Dysphaea ulu, Hämäläinen, Matti, Dow, Rory A. & Stokvis, Frank R., 2015

Hämäläinen, Matti, Dow, Rory A. & Stokvis, Frank R., 2015, Revision of the Sundaland species of the genus Dysphaea Selys, 1853 using molecular and morphological methods, with notes on allied species (Odonata: Euphaeidae), Zootaxa 3949 (4), pp. 451-490 : 472-476

Dysphaea ulu spec. nov.

( Figs. 11 , 15 , 28 , 33 , 39 , 43 , 47 , 51 , 55 , 61 , 67 , 73 , 80 , 85 )

Dysphaea dimidiata [nec Selys, 1853];— Schmidt (1934: 330–331, part: Baram, plate 16, fig. 8);— Lieftinck (1954: 19–20, part: Borneo);— Orr (2001: 178, specimens from Kuala Belalong Field Studies Centre, Sites 1,2; except one male which was D. dimidiata Selys, 1853 );— Orr (2003: 37, part; 58 part; p. 59, fig. 72; plate 3h);— Orr (2005: 26, illustration);— Dow (2005: 11, Gunung Mulu National Park, part);— Dow & Reels (2008: 3, Gunung Mulu National Park, part);— Dow & Reels (2009: 13, Mount Dulit);— Dow & Reels (2010: 15, Kubah National Park, part);— Dow, Reels & Butler (2013a: 9, Dulit range);— Dow, Reels & Butler (2013b: 11, Kubah National Park, part).

Dysphaea limbata semilimbata [nec Selys, 1873];— Laidlaw 1920: 328 (specimen from Baram).

Dysphaea species;— Dow, Reels & Ngiam (2015: 11, Usun Apau).

Material studied: Holotype ♂ (SAR13_14_EUP15): Borneo, Sarawak, Miri division, Upper Baram, Sungai Pejelai, Ulu Moh, 24 viii 2014, leg. M. Tegong. Deposited at RMNH, Leiden. Paratypes (47 ♂, 2 ♀; all from Borneo; deposited in Coll. Dow unless otherwise stated).—From Sarawak: 1 ♀ (SAR13_14_EUP15, in tandem with holotype), Sarawak, Miri division, Upper Baram, Sungai Pejelai, Ulu Moh, 24 viii 2014, leg. M. Tegong; 3 ♂ (SAR13_14_EUP24–26), same location and date as above, leg. N. Sigau; 1 ♂ (SAR13_14_EUP23), same location as above, 26 viii 2014, leg. M. Tegong, E. Aran & N. Sigau; 1 ♂ (SAR13_14_EUP29), same location as above, 27 viii 2014, leg. E. Aran & O. Udau; 1 ♂ (SAR13_14_EUP19), Sarawak, Miri division, Upper Baram, tributary of Sungai Pejelai, Ulu Moh, 23 viii 2014, leg. Y. Tegong; 1 ♂ (SAR13_14_EUP35), same area as above, Sungai Moh, 24 viii 2014, leg. R.A. Dow; 1 ♂ (SAR13_14_EUP27), same area as above, Sungai Sii, 10 vii 2014, leg. R.A. Dow; 1 ♂ (SAR13_14_EUP34), same location as above, 12 vii 2014, leg. R.W.J. Ngiam; 1 ♂ (SAR13_14_EUP36), same location as above, 13 vii 2014, leg. N. Pawi; 1 ♂ (SAR13_14_EUP31), same location as above, 15 vii 2014, leg. O. Udau; 1 ♂ (SAR13_14_EUP17), same location as above, 17 vii 2014, leg. O. Udau; 1 ♂ (SAR09_10_EUP142, in RMNH: RMNH.INS.503424), Sarawak, Miri division, Upper Baram, Sungai Pawan, 19 vii 2010, leg. L. Southwell; 2 ♂ (SAR06_EUP18–19), Sarawak, Miri division, foot of Mount Dulit, Sungai Nuam, 30 iii 2006, leg. R.A. Dow; 1 ♂ (SAR06_EUP17, in RMNH: RMNH.INS.509905), same area as above, Sungai Long Aton, 31 iii 2006, leg. G.T. Reels; 1 ♂ (SAR09_10_EUP100; Coll. Hämäläinen), Sarawak, Miri division, Upper Tinjar River, Sungai Lobang, 4–5 v 2010, leg. L. Southwell; 1 ♂ (SAR11_12_EUP159), Sarawak, Miri division, Usun Apau National Park, Sungai Julan, 1 v 2012, leg. C. Wilfred; 1 ♂ (SAR07_8_EUP48, in RMNH: RMNH.INS.509906), Miri division, Gunung Mulu National Park, foot of Gunung Mulu, stream on old trail to Sarawak Chamber, 10 ix 2008, leg. R.A. Dow; 1 ♂ (SAR05-EUP57; Coll. Hämäläinen), Sarawak, Miri division, Gunung Mulu National Park, Long Lansat, 19 iv 2005, leg. R.A. Dow; 1 ♂ (SAR05-EUP31), Sarawak, Miri division, Kelabit Highlands, Bario area, Pa’Dapur river, 5 iv 2005, leg. G.T. Reels; 1 ♂ (SAR06_EUP33), Sarawak, Kuching division, Matang Range, Kubah National Park, Sungai Rayu at Matang Wildlife Centre, 25 i 2006, leg. G.T. Reels; 2 ♂ (SAR07_8_EUP37; SAR07_8_EUP 49 in RMNH: RMNH.INS.509907), Sarawak, Sri Aman division, Batang Ai National Park, Sungai Bebiong Besar, 4 xii 2007, leg. R.A. Dow; 1 ♂ (SAR07_8_EUP12), same location and date as above, leg. G.T. Reels; 3 ♂ (SAR13_14_EUP7–8, SAR13_14_EUP 2 in RMNH: RMNH.INS.557687), Sarawak, Kapit division, Lanjak Entimau Wildlife Sanctuary, Ulu Katibas, Sungai Datai, 22 viii 2013, leg. R.A. Dow; 1 ♂ (SAR13_14_EUP12), same location and date as above, leg. J. ak Awan & M. ak Adau.—From Brunei: 2 ♂ ( RMNH, ex. collection P. Mill), (Temburong district), Ulu Temburong, 1978 , leg. S.L. Sutton; 1 ♂ ( RMNH), 1 ♀ (Coll. A.G. Orr) Brunei (Temburong district), Kuala Belalong Field Studies Centre, iii 1991, leg. A.G. Orr; 1 ♂ ( RMNH), same data, x 1991; 1 ♂ ( RMNH), same data, vii 1992; 1 ♂ ( RMNH), same data, 29 ix 1992; 1 ♂ ( RMNH), same data, 23 i 1993; 1 ♂ ( RMNH), same data, vii 1993; 1 ♂ ( CUMZ), Brunei, Temburong, 10 x 1978, leg. T. Harman (ex coll. D.A.L. Davies); 3 ♂ ( CUMZ), Brunei, Snake Creek, 16 ii 1982, leg. T. Harman (ex coll. D.A.L. Davies).—From Sabah: 3 ♂ ( RMNH), Sabah, 105 km S of Beaufort, Long Pasia area, Sungai Padas, S. of Kampong Long Pasia, 1030 m asl., 4 24’N, 115 44’E, 12–13 iv 1987, leg. J. van Tol; 1 ♂ ( RMNH), Sabah, surr. Long Pasia, Sg. Matang, leg. J. van Tol.; 2 ♂ ( CUMZ), Borneo, Sabah, Maliau Basin, tributary to Sungai Maliau near Maliau Basin Field Centre, 14 v 2013, leg. S. Luke.

