Coeliccia matok, Dow, Rory A., 2016

Dow, Rory A., 2016, Revision of the genus Coeliccia Kirby in Borneo part II: Two new species from the membranipes- group, with a redescription of C. macrostigma Laidlaw (Odonata: Zygoptera: Platycnemididae), Zootaxa 4184 (1), pp. 79-103 : 88-94

Coeliccia matok spec. nov.

( Figs. 1 , 8, 9, 10 , 17, 18, 19 , 25, 26 , 31, 32 , 37, 38 , 43, 44 , 49, 50 , 55, 56 , 61, 62 , 66 , 69 , 72 )

Coeliccia new species;— Dow & Reels 2011: 51 (Matang Road near Kuching);— Dow & Reels 2013: 19, Fig. 10 (UNIMAS campus).

Coeliccia species;— Dow 2016a: 6 (Samunsam Wildlife Sanctuary).

Coeliccia sp.;— Laidlaw 1918: 230 (male “Platang Rd.”, brief description).

Coeliccia octogesima [nec Selys, 1863];— Laidlaw 1918: 231 (female, Matang Road)?

Type material: Holotype: Ƌ ( SAR 07 _8_PCD199), peat swamp forest, Universiti Malaysia Sarawak ( UNIMAS) campus, Kota Samarahan, Samarahan Division , 25 ii 2008, leg. R.A. Dow, to be deposited in BMNH .

Paratypes. All Sarawak, Malaysia, in collection Dow unless noted otherwise (42 Ƌ, 12 ♀): 2 Ƌ ( SAR 15 _PCD31–32), small tributary of Sungai Samunsam in kerangas forest, Samunsam Wildlife Sanctuary, Kuching Division, 15 viii 2015, leg. RAD; 3 Ƌ ( SAR 15 _PCD3–5), small streams in mixed dipterocarp forest, same area, 20 viii 2015, leg. RAD; 1 Ƌ ( SAR 06 _PCD133) , 1 ♀ (SAR06_PCD134, in tandem with Ƌ), remnant peat swamp forest, Matang Road, just outside Kuching, Kuching Division, 22 i 2006, leg. R.A. Dow; 2 Ƌ ( SAR 06 _PCD135–136), same location, 23 i 2006, leg. R.A. Dow ; 1 Ƌ ( SAR 06 _PCD157), same location, 23 i 2006, leg. G.T. Reels ; 1 Ƌ (SAR09_10_PCD252, RMNH.INS.503574, in RMNH), 2 Ƌ (SAR09_10_PCD250, 253), 2 ♀ ( SAR 09 _10_PCD251, 254), same location, 30 v 2010, leg. R.A. Dow ; 1 ♀ ( SAR 09 _10_PCD256, RMNH.INS.503575), same location and date, leg. G.T. Reels ; 1 Ƌ (SAR05_PCD111), location as holotype, 30 v 2005, leg. R.A. Dow; 1 ♀ ( SAR 05 _PCD110), same location, 2 vi 2005, leg. R.A. Dow ; 1 Ƌ ( SAR 06 _PCD137), same location, 24 i 2006, leg. R.A. Dow ; 1 Ƌ (RMNH.INS.229066, in RMNH), 20 Ƌ (SAR07_8_PCD44, 193 – 198, 200–210, 325–326), 1 ♀ (SAR07_8_PCD45, in tandem with SAR07_8_PCD44), 2 ♀ (SAR07_8 _PCD191– 192), 1 ♀ (SAR07_8_PCD211, in tandem with SAR07_8_PCD210), 1 ♀ SAR 07 _8_PCD327, in tandem with SAR 07 _8_PCD326), same location, 25 ii 2008, leg. R.A. Dow ; 1 Ƌ ( SAR 09 _10_PCD192, RMNH. INS.503572, in RMNH), 1 Ƌ ( SAR 09 _10_PCD193, RMNH.INS.503571, in RMNH), 1 Ƌ ( SAR 09 _10_PCD194, RMNH.INS.503573, in RMNH), 1 Ƌ ( SAR 09 _10_PCD195), same location, 7 vi 2010, leg. R.A. Dow ; 4 Ƌ ( SAR 11 _12_PCD445–448); 2 ♀ ( SAR 11 _12_PCD449–450), same location, 2 iv 2012, leg. R.A. Dow .

