Adult male (WII-ADR964; Fig. 4A–E), from a trail beside Forest Colony (25.8994°N; 94.7694°E; elevation 1300 m a.s.l.; Fig. 3A), Kiphire Forest Division, Kiphire District, Nagaland, India; collected by Abhijit Das and Bitupan Boruah on 1 August 2021.

An adult male (WII-ADR963; Fig. 4F, G) collection locality details are same as the holotype.

A medium-sized gecko (SVL up to at least 64.7 mm); 10 supralabials; 9–12 infralabials; 16 bluntly conical and feebly keeled dorsal tubercles; 26–29 paravertebral tubercles; 35 or 36 midventral scales rows between the weak ventrolateral folds; no precloacal groove; six or seven precloacal pores in continuous series; 12–15 total subdigital lamellae beneath digit IV of pes; dark brown irregular cross bars or reticulation on dorsum.

Holotype well preserved except single incision below left axilla ventrolaterally. Snout-vent length 63.9 mm. Head moderately large (HL/SVL = 0.26), dorsoventrally depressed, longer than width (HW/HL = 0.68), distinct from neck, broader at occipital region; snout tip rounded in both dorsal and lateral view; loreal region convex; canthus rostralis rounded, indistinct; interorbital space flat; a longitudinal furrow on dorsal surface of the snout; snout short (SO/HL = 0.4), longer than orbit (OD/SO = 0.7); nostril nearly rounded, opening directed posterolaterally; ear opening oval and oblique; scales on head heterogeneous, largest on snout and loreal region, posteriorly smaller in upper eyelid, interorbital space and occipital region, granular juxtaposed; scales on upper eyelids heterogeneous, supraciliaries outwardly sharp giving serrated appearance in dorsal view, size anteriorly and posteriorly decreases, largest at approximately anterior one third of it; rostral wide, a short groove at the middle on top; rostral connected with nasals, supranasals, internasal and first supralabials; two scale between the supranasals, larger than the rest of the granular snout scales; granular scales at parietal region and occipital region intermixed with slightly large rounded granular tubercles starting from the level of posterior margin of the upper eyelids, size increases towards nape; supralabial 10 on both side, seven supralabials upto midorbit, size decreases towards angle of jaw; a series of narrow, enlarged scales above the supralabials between nostril and anterior orbital border; mental slightly wider than rostral (RW/MW = 0.8), triangular, connected with first infralabials, inner postmentals; nine infralabials on both sides, size decreases towards angle of jaw; first infralabials connected with mental, second infralabial, inner and outer postmentals; inner pair of postmentals are larger than the outer postmentals; posterior margin of the inner postmentals bordered by eight granular scales of different size; two rows of enlarged scales along the infralabials starting below the outer postmentals, posteriorly size decreases, elongated and narrow; rest of the gular scales are small, granular juxtaposed, homogeneous, size increases towards the throat where they become imbricate.

Habitus slender (BW/SVL = 0.2, TRL/SVL = 0.35), dorsoventrally depressed; dorsal scales granular, rounded, heterogeneous, intermixed with rounded, weakly keeled and bluntly conical tubercles irregularly arranged, up to fourth segment of the tail, size increases towards posterior body and pronounced; 16 dorsal tubercles across mid dorsum; 26 paravertebral tubercles; ventrolateral fold weak; ventral scales larger than those of dorsal, flat, smooth, cycloid subimbricate to imbricate, largest on belly; 36 mid-ventral scales between ventrolateral fold; seven precloacal pores arranged in an inverted “V” shaped continuous series, followed by five unpored, large scales below it, largest at the apex.

Forelimbs and hindlimbs slender (FL/SVL = 0.13, CL/SVL = 0.16); digits strongly inflected at the joints, all bearing large recurved claw, enlarged subdigital lamellae; lamellae beneath digit IV of right and left manus (given as basal + distal) is 5+8 and 5+9 respectively; lamellae beneath digit IV of right and left pes (given as basal + distal) is 5+10 and 5+9 respectively; dorsal scales on forelimbs smooth and subimbricate, small and granular at elbow and proximal end of forearm; forearm scales intermixed with rounded large tubercles; dorsal scales of hindlimbs heterogeneous, intermixed with large, rounded and bluntly conical tubercles; horizontally upper half of the thigh scales are smooth, large and subimbricate, those on lower half small granular; scales on tibia are small, granular juxtaposed; ventral scales of forelimbs granular, juxtaposed, mostly homogeneous; scales on palm heterogeneous in shape and size, granular juxtaposed; scales on ventral side of hindlimbs smaller than those of belly, smooth, cycloid and subimbricate, but on the knee, above cloaca and on thigh below the level of precloacal pores are smaller and granular; scales on soles heterogeneous, granular, juxtaposed to subimbricate.

Tail regenerated (TL = 68 mm), slender, gradually tapering towards tip, segments indistinct, dorsal scales small, granular, juxtaposed at the base, posteriorly size increases, flat, smooth, subimbricate, heterogeneous in shape and size; large feebly keeled scales upto fourth segment of the tail, those on basal segment are pronounced; subcaudal scales smooth, subimbricate, wider than that of dorsal and widest at the middle, heterogeneous in shape and size; no enlarged plate like series of subcaudal scales; two and three bluntly conical spurs on right and left side of the tail base respectively.

Top of head intermixed with pale-yellow and pale-brown; dark-brown spots of irregular shape and size on posterior part of head; short dark-brown postorbital streak present; another streak along loreal region; supraciliary yellow; four dark-brown stripes on neck interspaced with cream colour; large spot at the middle on the anterior part of the nape; six dark-brown cross bars of irregular size and shape on dorsum between the level of axilla and groin (Fig. 4J); these cross bars intersected on mid dorsum and posteriorly bordered with cream coloured patches; limbs with slightly dark-brown indistinct reticulations intermixed with pale-yellow or cream coloured patches; scales on axillary region and base of forelimbs are with pinkish tinge; digits with alternating brown and pale-yellow bands; anterior half of the tail with alternating dark-brown and light-cream coloured cross bands of irregular shape and size; posterior half (regenerated part) of the tail pale-brown; ventrally head, trunk and limbs whitish; tail with brown mottling.

Top of head, back, limbs and tail pale-greyish-brown with dark-brown markings; upper eyelids grey; dorsal marking pattern as it was in life; and ventrally whitish.

Detailed morphometric variations are given in Table S2. Apart from those, the dorsal colour and marking pattern of the paratype varied from that of holotype. Paratype has two post orbital streaks on each side, upper one shorter than the lower one; no other spots on occipital region as in the holotype; dorsal cross bands somewhat reticulated (Fig. 4F).

