Devadatta podolestoides Laidlaw, 1934

Dow, Rory A., Hämäläinen, Matti & Stokvis, Frank R., 2015, Revision of the genus Devadatta Kirby, 1890 in Borneo based on molecular and morphological methods, with descriptions of four new species (Odonata: Zygoptera: Devadattidae), Zootaxa 4033 (3), pp. 301-349 : 333-334

Devadatta podolestoides Laidlaw, 1934

( Figs. 1 , 2 , 3 , 4 , 5 , 6 , 7 , 11 , 17 , 23 , 31 , 37 , 39 , 43 , 49 , 55 , 61 , 71 , 74 )

Devadatta podolestoides podolestoides Laidlaw 1934a: 101 –103, Fig. 1 (original description, part: specimens from Gunung Poteng);— Lieftinck 1954: 2 (part);— Lieftinck 1971: 107 (note on holotype and first described female); van Tol 1992: 182 (summary occurrences of name in publications by Lieftinck).

Devadatta podolestoides ;— Dow & Reels 2010: 15 (Kubah: part);— Dow 2012a: 4 (Gunung Penrissen);—Dow & Reels 2013: 12 (records western Sarawak);— Dow, Reels & Butler 2013b: 11 (Kubah National Park).

Material. Type material. Holotype 1 ♂ (JvT number 2084), Gunung Poteng, 550m, Singkawang, W. Borneo [Kalimantan Barat, Kalimantan, Indonesia], 21 viii 1932, leg. L.C. de Ruiter, in RMNH. Paratypes (all Kalimantan Barat, Indonesia): 1 ♂, same location and collector as holotype, 550m, 5 xi 1931; 1 ♂, same location and collector, 7 xi 1931, in BMNH; 1 ♂, all data as holotype; 1 ♂, same location and collector, 400m, 28 ii 1932; 1 ♂, same location and collector, 28 ii 1932, in BMNH; 1 ♂, same location and collector, 550m, 13 iii 1932; 3 ♂, same location and collector, 13 iii 1932, in BMNH (two ex. Cowley collection); 1 ♂ 1 ♀, same location and collector, 17 vii 1932, in BMNH; 1 ♂, same location and collector, 300m, 21 viii 1932; 1 ♂, same location and collector, 21 ix 1932; 1 ♂, same location and collector, 16 x 1932, in BMNH; 1 ♀, same location and collector, 18 x 1932, in CUMZ.

Other material. Kalimantan Barat, Indonesia (all leg. L.C. de Ruiter, all in RMNH): 1 ♀, Bagak, Gunung Raja Complex, Singkawang, 26 ii 1932; 2 ♂, location as holotype, 1 iv 1934; 1 ♀ (prothorax destroyed), same location, 550m, 1 iv 1934; ♂, same location, 30 vi 1939.

Sarawak, Malaysia: 2 ♂ 1 ♀, tributary to Sungai Annah Rais, Annah Rais, Gunung Penrissen, Kuching division, 19 v 2005, leg. R.A. Dow, in coll. Hämäläinen; 1 ♂, tributary to Sungai Annah Rais, Annah Rais, Gunung Penrissen, Kuching division, 26 i 2006, leg. G.T. Reels; 1 ♂ ( RMNH.INS.506808), forest stream running onto golf course, Borneo Highlands Resort, same mountain, 21 vii 2012, leg. R.A. Dow; 2 ♂, same data; 2 ♂, same location and collector, 28 vi 2014; 1 ♂ ( RMNH.INS.506809), stream near forest trail, same resort, 22 vii 2012, leg. R.A. Dow; 5 ♂ 1 ♀, same data; 3 ♂, forest stream, same resort, 23 vi 2012, leg. R.A. Dow; 1 ♂, forest stream near trail, 25 vii 2012, leg. R.A. Dow; 3 ♂ 1 ♀, forest streams below Penrissen peak, same resort, 26 vii 2012, leg. R.A. Dow, in coll. Hämäläinen; 2 ♂, same location and collector, 1 vii 2014; 1 ♂ (SAR05_ AMP 8), 1 ♀ (SAR05_ AMP 9; in tandem with the male), small stream, Gunung Serapi, Kubah National Park, Matang Range, Kuching division, 13 iv 2005, leg. R.A. Dow; 1 ♂ ( RMNH.INS.506258), drain by road on Gunung Serapi, same national park, 30 iii 2012, leg. R.A. Dow; 1 ♀, same data; 1 ♂ ( RMNH.INS.506910), ♀ ( RMNH.INS.506909), stream at ca 400m, Gunung Serapi, same national park, 30 ix 2012, leg. R.A. Dow; 2 ♂, same data; ♂, stream at 600–700m, Gunung Serapi, same area, 1 vii 2013, leg. R.A. Dow; 1 ♂, waterfall trail, same national park, 13 ii 2008, leg. G.T. Reels; 4 ♂ (PomS 7, 12, 13, 19a-2014), waterfall trail and waterfall at end of trail, same national park, 13 iii 2014, leg. P.O.M. Steinhoff, in coll. Steinhoff; 1 ♂, stream above waterfall at end waterfall trail, same national park, 16 ix 2009, leg. R.A. Dow; 1 ♂ 1 ♀, stream above church, Gunung Singgai, same range, 2 x 2008, leg. R.A. Dow; ♂, stream by path to site of colonial era bungalow, Mount Matang, same range, 31 iii 2014, leg. R.A. Dow; 1 ♂, Ranchan Recreational Park, Serian, Samarahan division, 14 ii 2008, leg. R.A. Dow.