Etymology. Matok , a noun in apposition, named for Marijan Matok in gratitude for his continued financial support of the International Dragonfly Fund.

Description of holotype male. Head: Labium pale cream except hooks of labial palps, which are black. Labrum shining black with small, obscure basal pale marks. Mandible bases shining black, pale along most of lower and anterior margins. Clypeus and lower part of genae shining black, pale blue band running from eye margin to clypeus and narrowly for short distance above clypeus ( Fig. 9 ). Antenna with top part of scape, base of pedicel white, remainder dark brown and black. Frons and vertex mostly black, with small pale mark between each lateral ocellus and median ocellus, short stripe based on outer edge of lateral ocellus, directed towards rear of antenna base, just joined to small spot behind antenna base, small pale mark at eye margin, faintly joined on left side to mark behind antenna base ( Fig. 8 ). Ocelli white. Yellow, elongate oval, transverse postocular spots. Underside of head ( Fig. 10 ) black with large bluish white marks at eye margin, at same level as, and narrowly connected with, pair of pale markings below point of attachment of prothorax, behind mandible bases.

Thorax: Prothorax with propleuron entirely pale except narrow black band at rear in upper part. Posterior and anterior pronotal lobes black, except narrowly laterally on anterior lobe, where pale. Anterior carina of anterior pronotal lobe lower than main part, which is relatively narrow, rounded at top and slightly sloping to rear ( Fig. 26 ). Upper part of notopleural projection present as small rounded lump fused with anterior pronotal lobe, small clump of setae on upper part. Middle lobe mostly pale laterally, widely black dorsally ( Fig. 25 ). Posterior pronotal lobe simple, collar like, raised along free margin. Synthorax with mesepisternum black with large blue antehumeral markings ( Fig. 37 ), extending from mesostigmal plates to ca apex of antealar triangle, occupying most of width of mesepisternum near prothorax, outer margin following mesepleural suture for most of length, inner margin ca semicircularly but irregularly excised centrally, marking narrower after excision. Mesepimeron black except narrowly blue above the interpleural suture for some distance and tiny pale mark in upper corner adjacent to antealar carina ( Fig. 38 ). Metepisternum largely blue with black stripe running from antealar carina most of way towards spiracle. Metepimeron pale. Venter of synthorax pale. Mesinfraepisternum black except lower corner adjacent to metepisternum, metinfraepisternum entirely pale. Legs with coxae and trochanters entirely pale, femur mostly pale with black stripe along extensor surface, black stripe along outer flexor surface in distal ca 2/3 of anterior femur only, tibia mostly pale, dark along flexor surface, tarsi pale with obscure darker areas, claws brown. Wings with arc situated slightly distal to Ax 2 (Fw) or at Ax2 (Hw). Fw with 15 Px, Hw with 14 Px. Three post quadrilateral cells in all wings. R4 at (Fw) or very slightly proximal (Hw) to Sn. Pt dark brown with narrow white margin except along costal edge, broad, almost a rhombus, covering one underlying cell entirely and ca 1/2 to all of another.

Abdomen: S1 whitish except black in narrow apical annulus including posterior carina and area behind, narrowly on dorsum. S2 pale laterally, dark brown dorsally, this widest apically; narrow pale stripe centrally on dorsum basally. S3–7 pale lower laterally, becoming brown dorsally, this darkening on successive segments so black on S7, pale basal annulus, interrupted dorsally, on each segment. S8 pale blue lower laterally, this extending dorsally subapically as irregular transverse band, very narrowly divided centrally, otherwise black ( Fig. 49 ). S9 almost entirely pale blue, except small black basal dorsal triangle. S10 pale blue. Cerci mostly blue laterally and dorsally in outer half, black interiorly except apically where pale, black at tip; short basal tooth visible in dorsal view, central tooth after half-length visible in lateral view. In dorsal view ( Fig. 61 ) cercus subrectangular, apex at outer side, distinct ridge running diagonally forwards from apex almost to inner margin. In lateral view ( Fig. 62 ) almost rectangular. Paraprocts mostly pale in visible parts, relatively short, tips a little short of those of cerci; in ventral view narrowing gently after base along outer margin, then curving gently out again before turning in just before apices, where turn sharply upward, along inner margin gently curved in basal half, then with an excised appearance ( Fig. 66 ). In lateral view paraprocts narrow after base, directed gently upwards towards cerci, almost straight along lower margin, rounded where turning inwards, actual upturned apices just visible. Genital ligula typical for membranipes -group.