Cyrtodactylus kiphiresp. nov. differs from C. aaronbaueri by having larger body size in male, SVL 63.9–64.7 mm (vs. SVL 54.2–62 mm in male), fewer dorsal tubercle rows, DTR 16 (vs. DTR 22–28), fewer paravertebral tubercles, 26–29 (vs. PVT 36–39); differs from C. aunglini by fewer dorsal tubercle rows, DTR 16 (vs. DTR 21–26), fewer precloacal pores, PcP 6 or 7 (vs. PcP 12–13), fewer mid-ventral scale rows, MVSR 35 or 36 (vs. MVSR 41–49), fewer paravertebral tubercles, PVT 26–29 (vs. 36–45); differs from C. bengkhuaiai by fewer dorsal tubercle rows, DTR 16 (vs. DTR 22–26), fewer paravertebral tubercles, 26–29 (vs. PVT 35–41), fewer mid-ventral scale rows, MVSR 35 or 36 (vs. MVSR 37–42); differs from C. brevidactylus by fewer dorsal tubercle rows, DTR 16 (vs. DTR 27), fewer mid-ventral scale rows, MVSR 35 or 36 (vs. MVSR 45), fewer precloacal pores, PcP 6 or 7 (vs. PcP 8); differs from C. chrysopylos by fewer precloacal pores, PcP 6 or 7 (vs. PcP 8–13), mid ventral scales, MVSR 35 or 36 (vs. MVSR 37–55), dorsal tubercle rows, DTR 26–29 (vs. DTR 30–35); differs from C. dianxiensis by smaller body size, SVL 63.9–64.7 mm (vs. SVL 73.8–79.9 mm), fewer dorsal tubercle rows, DTR 16 (vs. DTR 18 or 19), 35 or 36 mid ventral scales (vs. MVSR 37–41), 26 or 29 paravertebral tubercles (vs. PVT 31 or 32); differs from C. gansi by fewer dorsal tubercle rows, DTR 16 (vs. DTR 20–25), having fewer precloacal pores, PcP 6 or 7 (vs. PcP 16–29); differs from C. jaintiaensis by smaller body size in male, SVL 63.9–64.7 mm (vs. SVL 87–88.3 mm), fewer dorsal tubercle rows, DTR 16 (vs. DTR 19–20), fewer mid-ventral scale rows, MVSR 35 or 36 (vs. MVSR 40–42), fewer precloacal pores, PcP 6 or 7 (vs. PcP 11–12); differs from C. lungleiensis by smaller body size in male, SVL 63.9–64.7 mm (vs. SVL 65–68.1 mm), higher precloacal pores, PcP 6 or 7 (vs. PcP 3–5), fewer mid-ventral scale rows, MVSR 35 or 36 (vs. MVSR 37–43), fewer dorsal tubercle rows, DTR 16 (vs. 24–28), fewer paravertebral scales, PVT 26–29 (vs. 32–40), fewer subdigital lamellae beneath toe IV, TIVLam 12–14 (vs. TIVLam 16–18); differs from C. montanus by larger body size in male, SVL 63.9–64.7 mm (vs. SVL 53.6–55 mm), fewer dorsal tubercle rows, DTR 16 (vs. DTR 21–23), fewer precloacal pores, PcP 6 or 7 (vs. PcP 8–10); differs from C. myaleiktaung by fewer mid-ventral scale rows, MVSR 35 or 36 (vs. MVSR 57); differs from C. namtiram by fewer dorsal tubercle rows, DTR 16 (vs. DTR 21), fewer number of paravertebral scales, PVT 26–29 (vs. PVT 33), fewer precloacal pores, PcP 6 or 7 (vs. PcP 12); differs from C. ngopensis by fewer dorsal tubercle rows, DTR 16 (vs. 19–20), fewer paravertebral tubercles, PVT 26–29 (vs. 32–36); differs from C. siahaensis by fewer dorsal tubercle rows, DTR 16 (vs. DTR 22–24), fewer paravertebral tubercles, 26–29 (PVT 36–39); differs from C. vairengtensis by fewer dorsal tubercle rows, DTR 16 (vs. 22–23), fewer paravertebral tubercles, PVT 26–29 (vs. 34–39), fewer precloacal pores in male, PcP 6 or 7 (vs. PcP 9–11). The new species is morphologically close to C. nagalandensis; however, the new species differs from it by fewer paravertebral tubercles, PVT 26–29 (vs. 35–37). We could not compare with male specimens as data for C. nagalandensis is not available. Morphological differences with other members of khasiensis group is presented in Table 2.

Cyrtodactylus kiphiresp. nov. has a high genetic divergence of 12% from its closely related C. nagalandensis, 16.3% from C. dianxiensis, 16.9% from C. gansi and 17.9% from C. jaintiaensis. With other members of the clade, C. kiphiresp. nov. has a genetic divergence of 16.4% and 22.5% in the ND2 gene (Table S3).

The specific epithet is derived from Kiphire, a District of Nagaland from where the type series of this species were collected.

Kiphire bent-toed gecko.

We recorded Cyrtodactylus kiphiresp. nov. from Kiphire forest colony, Kiphire District, Nagaland, India. The area is characterised as subtropical forest with regenerating jhum forest dominated by Castanopsis indica (Roxb. ex Lindl.) A. DC., Quercus sp., Itea macrophylla Wall. ex Roxb., Albizia chinensis (Osbeck) Merr.. The holotype was collected from on a forest trail shrub (approximately 2.0 m from the ground) at approximately 23:00 hrs. The paratype was collected from a rocky slope along the edge of a stream.

Adult female (WII-ADR971), from Athibung (25.54199°N; 93.6307°E; elevation 740 m a.s.l.) (Fig. 3A), Peren District, Nagaland, collected by Abhijit Das and Bitupan Boruah on 14 August 2021.

Medium-sized gecko, SVL at least 68.8 mm in adult female; supralabials 9–12 and infralabials nine; 12 or 13 lamellae beneath the digit IV of manus; 17 lamellae beneath digit IV of pes; 17 feebly keeled and bluntly conical tubercles across mid dorsum and 32 paravertebral tubercles; 36 smooth mid-ventral scales between ventrolateral folds; at least 10 small precloacal pores in female; head on top pale-brown with purplish tinge; dark-brown postorbital stripe continuing to above the ear opening; a dark-brown cross bar with irregular edges on nape; indistinct pale-yellow patch on occipital region; dorsally and laterally neck and back pale yellowish-brown with dark-brown irregular reticulation.

