Devadatta aran Dow, Hämäläinen & Stokvis, 2015, spec. nov.

Devadatta aran spec. nov.

(Figs. 1, 2, 3, 4, 5, 6, 7, 8, 14, 20, 26, 38, 40, 46, 52, 58, 64, 65, 68, 74)

Devadatta argyroides (sic) [nec (Selys, 1859)];— Laidlaw 1915: 25, 33 (Mount Kinabalu);— Laidlaw 1920: 332.

Devadatta species B;— Dow, Reels & Butler 2013a: 9, 12 (Mount Dulit).

Devadatta species C;— Dow & Ngiam 2014: 9, 42, 44 (Ulu Balui, Ulu Baleh);— Dow et al. 2015a: 22 (Kapit);— Dow et al. 2015b: 11 (Usun Apau).

Devadatta sp.;— Laidlaw 1934b: 550 (Part: Mount Kinabalu);— Dijkstra et al. 2014: Table 5 (Sarawak: RMNH.INS.503484).

Devadatta podolestoides [nec Laidlaw, 1934];— Kimmins 1936: 79 (Mount Dulit: part);— Dow 2006: 10 (Tama Abu Range);— van Tol 2006: 13 (Gunung Lumut, East Kalimantan);— Dow & Reels 2009: 13 (Mount Dulit: part).

Devadatta podolestoides podolestoides [nec Laidlaw, 1934];— Lieftinck 1954: 2 (part: Mount Kinabalu);— Hämäläinen 1994: 79 (part).

Devadatta cf podolestoides;— Dijkstra et al. 2014: Figs. 2 b, 5b (RMNH.INS.503484)

Devadatta spp. cf podolestoides;— Dow & Ngiam 2012: 2, 7 (Hose mountains: part).

Type material. Holotype: 1 ♂ (SAR11_12_ AMP 74, RMNH.INS.506316), Pa’Gelawat and tributaries at 1400– 1500m, Pulong Tau National Park, Miri division, Sarawak, Malaysia, 16 iv 2012, leg. R.A. Dow, in RMNH.

Paratypes (139 ♂ 30 ♀). Female used in description: 1 ♀ (SAR06_ AMP 21), trail to waterfall, Bario, Tama Abu Range, Miri division, 14 iii 2006, leg. RAD. See appendix for remainder.

Etymology. aran, a noun in apposition; named in memory of the late Reddish (pronounced Raddish) Aran (born 6 July 1963, died 27 November 2012) of Bario in the Tama Abu Range, who accompanied the first author on his first trip to the type locality.

Diagnosis. A dark coloured Devadatta species, in most known populations it can be separated from other species from the podolestoides -group by having the venter of the synthorax mostly pale in both sexes and at least vestigial, but not long, horns on the middle pronotal lobe in females. Additionally, easily separated from D. clavicauda by the less expanded ends of the cerci in the male of D. aran.

Description of holotype male. Head: Labium mostly pale. Mandible bases and 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 pale area on protruding postocular lobes; no pale band joining these areas (Fig. 14); underside extensively pale. Antennae almost entirely black.

Thorax: Pronotum and propleuron largely black, with pale streak below notopleural suture, pale lateral marks on anterior pronotal lobe, rear of middle pronotal lobe, laterally on posterior pronotal lobe. Dorsum of synthorax matte black with small pale mark between arms of anterior bifurcation of middorsal carina. Synthorax laterally with bright pale marks as follows (Fig. 20): along lower side of mesopleural suture a long streak and short streak near antealar carina, longer streak running whole length of interpleural suture, larger subtriangular area and smaller mark under metapleural suture, joined by fainter pale marking. Area between metapleural carina and venter pale. Venter of synthorax entirely pale with areas of greyish shadowing (Fig. 26). Legs (posterior pair removed for DNA extraction): Coxae and trochanters pale with brown areas. Femora obscurely brown and grey. Tibiae and tarsi dark brown. Wings with arculus ca level of Ax 4 (5 in left Hw), quadrilateral with 1 crossvein in Fw, 2 in Hw, cubitoanal space free in all wings, pt brown. Wing membrane with brownish tint, extreme tips of both wings with slightly heavier brown tint.

