Dysphaea lugens Selys, 1873

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 : 468-472

Dysphaea lugens Selys, 1873

Selected references:

Dysphaea lugens Selys, 1873 :— Selys Longchamps (1873: 485–486, reprint p. 21 (description of male, South Borneo, as ‘ Dysphaea lugens, De Selys ; Race de dimidiata ?’);— Ris (1911: 232–233; Sintang, Borneo; description of ♀);— Kennedy (1920: pl. 1, Figs. 32–33 , penis);— Laidlaw (1924: 300, characters discussed);—Laidlaw (1931: 241, West Central Borneo);—Coomans de Ruiter (1936: 74–76, Fig. 3, W. Borneo, notes);— Lieftinck (1953: 382, S. Borneo);— Lieftinck (1954: 20, distribution, habitat, references);— Asahina (1985: 31, 33, 34, 36, Figs. 48–50 penis, Fig. 68 (male wings, Sarawak);— Orr (2003: 58, part).

Dysphaea dimidiata [nec Selys, 1853]:— Orr (2002: 291, Central Kalimantan).

Dysphaea limbata ‘race semilimbata ’ [nec Selys, 1873]:— Kimmins (1936: 78, specimens from River Kapah and junction of rivers Tinjar and Lejok; Mt Dulit area).

Study of the type material. Holotype. D. lugens was described from a single male specimen, which was sent to Selys by Robert McLachlan. Coll. Selys at IRSN (Brussels) includes nine pinned male specimens of D. lugens from Borneo (see below). Only one of them ( Fig. 7 ) has the same measurements given in the original description, i.e. abdomen (incl. cerci) 40 mm, Hw 31. It is also the only specimen with the cerci in a crossed position, which matches the single illustration of D. lugens in the portfolio of coloured paintings of Odonata species in Coll. Selys, executed in the late 1800’s. In September 2014 only a single small yellow handwritten label ‘Lansbg.’ was attached to this specimen. ‘Lansbg.’ refers to Johan Wilhelm van Lansberge (1830–1903), who resided in the Netherlands East Indies from 1875 to 1881. He could not have been the collector of the holotype of species described in 1873. No doubt a mix-up in the labels had taken place at some phase after Selys’ time. This is evident also by the fact that the only identification label in Selys’ handwriting was attached to a specimen which does not match the measurements of the holotype. This label reads: “ Dysphaea var. lugens Selys / ♂ / Sintang”. Later someone (not Selys) has crossed out the locality name ‘Sintang’ and replaced it with ‘W.K.’, which means West-Kust (an area presently known as West Kalimantan). We have removed the wrong collector label and restored the Selysian identification label to the holotype, which obviously was collected in Sintang in West Kalimantan, although in the original description reads only ‘Le sud de Bornéo’.

Other material studied. IRSN: 2 ♂, Borneo W.[est-]K[ust]., Sintang, Coll. Clément; 2 ♂, Borneo W.K., Coll. Clément; 1 ♂, Borneo, N. Bartan, Coll. Clément; 2 ♂, [Borneo], leg. J. W. van Lansberge; 1 ♂, N. Borneo, D. limbata S. [nec Selys 1859].— RMNH: 2 ♂, S. Borneo, Sampit (0–50 m), Pemantang, 28 vii 1953, leg. M.A. Lieftinck; 3 ♂, S.E. Borneo, Ampah (0–20 m), iv–v 1948, leg. L.S. Liong; 1 ♀, E. Borneo, Gunung Sari, alt. 95 m, 2 viii 1956, leg. A.M.R. Wegner; 7 ♂, E. Borneo, Tabang, Bengen River, 20 viii–26 x 1956, leg. A.M.R. Wegner; 1 ♂, E. Borneo, Kutai, Damay Kendeng Pau, 17 ix 1938, leg. B.M. Hoeks; 1 ♂, W. Borneo, M., Singkawang, Bakuan, Selakau River, 16 ix 1932, leg. L. Coomans de Ruiter; 3 ♂, as above, but date 9 iv 1934; 1 ♂, 1 ♀, Borneo Exped. Dr Nieuwenhuis, Bloe-oe, Nov. 1898; 2 ♂, N. Borneo, Sei Bekan, 16 viii 1961, Exp. P. Jauffret & R. Pujol, (ex coll. A. Heymer); 1 ♂, Centre Borneo, Kapuas River, Sepauk, 20 viii 1961, Exp. Jauffret; 1 ♂, Centre Borneo, Kapuas River, Environs de Silat, 24 viii 1961, Exp. Jauffret; 2 ♂, N. Borneo, E. Sabah, Lahad Datu 60 km W, Danum Valley Field Centre, Sungai Segama, 160 m asl., 3 v 1987, leg. J. van Tol.— BMNH: 4 ♂, [ Sarawak, foot of Mt Dulit], R. Tapah, tributary of Tinjar, 21–22 ix 1932, leg. B.M. Hobby and A.W. Moore, [published by Kimmins (1936) as ‘ D. limbata semilimbata ’]; 1 ♀, [ Sarawak, foot of Mt Dulit), Junction of rivers Tinjar and Lejok, 2 x 1932, leg. B.M. Hobby and A.W. Moore; 2 ♂, N. Borneo [?Sabah], Coll. McLachlan.— CUMZ: 1 ♂, 1 ♀, C. Borneo, Kalimantan, vii–viii 1992, leg. C. Jiggins, [ex coll. D.A.L. Davies].— Coll. Hämäläinen: 1 ♂, E. Borneo, Tabang, Bengen River, 10 x 1956, leg. A.M.R. Wegner [ex coll. RMNH].