Holotype well preserved except an incision below left axilla ventrolaterally. Snout-vent length 68.8 mm. Head moderately large (HL/SVL = 0.26), dorsoventrally depressed, longer than width (HW/HL = 0.65), oval, distinct from neck, broader at occipital region; snout tip rounded in both dorsal and lateral view; loreal region convex; canthus rostralis rounded, indistinct; interorbital space flat; a longitudinal furrow on dorsal surface of the snout; snout short (SO/HL = 0.39), longer than orbit (OD/SO = 0.59); nostril rounded, opening directed posterolaterally; ear opening oblique; scales on head heterogeneous, largest on snout and loreal region, posteriorly smaller in upper eyelid, interorbital space and occipital region, granular juxtaposed; scales on upper eyelids heterogeneous, supraciliaries outwardly sharp giving serrated appearance in dorsal view, size anteriorly and posteriorly decreases, largest at the anterodorsal region; rostral wide, a short groove at the middle on top; rostral connected with nasals, supranasals and first supralabials; a single scale between the supranasals, larger than the rest of the granular snout scales; granular scales at parietal, occipital and temporal region intermixed with slightly larger rounded and bluntly conical granular tubercles starting from the level of posterior margin of the upper eyelids, smaller in parietal region, size increases towards nape; supralabial 12 on right side and nine on left side, supralabials count up to midorbit is eight on right side and six on left side, size decreases towards angle of jaw; a series of narrow, enlarged scales above the supralabials between nostril and anterior orbital border; mental triangular, connected with first infralabials and inner postmentals; nine infralabials on both side, size decreases towards angle of jaw; inner pair of postmentals are larger than the outer postmentals; posterior margin of the inner postmentals bordered by eight granular scales of different size; two rows of slightly enlarged, narrow scales, larger than gular scales present along the infralabials starting below the outer postmentals, posteriorly size decreases; gular scales granular, juxtaposed, homogeneous, size increases towards the throat where they become imbricate.

Habitus slender (BW/SVL = 0.16, TRL/SVL = 0.47), dorsoventrally depressed; dorsal scales granular, rounded, heterogeneous, intermixed with densely placed rounded, weakly keeled and bluntly conical tubercles, irregularly arranged, continues to seventh segment of the tail, size increases towards posterior body and pronounced at the base of tail; 17 dorsal tubercles across mid dorsum; 32 paravertebral tubercles; ventrolateral fold weak; ventral scales larger than those of dorsal, flat, smooth, cycloid, subimbricate to imbricate, largest on posterior part of the belly, 36 mid-ventral scales between ventrolateral folds; 10 small precloacal pores arranged in an inverted “V” shaped continuous series, followed by a series eight unpored, enlarged scales below it.

Forelimbs and hindlimbs slender (FL/SVL = 0.14, CL/SVL = 0.18); digits strongly inflected at the joints, all bearing large recurved claw and enlarged subdigital lamellae; lamellae count beneath digit IV of right and left manus (given as basal + distal) is 5+7 and 5+8 respectively; lamellae count beneath digit IV of both right and left pes (given as basal + distal) is 8+9; dorsal scales on forelimbs heterogeneous in size; hind arm scales smooth and subimbricate; forearm scales small and granular towards proximal and towards distal end it is smooth, cycloid and imbricate; forearm scales intermixed with enlarged feebly keeled tubercles; dorsal scales of hindlimbs heterogeneous, intermixed with densely placed large, rounded and bluntly conical tubercles; scales on inner lateral and dorsolateral side of the thighs smooth, large and imbricate, rest of the scales are granular; scales on tibia are small, granular juxtaposed; ventral scales of forelimbs granular, juxtaposed, mostly homogeneous; scales on palm heterogeneous in shape and size, granular juxtaposed; ventral scales of hind limbs smaller than those of belly, smooth, cycloid and subimbricate; scales on the knee, above cloaca and on thigh below the level of precloacal pores smaller and granular; scales on soles heterogeneous, granular, juxtaposed to subimbricate.

Tail complete, posterior 6 mm regenerated (TL = 64 mm), slender, gradually tapering towards tip; tail segments indistinct; dorsal scales small, granular, juxtaposed at the base, posteriorly size of the scales increases, flat, smooth, subimbricate, heterogeneous in shape and size; enlarged feebly keeled scales up to seventh segment of the tail, those on basal segment are pronounced; subcaudal scales smooth, subimbricate, wider than that of dorsal, heterogeneous in shape and size; no enlarged plate like series of subcaudal scales; four bluntly conical spurs on both sides of the tail base.

Head on top pale-brown with purplish tinge, indistinct brown irregular spots on occipital region; a short brown streak behind the posterior corner of the upper eyelid; a broad, dark-brown postorbital stripe continuing to above the ear opening; area between these two postorbital streaks is paler than dorsal head colour; an indistinct brown loreal stripe covering the nasal and an indistinct pale stripe above it; indistinct pale-yellow patch on occipital region; lips and mandibular region paler with indistinct yellow spots; dorsally and laterally neck and back pale yellowish-brown with dark-brown irregular reticulation starting from neck to sacrum; a dark-brown cross bar with irregular edges on nape; limbs pale-yellowish-brown with dark-brown reticulation; digits with alternative dark-brown and pale-yellow bars; tail with alternative 11 dark and 10 light bars in the original part, dark bars posteriorly more darker and broader; light bars posteriorly more whitish, first bar at the base of tail broken into two spots; fourth and fifth bar connected on left side; ventrally head, trunk and limbs whitish; tail with irregular brown and pale-yellow patches (Fig. 5I).

Top of head pale-greyish-brown; neck, back, limbs and tail dorsally light-grey with dark-brown markings; marking pattern visible as those in life condition; ventrally whitish with brown specks on tail.