Description of male (based on SAR05_AMP8). Head ( Fig. 17 ). Labium mostly pale. Mandible bases, genae shining dark brown. Labrum, clypeus, frons and vertex to level of lateral ocelli black with metallic green reflection, remainder of dorsum of head matte black with indistinct brown area to rear; underside extensively pale. Antennae almost entirely black.

Thorax: Most of propleuron and pronotum black, with pale lateral marks on anterior pronotal lobe and rear of middle pronotal lobe, bronzy brown area centrally on posterior pronotal lobe. Dorsum of synthorax matte black with pair of small pale marks in antealar triangle and small pale mark between arms of anterior bifurcation of middorsal carina. Synthorax laterally with pale marks as follows ( Fig. 23 ): 2 short streaks along lower side of mesopleural suture, longer streak under interpleural suture for almost its entire length, a larger subtriangular area and smaller mark under metapleural suture. Area between metapleural carina and venter brown. Venter of synthorax largely pale ( Fig. 31 ). Legs brownish grey, with indistinct darker and paler areas. White pruinosity sparsely present on coxae and adjacent parts of pro- and synthorax and venter of synthorax. Wings with arculus ca level of Ax 5, quadrilateral with two crossveins, cubito-anal space free in all wings, pt dark brown. Membrane with brownish tint, darker at extreme tips of both wings.

Abdomen: Black above and very dark brown laterally, becoming almost entirely black on terminal 3 segments, S3–8 with a broad pale basal ring, widely interrupted dorsally. Genital ligula ( Fig. 39 ) typical for the group, seminal vesicle pale. Anal appendages dark brown. Paraprocts extremely short. Cerci longer than S10, in dorsal view ( Fig. 43 ) broad at base, narrowing rather abruptly at ca one-quarter of their length from base, in-turned at less than a right angle at half length, apical half moderately expanded ( Fig. 49 ). Interiorly deeply excavated from point where narrows in dorsal view to tip, brown inside excavation.

Measurements [mm]: Abdomen without anal appendages 32, cerci 1, Hw 28.5.

Description of female (based on SAR05_AMP9). As male except as noted. Head: Labrum mostly creamy brown. Most of frons dark brown, remainder of dorsum of head matte black with postocular lobes and a band joining them pale ( Fig. 55 ). Antennae with pedicel pale brown.

Thorax: Pronotum ( Fig. 61 ) mottled brown, with raised part of anterior lobe extensively pale yellow, pale yellow band along free margin of posterior lobe, widest laterally, narrowly divided centrally. No horns on middle pronotal lobe. Dorsum of synthorax brown with middorsal carina black, small pale blue mark between arms of anterior bifurcation of middorsal carina. Synthorax laterally brown with pale marks as follows ( Fig. 71 ): 3 short streaks along lower side of mesopleural suture, one of these alongside mesokatepisternum, streak under interpleural suture broken and irregular. Wings with arculus ca level of Ax 4 (3 in right Fw), quadrilateral with two crossveins (one in left Fw), pt dark brown with pale area apically. Extreme tips of both wings tinted brown, darker in Hw.

