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Drosophila (Drosophila) Robusta Species Group (Diptera, Drosophilidae) from Southeast Asian Tropics, with The Entomological Science (2010) 13, 381–391 doi:10.1111/j.1479-8298.2010.00398.x ORIGINAL ARTICLE Discovery of the Drosophila (Drosophila) robusta species group (Diptera, Drosophilidae) from Southeast Asian tropics, with the descriptions of six new speciesens_398 381..391 Awit SUWITO1 and Hideaki WATABE2 1Zoology Division (Museum Zoologicum Bogoriense), Research Center for Biology-LIPI, Cibinong, Bogor, Indonesia; and 2Biological laboratory, Sapporo College, Hokkaido University of Education, Sapporo, Japan Abstract Six new species of the Drosophila robusta species group are described from Southeast Asian Islands. Kalimantan and Sunda Islands lie east of Bali, from which Drosophila barobusta sp.nov and D. uncinata sp.nov belong to the lacertosa subgroup, and D. sungaicola sp.nov, D. baliensis sp.nov, D. hitam sp.nov and D. subaquatica sp.nov to the okadai subgroup. The robusta group from Southeast Asian tropics exclusively inhabits streamsides in mountainous highlands with an elevation of more than 600 m from the sea level. Key words: East Malaysia, geographic distribution, habitats, Indonesia, species subgroup. INTRODUCTION restricted to riparian environments with high humidity and relatively constant temperature, and we have The Drosophila robusta species group of the Drosophila carried out bait-trap collections along streams in Borneo virilis section established by Hsu (1949) is one of major and Sunda Islands in tropical Southeast Asia. We have lineages within a huge subgenus Drosophila, and it has discovered many members of the Drosophila virilis been extensively studied from macro- and micro- section, and the present paper describes six new species evolutionary points of view (Stalker 1972; Narayanan of the robusta group as a first report, along with 1973; Throckmorton 1975; Levitan 1982; Watabe et al. brief information on their altitudinal and horizontal 1997; Wang et al. 2006; He et al. 2007). The group is distributions. geographically distributed in temperate forests of the Nearctic and the Palarctic Regions including adjacent MATERIALS AND METHODS islands (Levitan 1982; Toda et al. 1996; Zhang et al. 1996; Brake & Bächli 2008). In lower latitudes of East Most specimens were collected by bait traps with fer- Asia, e.g. Yunnan Province and Guangdong Province of menting bananas set up along streams, and several flies mainland China, habitats of the robusta group are sifted from resting sites among rocks by insect nets, both just to mountainous areas with cool climates (Watabe et al. beside (within 50 cm from) water. All samples were 1990; Watabe & Peng 1991; Chen & Watabe 1993; He preserved in 70% alcohol. External morphology was et al. 2003), and any member has not been recorded observed under a stereoscopic microscope, and metric from tropical regions. characters were measured with an ocular micrometer. We have engaged in the faunal survey of Drosophil- Male and female terminalia were detached from the idae of Indonesia and Malaysia with numerous islands body, and treated with 10% KOH solution at 80°C for surrounding the equator from 2003. Micro-habitats of several minutes, and observed in a droplet of glycerin most of the Drosophila virilis section flies are quite under a compound microscope. Drawings were carried out using a camera–lucida (Olympus Optical Co.Ltd., Tokyo, Japan) and a DinoLite Camera (AnMo Elec- Correspondence: Awit Suwito, Division (Museum Zoologicum Bogoriense), Research Center for Biology-LIPI, tronic Corporation, Taipei, Taiwan). We followed Cibinong, Bogor Cibinong 16911, Indonesia. McAlpine (1981) for the morphological terminology, Email: [email protected] and Zhang and Toda (1992) for the definitions of mea- Received 28 December 2009; accepted 18 April 2010. surements and indices with the addition of two new © 2010 The Entomological Society of Japan A. Suwito and H. Watabe indices: pprnl (lower postpronotal seta/upper post- Characters (common to the four species described pronotal seta in length) and sterno2 (mid katepisternal here). Epandrial ventral lobe bent inward. Surstylus seta/posterior katepisternal seta in length). roundish on ventral margin. Cercus oval. Spermathecal The specimens are deposited in Museum Zoologicum capsule brown, wrinkled basally. Bogoriense (MZB), Research Center for Biology, Indo- nesian Institute of Science, Cibinong, Indonesia; System- Drosophila (Drosophila) sungaicola sp. nov. atic Entomology, the Hokkaido University Museum, (Figs 1–8) Hokkaido University, Sapporo, Japan (SEHU); and Diagnosis. C-index 5.64 in holotype (range in 18ǩ and Kinabalu Park, Sabah Parks, Sabah, Malaysia 30Ǩ paratype: 3.85–5.36). Distance between antennal (KPSP). sockets wider than socket width. Aristal branches shorter than upper seta on pedicel (Fig. 1). Facial carina twice as wide as 1st flagellomere. Male sternite V pumpkin- RESULTS AND DISCUSSION shaped, gently rounded laterally, slightly protruded at middle on distal margin (Fig. 3). Epandrium with hook- Drosophila (Drosophila) robusta