Cyrtodactylus barailensissp. nov. differs from C. aaronbaueri by fewer dorsal tubercle rows, DTR 17 (vs. DTR 22–28), 10 precloacal pores (vs. 6–8 pitted precloacal scales in female); differs from C. agarwali by fewer dorsal tubercle rows, DTR 17 (vs. 21–25); differs from C. aunglini by fewer mid-ventral scale rows, MVSR 36 (vs. MVSR 47–49), 32 paravertebral tubercle (vs. PVT 36–45), 17 dorsal tubercle rows (vs. DTR 21–26); differs from C. bengkhuaiai by having fewer dorsal tubercle rows, DTR 17 (vs. DTR 22–26), 10 precloacal pores (vs. 6–8 pitted precloacal scales in females); differs from C. brevidactylus by fewer dorsal tubercle rows, DTR 17 (vs. DTR 27–30), fewer paravertebral tubercles, PVT 32 (vs. PVT 38–42), enlarged dark blotches on head and dorsum absent (vs. large dark blotches on dorsum between nape and sacrum); differs from C. chrysopylos by presence of precloacal pores (vs. PcP absent in female), fewer mid-ventral scales, MVSR 36 (vs. MVSR 37–55); differs from C. dianxiensis by smaller body size, SVL 68.8 mm (vs. SVL 75 mm), presence of precloacal pores (vs. PcP absent in female), fewer subdigital lamellae, 12 or 13 under fourth finger and 16 under fourth toe (vs. 16 or 17 under fourth finger and 19 or 20 under fourth toe); differs from C. gansi by fewer dorsal tubercle rows, DTR 17 (vs. DTR 20–25), ventrolateral fold present on trunk (vs. absent), fewer precloacal pores, PcP 10 (vs. PcP 13 in female); differs from C. jaintiaensis by much smaller body size, SVL 68.8 mm (vs. SVL 96.2 mm in female), fewer mid-ventral scale rows, MVSR 36 (vs. MVSR 40–42), fewer precloacal pores, PcP 10 (vs. 12 PcP in female); differs from C. kiphiresp. nov. by the number of paravertebral tubercles 32 (vs. PVT 26 or 29); differs from C. lungleiensis by fewer dorsal tubercle rows, DTR 17 (vs. 24–28), 10 precloacal pores (vs. 5–7 pitted precloacal scales in females); differs from C. montanus by fewer dorsal tubercle rows, DTR 17 (vs. DTR 21–23), fewer paravertebral tubercles, PVT 32 (vs. PVT 37–43), precloacal pores present (vs. PcP absent in female); differs from C. myaleiktaung by fewer mid-ventral scale rows, MVSR 36 (vs. MVSR 57), precloacal pores present (vs. PcP absent), broad regular dark bands absent on dorsum (vs. present); differs from C. nagalandensis by presence of 10 precloacal pores (vs. six pitted precloacal scales in female), 32 paravertebral tubercles (vs. PVT 35–37); differs from C. namtiram by fewer dorsal tubercle rows, DTR 17 (vs. DTR 21); differs from C. ngopensis by presence of 10 precloacal pores in female, (vs. 0–6 pitted precloacal scales in female); differs from C. septentrionalis by fewer dorsal tubercle rows, DTR 17 (vs. DTR 23 or 24), less paravertebral tubercles, PVT 32 (vs. PVT 38–42), 10 precloacal pores present (vs. 14 precloacal scales with indistinct depression in female); differs from C. siahaensis by fewer dorsal tubercle rows, DTR 17 (vs. DTR 22–24), presence of precloacal pores in female (vs. PcP absent in female); differs from C. vairengtensis by the number of dorsal tubercle rows, DTR 17 (vs. DTR 22–23), 32 paravertebral tubercles (vs. PVT 34–39), 10 precloacal pores present (vs. 5–9 precloacal pits in female). Morphological differences with other members of khasiensis group is presented in Table 2.

Cyrtodactylus barailensissp. nov. has a high genetic divergence of 10.9% from its closely related C. namtiram. With other members of the clade, C. kiphiresp. nov. has a genetic divergence of 10.7% and 22.3% in the ND2 gene.

The specific epithet is a toponym derived from the name of the hill range “Barail” where the type locality of the species lies.

Barail Hills bent-toed gecko.

During our two-day survey we only located a single individual of this species. Thus, the new species is currently known only from the type locality, in Peren District, Nagaland, India. We recorded this species on the trunk of a small tree at a height of approximately 2 m from the ground in the Athibung Reserve Forest at approximately 20:00 hrs on 14 August 2021. The forest type is semi-evergreen with relatively little anthropogenic pressure.

Adult male (WII-ADR1596), collected near Lamdan Kabui village (24.5954°N; 93.7085°E; elevation 1240 m a.s.l.) (Fig. 3A), Churachandpur District, Manipur, India collected by Bitupan Boruah on 25 July 2022.

Medium-sized gecko, SVL at least 59.5 mm in adult male; 10 supralabials; eight or nine infralabials; 21 bluntly conical and feebly keeled tubercles across midbody; 37 paravertebral tubercles; 36 mid-ventral scales between ventrolateral folds; 11 or 12 subdigital lamellae beneath digit IV of manus; 13–16 subdigital lamellae beneath digit IV of pes; seven precloacal pores arranged in a continuous series; six irregular shaped dark-brown cross bands on back between axilla and groin; tail dorsally with eight dark-brown and seven pale-brown bands arranged alternatively.

Specimen well preserved except an incision below left axilla ventrolaterally. Snout-vent length 59.5 mm. Head moderately large (HL/SVL = 0.26), oval, dorsoventrally depressed, longer than width (HW/HL = 0.71), distinct from neck, broader at occipital region; snout tip rounded in both dorsal and lateral view; loreal region convex; canthus rostralis rounded, indistinct; interorbital space flat; a longitudinal furrow on dorsal surface of the snout; snout short (SO/HL = 0.39), longer than orbit (OD/SO = 0.68); nostril nearly rounded, opening directed posterolaterally; ear opening rounded; scales on head heterogeneous, largest on snout and loreal region, posteriorly smaller in upper eyelid, interorbital space and occipital region, granular juxtaposed; scales on upper eyelids heterogeneous; supraciliaries outwardly sharp giving serrated appearance in dorsal view, size anteriorly and posteriorly decreases, largest at the anterodorsal region; rostral wide, a short groove at the middle on top, rostral connected with nasals, supranasals, an internasal and first supralabials, two scales between the supranasals paced longitudinally, granular scales at parietal, occipital and temporal region intermixed with slightly large rounded granular tubercles starting from the level of posterior margin of the upper eyelids, those on temporal region are slightly larger than that of occipital region, size of the tubercles increases towards nape; 10 supralabial scales on both sides, size decreases towards angle of jaw; 8 supralabials up to midorbit on both sides; mental nearly triangular, connected with first infralabials, inner postmentals; nine infralabials on right side and eight on left side, size decreases towards angle of jaw; first infralabials connected with mental, second infralabial, inner and outer postmentals; inner pair of postmentals are larger than the outer postmentals; two rows of slightly enlarged scales along the infralabials starting below the outer postmentals, posteriorly size of those decreases; rest of the gular scales are small, granular juxtaposed, nearly homogeneous, size increases towards the throat where they become imbricate.

