Drosophila (Sophophora) pandora, McEvey & Schiffer, 2015

McEvey, Shane & Schiffer, Michele, 2015, New Species in the Drosophila ananassae Subgroup from Northern Australia, New Guinea and the South Pacific (Diptera: Drosophilidae), with Historical Overview, Records of the Australian Museum 67 (5), pp. 129-161 : 138-142

Drosophila (Sophophora) pandora sp.nov.

Figs. 2–5, 14–21, 35–36, 38–41, 46, 54–59, 90, 96

Drosophila “papuensis-like” of Tomimura et al. (1993); Matsuda et al. (2009).

Drosophila nr ironensis, van Klinken, 1996, p. 247 ; van Klinken & Walter, 2001, pp. 168, 176 [north Queensland and Northern Territory].

Drosophila nr ananassae, van Klinken et al., 2002, p. 238 [Northern Territory].

Distinguishing features

Drosophila pandora sp.nov. can be distinguished by reference to the elongate, straight, basal extension of the anterior paramere, the “viking-helmet” shape of the convexity of the caudal margin of the novasternum, the overall quadrate and relatively wide profile of the combined aedeagus plus posterior parameres, the presence of an acuminate kink on the bend in the anterior paramere, and the configuration, and number, of teeth in sex combs on the male foreleg.

Description (♂)

Types. Holotype ♂, AMS K357344 , McEvey 31861, from type culture CAQ408: “QLD Lake Placid , alt. 17m | -16.870° 145.676° WGS84 | [culture established by M. Schiffer from rainforest, fruit-baited, iso- ♀] ix.2011, [specimens eclosed, generation F 41] ii.2014 | Schiffer, ex culture CAQ408 [culture maintained at the University of Melbourne]” . Paratypes (70♂♂ and 35♀♀, registered McEvey 31855–860 and McEvey 31862–960, all from type culture with same labeldata as holotype): 35♂♂ (K357339–343, K357345–374), and 21♀♀ (K357375–395) in Australian Museum , Sydney ; 5♂♂ and 2♀♀ in each of the following museums (with McEvey’s registration numbers): American Museum of Natural History , New York ( AMNH, McE31855, 31892– 895♂♂, McE31937– 938♀♀); Australian National Insect Collection , Canberra ( ANIC 29031663– 667♂♂ [McE31896– 900♂♂], ANIC 29031668– 669♀♀ [McE31939– 940♀♀]); Museum Zoologicum Bogoriense , Bogor ( MZB, McE31901– 905♂♂, McE31941– 942♀♀); Kunming Institute of Zoology , Chinese Academy of Science , Kunming ( KIZ, McE31906– 910♂♂, McE31943– 944♀♀); National Science Museum , Tokyo ( NSMT, McE31911– 915♂♂, McE31945 ♀, McE31956 ♀); Queensland Museum , Brisbane ( QMB, McE31946– 950♂♂, McE31957– 958♀♀); and United States National Museum , Washington, D.C. ( USNM, McE31951– 955♂♂, McE31959– 960♀♀). Living flies from type culture CAQ408 have been sent to the Drosophila Genetic Resource Center (Kyoto) and the Drosophila Species Stock Center (San Diego) .

Body length. 2.2 mm ♂.

Head. Arista with four rays above and two to three below, plus terminal fork. Orbital setae in ratio 2:1:2. Measurements of holotype ♂ —BL(McE) = 2.17 mm, BL(Z&T) = 1.63 mm, hw/fw(ov) = 1.91, hw/fw(iv) = 2.29, hw/fw(vt) = 1.99, hw/ fw(a. r.orb ) = 2.20, hw/fw(a.oc) = 2.04, hw/fw(ptl) = 2.61, fw(ov)/fl = 1.57, fw(iv)/fl = 1.31, fw(vt)/fl = 1.51, fw(a.oc)/fl = 1.48, fw(a. r.orb )/fl = 1.36, p. r.orb = 0.99, rc.orb = 0.48, proc. orb/a. r.orb = 2.05, oc/proc.orb = 1.08, pv/oc = 0.76, p. r.orb /iv = 0.73, orbito-index = 0.76, vt-index = 1.05, oc-gap/pv-gap = 0.44, o/j = 12.00, ch/o = 0.10, o/ow = 1.32, svb/vb = 0.65, flw = 1.51, avd = 0.93, adf = 1.96, arista free ends = 8–9.