Abdomen: Brown, becoming darker on successive segments; basal ring, widely interrupted dorsally, also present on S2. Cerci black, subtriangular, shorter than S10. Ovipositor short, only reaching beyond S10 to ca level of tips of cerci.

Measurements [mm]: Abdomen without ovipositor or cerci 29.5, Hw 28.5.

Variation: Apart from minor differences in the extent of pale markings, there is little significant variation in both sexes apart from size and the following characters. In males occasionally there is no darkened area at all at tip in one or both pairs of wings; the degree of the general tinting of the wing membrane is variable. The arculus typically arises at Ax 4, occasionally at Ax 3 or Ax 5 in one or more wings ( Fig. 37 shows the wings of a paratype). Very rarely there is a single crossvein in the cubito-anal space of one wing. In females the pterostigma is sometimes entirely dark; this variation is associated with maturity.

The male described differs from the holotype by being generally browner, in particular there are irregular brown marks between antennae and lateral ocelli; the brown patches at the rear of the head are joined by a brown band along the occipital ridge; the abdomen is more brown than in the male described, but the pale marks are faded and not very distinct. Additionally the holotype lacks pale marks in the antealar triangle and the pale mark between the arms of the anterior bifurcation of the middorsal carina are very faint. The cerci of all males (including the holotype) bear small spines; these are highly variable in size and position and non-diagnostic.

Measurements [mm]: Males: Abdomen without anal appendages 32–40, Hw 28.5–32.5. Females: Abdomen without ovipositor or cerci 29.5–31.5, Hw 28–29.

Remarks. The type series of D. podolestoides is stated by Laidlaw (1934a) to consist of 19 males (including the holotype) and four females collected at “Cjn” (presumably a misreading of Gn [Gunung] by Laidlaw) Poteng near Singkawang in present day Kalimantan Barat, “between Nov. 1931 and Oct. 1932 ”, five males and a female from “Kinakok river, [Mount] Kinabalu”, and two males collected at “Mampawa by Andjoengan” (also in the Singkawang area) by Coomans de Ruiter in March 1932. A total of 15 males (including the holotype) and one female from Gunung Poteng collected within the date range stated for the type series were found in RMNH, BMNH and CUMZ, but not all were marked as types; all have been listed as types here, leaving four males and three females unaccounted for. None of the specimens from Mount Kinabalu are D. podolestoides . Only one male Devadatta from Mampawa was located, in RMNH, collected on 19 March 1932; this is presumably one of the paratypes, however it actually belongs to D. clavicauda and is listed as a paratype of that species above.

Devadatta podolestoides is locally common at some locations in southwestern Sarawak, and judging from the type series, on Gunung Poteng near Singkawang in western Kalimantan. Above 800m on Gunung Penrissen in Sarawak, at the Indonesian border, it is the only Devadatta species that has been found, but at lower altitudes in Sarawak it overlaps with D. clavicauda . The habitat of this species is typically small forest streams in hilly or mountainous terrain.

Figures

FIGURE 1. COI gene tree for 73 specimens of Devadatta and one outgroup taxon, from Bayesian Inference analysis. Posterior probability values are shown (as percentages) if less than 100 %. RMNH collection codes are shown for each specimen, with the RMNH. INS. prefix omitted for clarity.

FIGURE 2. COI gene tree for the 33 specimens of Devadatta also used in the analysis with other markers and one outgroup taxon. Posterior probability values are shown (as percentages) if less than 100 %. RMNH collection codes are shown for each specimen, with the RMNH. INS. prefix omitted for clarity.

FIGURE 4. ITS gene tree for 33 specimens of Devadatta and one outgroup taxon, from Bayesian Inference analysis. Posterior probability values are shown (as percentages) if less than 100 %. RMNH collection codes are shown for each specimen, with the RMNH. INS. prefix omitted for clarity.

FIGURE 6. Phylogenetic reconstruction for 33 specimens of Devadatta and one outgroup taxon from the combined COI + 16 S + ITS + 28 S data, using Bayesian Inference analysis. Posterior probability values are shown (as percentages) if less than 100 %. RMNH collection codes are shown for each specimen, with the RMNH. INS. prefix omitted for clarity.