Measurements (mm): Abdomen without anal appendages 34 mm, cerci 0.8 mm, Hw 21 mm.

Description of female paratype (SAR09_10_PCD195). As male except as noted. Head: Pale markings on labrum larger, mandible bases almost entirely pale except for narrowly adjacent to clypeus and gena ( Fig. 18 ). Pale markings around ocelli more extensive, continuous ( Fig. 17 ). A very narrow, short pale stripe along eye margin in line with postocular markings.

Thorax: Cervical spurs well developed, lower one protruding outwards and to rear, barely overlapping propleuron in lateral view. Large notopleural projections ( Figs. 31, 32 ), lower part with narrow nipple-like structure at top, directed outwards and upwards, upper part a ridge running to and fused to rear of lateral part of anterior pronotal lobe, clump of short setae on upper part of ridge. Posterior pronotal lobe with short lapels ( Fig. 32 ), dorsally flattened, apex just below level of highest part of middle lobe, in dorsal view square-sided interiorly, rounded at apex ( Fig. 31 ); horn narrow, longer than middle lobe, curved forward so apex just to rear of, but well above, anterior lobe ( Fig. 32 ) in lateral view. Synthorax with subrectangular mesostigmal plates, depressed at front, with fringe of long setae on low ridge at rear. A few long, erect, setae on quarter of mesepisternum behind prothorax. Antehumeral markings similar to male but more widely separated from each other anteriorly, separated into two at the position of the excision in male. Dark stripe along metepleural suture slightly expanded compared to male. Wings with arc situated slightly distal to Ax 2. Fw 16 Px (left), 15 Px (right), Hw 15 Px. R4 slightly proximal to Sn. Pt brown with very narrow white margin, partial on costal side, broad, ca rhomboidal, covering more than one but less than one and a half underlying cells in all wings.

Abdomen: Coloration overall similar to male but less dark above. S8 pale except subtriangular basal dark brown dorsal mark, apex just over half segment length, and small brown mark behind posterior carina lower laterally. Apical part of pale colour on S8 tergite blue, including behind posterior carina. S9 mostly brown to black laterally, blue above, with narrow dark central dorsal mark in basal half. Narrowly dark apically on dorsum. S10 black. Cerci shorter than S10, brown. Ovipositor mostly black, basal pale patch, brown along lower margin, pale mark dorsally near apex, extending well beyond cerci.

Measurements (mm): Abdomen with anal appendages or ovipositor 34 mm, Hw 21 mm.

Variation in paratypes. Males. The markings on the labrum frequently reduced to faint brown smudges, but sometimes larger and clearer than in the holotype. The markings around the ocelli are frequently reduced compared to the holotype; in particular, the marks between lateral and median ocelli are only rarely present. However, in a few individuals the markings from lateral ocellus to eye margin are almost continuous. In some individuals the postocular markings are narrowly connected to the large markings on the underside of the head. Upper part of notopleural projection often barely apparent and clump of setae often absent. Black on pronotal middle lobe sometimes more extensive, resembling that seen in the holotype of C. paludensis ( Fig. 27 ). Arc at or distal to Ax 2, R4 at or proximal to Sn, rarely less than 3 postquadrangular cells. Pt colour varies with maturity, so can be paler in immature individuals and darker in very mature individuals. Markings on S8 very variable, completely absent or highly reduced on many individuals, but more extensive in others. The basal tooth on the cerci is always present and visible in dorsal view.