Thorax. Acrostichal hairs in 8 rows in front of dorsocentral bristles, 4 rows between dorsocentrals. Ratio anterior/ posterior dorsocentrals 0.6. Preapical bristles on all tibiae; apicals on first and second tibiae. Sex comb of male foreleg ( Figs. 54–59 , Tables 1 and 2) in 2–6 transverse rows on tarsomere I of (from above down) 0–1, 0–2, 0–3, 0–6, 2–8, and 3–8 teeth; and 2–4 rows on tarsomere II of 0–2, 0–4, 1–6, and 2–8 teeth. Other thoracic measurements: bsc/asc = 0.90, sterno-index = 0.55, m/a.kepst = 0.60, p.kepst/pdc = 1.06, pdc/asc = 0.93, asc–bsc/asc–asc = 1.21, a–pdc/dc-gap = 0.39, adc/pdc = 0.61, fw(a.oc)/dc-gap = 1.32.

Wing. Hyaline, wing length c. 1.8 mm. L(Ax) = 1.76 mm, WL = 1.51 mm, L 1 = 1.33 mm, L(Ax)/WW = 2.31, WL/WW = 1.98, L 1 /WW = 1.75, C-index = 1.54, 4v-index = 2.56, 4c-index = 1.75, 5x-index = 2.29, M-index = 0.89, ac-index = 3.26, C3 fringe = 0.54.

Abdomen (Fig. 96). Tergites of both sexes with diffuse, dark, narrow bands posteriorly, fainter posterolaterally. Setae of T5 and T6 generally pointing caudally (cf. D. ironensis , Figs. 94–95).

Male terminalia. Epandrium (periphallic organs) ( Figs. 38–41 ). Genital arch narrow dorsally and broad laterally; toe (ventral epandrial lobe) elongate as in D. ananassae , with about 6–8 setae. Primary and secondary claspers present. Primary clasper (surstylus) large with an inner or median row of 5–6 strong setae that merges into a cluster of an additional 8–9 setae (one large) and two series of short, blunt, thick teeth (prensisetae) laterally. The upper series has 5–6 prensisetae, the lower series has 2–3 of similar form. The secondary clasper (ventral cercal lobe) is very small with a very large curved, black, medial tooth, and with several small setae.

Hypandrium (phallic organs). (Figs. 2–5, 14–21, 35, 36). The medial expansion of the novasternum (n in Fig. 4) resembles a “viking helmet”—rounded like the crown of a human head with the submedian spines arising like “horns” laterally. Aedeagus non-bifid, apically hirsute. Anterior parameres small, digitiform with large apical seta and several medial sensilla (ap in Fig. 4) and with a hugely extended and recurved structure at its base—the basal extension (bx in Fig. 4). The basal extension terminates as a very long, pointed and sclerotized appendage (Figs. 2–4); in ventral view, the lateral side of this caudally extended structure is almost straight, the other side curves near the tip (cf. D. ananassae —basal extension tapers on both sides toward a pointed tip, e.g., Figs. 6, 8). The basal extension pivots outwards (e.g., Fig. 16 ) when the aedeagus extends. It is figured in the resting or not-outwardly-pivoted position in Fig. 4. An acuminate kink in the “bend” of the anterior paramere (ak, Fig. 4) is visible in ventral view (Figs. 2–5, 14–21, this kink is absent in D. ananassae e.g., Figs. 6–9, 22–25). Posterior paramere long, extending beyond aedeagus; slender, tapering to a point, bending abruptly (cf. gradually in D. ananassae and D. pallidosa ) at apex of aedeagus and sheathing it. The overall “sheathed” phallus width is about 0.4 of the hypandrium width in D. pandora and smaller, c. 0.3, in D. ananassae (compare Figs. 2–5 and Figs. 6–9). The transverse band (tb Fig. 4) of the ventral phragma is short (cf. long in D. ananassae ) and opens into a lateral deltoid-shaped expansion (lx Fig. 4, versus not expanded in D. ananassae , e.g. Figs. 6, 23).

Female. Difficult to identify except by extrapolation from male siblings or progeny.

Female terminalia. Oviscapt concolorous with tergite VI ( Fig. 90 ).

