Dysphaea dimidiata Selys, 1853

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 : 459-467

Dysphaea dimidiata Selys, 1853

Selected references:

Dysphaea dimidiata Selys, 1853 :— Selys Longchamps (1853: 54, original description of ‘ Dysphaea dimidiata, De Selys’, male, Java);— Selys Longchamps & Hagen (1854: 185–187, pl. 5, Fig. 4, pl. 14, Fig. 4, extended description of male, Java);— Selys Longchamps (1879: 378–379, reprint p. 32–33, description of female, Sumatra);— Albarda (1887: 6–7: pl. 2, Figs. 1–2, ♂, ♀ from C. Sumatra); Krüger (1898: 78, Sumatra, Silago);– Laidlaw (1924: 300, Pahang);— Laidlaw (1931a: 180, Pahang);— Schmidt (1934: 330–331,S. Sumatra, Borneo);— Lieftinck (1934: 385, Java; no new records from Java);— Lieftinck (1935: 4, Sumatra);—Coomans de Ruiter (1936: 74–75, Fig. 3, W. Borneo, W. Java);— Lieftinck (1936: 109–110, Fig. 14 ; W. Java);— Lieftinck (1949: ix–xi, Java, notes on larva, bionomics);— Lieftinck (1950: 664–665, Java, phenology);— Lieftinck (1954: 19–20, distribution, habitat, references);— Lieftinck (1959: 101–103, notes, drawings ( Figs. 4–6), behaviour of both sexes);— Furtado (1969: 866, 874, 876, habitats, Peninsular Malaysia);— Paulson (1981: 134–135, Sarawak, behaviour;— Asahina (1981: 6, South Thailand, Trang);— Asahina (1985: 31–34, Figs. 35–44 , 64 , S. Thailand, descriptions of both sexes);— Tsuda & Kitagawa (1988: 3, Sumatra);— Kemp & Kemp (1989: 39, Pahang);— Dunkle (1993: 75; Songkla);— Norma-Rashid (1995: 81, Perak);—Hämäläinen & al. (1996: 5, Kelantan);—Norma- Rashid & al. (1996: 134, Kelantan, Pahang);— Hämäläinen & Pinratana (1999: 29, 123, part; Thailand, records from the southermost provinces);— Orr (2001: 178 (part, 1 male specimens from Kuala Belalong Field Studies Centre; sites 1,2); Sites 30–31, all specimens);— Norma-Rashid & al. (2001: 143, Pahang, Tasek Bera);— Orr (2003: 37, part; 58 part);— Kalkman (2004: 28, Pahang);— Cleary & al. (2004: 445);— Dow (2005: 11, Gunung Mulu N.P.);— Wilson & Gibert (2005: 27, Johor, Endau-Rompin);— Orr (2005: 26, text);—van Tol (2006: 13, East Kalimantan);—Wahizatul & al. (2006: 101, 102, 103, Terengganu);— Kitagawa & Katatani (2007: 31, Johor);— Dow & Reels (2008: 3, Gunung Mulu N.P., part);— Choong (2009: 226, Johor, Panti Forest Reserve);—Norma-Rashid (2009: 68, Negeri Sembilan);— Choong (2010: 230– 231, Fig. 2, Perak, Bintang Hijau Forest Reserve);— Dow & Reels (2010: 15, Sarawak, Kubah National Park, part);— Dow & Unggang (2010: 1353, Sarawak, Binyo Penyilam);— Tang & al. (2010: 41–42, Singapore, past records, photos of Johor specimens);— Dolný & al. (2011: 72), Kalimantan, Sg. Wain);—Choong (2011: 237, Perak, Royal Belum State Park);— Choong (2013: 207, Kelantan, Chamah Highlands);—Ng & al. (2011: 10, 11, Perak);—Choong & al. (2012: 97, Terengganu);—Ng & al. (2012: 83, Pahang);—Dow (2012: 9, Sarawak, Annah Rais);— Dow & al. (2013b: 11, record Kubah N.P.);— Bárta & Dolný (2013: 70, 138, Kalimantan, Sg. Wain).

Dysphaea dimidiata limbata Selys, 1859 :— Selys Longchamps (1859: 443–444, reprint p. 9, description of male and female from Mt Ophir (Malay peninsula), Singapore and Sarawak as ‘ Dysphaea dimidiata, De Selys ; Race Dysphaea limbata, De Selys’);— Fraser (1942: 99, Perak, behaviour);— Lieftinck (1949: xi, limbata ranked as subspecies of D. dimidiata ).

Dysphaea limbata Selys, 1859 :— Selys Longchamps (1869: 660, reprint p. 16, upgraded to full species);— Selys Longchamps (1873: 487, reprint p. 23, correction of original description, discussion of variability);— Krüger (1898: 78, Sumatra, Soekaranda);— Laidlaw (1902: 88, Kelantan);— Laidlaw (1903: 194, Jalor [Yala];— Williamson (1904: 182–183, Lower Siam, Trang);— Ris (1911: 232–233, Malacca);— Laidlaw 1920: 328 (specimens from Tatau, Sarawak);— Kennedy 1920: pl. 1, Figs. 40–41 , penis);— Laidlaw (1931b: 241, West Central Borneo);— Lieftinck (1935: 4, mutual status of limbata and dimidiata discussed);— Lieftinck (1954: 19–20, listed as synonym of dimidiata ).

