abdominal sternum V with a slight concavity on posterior margin. Tergum I and sterna I and II wider than long. Tergum I dark fuscous with a narrow transverse fulvous band across posterior margin but not reaching lateral margins; tergum II dark fuscous with a transverse posterior fulvous band which just reaches the narrow black posterolateral corners; tergum III dark fuscous; terga IV–V orange–brown with broad medial dark fuscous band reaching the apex of abdomen and broad lateral dark fuscous bands narrowed in apical half of tergum V. Orange–brown markings and medial band frequently extended to apical portion of tergum III. Ceromata (shining spots) on tergum V orange–brown and indistinct. Abdominal sterna dark, except for fuscous sternite II.

The name is an adjective that refers to the absence of dark markings on the femora.

The characters distinguishing B. clarifemur and B. pendleburyi were noted as variation of B. pendleburyi by Drew and Romig (2013), genetic evidence hereby confirms that they indicate distinct species.

Male. Labelled: “Vietnam: Lâm Đồng Province, Cát Tiên National Park, Ranger station Road, 11.4485N, 107.4416E, 16–18-x-2015, M. San Jose and D. Rubinoff, FF581, Zingerone trap. Molecular voucher ms6092”. Deposited in UHIM. Paratypes: 3 males. Vietnam, Lâm Đồng Province, Cát Tiên National Park, 16–18-x-2015, at the following sites, identified by their geographical coordinates: 11.4539N, 107.4430E, (1 pinned, molecular voucher ms6093), 11.4485N, 107.4416E, (1 pinned, molecular voucher ms6236), 11.4472N, 107.4392E, (1 in ethanol). All specimens collected by Michael San Jose and Daniel Rubinoff, in zingerone-baited traps. One paratype deposited in UHIM, one in WFBM and one in VNMN.

Bactrocera adamantea belongs to the (polyphagous) B. dorsalis complex of notoriously difficult to identify species (Leblanc et al. 2015c), defined by having a mostly dark scutum, a costal band that does not expand apically, and a black T shaped pattern on the abdomen. It can easily be differentiated from all congeners however by the yellow diamond shaped medial vitta on the scutum (Figure 12B).

Bactrocera adamantea is easily distinguished from all other Bactrocera using either section of COI. The closest species is B. nigrita, with a minimum intraspecific pairwise distance of 7.82 % in 1,496 bp of COI [8.66 % in COI5P, 7.18 % in COI3P] (Figure 11).

Figure 11. 

Maximum likelihood tree based on COI sequences of B. adamantea sp. n. and several of its genetically closest congeners. Bootstrap branch supports shown for intraspecific relationships. HT = holotype.

Figure 12. 

Bactrocera (Bactrocera) adamantea. A head B head and scutum C, D abdomen E wing F lateral view.

Head (Figure 12A). Vertical length 1.66 ± 0.09 (SE) (1.55–1.73) mm. Frons, of even width, length 1.38 ± 0.02 (1.36–1.40) times as long as broad; fulvous with light fuscous (may be absent) around orbital setae and on anteromedial hump; latter covered by short red–brown hairs; orbital setae dark fuscous: one pair of superior and two pairs of inferior fronto-orbital setae present; lunule yellow. Ocellar triangle black. Vertex light fuscous. Face fulvous with medium sized oval black spots in each antennal furrow; length 0.58 ± 0.04 (0.55–0.63) mm. Genae fulvous, with fuscous sub-ocular spot; red–brown seta present. Occiput fulvous to light fuscous and yellow along eye margins; occipital row with 6–8 dark setae. Antennae with segments 1 (scape) and 2 (pedicel) fulvous and segment 3 (first flagellomere) fulvous with fuscous outer surface; a strong fulvous dorsal seta on segment 2; arista black (fulvous basally); length of segments: 0.23 ± 0.01 (0.23–0.25) mm; 0.28 ± 0.03 (0.25–0.30) mm; 0.82 ± 0.01 (0.80–0.83) mm.

Thorax (Fig. 12B, F). Scutum black except red–brown as two parallel bands running from anterior margin to mid-length and as markings around notopleural suture and medial to postpronotal lobes. Pleural areas black except red–brown proepisternum and anterior portion of anepisternum below postpronotal lobes. Yellow markings as follows: postpronotal lobes; notopleura (notopleural callus); broad mesopleural (anepisternal) stripe, reaching level of anterior notopleural seta dorsally, continuing to katepisternum as a transverse spot, anterior margin slightly convex; anatergite (posterior apex black); anterior 80 % of katatergite (remainder black); two broad parallel-sided lateral postsutural vittae ending at level of intra-alar setae; large diamond-shaped medial marking on posterior end of scutum. Postnotum red–brown medially and black laterally. Scutellum yellow except for narrow black basal band. Setae (number of pairs): 1 scutellar; 1 prescutellar; 1 intra-alar; 1 posterior supra-alar; 1 anterior supra-alar; 1 mesopleural; 2 notopleural; 4 scapular; all setae well developed and red–brown.

Legs (Figure 12F). Femora fulvous, except for small fuscous subapical spot on outer surface of fore femur; tibiae dark fuscous; mid-tibiae each with an apical black spur; tarsi fulvous.

Wings (Figure 12E). Length 5.78 ± 0.11 (5.67–5.89) mm; basal costal (bc) and costal (c) cells colorless; microtrichia in outer corner of cell c only; remainder of wings colorless except fuscous subcostal cell, moderately broad fuscous costal band overlapping with R₂₊₃ and widening slightly as it crosses apex of R₂₊₃ to end between extremities of R₄₊₅ and M, a narrow fuscous anal streak ending at apex of posterior cubital cell; dense aggregation of microtrichia around A₁ + CuA₂; supernumerary lobe of medium development.

Abdomen (Figure 12C, D, F). Elongate oval; terga free; pecten present on tergum III; posterior lobe of surstylus short; abdominal sternum V with a deep concavity on posterior margin. Tergum I and sterna I and II wider than long. Tergum I fulvous with dark fuscous along base medially and black along lateral margins; tergum II fulvous except for a narrow transverse black band across anterior margin which extends to cover narrow lateral margins. Terga III–V orange–brown with a dark T-shaped pattern consisting of a broad transverse black band across anterior margin of tergum III expanding broadly over lateral margins and a broad medial longitudinal black band over all three terga, broad black lateral markings on tergum IV and narrowly black along lateral margins of tergum V. A pair of dark fuscous ceromata (shining spots) on tergum V. Abdominal sterna dark fuscous except for fulvous sternite II.

Etymology. The name adamantea is an adjective that refers to the diamond-shaped marking on the scutum, uniquely distinctive to this species.

Notes. It was referred to as Bactrocera species 69, sister to B. (Bactrocera) fuscitibia Drew and Hancock, in the phylogeny presented in San Jose et al. (2018). It keys as far as couplet 80, page 240, in Drew and Romig (2016), and differs from B. lateritaenia by the presence of the conspicuous diamond-shaped marking on the scutum (Figure 12B), which is also uniquely distinctive among Dacine fruit flies.