Specimens examined. Australia (Western Australia): 1♂, … - 16.3604°S 124.7684°E | Northwest Kimberley | MALAISE-trap sample (7 days) | Coll : M21/2E2rb (29Jan2013) | O.R. Edwards & R.K. Didham CSIRO ( WAM) . Australia, Northern Territory: 3♂♂, NT Holmes Jungle | -12.3978° 130.9345° ± 50m | 21.v.2013 swept | S.F. McEvey & G.R. Brown | AMS K357515–517 ; 3♂♂, ibid | 22.v.2013 swept | S.F. McEvey & G.R. Brown | AMS K357518–520 . Australia (Queensland): 2♂♂, | AMS K194455–456 | Gordon Ck, Iron Range | 12°43'S 143°19'E N.Qld | swept over flowers at | rainforest fringe | 12.v.1981 S.F. McEvey; 1♂, QLD Lake Placid | -16.8678° 145.6731° ± 50m | est. ix.2011, em. 22.i.2013 | M. Schiffer culture CAR 274; 2♂♂, ibid | M. Schiffer culture CAQ408; 1♂, ibid | M. Schiffer culture CAQ425; 1♂, | AMS K274605 | McE9362 | QLD, Heathlands | Bertie Creek pump | 11:46S 142:36E fruit | 13.3.92 S.F. McEvey ; 2♂♂, | AMS K274602–603 | McE9156, McE9162 | QLD, Heathlands | Bertie Creek pump | 11:46S 142:36E fruit | 10.iii.92 S.F. McEvey; 1♂, QLD Townsville | Tucson stock 14024-0371.11 | Kyorin University stock k-aat001; 1♂, QUEENSLAND | culture AUS52 | Kyorin University stock k-aat002 | M. Hatsumi | iso- ♀ line; 1♂, QLD Black River | … Vivian Voss Crt | 11 April 2014 fruit bait | Michele Schiffer; 1♂, QLD Mango Tree Tourist Pk | Innisfail, orange trees | 13–14 April 2014 fruit bait | Michele Schiffer; 4♂♂, QLD Serina Beach , fruit shed | 6–7 April 2014 | Michele Schiffer CGZ MSRC ; 32♂♂, QLD Rockhampton ,orchard | 2April 2014 | Michele Schiffer CGW MSRC . Australia ( Torres Strait islands ): 1♂, ANIC 29031424 | Thursday Is. , NQ | fruit (domestic) | 15.i.1980 | S.F. McEvey; 2♂♂, ANIC 29031431–432 | Moa Island , N. Qld | fruit (domestic) | 2.ii.1980 | S.F.McEvey . Papua New Guinea: 2♂♂ , PNG Wanigela | 9°16'S 149°08'E | 12–28 Feb 2003 | Shane F. McEvey; 2♂♂ , PNG Tabubil 570 m | 5.258°S 141.220°E | 3.ii.2009 fruit | Shane F. McEvey; 1♂, Wau , Papua New Guinea | culture WAU142 [1981] | Kyorin University stock k-aat003 | E. Takanashi, Y. N. Tobari | iso- ♀ line .

Distribution ( Fig. 1 ). This species is known from across tropical Australia (to as far south as Rockhampton in the east and The Kimberley in the west) and eastern New Guinea (from Tabubil, Wanigela, Wau, Lae and Port Moresby).

Etymology. The proposed name pandora is a noun from the contemporary phrase to open Pandora’s Box which, in turn, is from Ancient Greek mythology. By investigating the possibility that two or more species co-exist where previously it was thought only Drosophila ananassae occurred, we felt that we were opening a taxonomic Pandora’s Box, replacing simplicity with complexity.

Remarks

This species has, since the 1970s, been confused in Australia with Drosophila ananassae , see, for example, Bock (1977); Parsons & Bock (1979, p. 230); Tribe & Bock (1981); McEvey (1982); McEvey & Bock (1982); and Schug et al. (2007). The two species are exceedingly similar in overall coloration but with experience one can detect that D. ananassae is slightly larger and darker. The sex combs of male forelegs are a useful indicator ( Figs. 54–65 ). Table 2 shows that the average total number of teeth in all combs of one leg is about 22 in D. pandora and about 37 in D. ananassae . However, the most reliable diagnostic characters are found in the male hypandrium.

By examination of terminalia we have determined that D. pandora is not synonymous with Drosophila ananassae from Norfolk Island (Bock & Parsons, 1981), with Drosophila “ pallidosa -like” of Tomimura et al. (1993, p. 147), or with Drosophila “ pallidosa -like Wau” of Matsuda et al., 2009:159.