Dysphaea dimidiata semilimbata Selys, 1873 :— Selys Longchamps (1873: 486, reprint p. 22, description of male from Labuan as ‘ Dysphaea semilimbata, De Selys ; Variété ou race de dimidiata ?’);— Lieftinck (1954: 19–20, listed as synonym of dimidiata ).

Dysphaea lugens [nec Selys, 1873]:— Laidlaw (1920: 327, specimens from Limbang River and Retuh);— Matsuki & Kitagawa (1993: 2, specimens from Mulu N.P. are obviously D. dimidiata );— Orr (2001: 178, Brunei, all specimens from sites 15, 29–31, 35);— Orr (2003: 58, part);— Dow & Reels (2008: 3, Sarawak, Gunung Mulu N.P.);—Dow (2011: 48, Sarawak Planted Forest Project).

Study of the type material. Selection of lectotype of Dysphaea dimidiata Selys, 1853 . The type series preserved in Coll. Selys at IRSN (Brussels) consists of five pinned male specimens, all from West Java. To fix the identity of the nominal taxon Dysphaea dimidiata based on one specimen, we herewith designate the best preserved specimen of this series as lectotype. For the selected lectotype and its labels, see Fig. 5 . The measurements of the lectotype are Hw 32 mm, abdomen (apps. excl.) 35 mm, cerci 2 mm. The ratio of the length of abdomen (apps. excl.) and Hw is 1.10. The corresponding measurements of the paralectotypes are 30–33.5 and 34–38, the ratio being 1.13–1.17.

Selection of lectotype Dysphaea limbata Selys, 1859 . The type series of limbata consists of at least five pinned male specimens and one female. The males originate from Mt Ophir [Johor, Peninsular Malaysia], Singapore and Sarawak and the female specimen from Singapore, all collected by Alfred Russell Wallace (1823– 1913). The specimens from Sarawak and Singapore are in Coll. Selys. There are three known male specimens from Mt Ophir, one in Coll. Selys at IRSN (Brussels), one at the Übersee-Museum Bremen (see Seehausen 2014) and one at BMNH (B.M. 1938-674; ex. coll. McLachlan); the latter bears the labels “ D. limbata ”, “ Malacca ”, “ Paratype ”. To fix the identity of the nominal taxon Dysphaea limbata based on one specimen, we herewith designate the male specimen from Mt Ophir, kept in Coll. Selys at IRSN (Brussels), as lectotype. The lectotype and the attached labels are presented in Fig. 6 . The measurements of the lectotype are: Hw 30.5 mm, abdomen (apps. excl.) 36 mm, cerci 2 mm. The ratio of the length of abdomen (apps. excl.) and Hw is 1.18. The Hw length of the male paralectotype from Singapore is 28.5, the length of abdomen 34; the abdomen/Hw length ratio is 1.19. Corresponding figures for the Sarawak male paralectotype are 29, 34.5 and 1.19. The female paralectotype is a small broken specimen; its Fw is 29 mm.

Holotype of Dysphaea semilimbata Selys, 1873 . Described from a single male specimen from Labuan ( Fig. 8 ). This (pinned specimen) is preserved in Coll. Selys at IRSN (Brussels). Measurements: Hw 32, abdomen (apps. excl.) 37 mm, cerci 2 mm. Abdomen/Hw length ratio 1.16. The real type locality of semilimbata in Borneo is uncertain, it is possible that ‘Labuan’ [island off the west coast of Sabah] merely refers to the port from where the specimen was shipped to Europe.

Other material studied. Java.— RMNH: 39 ♂, 9 ♀, W. Java, 7 km SW of Djasinga, Tjibarangbang River, alt. 150 m, leg. M.A. Lieftinck; various dates in 1935–1939, in April–June, September–February: 28 iv 1935, 30 iv 1935, 12 v 1935, 14 vi 1935, 16 x 1935, 25 x 1935, 24 xi 1935, 6 xii 1935, 8 xii 1935, 20 ix 1936, 30 ix 1936, 1 xi 1936, 15 xi 1936, 7 ii 1937, 15 i 1939, 14 xi 1939, 26 xi 1939; 1 ♂, W. Java, 10 km NW of Djasinga, Tjibeurum River, 25 x 1935, leg. M.A. Lieftinck; 1 ♂, [Java], Soerabaja [= Surabaya], D. [= Danau] Ranau, 28 x [no year, no collector].— BMNH: 3 ♂, W. Java, Djasinga, Tjibarangbang River, 16 x 1935, leg. M.A. Lieftinck.

Billiton.— RMNH (all specimens in 1935–1937, leg. F.J. Kuiper unless noted otherwise): 2 ♂, Begantung, 17 iii 1936; 2 ♂, as above, 20 iv 1937; 1 ♂, as above, 27 v 1937; 1 ♂, O. Billiton, Kp. Ajer Lautjie, 21 x 1935; 2 ♂, G.