Habitus slender (BW/SVL = 0.16, TRL/SVL = 0.49), dorsoventrally depressed, dorsal scales granular, rounded, heterogeneous, intermixed with rounded, weakly keeled and bluntly conical tubercles irregularly arranged, up to third segment of the tail, size increases towards posterior body and pronounced at the base of tail; 21 dorsal tubercles across mid dorsum; 37 paravertebral tubercles; ventrolateral fold weak; ventral scales larger than those of dorsal, flat, smooth, cycloid subimbricate to imbricate; 36 mid-ventral scales between ventrolateral fold; seven precloacal pores arranged in an inverted “V” shaped continuous series, followed by a series of five unpored enlarged scales below it, largest at the apex, an unpored scale equal size to PcP present in continuous with the PcP in both ends; scales above the PcP larger than those of belly scales.

Forelimbs and hindlimbs slender (FL/SVL = 0.14, CL/SVL = 0.17), digits strongly inflected at the joints, all bearing large recurved claw, enlarged subdigital lamellae; lamellae beneath digit IV of right and left manus (given as basal + distal) is 6+5 and 6+6 respectively; lamellae beneath digit IV of both right and left pes (given as basal + distal) is 6+7 and 7+9 respectively; dorsal scales on forelimbs heterogeneous is size, mostly granular; proximal scales on upper arm are smaller than that of lower arm; scales near elbow on lower arm smooth and subimbricate; upper arm scales granular at proximal part and at the distal end it is smooth, cycloid and imbricate; dorsal scales of hindlimbs heterogeneous, intermixed with slightly enlarged, feebly keeled and bluntly conical tubercles; scales on inner lateral side of the thighs are smooth, large and subimbricate, those on dorsal side are small granular; scales on tibia are small, granular juxtaposed; ventral scales of forelimbs granular, juxtaposed, heterogeneous; scales on palm heterogeneous in shape and size, granular juxtaposed; ventral scales on hindlimbs heterogeneous; most of the thigh scales are smooth, cycloid and subimbricate, but on the knee and below the level of precloacal pores, scales are smaller and granular; scales above the vent granular; tibia scales smooth, nearly homogeneous, cycloid and subimbricate; scales on soles heterogeneous, granular, juxtaposed to subimbricate.

Tail complete (TL = 67 mm), slender, gradually tapering towards tip, segments indistinct, dorsal scales small, granular, juxtaposed at the base, posteriorly size increases, flat, smooth, subimbricate, heterogeneous in shape and size, large feebly keeled scales up to third segment of the tail, those on basal segment are pronounced; subcaudal scales smooth, subimbricate, wider than that of dorsal, heterogeneous in shape and size; no enlarged plate like series of subcaudal scales; three bluntly conical spurs on both sides of the tail base.

Head on top and laterally brown; upper eyelids grey; dorsal ground colour of neck and back slightly paler than that of head; dark-brown irregular patches on neck up to fore limb insertion level; six irregular shaped dark-brown cross bands on back between axilla and groin, mid-dorsally interrupted; one dark-brown band on sacrum; tail dorsally with eight dark-brown and seven pale-brown bands arranged alternatively; dark bands are comparatively broader than the pale bands; the first dark-brown band on tail base broken mid-dorsally into two enlarged elongated spots. Ventrally head, neck, trunk and limbs light-brown; tail with irregular dark-brown specks.

Cyrtodactylus manipurensissp. nov. differs from C. aaronbaueri by fewer dorsal tubercle rows, DTR 21 (vs. DTR 22–28); differs from C. aunglini by fewer mid-ventral scale rows, MVSR 36 (vs. DTR 47–49), by fewer precloacal pores, PcP 7 (vs. PcP 12 or 13); differs from C. barailensissp. nov. by number of dorsal tubercle rows, DTR 21 (vs. DTR 17), number of paravertebral tubercle rows, PVT 37 (vs. PVT 32); differs from C. bengkhuaiai by fewer dorsal tubercle rows, DTR 21 (vs. DTR 22–26); differs from C. brevidactylus by having fewer dorsal tubercle rows, DTR 21 (vs. DTR 27–30), enlarged chocolate-brown patches on head and back absent (vs. present); differs from C. chrysopylos by much smaller body size, SVL 59.5 mm (vs. SVL 64.9–79.1 mm in male), fewer precloacal pores, PcP 7 (vs PcP 8–13); differs from C. dianxiensis by smaller body size, SVL 59.5 mm (vs. SVL 73.8–79.9 mm), 21 rows of dorsal tubercle rows (vs. DTR 18 or 19); differs from C. gansi by fewer precloacal pores, PcP 7 (vs. PcP 16–29), ventrolateral fold present on trunk (vs. absent); differs from C. jaintiaensis by much smaller body size, SVL 59.5 mm (vs. SVL 87.0–88.3 mm in male), fewer precloacal pores in male, PcP 7 (vs. PcP 11 or 12); differs from C. kiphiresp. nov. by smaller body size, SVL 59.5 mm (SVL 63.9–64.7 mm), number of dorsal tubercle rows, DTR 21 (vs. DTR 16), number of paravertebral tubercle rows, PVT 37 (vs. PVT 26 or 29); differs from C. lungleiensis by higher number of precloacal pores, PcP 7 (vs. PcP 3–5 in male), fewer dorsal tubercle rows, DTR 21 (vs. 24–28); differs from C. montanus by larger body size, SVL 59.5 mm (vs. 53.6–55.0 mm in male); differs from C. myaleiktaung by fewer mid-ventral scale rows, MVSR 36 (vs. MVSR 57), precloacal pores present (vs. PcP absent), broad regular dark bands absent on dorsum (vs. present); differs from C. nagalandensis by higher number of dorsal tubercle rows, DTR 21 (vs. DTR 16–18); differs from C. namtiram by having fewer precloacal pores, PcP 7 (vs. PcP 12); differs from C. ngopensis by the number of precloacal pores, PcP 7 (vs. PcP 6); differs from C. siahaensis by be fewer subdigital lamellae, 11 or 12 lamellae under fourth finger (vs. 13 or 14 lamellae under fourth finger), dark-brown broad bands consists of two series of enlarged spots on dorsum (vs. dark-brown blotches in the form of reticulation on dorsum); differs from C. vairengtensis by fewer precloacal pores, PcP 7 (vs. PcP 9–11). Morphological differences with other members of khasiensis group is presented in Table 2.

Cyrtodactylus manipurensissp. nov. has a high genetic divergence of 11.2–11.3% from its closely related C. ngopensis, 10–10.3% from C. aaronbaueri, 10.2–10.5% from C. vairengtensis and 9.7–11.1% from the C. montanus. With other members of the clade, C. kiphiresp. nov. has a genetic divergence of 8.6% and 23% in the ND2 gene.