Abdomen: Black above and very dark brown laterally, becoming almost entirely black on terminal 3 segments, S3–7 with broad pale basal ring, brown dorsally, small remnant of the basal ring present laterally on S8. Genital ligula (Fig. 38) typical for podolestoides -group, seminal vesicle brown. Anal appendages black. Paraprocts extremely short, rounded, dark basally, paler apically. Cerci longer than S10, in dorsal view (Fig. 40) abruptly curved inward at ca mid length, apical half at just more than a right angle to proximal half, left cercus lying over right. In both dorsal and lateral (Fig. 46) views widest at base, narrowing towards inward turn then moderately expanded in apical half. Interior surface with deep slot-like excavation around middle part, brownish inside excavation.

Measurements [mm]: Abdomen without anal appendages 39.5, cerci ca 1, Hw 34.

Description of female (SAR06_AMP21). As male except as noted. Head. Basal ca 2/3 labrum brown. Most of mandible bases, genae and postclypeus brown, remainder of dorsum of head matte black with postocular lobes brown, joined by very distinct pale band along occipital ridge (Fig. 52). Antennae with anterior yellowish streak on scape.

Thorax: Pronotum (Fig. 58) mottled brown, larger pale marks on anterior lobe, middle lobe bearing small projections, stump-like in lateral view (Fig. 64). Posterior lobe with yellow lateral patches. Dorsum of synthorax brown with middorsal carina black, small bluish pale mark between arms of anterior bifurcation of middorsal carina. Synthorax laterally brown with pale marks as in male (Fig. 68). Wings with arculus at Ax 4 in all wings, quadrilateral with two crossvein in all wings, pt black proximally apical ca 2/3 mostly pale. Extreme tips of both wings slightly tinted brown, a little darker in Hw.

Abdomen: Small pale lateral basal spot on S2. Cerci black, subtriangular, ca same length as S10. Ovipositor short, only reaching beyond S10 to ca level of tips of cerci.

Measurements [mm]: abdomen without ovipositor and appendages 35, Hw 33.

Variation in paratypes. Males: Apart from small differences in the brightness and extent of pale markings on the head, thorax and abdomen, there is little variation in most paratypes from Sarawak and Sabah except in size. Teneral males frequently have a pale band joining the pale areas on the postocular lobes. Pale marks on the posterior pronotal lobe are absent in some individuals and more extensive in others. The arculus is at Ax 3–5, but in almost all individuals it is at Ax 4 or 5 in at least 2 wings; in only one specimen from the Hose Mountains it is at Ax 3 in all wings. Up to three crossveins in the quadrilateral. Rarely there is a crossvein in the cubito-anal space of one wing; in one individual from the Tama Abu range there are crossveins in two wings, and in one other there are single crossveins in two wings and two crossveins in one wing. The colour of pt varies, sometimes darker than in the holotype, sometimes with a pale central area; this variation appears to be associated with maturity. The wing tips are sometimes entirely hyaline. Rotation of one of the cerci, in one case through ca 90 degrees, is not uncommon and can make them appear different on a superficial view. The two male specimens from southeast Kalimantan have a slightly darker appearance than the others, with some shadowing on the venter of the synthorax near the abdomen. The cerci of all males (including the holotype) bear small spines; these are highly variable in size and position and non-diagnostic.

Females: The other female paratypes show a range of variation in markings similar to that of males. In all specimens examined the arculus is at Ax 4 or 5, except one from the Hose Mountains where it is at Ax 3 in all wings. Rarely with a crossvein in the cubito-anal space of one wing. In some specimens there is no or virtually no darkening of the wings tips, in a few it is more extensive and darker than in the female described. In some individuals the wing is entirely un-tinted; all of these appear teneral or just post-teneral. The most significant variation is in the horns on the middle pronotal lobe, which vary from the condition seen in the type to upright nipple-like horns (e.g. Fig. 65), which slope back slightly in a few individuals. Most individuals from the Tama Abu range have projections like that in the female described, but one is intermediate between this and the one shown in Fig. 65, while both extremes are present in other areas; a rearward slant is more common in specimens from Kapit division.

Measurements [mm]: Males: abdomen without anal appendages 33.5–41.5, Hw 28.5–36.5. Females: abdomen without anal appendages or ovipositor 29–37, Hw 28.5–36.5.

Remarks. Across most of its known range D. aran is scarce at lower altitudes and more common above ca 900m. Although its distribution (Fig. 74) broadly or entirely overlaps those of D. clavicauda, D. somoh and D. tanduk, above ca 1000m in the Tama Abu Range in Sarawak and the adjacent Long Pasia highland area in Sabah it appears to be the only Devadatta species present. Devadatta aran occurs up to at least 1500m and is typically found at small forest streams.