Tadjem, 20 xi 1936; 1 ♂, N.W. Billiton, Ajer Gelarak, 30 iv 1937; 1 ♂, 1 ♀, Tandjong Pandan, 7 xii 1949, leg. Cardinaal.

Sumatra.— RMNH: 1 ♂, [N. Sumatra], Atjeh [= Aceh], Losten, 400m, 22 v 1957, leg. A. Hoogerwerf; 1 ♂, N.E. Sumatra, Sg. Radja, 9 ix 1928, [leg. J.C.] v.d. Meer Mohr; 1 ♂, N.E. Sumatra, Deli, Dolok Ilir, Serbalawan, 20 ii 1948, leg. R. Straatman; 1 ♂, N.E. Sumatra, Serbalawan, Dolok Ilir, 90m, 15 iv1948, leg. R. Straatman; 4 ♂, N.E. Sumatra, Tandjong Morawa, Serdang, (no date), leg. Dr B. Hagen; 2 ♂, N.E. Sumatra, Deli, Laut Tador, 5 ii & 6 v 1951, leg. R. Straatman; 2 ♂, as above, alt. 60 m, 29 iv 1948; 1 ♀, as above, alt. 90 m, 10 vii 1948; 1 ♂, as above, 24 i 1950; 1 ♂, E. Sumatra, Kampar Kanan, 22 x 1925, leg. Fulmek & Karny; 2 ♂ ( RMNH.INS.557835, 557839), Riau province, Pekan Baru, Rama Rama, 18 ii 2014, leg. R.A. Dow; 4 ♂, C.W. Sumatra, Bengkulen, Ds. Bindjai, Air Musi, alt. 100 m, x 1941, leg. W.C. Verboom; 1 ♂, [C.W. Sumatra], Bengkulen, Lebong area, Air Kelemong, alt. 600m, 11 ix 1937, leg. F.W. Rappard; 1 ♀, [C.W. Sumatra], Silago, tot.12/777; 2 ♂, S. Sumatra, S.W. Lampongs, Mt Tanggamoes Giesting, Waiteboe [Wai Tebu], 29 xii 1934, leg. Lieftinck & Toxopeus; 3 ♂, S. Sumatra, Mt. Tanggamoes, Wai Tebu, 300m, 19–31 iii 1940, leg. M.A. Lieftinck; 1 ♂, S. Sumatra, Lampung distr., Borgen Est., Tg. Karang, 10 ix 1952, leg. A. Sollaart; 1 ♂, S. Sumatra, S.W. Lampongs, Terbanggi hilir (Menggala), 14 viii 1936, leg. M. Bartels.— BMNH: 1 ♂, W. Sumatra, Lebong Tandai, Benkoelen Residency, vii 1922, leg. C.J. Brooks; 1 ♂, as above, 1912–1919; 1 ♂, W. Sumatra, leg. C.J. Brooks; 1 ♂ Sumatra, McLachlan collection, ‘ limbata ’.— Coll. Dow: 2 ♂, Rama Rama, Pekan Baru, Riau province, 18 ii 2014, leg. R.A. Dow.— Coll. Hämäläinen: 1 ♂, Aceh, v 1990, ex. coll. Haruki Karube.