This species is named after Manipur state in India.

Manipur bent-toed gecko.

Cyrtodactylus manipurensissp. nov. is currently only recorded from the type locality. We collected a single individual at 18:00 hrs on 25 July 2022 near Lamdan Kabui village. It was perched on a shrub at a height of approximately 1.5 m, on the road connecting Leimatak and Charoikhullen. The habitat is secondary forest, with Zingiber sp. cultivation and settlements.

Adult male (WII-ADR1057; Fig. 7A–J), from Ngengpui Wildlife Sanctuary (22.4906°N; 92.7575°E; elevation 160 m a.s.l.) (Fig. 3A), Lawngtlai District, Mizoram, India collected by Abhijit Das, Bitupan Boruah and Samuel Lalronunga on 8 September 2021.

Two adult females (WII-ADR991 and WII-ADR1058) and one subadult female (WII-ADR1059) collected from the same locality in Ngengpui Wildlife Sanctuary by the same team on 7 and 8 September 2021.

Medium-sized gecko (SVL at least 61.2 mm in adult male and 72–74.1 mm in adult females); 9–12 supralabials; 8–11 infralabials; 18–20 bluntly conical and feebly keeled dorsal tubercles; 29–34 paravertebral tubercles; 38 or 39 midventral scale rows between the weak ventrolateral folds; no precloacal groove; at least 27 precloacofemoral pores in continuous series in male and 10–16 small precloacal pores in females; short dark-brown bars between a pair of dorsolateral stripes or enlarged irregular dark-brown spots present on dorsum.

Holotype well preserved except an incision below left axilla ventrolaterally. Snout-vent length 61.2 mm. Head moderately large (HL/SVL = 0.27), dorsoventrally depressed, longer than width (HW/HL = 0.68), distinct from neck, broader at occipital region, snout tip rounded in both dorsal and lateral view, loreal region convex, canthus rostralis rounded, indistinct, interorbital space flat, a longitudinal furrow on dorsal surface of the snout, snout short (SO/HL = 0.4), longer than orbit (OD/SO = 0.66), nostril nearly rounded, opening directed posterolaterally, ear opening oval and oblique, scales on head heterogeneous, largest on snout and loreal region, posteriorly smaller in upper eyelid, interorbital space and occipital region, granular juxtaposed, scales on upper eyelids heterogeneous, supraciliaries outwardly sharp giving serrated appearance in dorsal view, size anterior and posterior end decreases, rostral wide, a short groove at the middle on top, rostral connected with nasals, supranasals, an internasal and first supralabials, two scales between the supranasals, larger than the rest of the granular snout scales, granular scales at parietal region and occipital region intermixed with slightly large rounded granular tubercles, dense in occipital and temporal region and size increases towards nape, nine supralabials on right and 10 on left side, size decreases towards angle of jaw, supralabials up to midorbit seven on right and eight on left side, a series of narrow and slightly elongated scales above the supralabials between nostril and anterior border of the orbit, mental as wide as rostral, triangular, connected with first infralabials, inner postmentals, nine infralabials on both side, size decreases towards angle of jaw, first infralabials connected with mental, second infralabial, inner and outer postmentals, inner pair of postmentals are larger than the outer postmentals, posterior margin of the inner postmentals bordered by seven granular scales of different size, two or three rows of enlarged scales along the infralabials starting below the outer postmentals, posteriorly size of those decreases, elongated and narrow, rest of the gular scales are small, granular juxtaposed, homogeneous, size increases towards the throat where they become imbricate.

Habitus slender (BW/SVL = 0.17, TRL/SVL = 0.41), dorsoventrally depressed, dorsal scales granular, rounded, heterogeneous, intermixed with rounded, weakly keeled and bluntly conical tubercles irregularly arranged, starting from occipital region to seventh segment of the tail, size increases towards posterior body, pronounced at sacrum and base of the tail, 18 dorsal tubercles across mid dorsum, 29 paravertebral tubercles, ventrolateral fold weak, ventral scales larger than those of dorsal, flat, smooth, cycloid subimbricate to imbricate, largest on belly, 39 mid-ventral scales between ventrolateral fold, 27 precloacal femoral (PcFP) pores in a continuous series (Fig. 7I), followed by eight unpored, large scales below the PcFP at the middle.

Forelimbs and hindlimbs slender (FL/SVL = 0.14, CL/SVL = 0.19); digits strongly inflected at the joints, all bearing large recurved claw, enlarged subdigital lamellae; lamellae beneath digit IV of right and left manus (given as basal + distal) is 5+8 and 5+9 respectively; lamellae beneath digit IV of right and left pes (given as basal + distal) is 4+10 and 5+10 respectively; dorsal scales on forelimbs heterogeneous, granular juxtaposed; dorsal scales of hindlimbs heterogeneous, granular, intermixed with densely placed large, rounded and bluntly conical tubercles, scales on inner lateral side of the thighs near knee are subimbricate; ventral scales of forelimbs granular, juxtaposed, mostly homogeneous; scales on palm heterogeneous in shape and size, granular; scales on ventral side of hindlimbs nearly equal to those of belly, smooth, cycloid and subimbricate, but on the knee, above cloaca and on thigh below the level of precloacal pores are smaller and granular; scales on soles heterogeneous, granular, juxtaposed to subimbricate.

Tail complete (TL = 70 mm), slender, gradually tapering towards tip, segments indistinct, dorsal scales granular, juxtaposed, flat, smooth, heterogeneous in shape and size; enlarged feebly keeled scales up to eight segments of the tail, those on basal segment are pronounced; subcaudal scales smooth, subimbricate, wider than that of dorsal, heterogeneous in shape and size; no enlarged plate like series of subcaudal scales.

Fig. 7J. Top of head pale-brown with indistinct pale-cream coloured spots on anterior part, a few slightly dark-brown patches on upper eyelids, temporal and occipital region; supraciliary yellowish-brown; dark-brown postocular streak on each side which continues dorsolaterally along neck and trunk to tail base, on right side this stripe is disjunct; a pale-cream coloured steak on each side of the dark-brown post orbital streak; an indistinct brown stripe along loreal region and a pale-cream coloured streak above it; upper lip with irregular brown and pale-yellow spots; a pair of dark-brown stripe with irregular edge on neck interspaced by an elongated pale-cream coloured patch, posterior ends of these stripes are connected; an enlarged dark-brown spot at the middle of nape; a pair of narrow pale-cream coloured stripe on neck above the lateral dark-brown stripe running up to slightly behind the level of axilla; dorsum pale-brown with seven irregular shaped and sized, broad dark-brown cross bars, these bars posteriorly edged with narrow cream coloured patch, first two cross bars are shorter than the rests; a few enlarged whitish spots along lateral side of the trunk; dorsally limbs pale-brown intermixed with slightly darker spots and light patches; dorsally tail with alternating slightly dark-brown broad bands and narrow pale-cream coloured bands of irregular shape and size; spurs cream coloured; ventrally head and trunk whitish with pinkish tinge and scales with brown marbling; some scales above and below the cloaca are pale-yellow; tail with slightly dense brown marbling.