FIGURE 7. Maximum Likelihood best tree for 33 specimens of Devadatta and one outgroup taxon from the combined COI + 16 S + ITS + 28 S data set. Bootstrap support values below 100 are superimposed on the tree. RMNH collection codes are shown for each specimen, with the RMNH. INS. prefix omitted for clarity.

FIGURE 74. Distribution of D. aran (squares) and D. podolestoides (triangles).

FIGURE 3. 16 S gene tree for 33 specimens of Devadatta and one outgroup taxon, from Bayesian Inference analysis. Posterior probability values are shown (as percentages) if less than 100 %. RMNH collection codes are shown for each specimen, with the RMNH. INS. prefix omitted for clarity.

FIGURE 5. 28 S gene tree for 33 specimens of Devadatta and one outgroup taxon, from Bayesian Inference analysis. Posterior probability values are shown (as percentages) if less than 100 %. RMNH collection codes are shown for each specimen, with the RMNH. INS. prefix omitted for clarity.

FIGURES 8 – 13. Habitus of males of Devadatta species: (8) D. aran holotype; (9) D. basilanensis Davao City, Mindanao; (10) D. clavicauda holotype; (11) D. podolestoides male described in this paper; (12) D. somoh holotype; (13) D. tanduk holotype.

FIGURES 14 – 19. Heads of male Devadatta species: (14) D. aran holotype; (15) D. basilanensis Davao Oriental, Mindanao; (16) D. clavicauda holotype; (17) D. podolestoides male described in this paper; (18) D. somoh holotype; (19) D. tanduk holotype.

FIGURES 20 – 25. Synthorax of male Devadatta species, lateral view: (20) D. aran holotype; (21) D. basilanensis Davao Oriental, Mindanao; (22) D. clavicauda holotype; (23) D. podolestoides male described in this paper; (24) D. somoh holotype; (25) D. tanduk holotype.

FIGURES 30 – 33. Venter of synthorax of male Devadatta species: (30) D. basilanensis Davao Oriental, Mindanao; (31) D. podolestoides male described in this paper; (32) D. clavicauda holotype; (33) D. clavicauda paratype (SAR 09 _ 10 _ AMP 40, Gunung Serapi).

FIGURES 34 – 37. Wings of some Devadatta species: (34) D. basilanensis male Zamboanga, Mindanao; (35) D. basilanensis female Cotabato, Mindanao; (36) D. clavicauda paratype, Gunung Santubong, Sarawak; (37) D. podolestoides paratype, Gunung Poteng, Kalimantan Barat.

FIGURES 38 – 39. Genital ligula of two Devadatta species, scale bars = 0.5 mm: (38) D. aran holotype; (39) D. podolestoides male described in this paper.

FIGURES 40 – 45. Male anal appendages, dorsal view: (40) D. aran holotype; (41) D. basilanensis Davao Oriental, Mindanao; (42) D. clavicauda holotype; (43) D. podolestoides Kubah NP, Sarawak; (44) D. somoh holotype; (45) D. tanduk holotype.

FIGURES 46 – 51. Male anal appendages, lateral view (scale bars 1 mm): (46) D. aran holotype; (47) D. basilanensis Davao Oriental, Mindanao; (48) D. clavicauda holotype; (49) D. podolestoides Kubah NP, Sarawak; (50) D. somoh holotype; (51) D. tanduk holotype.

FIGURES 52 – 57. Heads of Devadatta female specimens described in this paper: (52) D. aran; (53) D. basilanensis; (54) D. clavicauda; (55) D. podolestoides; (56) D. somoh; (57) D. tanduk.

FIGURES 58 – 63. Prothorax of Devadatta female specimens described in this paper, dorsal view: (58) D. aran; (59) D. basilanensis; (60) D. clavicauda; (61) D. podolestoides; (62) D. somoh; (63) D. tanduk.

FIGURES 68 – 73. Synthorax of Devadatta female specimens described in this paper, lateral view: (68) D. aran; (69) D. basilanensis; (70) D. clavicauda; (71) D. podolestoides; (72) D. somoh; (73) D. tanduk.