Borneo.— IRSN: 6 ♂, Borneo, Labuan; 3 ♂, Borneo W.[est-]K.[ust]. [= West Kalimantan]; 5 ♂, [Borneo], Sintang, Coll. Clément; N. Borneo, 1 ♂, Bukau, (Juill.).— RMNH: 1 ♂ ( RMNH.INS.229164), Brunei Darussalam, Temburong, Belalong River at Kuala Belalong Field Studies Center, 19 xi 2004, leg. K.D.B. Dijkstra & V.J. Kalkman; 4 ♂, Brunei Darussalam, Belait River, caught from boat, 11 ix 1992, leg. D. Thompson; 1 ♂, as above, 29 ix 1992; 1 ♂, Brunei Darussalam, 15 minutes walk south of Ingei base camp, 50 m E of Ingei River, ca. 4° 09’ N, 114° 43’ E. 6 x 1992, leg. D. Thompson; 3 ♂ ( RMNH.INS.557872, 557875, 557876), Brunei, Belait district, Sungai Ingei, 2 iii 2014, leg. R.A. Dow; 4 ♂, N. Borneo, E. Sabah, Lahad Datu 60 km W, Danum Valley Field Centre, Sungai Segama, 160–180 m asl., 4° 58’N, 117° 48’E, 3 v 1987, leg. J. van Tol; 1 ♂, as above, Sungai Palum Tambun, 1 v 1987; 1 ♀, as above, 17 iii 1987, leg. J. Huisman; 1 ♂ ( RMNH.INS.509902), Sarawak, Kuching division, Lundu, Kampung Sebako, Sungai Sebako, 28 vii 2013, leg. R.W.J. Ngiam; 1 ♂ ( RMNH.INS.509903), Sarawak, Kapit division, Lanjak Entimau Wildlife Sanctuary, Ulu Katibas, 22 viii 2013, leg. J. ak Awan & M. ak Adau; 1 ♂ ( RMNH.INS.501316), Sarawak, Kapit division, stream in disturbed forest between Kapit town and the Hose Mountains, 14 v 2010, leg. G.T. Reels; 1 ♂ ( RMNH.INS.509664), Sarawak, Kapit division, between Kapit town and Hose Mountains, stream on main Rimbunan Hijau logging road ca one hour from the Baleh river crossing, 18 vi 2013, leg. R.A. Dow; 1 ♂ ( RMNH.INS.509904), Sarawak, Bintulu division, Bukit Sarang Proposed National Park, Sungai Sarang, 13 x 2008, leg. R.A. Dow; 1 ♂ ( RMNH.INS.500943), Sarawak, Bintulu division, Sarawak Planted Forest Project, stream at Kakus acacia nursery, 12 xi 2010, leg. S. Stone; 1 ♂ ( RMNH.INS.501315), Sarawak, Bintulu division, Tubau, Sarawak Planted Forest Project, stream at Kemena Camp, 11 vi 2010, leg. R.A. Dow; 1 ♂ ( RMNH.INS.509901), Sarawak, Miri division, Gunung Mulu National Park, Bat Observatory stream, 5 ii 2006, leg. L. Southwell; 2 ♂, Borneo Exped. Dr Nieuwenhuis, Bloe-oe, 26 ix 1894; 10 ♂, 1 ♀, S. E. Borneo, Balikpapan, Mentawir River, 2–25 x 1950, leg. A.M.R. Wegner; 1 ♂, 1 ♀, as above, 12 xi 1950; 2 ♂, E. Borneo, Sangkoelirang, Kariorang, v 1937, leg. M.E. Walsh; 4 ♂, 1 ♀, as above, Babi Djuton, alt. 0 m, vi 1937; 1 ♂, as above, Batau Besi, v 1937; 1 ♂, E. Borneo, Tabang, Bengen River, 26 x 1956, leg. A.M.R. Wegner; 4 ♂, 1 ♀, S.E. Borneo, Moam, 12 xi 1950, leg. A.M.R. Wegner; 3 ♂, East Kalimantan, Gunung Meratus Protected Forest—ICTI, 1998–2000, leg. D.F.R. Cleary; 1 ♂, as above, 26 x 2000; 1 ♂ ( RMNH.INS.509982), East Kalimantan, Pasir district, Prayan Base Camp area, Sungai Prayan, 14 xi 2005, leg. J. van Tol; 1 ♂ ( RMNH.INS.509981), East Kalimantan, same area, Sungai Seranum, 20.xi.2005, leg. J. van Tol; 1 ♀, W. Borneo, Singkawang, Bengkajang Rd, 20 i 1932, leg. L. Coomans de Ruiter; 1 ♂, as above, 8 iv 1932; 1 ♂, W. Borneo, Singkawang, Penaring, 8 ii 1932, leg. L. Coomans de Ruiter; 1 ♂, W. Borneo, Singkawang, Kali Piong San, 8 xii 1931, leg. L. Coomans de Ruiter; 2 ♂, W. Borneo, Singkawang, Bengkajang, 23 ii 1934, leg. L. Coomans de Ruiter; 1 ♂, W. Borneo, Singkawang, Bakuan, 22 i 1934, leg. L. Coomans de Ruiter; 1 ♂, West Borneo, weg naar Bengkajang, 18 iii 1932, leg. L. Coomans de Ruiter; 1 ♂, 97.53.14. Kalimantan Tengah, Sangai-C92, 1998-2000, 13 vi 1997, leg. D.F.R. Cleary.— BMNH: 2 ♂, “SAR” [ Sarawak], leg. Wallace, “Saunders 68.3”; 1 ♂, “SAR”, “57/ /55”; 1 ♂, Sarawak, (“ D. limbata ”); 2 ♂, 1 ♀, Labuan, McLachlan collection.— CUMZ: 1 ♂, Sabah, Maliau Basin, tributary to Sungai Maliau near Maliau Basin Field Centre, 15 vii 2012, leg. S. Luke; 1 ♂, Brunei, SW, 23–27 vi 1995, leg. D.A.L. Davies.