Females. Sometimes with much more extensive pale markings on the labrum, always with markings around lateral ocelli as in that described except one that has an additional pair of pale markings on either side of the median ocellus. The small mark at the eye margin is not always present; in one individual it is diffuse and brown but almost connected to a spur from the postocular mark; the postocular markings are occasionally connected narrowly to the large markings on the underside of the head. There is some variation in the shape of the cervical spurs. The nipples on the upper part of the notopleural projections are sometimes reduced and are entirely absent on the three females from the Matang Road site. However, the upper part of the notopleural projection is always present and always bears a clump of setae. The prothoracic horn is sometimes straight and slanted forwards and sometimes curved forwards almost through a right angle. In one individual the antehumeral marking is not separated into two. Variation in wing venation is the same as in the male except that the Pt only rarely covers two underlying cells. In some individuals S2–7 are darker and almost uniformly brown. There is considerable variation in the dark markings on the dorsum of S8–9, in semi-teneral females they are small and diffuse, in mature individuals they are occasionally reduced to small basal triangles, sometimes larger than in the female described; that on S9 is often subtriangular. The extent of blue colour behind the posterior carina of S8 is very variable. In one mature individual the pale marks at base and apically on the ovipositor almost meet; in semi-teneral individuals the ovipositor is entirely pale.

Measurements (mm): Males 15–18 Px in Fw, 13–17 in Hw; abdomen without anal appendages 31–38; Hw 19.5–22. Females with 14–16 Px; abdomen without anal appendages and ovipositor 33–36; Hw 21–23.

Diagnosis. The male of C. matok is readily separated from those of C. nemoricola (and allied forms), C. cyaneothorax , C. nigrohamata (and allied forms), C. resecta and C. octogesima by having the tips of the paraprocts a little before those of the cerci, whereas the tips of the paraprocts are clearly beyond those of the cerci in the others. It is separated from C. paludensis by its less inflated cercus with a basal tooth, and from C. macrostigma by its antehumeral markings and details of the anal appendages, e.g. position of the middle tooth of the cercus and shape of the apex of the cercus. The females of C. cyaneothorax , C. nemoricola , C. nigrohamata , C. octogesima and C. resecta have prothoracic horns shorter than the middle lobe of the prothorax, and additionally C. cyaneothorax , C. nemoricola and forms allied to it have much smaller notopleural projections. The females of C. macrostigma and C. paludensis have much larger notopleural projections than that of C. matok .

Remarks. The male Coeliccia sp. from “Platang” Road (most likely a typographical error for Matang Road) described briefly in Laidlaw (1918) appears to belong to C. matok , but I have not seen this specimen; notably it is not listed in Laidlaw (1932) although some other specimens not assigned to a named species in Laidlaw (1918) are listed again in Laidlaw (1932), suggesting that the specimen was no longer in Laidlaw’s possession at the time of the 1932 paper. Laidlaw describes this specimen as having abdominal S8 with “its terminal quarter ringed with blue”, essentially agreeing with C. matok . On the anal appendages Laidlaw states that “The upper anal appendages are brownish black, more abruptly truncate than is the case in C. nigrohamata , and distinctly, though very little longer than the lower pair” which, apart from the colour (which is presumably the result of poor preservation), agrees with C. matok . In the same 1918 paper Laidlaw lists a female Coeliccia from Matang Road, collected on the third of January 1910, under C. octogesima , stating that although it was “unfortunately much damaged” it agreed well with the description of the type female (from Singapore) and that “It differs from the other females of the genus I have seen in the greater length of the spine developed on the posterior margin of the prothorax, which projects forward at an angle of 45° to the main axis of the body and is approximately 1 mm. in length.” This specimen is not listed in Laidlaw (1932) under C. octogesima or separately, and I have not found it. The description of the prothoracic horn given does not match that of C. octogesima or that of C. nigrohamata , which could have been found at non-peat swamp forest sites along the Matang Road, especially near to the Matang Range, in 1910; in both of these species the prothoracic horn is shorter. It appears likely that this specimen belonged to C. matok .

Coeliccia matok is currently known from only three locations in south-western Sarawak. It was first discovered in peat swamp forest on the UNIMAS campus near Kota Samarahan in Samarahan division, and it is sometimes abundant at this location. Later it was found at a remnant peat swamp forest site, with many old rubber trees, on the Matang Road outside Kuching in the division of the same name. In 2015 it was found at two non-peat swamp forest sites in the Samunsam Wildlife Sanctuary in the extreme west of Kuching division. The habitats it occupied at this location, small streams and seepages, are more typical of C. nigrohamata , a species that was not found at Samunsam, although it occurs not much further east at the foot and on the lower slopes of Gunung Pueh (e.g. Dow 2012). It seems likely that in areas where C. nigrohamata occurs, C. matok is confined to peat swamp forest habitats, but outside of the range of C. nigrohamata , it occupies a broader range of habitats.