Etymology. The species epithet is based on the word ‘ulu’, the form generally in use in Borneo of the Bahasa Melayu/ Indonesia word ‘hulu’, which means upstream. The species epithet is used as a noun in apposition. The species typically inhabits ‘upstream’ habitats.

Diagnosis. A narrow-winged Dysphaea species with male wings broadly opaque at basal half and at wing tips. Cerci with lower border nearly straight in lateral view.

Description of holotype male ( Fig. 11 ). Head: Labium, labrum, base of mandibles and clypeus shining black, frons and vertex matt black.

Thorax: Matt black, with very faint obscure brown marking on metepisternum above second lateral suture and similar marking on metepimeron ( Fig. 15 ). Venter of thorax without tiny tubercles on metaposternum ( Fig. 18 ). Legs wholly black.

Wings: Basal half of wings opaque black with strong metallic blue reflections. In Fw opaque area extends to level of 3–4 cells before nodus, except in costal field where the opaque stripe extends 1 or 2 cells beyond the nodus. Otherwise costal field between nodus and pterostigma hyaline. In Hw basal opaque area extends more apicad, not quite reaching half way between nodus and proximal border of pterostigma; costal field between nodus and pterostigma opaque throughout. Tips of wings narrowly opaque, in Hw slightly more extensively than in Fw ( Fig. 11 ). Venation typical of genus. Fw with 37 antenodals in first row; Hw correspondingly with 27 antenodals. Quadrangle with 2 crossveins in Fw, 2–3 in Hw. Pterostigma long and narrow, broadest in middle; covering 14 underlying cells.

Abdomen: Matt black throughout. Appendages black; cerci in dorsal ( Fig. 33 ) and ventral view of typical shape for genus; in lateral view ventral margin of cercus almost straight ( Fig. 39 ). Paraprocts very short, rounded and featureless.