We have excluded D. atripex Bock & Wheeler, 1972 (Fig. 10) as a possible synonym by examining specimens from Thailand ( THAILAND Pak Chong | 14°41'N 101°24'E | 1–15.vii.1989 | coll. J.R. David) and Bali ( INDONESIA, BALI | Mumbul Inn, Ubud | 8.5057°S 115.2608°E | WGS 84±100m 15.iv.2011 | S. McEvey & J. Weiner). Drosophila pandora is not synonymous with the species under culture as “ pallidosa -like”(NEW GUINEA Lae | culture LAE345 | Kyorin University stock k-aau001 | 1981 | E. Takanashi, and Y.N. Tobari | iso- ♀ line), Takanashi, Tobari and others, treat this strain as not “papuensis-like” (i.e. not D. pandora ). We have examined males from two strains of Drosophila “ pallidosa -like WAU” ( Fig. 37 ) (NEW GUINEA Bulolo | culture Bulolo79-2 | Kyorin University stock k-aav002 | 1979 H.L. Carson, T. Okada | iso- ♀ line; and Wau, Papua New Guinea | culture WAU92 [1981] | Kyorin University stock k-aav001 | E. Takanashi, Y. N. Tobari | iso- ♀ line) and found it not to be synonymous with D. pandora .

The species is easily reared in the laboratory. Future studies of this and other species in this complex should consider depositing male voucher specimens in a museum in order that links can confidently be made between genetic, genomic, cytological, behavioural and other experimental findings and taxonomy which is still based largely on morphology.

At Iron Range ( Fig. 1 ) van Klinken reared “sp. nr D. ironensis ” (a likely synonym of D. pandora ) from fruits of the following plants: Mangifera indica , Ptychosperma elegans, Garcinia riparia, Momordica charantia, Elaeocarpus arnhemicus, Ficus nodosa, Syzygium bamagense, Morinda citrifolia, Nauclea orientalis and three other unidentified fruits (van Klinken & Walter, 2001). In rainforests around Cairns he reared D. pandora from fruit of the following species: Barringtonia calyptrata, Elaeocarpus angustifolius, E. bancroftii, Gmelina sp. , Polyalthia michaelii, Polyscias sp., Randia fitzalani, Syzygium cormiflorum, and two unidentified spp. In Northern Territory, around Darwin, D. pandora (det. as “sp. nr D. ironensis ”) was reared from fruit of: Averrhoa carambola, Citrus aurantifolia, C. reticulata, Citrus sp., Malphighia glabra , Mammea americana, Manilkara zapota, Nauclea orientalis, Psidium cattelianum, P. guajava, Spondia cytherea, S. mombin , and Terminalia sp. (van Klinken & Walter, 2001).

2 3

Lake Placid (type strain) Lake Placid (type strain) D. pandora sp.nov. D. pandora sp.nov.

4 5

Lake Placid, Qld Lake Placid, Qld D. pandora sp.nov. D. pandora sp.nov.

Supiori, West Papua Tabubil, PNG D. ananassae D. ananassae

10 11

Lihir, Bismarck Archipelago Thursday Island, Qld D. ananassae D. ananassae

12 13

Bali Moorea New Caledonia Samoa

D. atripex D. monieri D. ochrogaster D. schugi sp.nov.

Figures 2–13. Hypandria of species of the Drosophila ananassae subgroup . Drosophila pandora sp.nov. from Lake Placid, northern Queensland, (2–3) type strain = iso- ♀ strain CAQ408, (4) iso- ♀ strain CAQ425, (5) iso- ♀ strain CAR 274. Drosophila ananassae from (6) Supiori, West Papua; (7) Tabubil 750m, Western Province, Papua New Guinea; (8) Lihir, Bismarck Archipelago, New Ireland Province, PNG; (9) Thursday Island, Torres Strait, northern Queensland. Drosophila atripex (10) Ubud, Bali, Indonesia. Drosophila monieri (11) Belvédère 250 m, fruit bait, Moorea, Society Islands, French Polynesia. Drosophila ochrogaster (12) Mont Koghis, New Caledonia. Drosophila schugi sp.nov. (13) Malololelei, Upolu, Samoa (paratype AMS K282923).Abbreviations: a, aedeagus; aa, aedeagal apodeme; ak, acuminate kink of anterior paramere (cf. featureless curve, e.g. Figs. 7–9, 22–25); ap, anterior paramere; bx, basal extension of anterior paramere; lx, lateral deltoid expansion of transverse band; n, novasternum, medial expansion; pp, posterior paramere; tb, transverse band. Localities—see Fig. 1 and Appendix 1; all specimens in Australian Museum.