Figures

FIGURE 1. Genital ligula of a Coeliccia membranipes - group species: C. matok paratype (SAR 06 _ PCD 136) in ventral view. Scale bar 0.1 mm.

FIGURE 72. Map of Borneo showing the distributions of three Coeliccia species. White triangle— C. macrostigma; white square— C. matok; yellow circle— C. paludensis.

FIGURES 5 – 13. Heads of males of Coeliccia species. C. macrostigma male described: (5) dorsal view; (6) dorsal-frontal view; (7) ventral view. C. matok holotype: (8) dorsal view; (9) dorsal-frontal view; (10) ventral view. C. paludensis holotype: (11) dorsal view; (12) dorsal-frontal view; (13) ventral view.

FIGURES 14 – 22. Head of females of Coeliccia species. C. macrostigma female described: (14) dorsal view; (15) dorsalfrontal view; (16) ventral view. C. matok female described: (17) dorsal view; (18) dorsal-frontal view; (19) ventral view. C. paludensis female described: (20) dorsal view; (21) dorsal-frontal view; (22) ventral view.

FIGURES 23 – 28. Prothorax of males of Coeliccia species; all scale bars 0.2 mm. C. macrostigma male described: (23) dorsal view; (24) lateral view. C. matok holotype: (25) dorsal view; (26) lateral view. C. paludensis holotype: (27) dorsal view; (28) lateral view.

FIGURES 29 – 34. Prothorax of females of Coeliccia species; all scale bars 0.2 mm. C. macrostigma female described: (29) dorsal view; (30) lateral view. C. matok female described: (31) dorsal view; (32) lateral view. C. paludensis female described: (33) dorsal view; (34) lateral view, shows the right hand side, flipped for consistency with the other figures.

FIGURES 35 – 40. Synthorax of males of Coeliccia species. C. macrostigma male described: (35) dorsal view; (36) lateral view. C. matok holotype: (37) dorsal view; (38) lateral view. C. paludensis holotype: (39) dorsal view; (40) lateral view.

FIGURES 41 – 46. Synthorax of females of Coeliccia species. C. macrostigma female described: (41) dorsal view; (42) lateral view. C. matok female described: (43) dorsal view; (44) lateral view. C. paludensis female described: (45) dorsal view; (46) lateral view.

FIGURES 47 – 52. Terminal abdominal segments of males of Coeliccia species. C. macrostigma male described: (47) dorsal view; (48) lateral view. C. matok holotype: (49) dorsal view; (50) lateral view. C. paludensis holotype: (51) dorsal view; (52) lateral view.

FIGURES 53 – 58. Terminal abdominal segments of females of Coeliccia species. C. macrostigma female described: (53) dorsal view; (54) lateral view. C. matok female described: (55) dorsal view; (56) lateral view. C. paludensis female described: (57) dorsal view; (58) lateral view.

FIGURES 59 – 64. Anal appendages of males of Coeliccia species; all scale bars 0.5 mm. C. macrostigma male described: (59) dorsal view; (60) lateral view. C. matok holotype: (61) dorsal view; (62) lateral view. C. paludensis holotype: (63) dorsal view; (64) lateral view.

FIGURES 65 – 67. Paraprocts of Coeliccia species in ventral view, all scale bars 0.5 mm: (65) C. macrostigma male described; (66) C. matok holotype; (67) C. paludensis holotype.

FIGURES 68 – 71. Photos in life. (68) C. macrostigma male, Belait district, Brunei, photograph by C. Y. Choong; (69) C. matok female, UNIMAS campus, Sarawak, photograph by G. T. Reels; 70) C. paludensis male, Tuanan, Kalimantan Tengah, photograph by M. Silvius; (71) C. paludensis female, Tuanan, Kalimantan Tengah, photograph by M. Silvius.