species group like, ventrally curved projection at anteroventral corner Drosophila robusta species group, Sturtevant (1942, p. (Fig. 4). Anterior lobe of hypandrium medially deeply 31); Watabe et al. (1990, p. 133). notched, strongly convex ventrad, sculptured on medio- Diagnosis (modified from Watabe et al. 1990). Dark anterior to outer surface, pubescent on medio-inner brown/black, large (3.0–4.5 mm in body length) species surface (Figs 5,6). Aedeagus concaved on distal, outer with 2 pairs of dorsocentral setae. Palpus pale brown, margin in lateral view (arrow in Fig. 6). Spermatheca horizontally flattened, with several long setae besides capsule distally narrowed (Fig. 8). numerous tiny setulae. Acrostichal setulae in 6 regular Description (ǩ and Ǩ). Head: Frons dark brown, mat rows. Preapical dorsal setae present on all tibiae; apical black on frontal vittae. Arista with 3-4 dorsal and 1-2 ventral setae on fore and mid tibiae. Postpronotal lobe ventral branches besides terminal bifurcation (Fig. 1). with 2 prominent setae. Wing hyaline, slightly fuscous, Anterior reclinate orbital seta situated just outside of or with veins dark brown; cross veins clear, paler; R2+3 slightly behind proclinate orbital seta. Face dark brown. straight; R4+5 and M1 parallel; C1 setae 2, subequal. Palpus large, with 2-5 long setae along distal margin Paramere (anterior paramere) rudiment/absent; gono- (Fig. 2). pod (posterior paramere) absent. Aedeagus curved Thorax: Scutum entirely dark brown to black. Post- ventrally; apodeme shorter than aedeagus. pronotal (humeral) lobe dark brown, pale brown in Characters (common to the six species described lower part. Anterior scutellar setae parallel to slightly here). Eye dark red with thick interfacetal setulae. convergent. Legs dark brown; knee joints paler; femora Occiput and ocellar triangle dark brown. Scape (1st of mid- and hindlegs paler on basal half. antennal segment) dark to pale brown; pedicel dark Abdomen: Tergites nearly entirely dark brown, brown; 1st flagellomere grey brown. Pedicel with 2 slightly paler on I and II and anteromedially on III. prominent setae; upper one longer. Gena yellowish to Sternites grey brown; male IV and V dark brown. grey brown, darker on lower margin. Clypeus dark Male terminalia (Figs 4–6): Epandrium pointed at brown to black. Scutellum nearly entirely dark brown to caudoventral corner, pubescent except for anterior black. Halter white, basally grey. Posterior scutellar setae margin and most of ventral lobe, with c. 4 long setae on strongly convergent, usually cruciate. Epandrium and upper half and c. 25 on middle to lower half per side. cercus grey to dark brown. Surstylus yellowish to pale Surstylus with 8-9 prensisetae on distal margin and c. 12 brown. Aedeagus yellowish brown to yellow. Oviscapt trichoid setae caudoventrally. Cercus separated from yellowish brown to yellow, darker on distal margin. epandrium, with c. 36 long setae, pubescent except for 1/3 ventral portion. Hypandrium dark brown; apodeme Drosophila (Drosophila) okadai species narrow. Paramere absent. Aedeagal apodeme c. 1/5 as subgroup long as aedeagus, horizontally plate-like; aedeagal guide Drosophila okadai species subgroup, Watabe and dark brown, long, anteriorly curved. Nakata (1989, p. 28). Female terminalia (Figs 7,8): Tergite VIII grey brown, Diagnosis. Surstylus basally narrow, less than 1/2 pubescent on dorsal half, ventrally with c. 9 setae per maximum width. Hypandrium (novasternum) without side. Oviscapt with c. 3 discal and c. 25 marginal peg- paramedian setae. Aedeagus broadened distal half in like ovisensilla. Spermathecal capsule with spinules dis- lateral view, nearly entirely bilobed except ventrobasal tributed on medial portion, slightly indented at apex; portion; aedeagal guide (vertical rod) plate-like. introvert c. 0.9 as deep as capsule. 382 Entomological Science (2010) 13, 381–391 © 2010 The Entomological Society of Japan Drosophila from Southeast Asian Islands Figures 1–16 Drosophila (Droso- phila) sungaicola sp. nov. (1–8) and D.(D.) baliensis sp. nov. (9–16). 1,9 Antenna; 2,10 palpus; 3,11 male ster- nite V; 4,12 periphallic organs: epan- drium (epand), surstylus (sur) and cercus (cerc); 5,6,13,14 aedeagus and hypandrium: aedeagus (aed), aedeagal apodeme (aed apod), aedeagal guide (aed gd), hypandrium (hypd) and hypandrial apodeme (hypd apod) in ventral (5,13) and left lateral (6,14) views; 7,15 oviscapt; 8,16 spermath- eca. Scale lines: 0.5 mm (1–3,9–11) and 0.1 mm (4–8,12–16). Measurements: BL = 4.00 mm in holotype (range in Holotype. ǩ (MZB), “Citarik River, Sukabumi, West 18ǩ paratypes: 3.36–4.00 mm, range in 30Ǩ Java, Indonesia, 6°45′53.8″S, 107°21′49.4″E, 1006 m paratypes: 3.68–4.48 mm), ThL = 1.80 mm (ǩ: 1.44– above sea level (a.s.l.), 24.VII. 2007, A. Suwito and E. 1.68 mm, Ǩ: 1.40–1.84 mm), WL = 3.80 mm (ǩ: 3.32– Cholik”. 4.16 mm, Ǩ: 3.44–4.36 mm); WW = 1.28 mm (ǩ: Paratypes. Indonesia: 1ǩ1Ǩ, Gunung Tujuh, Kerinci 1.16–1.48 mm, Ǩ: 1.16–1.56 mm). Seblat, West Sumatra, 1°25′30.5″S, 101°12′54.0″E, Indices: FW/HW = 0.50 (ǩ: 0.46–0.56, Ǩ: 0.44– 1735 m a.s.l., 5.XII.2004, H.
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