Dorsal and ventral colour nearly the same as that of in life; dorsal markings visible as that of life condition.

Details of the variations in morphometric and meristic characters of the type series are provided in Table S2. Apart from these dorsal markings among the paratypes slightly varied. In the two paratypes (WII-ADR1058 and WII-ADR1059), dorsal bands are broken mid dorsally giving the appearance of enlarged spots (Fig. S2). Precloacal pores in female are smaller than that of male.

Cyrtodactylus ngengpuiensissp. nov. can be differentiated from the members of the C. khasiensis clade (except C. ayeyarwadyensis, C. guwahatiensis, C. karsticola and C. tripuraensis) by the presence of precloacofemoral pores in male (vs. no femoral pores in male of C. agarwali, C. exercitus, C. kazirangaensis, C. khasiensis, C. septentrionalis, C. urbanus). Cyrtodactylus ngengpuiensissp. nov. differs from C. ayeyarwadyensis by presence of tiny precloacal pores in female, PcP 10–16 (vs. PcP absent in female); differs from C. bapme by presence of 10–16 precloacal pores in female (vs. 0–13 pitted precloacal scales), dorsal tubercle rows, DTR 18 or 20 (vs. 21–24); differs from C. guwahatiensis by fewer preclocaofemoral pores, PcFP 27 (vs. PcFP 35–39), PcFP in a continuous series (vs. 26 PcFP interrupted by 11 unpored scales in holotype C. guwahatiensis), PcP present in females (vs. PcP absent in females), higher mid-ventral scale rows, MVSR 38–39 (MVSR vs. 30–35), fewer dorsal tubercle rows, DTR 18–20 (vs. DTR 21–24); differs from C. karsticola by fewer precloacal femoral pores in male, PcFP 27 (vs. PcFP 34–38), less number of dorsal tubercle rows, DTR 18–20 (vs. DTR 21–24), fewer paravertebral tubercles, PVT 29–34 (vs. PVT 34–39); differs from C. tripuraensis by fewer precloacal femoral pores in male, PcFP 27 (vs. PcFP 29–37) and less number of precloacal pores, PcP 10–16 in female (vs. PcP 19–29 in female). Morphological differences with other members of khasiensis group is presented in Table 2.

Cyrtodactylus ngengpuiensissp. nov. has a moderate genetic divergence of 4.1–6.6% from its closely related C. ayeyarwadyensis, 4.2–6.0% from C. tripuraensis. With other members of the clade, C. ngengpuiensissp. nov. has a genetic divergence between 8.6% and 15.7% in the ND2 gene. The intraspecific divergence is between the two samples of C. ngengpuiensissp. nov. is 0.8%.

The specific epithet is a toponym derived from the name “Ngengpui Wildlife Sanctuary” of Mizoram state from where the type series of the species were collected.

Ngengpui bent-toed gecko.

Cyrtodactylus ngengpuiensissp. nov. new species is currently only known from the type locality, the Ngengpui Wildlife Sanctuary, Lawngtlai District, Mizoram, India. The forest is characterised as tropical semi-evergreen to moist evergreen forest. The forest is dominated by Dipterocarpus spp., palms, canes and rattans. Individuals of C. ngengpuiensissp. nov. were collected during 6–9 September 2021 at 21:00–22:00 hrs. Individuals were recorded in bamboo thickets, on tree buttresses and trunks, and amongst ferns and rocks along the banks of evergreen forest streams.

Adult male (WII-ADR1416), collected from Kamala Valley (27.4595°N; 96.4279°E; elevation 650 m a.s.l.), Namdapha Tiger Reserve, Changlang District, Arunachal Pradesh, India by Abhijit Das and Bitupan Boruah on 18 May 2022 (Fig. 3A).

Two adult males (WII-ADR1415, WII-ADR1417) collected from the same locality as the holotype on the same date; one adult female (WII-ADR1404) collected near Deban (27.4942°N; 96.3701°E; elevation 400 m a.s.l.), Namdapha Tiger Reserve, Changlang District, Arunachal Pradesh, India by Abhijit Das and Bitupan Boruah on 11 May 2022; two adult females (WII-ADR3067, WII-ADR3068) collected from Motijheel trail (27.4899°N, 96.3348°E; elevation 470 m a.s.l.), Gibbons’ Land, Namdapha Tiger Reserve, Changlang District, Arunachal Pradesh, India by Abhijit Das and Bitupan Boruah on 11 September 2022; one adult female (WII-ADR1790) collected near Burma Nullah (40 mile point) (27.4878°N; 96.5416°E; elevation 480 m a.s.l.), Namdapha Tiger Reserve, Changlang District, Arunachal Pradesh, India by Abhijit Das and Bitupan Boruah on 18 September 2022.

An adult female (WII-ADR3060) collected from Sinabrai (27.7434°N; 96.3872°E; elevation 470 m a.s.l.), Kamlang Tiger Reserve, Lohit District, Arunachal Pradesh, India on 5 September 2022 by Bitupan Boruah and Abhijit Das; an adult female (WII-ADR3281) and an adult male (WII-ADR3282) collected from Hornbill (27.5381°N; 96.4403°E; elevation 670 m a.s.l.), Namdapha Tiger Reserve, Changlang District, Arunachal Pradesh, India on 10 May 2023 by Rajiv N.V.

Medium-sized gecko (SVL 57.5–70.7 mm in males and SVL 54.8–69.3 mm in females); supralabials and infralabials 8–11; dorsum with weakly keeled and bluntly conical tubercles, 29–36 paravertebral tubercles between the level of axilla and level of groin; 17–19 dorsal tubercle rows at mid body; 33–40 mid ventral scale rows; seven to nine precloacal pores in males and 8–10 small precloacal pores in females in a continuous series, PcP much smaller in females than that of males; 12–14 subdigital lamellae on finger IV and 11–17 subdigital lamellae on toe IV; irregular dark-brown spots or stripes on dorsum.