Mauritius Christmas Island D. ananassae D. ananassae Wé , Loyalty Islands Bali D. ananassae D. ananassae

Drosophila (Sophophora) parapallidosa Tobari , in Matsuda & Tobari, 2009

Fig. 34

Drosophila (Sophophora) parapallidosa Tobari, 2009:135– 140 , in Matsuda & Tobari (2009).

Drosophila (Sophophora) parapallidosa Tobari, 2009: 157–162 , 164, 166, 167 and unpaginated Appendix A, Supplementary material, in Matsuda et al. (2009). Unavailable.

A description of Drosophila parapallidosa has been published twice, once in the journal Fly and once in Drosophila Information Service. Both works have also been published online. If the print versions fulfilled the regulations of The Code (1999) the name proposed in the more recent print version would be treated as preoccupied. The Fly publication is probably the earlier or older print version, however, it lacks explicit fixation of a holotype for Drosophila parapallidosa (see The Code, Article 16.4.1) and the description of Drosophila parapallidosa in Fly is thus invalid—the name proposed therein is unavailable. The two online versions can both be discounted because they lack ZooBank registration. Of all four taxonomic treatments only one is valid: the print version of the paper in DIS Tobari , in Matsuda & Tobari, 2009:135 (print version). [It is assumed that the print version of DIS 92 was published before 31 December 2009, not after. If published in print in 2010, however, the correct authority and year for this species would be Tobari, 2010, in Matsuda & Tobari, 2010].

Drosophila parapallidosa is reported from Kota Kinabalu ( Malaysia), Lanyu (Taiwan) and Okinawa ( Japan) ( Fig. 1 ). Specimens have been obtained from the Kyorin University stock center and dissected. The terminalia is quite unlike D. pandora sp.nov. (Matsuda & Tobari, 2009: fig. 1A; and present work Fig. 34 ).

Distribution. Malaysia (Kota Kinabalu), Taiwan and Japan.

Specimens examined. The following males have been dissected and determined to be D. parapallidosa . Malaysia: MALAYSIA Kota Kinabalu | Kyorin University stock k-aas | 1979 see Tomimura et al., 1993 | isofemale lines | Y. Fuyama, F. Hihara, and T.K. Watanabe.

Figures

Figure 1. Distribution of 11 of the 12 Drosophila ananassae complex species, the authors have collected at red dot localities or examined specimens collected by others from red circle localities; these data are complete for the Australian Region and incomplete for the Oriental Region. Drosophila lachaisei Tsacas, 1984 from Africa is not shown. The Drosophila pandora sp.nov. type locality is Lake Placid near Cairns (see inset); the D. anomalata sp.nov. type locality is Deeragun near Townsville. Drosophila ananassae s.str. is pantropical within the pale blue lines, the Ambon type locality is shown; D. ananassae extends as far south as Corindi in Australia, it is absent in Sydney and New Zealand. Three or more additional, but undescribed species, occur in New Guinea (and perhaps also in northern Australia), these are not shown. Islands of Torres Strait:(1) Mabuiag, (2) Moa, (3) Thursday, and (4) Mount Adolphus; and localities on Cape York Peninsula: (5) Bamaga, (6) Bertie Creek and Heathlands homestead, (7) Doublemouth and Gunshot Creeks, (8) Gordon Creek, (9) Iron Range (nr airstrip), and (10) upper Peach Creek (McIlwraith Range) are indicated (see inset). Geospatial coordinates of all localities are given in Appendix 1.

Figures 54–71. Male fore-leg and sex comb of tarsomere I (metatarsus or basitarsus), tarsomere II and III of Drosophila pandora sp.nov. (54–59), D. ananassae (60–65)(from strains established by Schiffer at Lake Placid, northern Queensland) and D. schugi sp.nov.(66–71). Drosophila pandora sp.nov. (54–55) from iso-♀ strain CAR274 [the teeth of the sex comb in Fig. 51 are artificially enhanced and represented schematically to indicate how they are scored]; (56–57) from type strain = iso-♀ strain CAQ408; and (58–59) from iso-♀ strain CAQ425. Drosophila ananassae (60–61) from iso-♀ strain CBR57; (62–63) from iso-♀ strain CBR54; and (64–65) from iso-♀ strain CBR52. Drosophila schugi sp.nov. wild caught males from Malololelei, Upolu, Samoa, 14–17 June 2003, Schug, Gray-Smith, Kilon-Attwood, McEvey; AMS K356978 (66), AMS K356979 (67–68), AMS K356977 (69), and AMS K356976 (70–71).