Penis: Terminal segment with two apical arms directed upwards on either side of shaft, turning out and down for short distance at ends, slightly expanded in this part ( Figs. 43 , 47 ).

Measurements (mm). Fw 33, Hw 31, abdomen (apps. excl.) 35.5, cerci 2.

Description of female ( Fig. 51 ). Head: Labium shining black with lateral lobes largely yellow, with black hooks. Labrum shining black, with broadly yellow centre, indented by black basally. Base of mandibles largely yellow, with black incomplete stripe medially. Clypeus shining black with narrow yellow stripe along ridge. Frons matt black with sides below antennae yellow, yellow colour continuing over genae ( Fig. 55 ). Antennae black. Vertex and occiput matt black, with two tiny yellow spots on occiput.

Thorax: Prothorax black with rounded large yellow spots on either side of dorsum of middle lobe. Posterior part of hind lobe raised obliquely upwards to form an elongate rectangular flap, which is narrowly bordered by yellow, more broadly so at sides ( Fig. 61 ). Lateral parts of flap not prominent, lying in ca same plane as median part. Synthorax matt black, with moderately narrow yellow stripes as in Figs. 67 , 73 . Yellow stripes on mesepisternum not connected at wing base, but those on metepimeron forming a loop at wing base. Legs black, with yellow markings on both apical and posterior sides of coxae and obscure streaks on hind femora.

Wings: Hyaline with broad blackish opaque streaks in middle section of wings, at base from subcostal field to MA, costal field and much of lower part of wing hyaline ( Fig. 51 ). In Fw opaque streak extends to level of 12 cells before nodus, in Hw it extends gradually narrowing to proximal end of pterostigma. Apex of Fw narrowly darkened, slightly more extensively on Hw. Fw with 37 antenodals in first row; hindwings with 26–28. Quadrangle with 3 crossveins in Fw, 5 in Hw. Pterostigma long, covering 11–13 underlying cells.

Abdomen: Matt black, with yellow markings as follows: S1 with lateral spot, extending upwards and downwards at apex of segment. S2–7 with lateral stripe, broadest on S2–3 and gradually narrowing towards apical segments. Separate small lateral spots at base of S3–7. On S2–5 stripe occupies almost whole segment length, on S6–7 stripe is interrupted.

Measurements (mm). Hw 31.5–33, abdomen (apps. excl.) 30.5–32, cerci 1.

Variation in male paratypes. In some specimens the pale markings on the synthorax are slightly more distinct, in others they are entirely absent; clearly this is an age dependent character. The extent of the opaque area in wings is somewhat variable. In some specimens the opaque basal area in the Fw extends to the level of the nodus or even 1–4 cells beyond the nodus (up to 8 in costal field) and in Hw beyond half way between the nodus and proximal border of pterostigma. There is also some slight variability in venational details.

Measurements (mm): Hw 27.5–32.5, abdomen (apps. excl.) 32.5–37.

Distinguishing characters. Male: Superficially D. ulu male closely resembles D. dimidiata , a species with which it co-occurs in many locations in northern Borneo. However, these species are easy to separate by the shape of cercus as seen in lateral view; in D. ulu ( Fig. 39 ) the ventral margin of the cercus is almost straight, but distinctly arched in D. dimidiata ( Fig. 37 ). Other characters, although less consistent, include the colour of the costal field between nodus and pterostigma in the Fw; in D. ulu ( Fig. 28 ) the field is hyaline, but in most specimens of Bornean D. dimidiata the costal field in the Fw is opaque ( Fig. 22 ). In D. ulu the venter of the thorax is always without tiny tubercles on metaposternum ( Fig. 18 ), whereas in Bornean D. dimidiata they are almost always present ( Fig. 17 ). There are also differences in the shape of the apical arms of the penis: the terminal, upward directed part is distinctly shorter in D. ulu than in D. dimidiata , best seen in lateral view (cf. Figs. 45 and 47 ), and is more rounded in ventral view (cf. Figs. 41 and 43 ). Female: D. ulu is easy to separate by the distinct opaque streak in both wings ( Fig. 51 ). In D. dimidiata the wings are largely hyaline or semihyaline with brownish tinge, the tips being slightly darkened ( Fig. 49 ). In D. ulu the yellowish stripes on thorax are narrower ( Fig. 67 ) than in D. dimidiata ( Figs. 63– 65 ). D. ulu lacks the conspicuous yellow stripe on antefrons ( Fig. 55 ), which is often present in D. dimidiata ( Fig. 53 ).