Descriptive notes on D. lugens . Diagnosis. Male: A species with a proportionally longer abdomen than other Sundaland Dysphaea species. Opaque area on wings wider than in most D. dimidiata specimens, colour of wings paler than in D. dimidiata , brownish opaque rather than blackish opaque. Apical arms of terminal segment of penis shorter than in the other species occurring in Borneo, each with a small squarish, apical expansion

Male (for habitus see Figs. 7 , 10 ). Head: Labrum, base of mandibles and clypeus shining black, frons and vertex matt black, many individuals with two small faint brownish spots anterior to median ocellus; two specimens have an additional pair of small spots, placed outside the lateral ocelli. Thorax: Matt black, with obscure brownish stripes on synthorax as in Fig. 14 ; in many (more mature?) specimens there are stripes only on metepisternum and metepimeron. Narrow and usually very faint bronzy antehumeral stripes extending from ca 1/3 to 1/2 length are present on many specimens. Venter of thorax without tiny tubercles on metaposternum. Legs black, partly dark brownish.

Wings: In Fw basal brownish opaque area usually extends 5–7 cells beyond nodus but sometimes as few as 2 or as many 11 cells beyond, in Hw much further apicad, usually to level of 3–7 cells proximal to pterostigma ( Fig. 7 , 25 ) but sometimes reaches the pterostigma. The opaque patch at wing tips extends anteriorly to level of distal end of pterostigma, or a little more apicad. In some specimens there remains only a narrow hyaline area between the opaque areas ( Fig. 26–27 ).

Abdomen: Matt black; S1 with obscure brownish patch; S2–5 with narrow, brown lateral stripes, stripe on S2 placed midlaterally, those on S3–5 ventrolaterally. Appendages ( Fig. 32 ) black, similar to D. dimidiata ; in lateral view ventral margin of cercus distinctly arched at base ( Fig. 38 ). Penis: Terminal segment with two short apical arms, directed outwards, upwards, then downwards; transversely expanded at apex ( Fig. 42 , 46 ).

Measurements (mm): Hindwing 30–35 (usually 32–34), abdomen (apps. excl.) 37–43 (usually 39–42), cerci 2 mm. Abdomen/Hw ratio 1.20–1.25.

Female (for habitus see Fig. 50 ). 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 incomplete black stripe medially. Anteclypeus shining black with narrow yellow stripe on postclypeus along ridge. Frons matt black, yellow below antennae except for a transverse central stripe, yellow colour continuing over much of genae, and narrowly along eye margin almost to narrowest point of vertex. Antennae black and dark brown, yellow anterior mark on scape. Vertex and occiput matt black, with a transverse yellow stripe on either side of the median ocellus, and a yellow spot adjacent to each lateral ocellus ( Fig. 54 ), in the females from Bloe-oe and Mount Dulit these marks joined on one (Dulit) or both sides; two tiny yellow spots on occiput.

Thorax: Prothorax black with rounded large yellow spots on either side of dorsum of middle lobe, small yellow mark just anterior to central pit, pair of smaller dorsal spots on anterior lobe. Posterior part of hind lobe raised upwards to form an elongate rectangular flap; lateral parts almost perpendicular to base and central part at an oblique angle, appearing as a square tongue in dorsal view ( Fig. 60 ); flap narrowly bordered by yellow, more broadly so at sides, with a separate paler area centrally in the female from Bloe-oe (not illustrated). Ends of flap prominent laterally. Synthorax matt black, with yellow stripes as in Figs. 66 , 72 . Yellow stripes on mesepisternum and mesepimeron forming loops. Metepisternum and metepimeron mostly yellow. Legs black, yellow markings on both apical and posterior sides of coxae, small yellow mark upper anterior femur, large yellow mark upper ca 1/3 middle femur and ca 2/3 posterior femur.

Wings: Hyaline with slight brownish tint around costal and subcostal space from base to nodus in both wings and faintly darkened area at extreme wing tips ( Fig. 50 ). Fw with 32–36 antenodals in first row; Hw with 28–31. Quadrangle with 3–5 crossveins. Pterostigma long, covering 10–12 underlying cells in Fw, 9–10 in Hw.

Abdomen: Matt black, with pale markings as follows: S1 mostly yellow laterally, extending upwards and downwards at apex of segment. S2–7 with yellow lateral stripe, becoming orange on S6–7, broadest on S2–3 and gradually narrowing towards apical segments, narrowly interrupted at near base from S4 or S6. Separate small lateral spots at apex of S6–7. Faint, interrupted, narrow orange lateral stripe on S8, absent on one female, only apical part present on another, irregular orange lateral mark on S9. Appendages black.

Measurements (mm): Hw 30.5–34, abdomen (apps. excl.) 33.5–35, cerci ca 1.

Remarks. Unfortunately no fresh specimens of D. lugens were available for the molecular study. However, due to the considerable structural differences, the specific status of D. lugens is not in doubt. The association of female specimens with D. lugens is a supposition based on occurrence at the same sites and differences from D. dimidiata females. The fact that some males bear faint markings on the vertex in equivalent positions to the females (never seen in the other species occurring in Borneo) lends support to the association of the sexes. The description by Ris (1911: 232) of female D. lugens mentions “between antenna and ocelli weakly reddish-brown spots”. It seems likely that the markings on the vertex of males are age dependent, fading with maturity.

Figures

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 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 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 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 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 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.