Figures 38–53. Epandria of Drosophila ananassae complex species. Drosophila pandora sp.nov. (38) Lake Placid type strain CAQ408, lateral view; (39) type strain CAQ408, caudal view; (40) Lake Placid strain CAR274; (41) Lake Placid strain CAQ425. Drosophila ananassae (42) Beach Lane, Mauritius, Oct. 2012, J. Tann.Drosophila?ananassae (43) Kuranda, northern Queensland,AMS K275429; (44) Sigatoka, 35 km S Nadi, Fiji, AMS K275290; (45) Iron Range, fruit bait, 30.iv.1976, I.R. Bock, [possibly pinned on this date from a strain collected with P.A. Parsons in November 1975—there is no evidence that Bock returned to Iron Range in 1976], AMS K119308 (det. Drosophila ananassae by Bock in 1976). Drosophila pandora sp.nov.: (46) D. “papuensis-like” Kyorin University stock k-aat001, Townsville, Queensland.Drosophila pallidosa: (47) det. by?Wheeler, Pago Pago,American Samoa, DrosophilaSpecies Stock Center (San Diego), stock 14024-0433.01. Drosophila ananassae or D. pallidosa (48) Sigatoka, 35 km S Nadi, Fiji, June 2004, Schug, Gray-Smith, Kilon-Attwood, McEvey, AMS K282851. Drosophila schugi sp.nov. (49) Malololelei, Upolu, Samoa, 14–17 June 2003, paratype AMS K282923. Drosophila anomalata sp.nov. (50–52) ex type strain, Schiffer CHC221. Drosophila schugi sp.nov. (53) Malololelei, Upolu, Samoa, 14–17 June 2003, AMS K356977. Localities—see Fig. 1 and Appendix 1; all specimens in Australian Museum.

Figures 14–25. Hypandria ofDrosophila pandora sp.nov. and D.ananassae. Drosophila pandora sp.nov.(14–15) Holmes Jungle, Darwin, NorthernTerritory;(16) Tabubil750 m,Western Province, PapuaNew Guinea; (17) Wanigela, Oro Province,Papua New Guinea;(18) Bertie Creek pump, near Heathlands Homestead, northern Queensland; (19–20) Gordon Creek nr Cooks Hut, Iron Range, northern Queensland; (21)Thursday Island, Torres Strait, northern Queensland.Drosophila ananassae (22) Beach Lane, Mauritius;(23) Christmas Island, Indian Ocean nr Java; (24) Wé, Loyalty Islands; (25) Ubud, Bali, Indonesia. Localities mapped in Fig. 1; all specimens in Australian Museum.

Figures 89–91. Female oviposcapt (ovipositor) and setation of sixth abdominal tergite of Drosophila ananassae, D. pandora, D. anomalata and D. ironensis. Oviscapt form and colour: (89) Drosophila ananassae ex strain Schiffer CBR57 from Lake Placid; (90) D. pandora sp.nov. ex type strain Schiffer CAQ408 from Lake Placid; (91) D. anomalata sp.nov. ex type strain Schiffer CHC221 from near Deeragun, W of Townsville. Setation of sixth abdominal tergite: (92) D. ananassae ex strain Schiffer CBR54; (93) D. anomalata sp.nov. ex type strain; (94, 95) Drosophila ironensis ♀ and ♂ ex strain Schiffer CHH18 from Lake Placid, note the irregualr orientation of T6 setation in D. ironensis comparaed to the caudally oriented setation in other figured species; and (96) D. pandora sp.nov. ♂ ex type strain.All to same scale.

Figures 26–37. Hypandria of Drosophila ananassae complex species. Drosophila ananassae (26) 1.5 km NW Taipivai, Nuku Hiva, Marquesas Islands; (27) Belvédère 250 m, fruit bait, Moorea, Society Islands, French Polynesia.Drosophila ananassae orD.pallidosa (28) Sigatoka, 35 km S Nadi, Fiji; (29) Apia, Samoa. Drosophila pallidosa (30) det. by?Wheeler, Pago Pago, ... [continued on facing page]