— Coll. Dow: 2 ♂ (1 ♂ in collection Hämäläinen), Sarawak, Kuching division, foot of Gunung Penrissen, Annah Rais, 19 iii 2005, leg. R.A. Dow; 1 ♂, same location, 26 i 2006, leg. G.T. Reels; 1 ♂, Sarawak, Kuching division, Kubah National Park, Matang Wildlife Centre, Sungai Rayu, 25 i 2006, leg. G.T. Reels; 1 ♂, same location, 4 i 2015, leg. R.A. Dow; 1 ♂, ♀ (in tandem), Sarawak, Kapit division, Lanjak Entimau Wildlife Sanctuary, Ulu Katibas, 22 viii 2013, leg. J. ak Awan & M. ak Adau; 1 ♂, same area as above, Sungai Katibas at mouth of Sungai Satap, 23 viii 2013, leg. J. ak Awan & M. ak Adau; 1 ♂, same area as above, Sungai Pemanca, 24 viii 2013, leg. J. ak Awan & M. ak Adau; 1 ♂, Sarawak, Bintulu division, Tatau, Anap Muput Forest Management Unit, Sungai Sawih, 17 xi 2010, leg. R.A. Dow; 1 ♂, Sarawak, Bintulu division, Samarakan area, Sarawak Planted Forests Project, small muddy stream with narrow forest buffer in acacia, 28 ii 2008, leg. R. A. Dow; 1 ♂, 1 ♀ (in tandem), same area as above, Sungai Gagak (stream with acacia to bank), 3 iii 2008, leg. R.A. Dow; 1 ♂, Sarawak, Bintulu division, Sarawak Planted Forests Project, stream with forest buffer in block T1C, 9 v 2011, leg. R.A. Dow; 1 ♂, Sarawak, Bintulu division, Binyo Penyilam Proposed National Park, Sungai Penyilam, 9 iii 2006, leg A. Juhin; 1 ♂, Sarawak, Bintulu division, Sarawak Planted Forest Project, stream in block K2L, 23 vi 2010, leg. R.A. Dow; 1 ♂, same area as above, stream at Kakus acacia nursery, 30 vi 2010, leg. R.A. Dow; 1 ♂, ♀, same location and date as above, leg. O. Tateh; 1 ♂, Sarawak, Bintulu division, Sarawak Planted Forest Project, stream at Kapur Camp, 1 vii 2010, leg. R.A. Dow; 1 ♂, Sarawak, Bintulu division, Sarawak Planted Forest Project, Bukit Mina Wildlife Corridor, Sungai Mina , 27 iv 2011, leg. R.A. Dow; 1 ♂, same location and date, leg. O. Tateh; 1 ♂, Sarawak, Bintulu division, Tubau, Sarawak Planted Forest Project, stream at Kemena Camp, 19 i 2008, leg. R.A. Dow; 1 ♂, same location as above, 18 x 2008, leg. R.A. Dow; 1 ♂, same area as above, stream in block E2N, 16 vi 2010, leg. R.A. Dow; 2 ♂, Sarawak, Miri division, mid Baram, Sungai Suan, 11 xii 2007, leg. G.T. Reels; 1 ♂, same area, Sungai Lamah, 7 ii 2015, leg. R.A. Dow; 1 ♂, Sarawak, Miri division, Gunung Mulu National Park, Bat Observatory stream, 16 iv 2005, leg. R.A. Dow; 1 ♂, same location as above, 5 ii 2006, leg. G.T. Reels; 1 ♂, same location as above, 30 xii 2007, leg. R.A. Dow; 2 ♂, Sarawak, Limbang division, Gunung Mulu National Park, Lubang Cina, Headhunters Trail, 11 ii 2006, leg. R.A. Dow; 1 ♂, same area as above, Sungai Mentawei and tributaries, 13 ii 2006, leg. R.A. Dow; ♂, same location and date as above, leg. L. Southwell; 2 ♂, Brunei, Belait district, Sungai Ingei, 2 iii 2014, leg. R.A. Dow.— Coll. Hämäläinen: 3 ♂, Sabah, Danum Valley Field Centre, Sungai Palum Tambun, 150–160m, 23–28 iv 1994, leg. M. Hämäläinen; 1 ♂, Sarawak, Kuching division, foot of Gunung Penrissen, Annah Rais, 19 iii 2005, leg. R.A. Dow; 1 ♂, Sarawak, Kuching division, Kubah National Park, Matang Wildlife Centre, Sungai Rayu, 20 i 2006, leg. R.A. Dow; 1 ♂, Sarawak, Sri Aman division, Sekerang (Skrang) River, Bunu Longhouse, 1° 19.2’ N, 111° 39.4 E, 11 vii 1997, leg. T. Donnelly; 1 ♂, Sarawak, Miri division, Gunung Mulu National Park, Bat Observatory stream, 16 iv 2005, leg. R.A. Dow.—Coll. Steinhoff: 2 ♂, Sarawak, Miri Division, Gunung Mulu National Park, stream below Kenyalong loop start and Bat Observatory, 27 iii 2014, leg P.O.M. Steinhoff; 1 ♀, same area as above, swamp pool/stream, next to Waterfall Trail, 4 iv 2014, leg P.O.M. Steinhoff; 1 ♂, same area as above, Sg. Melinau, up stream from [park] HQ, 20 iv 2014, leg P.O.M. Steinhoff; 1 ♂, same area as above, Sg. Paku tributary upstream from conflux, 7 v 2014, leg P.O.M. Steinhoff.