Remarks. Some published records on Bornean Dysphaea species still remain uncertain as regards the real identity of the species. The D. dimidiata record from Lanjak Entimau Wildlife Sanctuary in Sarawak by Norma- Rashid & al. (2010, p. 326) could refer to either to D. dimidiata or D. ulu ; both species are known to occur there. The D. dimidiata record from ‘ Sarawak, Kampong Seku’ by Asahina (1966) might just as well refer to D. ulu .

Figures

FIGURE 85. Distribution map of Dysphaea ulu.

FIGURES 9 – 12. Habitus of male Dysphaea: 9) D. dimidiata West Java; 10) D. lugens Danum Valley, Sabah; 11) D. ulu holotype (flipped horizontally); 12) D. vanida holotype (flipped horizontally).

FIGURES 13 – 18. Synthorax male Dysphaea: 13) D. dimidiata West Java; 14) D. lugens Bakuan, Kalimantan Barat; 15) D. ulu holotype; 16) D. vanida holotype; 17) metaposternum, D. dimidiata Brunei; 18) metaposternum, D. ulu holotype.

FIGURES 25 – 30. Wings of Dysphaea males: 25) D. lugens Pemantang, Kalimantan Tengah; 26) D. lugens Ampah, Kalimantan Tengah; 27) D. lugens Selekau River, Kalimantan Barat; 28) D. ulu paratype, Temburong, Brunei; 29) D. vanida spec. nov. paratype, Khlong Nakha, Ranong; 30) D. vanida paratype, Khlong Nakha, Ranong.

FIGURES 31 – 36. Male anal appendages. Dorsal view: 31) D. dimidiata West Java; 32) D. lugens Bakuan, Kalimantan Barat; 33) D. ulu holotype; 34) D. vanida holotype. Dorso-lateral views: 35) D. dimidiata Terengganu; 36) D. vanida paratype Lam Khlong Gnu, Kanchanaburi.

FIGURES 37 – 40. Male anal appendages, lateral view (scale bars 1 mm): 37) D. dimidiata west Java; 38) D. lugens Bakuan, Kalimantan Barat; 39) D. ulu holotype; 40) D. vanida holotype.

FIGURES 41 – 44. Penis, ventral view: 41) D. dimidiata west Java; 42) D. lugens Selekau River, Kalimantan Barat; 43) D. ulu holotype; 44) D. vanida paratype, Khlong Nakha, Ranong.

FIGURES 45 – 48. Penis, lateral view: 45) D. dimidiata west Java; 46) D. lugens Selekau River, Kalimantan Barat; 47) D. ulu holotype; 48) D. vanida paratype, Khlong Nakha, Ranong.

FIGURES 49 – 52. Habitus of female Dysphaea: 49) D. dimidiata west Java; 50) D. lugens Mount Dulit, Sarawak, photograph by Benjamin Price; 51) D. ulu paratype; 52) D. vanida paratype Kanchanaburi.

FIGURES 53 – 56. Female head: 53) D. dimidiata Sarawak, dorsal view; 54) D. lugens Gunung Sari, Kalimantan Timur, dorsal view; 55) D. ulu paratype, dorsal, slightly frontal view; 56) D. vanida paratype, Kanchanaburi, dorsal, slightly frontal view.

FIGURES 57 – 62. Female prothorax, dorsal view: 57) D. dimidiata west Java; 58) D. dimidiata Sarawak; 59) D. dimidiata Tandjong Pandan, Billiton; 60) D. lugens Gunung Sari, Kaliamantan Timur; 61) D. ulu paratype; 62) D. vanida paratype, Kanchanaburi.

FIGURES 63 – 68. Female synthorax, dorsal view: 63) D. dimidiata west Java; 64) D. dimidiata Sarawak; 65) D. dimidiata Johor; 66) D. lugens Gunung Sari, Kaliamantan Timur; 67) D. ulu paratype; 68) D. vanida paratype Kanchanaburi.

FIGURES 69 – 74. Female synthorax, lateral view: 69) D. dimidiata west Java; 70) D. dimidiata Sarawak; 71) D. dimidiata Johor; 72) D. lugens Mount Dulit, Sarawak, photograph by Benjamin Price; 73) D. ulu paratype; 74) D. vanida paratype Kanchanaburi.

FIGURES 77 – 82. Dysphaea males in life: 77) D. basitincta Hainan (photo by Mo Shanlian) 78) D. dimidiata Sarawak (photo by Graham T. Reels); 79) D. dimidiata Negeri Sembilan (photo by C. Y. Choong); 80) D. ulu Sarawak (photo by Graham T. Reels); 81) D. vanida Ranong (photo by Matti Hämäläinen); 82) D. vanida Surat Thani (photo by André Günther).

FIGURES 19 – 24. Wings of D. dimidiata males: 19) West Java; 20) West Java; 21) Billiton; 22) Babi Djuton, Kalimantan Timur; 23) Perak; 24) Labuan.