Palawan.—Coll. André Günther: 1 ♂, Philippines, South Palawan, Quezon, 14 iii 1992, A. Günther leg.

Peninsular Malaysia.— RMNH: 1 ♂ ( RMNH.INS.503913), Terengganu, Sekayu Recreational Forest, Sungai Peres, 19 viii 2011, leg. R.A. Dow; 1 ♂, Perak, Jor Camp, alt. 1200m, leg. A. Grubauer; 2 ♂, Perak, Sg. Ternom/S. Plus, 22 vii 1961, leg. H.T. Pagden; 1 ♂, C. Perak, 10 mi N. of Ipoh, Sungai Chepor, 6 iii 1963, leg. M.A. Lieftinck; 1 ♂, C. Perak, Phu River area, Sei Chior, 7 iii 1963, leg. M.A. Lieftinck; 1 ♂, Selangor, Ampang Intake Res., 24 iv 1961, H.T. Pagden & M.?; 2 ♂, Selangor, Ulu Gombak, alt. 200m, 16–17 iii 1963, leg. M.A. Lieftinck; 2 ♂, Selangor, Kuala Lumpur, Templer Park, 21 iii–11 iv 1963, leg. M.A. Lieftinck; 1 ♂, Pahang, Bilut Valley, 7 x 1960; 1 ♂, Pahang, 55 km N of Jerantut, surroundings of Park Headquarters of Taman Negara near village of Kuala Tahan, 13 vii 2002, leg. V.J. Kalkman; 1 ♂ ( RMNH.INS.501089), Pahang, Kuala Tahan, tributary of Sungai Tembeling, 10 xii 2010, leg. R.A. Dow; 1 ♂, [Johor], Mt Ophir; 1 ♂, S. Johor, Bahru, Sei, Mapor, K. Tinggi, Mawai Rd, 17 iv 1963, leg. M.A. Lieftinck; 1 ♂ ( RMNH.INS.506832), Johor, Gunung Belumut, 2 viii 2012, leg. R.A. Dow.— BMNH: 1 ♂, Malacca (?); 2 ♂, [Pahang], Kuala Teku, alt. 500–1500 ft., 31 vii 1905, leg. H.C. Robinson; 1 ♂, Pahang, Kuala Teku, Pahang, 7 xii 1921; 1 ♂, Pahang, Kuala Tahan, 300 ft, 22 xi 1921, leg. H.M. Pendlebury; 1 ♂, 1 ♀, Perak, Kuala Kangsar, Kenas dam, 9 i 1938.— CUMZ: 2 ♂, Selangor, Templer Park, 16 x 1995, (ex coll. Davies).— Coll. Dow: 1 ♂, Terengganu, Sekayu Recreational Forest, Sungai Peres, 19 viii 2011, leg. F-N. Yong; 1 ♂, Terengganu, Bukit Kesing Recreational Forest, 23 viii 2011, leg. R.A. Dow; 1 ♂, Perak, Kuala Woh, 27 ix 2008, leg. R.A. Dow; 2 ♂, ♀ (in tandem with one of the males), Johor, Gunung Belumut, 4 viii 2012, leg. R.A. Dow.— Coll. Hämäläinen: 1 ♂, Perak, on road up to Cameron Highlands, 1986, leg. Brandstetter, ex coll. D.A.L. Davies; 6 ♂, Kelantan, Sungai Galas, 13 vi 1994, leg. Mohd. Zakaria-Ismail; 2 ♂, Kelantan, Sungai Selieh (tributary of Sg. Berok), near Kampung Selieh, 4° 42’ 55’’N, 101° 40’ 15’’E, 14 vi 1994, leg. Mohd. Zakaria-Ismail; 3 ♂, Kelantan, 31 km E of Fort Brooke, Sungai Selieh, 4°43’12”N; 101°39’22”E, alt. 365 m, 10–13 iv 1995, leg. M. Hämäläinen; 1 ♂, Kelantan, 58 km E of Fort Brooke, Sungai Lasau, 4°44’51”N; 101° 45’14”E, 10–13 iv 1995, leg. M. Hämäläinen; 2 ♂, Pahang, near Kuala Pian, Sungai Krau, 3°47’20”N; 102° 14’10”E, 22 vi 1994, leg. Mohd. Zakaria-Ismail; 1 ♂, Pahang, Sungai Krau, 3°49’20”N; 102° 13’25”E, 24 vi 1994, leg. Mohd. Zakaria-Ismail; 2 ♂, Pahang, Sungai Perlok, 3°50’45”N; 102°13’ 00”E, 21 vi 1994, leg. Mohd. Zakaria-Ismail; 1 ♂, Pahang, Sungai Lompat, 3°41’40”N; 102° 12’40”E, 24 vi 1994, leg. Mohd. Zakaria-Ismail; 1 ♂, Pahang, Krau Wildlife Reserve, Kuala Lompat, Sg. Lompat, 22–27 xi 1996, leg. M. Hämäläinen & H. Olsvik; 5 ♂, 1 ♀, Pahang, Krau Wildlife Reserve, Bukit Rengit, Sg. Rengit, 18–23 iv 1995, leg. M. Hämäläinen; 3 ♂, 1 ♀, as above, 21–23 i 1996; 2 ♂, 1 ♀, as above, 15–20 xi 1996; 3 ♂, as above, 5–8 vi 1997; 1 ♂, as above, 24–27 xi 1998; 2 ♂, as above, 18–19 xii 2003; 1 ♂, as above, 7–8 ii 1997, leg. H-S. Yong; 1 ♂, Pahang, Krau Wildlife Reserve, Kuala Gandah, Sg. Gandah, 22 iv 1995, leg. M. Hämäläinen; 1 ♂, as above, 18 xi 1996; 3 ♂, Pahang, Krau Wildlife Reserve, Pos Perlok, Sg. Perlok, 28 xi 1996, leg. M. Hämäläinen & H. Olsvik; 1 ♂, Pahang, river 1 km south of main Temerloh/ Kuantan road, on road to Kampung Rambai/Lake Chini, 15 ii 1988, leg. R.G. Kemp; 1 ♂, Selangor, Kerling, Bukit Tarek, Sungai Jerneh, 4 vi 1997, leg. M. Hämäläinen; 1 ♂, Johor, Panti Forest, alt. 66m, 25–26 iv 2009, leg. M. Hämäläinen.

Singapore.— BMNH: 1 ♂, “ Singapore ”, “96-163”. It is uncertain whether this specimen is part of the type series.

Thailand.— RMNH: 1 ♂ ( RMNH.INS.509983), Narathiwat, Sungai Ko-Lok, Pru Tuh Daeng peat swamp forest, 5 vi 2003, ex. coll. Amnuay Pinratana.— BMNH: 1 ♂, Malacca, Mabek, Jalor [= Siam, Yala], 24.7.1901, leg. Annandale, “ D. limbata ”.— CUMZ: 1 ♂, Yala, 18 iii 1985. [Rest of labels illegible]. (Ex coll. D.A.L. Davies).— Coll. Hämäläinen: 1 ♂, Songkhla, Ton Nga Chang waterfall, 28 iii 1993, ex coll. Amnuay Pinratana; 2 ♂, as above, 25 iv 1996; 1 ♂, Songkhla, Boripat waterfall, 27 i 1995, ex coll. Amnuay Pinratana; 1 ♂, Narathiwat, Sungai Ko-Lok, Pru Tuh Daeng peat swamp forest, 5 vi 2003, ex. coll. Amnuay Pinratana.

Descriptive notes on D. dimidiata . Topotypical specimens from Java.

Diagnosis. Male: Black-bodied species with much of the wing surface opaque blackish; hyaline area longer in Fw. Cerci with lower border distinctly arched near base in lateral view. Terminal segment of penis with two long apical arms, appearing sub-rectangular in ventral view.

Male (for habitus see Figs. 5 , 9 ). Head: Labrum, base of mandibles and clypeus shining black, frons and vertex matt black. Thorax: Matt black, with pale brownish stripe on mesepimeron bordering first lateral suture; much of metepisternum pale brownish; metepimeron with small apical pale brownish patch near wing base and with ventrolateral patch in the apical half of metepimeron ( Fig. 13 ). Venter of thorax with or without sparse, tiny tubercles on metaposternum, the tubercles never as dense as in Bornean specimens (cf. Fig. 17 ). Legs wholly black.

Wings: Most of wing surface opaque black; extent of hyaline area between dark base and apex somewhat variable. Opaque areas of wings with a distinct bluish-violet reflections (cf. Fig. 79 ). In Fw of lectotype ( Fig. 5 ) basal opaque area extends to level of nodus; in costal field one cell beyond nodus. In other specimens ( Figs. 19–20 ) this character is somewhat variable, extending to level of 3–4 cells before or beyond nodus (rarely 5 cells before or 6–7 cells beyond); but in costal field always at least to level of nodus, although occasionally the colour is incomplete in the last 1–6 cells. In Hw of lectotype opaque area extends to level of 10 cells beyond nodus; in other specimens to level of 1/3–1/2 of distance between nodus and pterostigma. Costal field between the opaque areas is hyaline in both wings, or at most only slightly darkened at proximal and distal ends.

Abdomen: Matt black throughout in dead mature specimens, rarely with very faint and incomplete intersegmental rings on one or two segments; these more developed in living specimens. Appendages black; cerci twice as long as S10, subcylindrical, widely separated at base, hollowed out interiorly in the apical half; apices curling and meeting or overlapping ( Fig. 31 ); in lateral view ventral margin of cercus distinctly arched near base ( Fig. 37 ). Paraprocts very short, rounded and featureless.

Penis: Terminal segment with two long, flat, rectangular, apical arms directed straight outwards and upwards, then turned downwards towards apex ( Figs. 41 , 45 ).

Measurements (mm): Hw 29.5–34 (in lectotype 32), abdomen (apps. excl.) 34–38 (in lectotype 35); cerci 1.5– 2. Abdomen/Hw ratio 1.10–1.17.

Female (for habitus see Fig. 49 ). Head: Labrum shining black, with broadly yellow centre, indented with black basally. Base of mandibles largely yellow with a black incomplete stripe medially. Clypeus shining black with obscure yellow stripe or marking anteriorly. Frons matt black with broad yellow stripe throughout antefrons, stripe narrower in the middle, connected to yellow genae (cf. Fig. 53 ). 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 and with yellow spots on sides and centrally on hind lobe. Hind lobe largely black, usually with lateral ends yellow and a yellow marking medially. Posterior part of hind lobe raised obliquely upwards to form an elongate rectangular flaplike process, the lateral ends of which are often curled ( Figs. 57 , 63 ). The median part between the curled lateral parts is typically less raised, usually appearing at least slightly concave in dorsal view, occupies little more than 1/2 length of hind lobe. Synthorax matt black, with extensive yellowish markings as in Figs. 63 , 69 . Broad stripes on mesepisternum always forming complete loop, those on mesepimeron not joining to form loop at wing base. Metepisternum and metepimeron nearly completely yellowish, black stripe covering both sides of second lateral suture in apical half; a small black area in middle of metepimeron. Legs black, coxa broadly yellow on sides; short, basal pale stripes on flexor surface of middle and hind femora.

Wings: Hyaline, slightly brownish tint in basal half of Fw (to the level of nodus) and much more extensively on Hw, typically to level of halfway between nodus and pterostigma, occasionally almost to pterostigma. Tips of both wing tinted slightly darker brown; in Hw dark apical area larger than in Fw.

Abdomen: Matt black, with yellowish lateral markings. S1 with lateral spot covering much of segment. S2–7 with lateral stripes, rather broad on S2–3, gradually becoming narrower towards apical segments; on S3–7 pale area projecting dorsally to form a “tooth” at base of each segment, that on S6–7 being disconnected. S8 usually with only a tiny, pale spot apically but occasionally also with a tiny basal spot, S9 with distinct lateral marking at apical half, ventrolateral edge of S8 obscurely pale at apical half to two-thirds, that of S9 typically pale at basal third, sometimes along whole segment length and occasionally fused with the lateral marking. The markings on S8 and S9 often differ between left and right sides of the same individual. Appendages black.

Measurements (mm): Hw 30–33.5; abdomen (apps. excl.) 30.5–33, cerci 1.

Variation in specimens from outside Java. Male. Thorax: Mature specimens from other areas of its range usually have fewer distinct pale markings on synthorax than those from Java, synthorax being either wholly black or having only obscure pale markings: stripe on metepisternum, extending from below stigma apicad above the second lateral suture, not reaching wing base; metepimeron obscurely pale ventrolaterally. Venter of synthorax either with or without tiny tubercles on metaposternum; tubercles present in all Bornean specimens studied ( Fig. 17 ), with the exception of one male ( RMNH) from Sungai Palum Tambun in the Danum Valley. Wings: In Fw basal opaque area extends at least to level of 3–4 cells before nodus, but usually extending a few cell rows or more beyond nodus (see Figs. 21–24 ). In Hw opaque area extends always beyond nodus, typically to level of 1/3–1/2 of distance between nodus and pterostigma. Extent of opaque colouration on wings is often subject to individual variation in same populations. Geographical variability is less clear, but in the northernmost populations in southern Thailand the opaque area is usually similar to that illustrated in Figs. 20–21 . In a specimen (labelled ‘Labuan’) in Coll. Selys ( IRSN), opaque area in Hw extends almost to proximal end of pterostigma and in Fw almost to halfway between nodus and pterostigma ( Fig. 24 ). Costal field between nodus and pterostigma is usually opaque in both wings (therefore named ‘ limbata ’ by Selys), but there is rare individual variability, in some specimens the costal field in Fw hyaline. In some of these ‘ semilimbata ’ specimens the costal field in Hw is only slightly darkened. Abdomen: usually entirely black, occasionally with very faint dorsal lateral marks at base of S4– 7, slightly more distinct in living specimens ( Figs. 78–79 ). Measurements (mm): Hw 27–33.5, abdomen (excl. cerci) 31–38, cerci 1.5 –2. Abdomen/Hw ratio 1.10–1.19. Female. Thorax: Pale markings on synthorax are less extensive than in Javan specimens; especially on metepisternum and metepimeron the extent of black colour is distinctly greater ( Figs. 64–65 , 70–71 ). Wings: Quite similarly coloured to Javan specimens, but in many specimens whole surface of Hw is slightly brown tinted, with at most only a narrow irregular untinted area at proximal end of pterostigma; tint darker at wing tip. Abdomen: Yellow lateral stripes slightly narrower than in Javan specimens, dorsally projecting basal “tooth” on S3–7 is usually longer and more distinct. Measurements (mm): Hw 29.5–33.5, abdomen 29–33, cerci 1.

Remarks. The lateral ends of the ‘flap’ of the hind lobe of the prothorax are often but not always curled upwards (cf. Figs. 57–59 ). The curvature of the ends of the flap might be the result of tandem formation with males; the female from Silago, Sumatra, is teneral in appearance and among the minority of specimens where the ends of the flap are uncurled. Fig. 59 shows a female from Billiton where the ends of the flap are not curled.

Figures

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 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 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 5 – 8. Selysian type specimens and labels: 5) D. dimidiata lectotype; 6) D. limbata lectotype; 7) D. lugens holotype; 8) D. semilimbata holotype.

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 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.

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 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.