Holotype well preserved except a ventrolateral incision below left axilla. Snout-vent length 65.3 mm. Head moderately large (HL/SVL = 0.27), dorsoventrally depressed, ovoid in shape, longer than width (HW/HL = 0.66), distinct from neck, broader at occipital region; snout rounded in both dorsal and lateral view; loreal region convex; canthus rostralis rounded, indistinct; interorbital space flat, a longitudinal furrow on dorsal surface of the snout, snout short (SO/HL = 0.4), longer than orbit (OD/SO = 0.71); nostril semicircular, opening directed posterolaterally; ear opening oval and oblique; scales on head heterogeneous, largest on snout and loreal region, posteriorly smaller, interorbital space and occipital region, granular juxtaposed; scales on upper eyelids nearly homogeneous, granular juxtaposed; supraciliaries outwardly sharp giving serrated appearance in dorsal view, size anteriorly and posteriorly decreases, largest at the anterodorsal region; rostral wide, a short groove at the middle on top, rostral connected with nasals, supranasals, internasals and first supralabials; granular scales at parietal, occipital and temporal region intermixed with slightly large, rounded and bluntly conical tubercles, dense in occipital and temporal region and size increases towards nape; supralabial 11 on right and 10 on left side, supralabials up to midorbit eight on right and seven on left side, size decreases towards angle of jaw; a series of scales slightly larger than the loreal scales present above the supralabials, posteriorly size decreased; mental as wide as rostral, nearly triangular, connected with first infralabials, inner postmentals; nine infralabials on right and eight on left side, size decreases towards angle of jaw; inner pair of postmentals are larger than the outer postmentals; two rows of enlarged scales along the infralabials starting below the outer postmentals, posteriorly size of those decreases; rest of the gular scales are small, granular juxtaposed, homogeneous, size increases towards throat where they becomes imbricate.

Habitus slender (BW/SVL = 0.17, TRL/SVL = 0.47), dorsoventrally depressed; dorsal scales granular, rounded, heterogeneous, intermixed with rounded, irregularly arranged weakly keeled and bluntly conical tubercles, these tubercles continues to fourth segment of the tail, size increases towards posterior body and pronounced; 18 dorsal tubercles across mid dorsum; 36 paravertebral tubercles; ventrolateral fold weak; ventral scales larger than those of dorsal, flat, smooth, cycloid subimbricate to imbricate, largest towards belly; 40 mid-ventral scales between ventrolateral fold; seven precloacal pores arranged in an inverted “V” shaped continuous series, followed by five unpored, large scales below it, one enlarged unpored scale present on right end of the PcP series.

Forelimbs and hindlimbs slender (FL/SVL = 0.15, CL/SVL = 0.18); digits strongly inflected at the joints, all bearing large recurved claw, enlarged subdigital lamellae; lamellae beneath digit IV of right and left manus (given as basal + distal) is 5+8; lamellae beneath digit IV of right and left pes (given as basal + distal) is 6+8 and 5+8 respectively; dorsal scales on forelimbs smooth, subimbricate and heterogeneous, small and granular at elbow, scales on forearm nearly rounded while those on hind arm are posteriorly tapering; forearm scales intermixed with enlarged rounded and bluntly conical tubercles; dorsal scales of hindlimbs heterogeneous, intermixed with large, rounded and bluntly conical tubercles, dense than those on forelimbs; horizontally upper half of the thigh scales are smooth, large and subimbricate, those on lower half small granular; scales on tibia are small, granular juxtaposed; ventral scales of forelimbs granular, juxtaposed, mostly homogeneous; scales on palm heterogeneous in shape and size, granular juxtaposed; scales on ventral side of hindlimbs smaller than those of belly, smooth, cycloid and subimbricate, but on the knee, above cloaca and on thigh below the level of precloacal pores are smaller and granular; scales on soles heterogeneous, granular, juxtaposed to subimbricate.

Tail regenerated (TL = 79 mm), slender, gradually tapering towards tip, segments indistinct, in the original part of the tail dorsal scales small, granular, juxtaposed at the base, posteriorly size increases, flat, smooth, subimbricate, heterogeneous in shape and size; in the regenerated part of the tail, scales are irregular in shape and size; large feebly keeled scales upto fourth segment of the tail, those on basal segment are pronounced; subcaudal scales smooth, subimbricate, wider than that of dorsal, heterogeneous in shape and size; no enlarged plate like series of subcaudal scales; two and four bluntly conical spurs on right and left side of the tail base respectively.

Top of head pale-brown, upper eyelids grey; neck and dorsum dorsally greyish-brown; two brown stripes with irregular edge on dorsal side of the neck continuing to the level of forelimb insertion, and another stripe on each lateral side of the neck; seven pairs of slightly dark-brown elongated spots of irregular shape and size on dorsum, these spots are outwardly connected to a narrow, brown dorsolateral stripe; tail dorsally pale-brown with broad dark-brown cross bars of irregular shape and size on the original part, the first bar broken into two elongated spots, regenerated part of the tail is plain pale-brown; ventrally head, trunk and tail pale-cream coloured, a few irregular brown spots on tail.

(Based on paratype WII-ADR1417) (Fig. 8I); head dorsally brown with irregular dark-brown patches, a pair of pale-yellowish spots on loreal in front of anterodorsal corner of eyes, lips slightly paler than dorsal head colour with irregular pale-yellowish spots; a pale-brown postorbital streak; pale-brown irregular spots on dark-brown background on neck; upper eyelids greyish-brown; slightly dark-brown large spots of irregular size and shape on dorsal and lateral side of the trunk, interspaced with pale-brown patches, dark-brown spots on dorsal side of the trunk giving appearance of continuous stripes up to middle of the trunk; anterior two third of dorsal side of the tail with alternative broad dark-brown and narrow pale-brown bands, those dark-brown bands anteriorly diffused and posterior edge zigzagged, posterior one third brown with dark-brown marbling; limbs brown with irregular pale spots and dark-brown reticulation; digits with alternative dark-brown and pale-brown bands.

Morphological variations are given in Table S2. Except for those, the dorsal marking pattern varied among the collected specimens. The dorsal spots of WII-ADR1417 are in the form of two continuous stripes starting from neck, posteriorly broken; a spot at the middle of occipital region followed by a dark cross bar on the nape; WII-ADR3067 and WII-ADR3281 has four dorsal stripes starting from neck to the level of hind limb insertion; WII-ADR3068 has four stripes on neck, followed by five irregular and zigzag cross bars on dorsum; in WII-ADR1790, dorsal spots are indistinct and irregular; WII-ADR1415 has three pairs of dark spots posteriorly bordered with white spots on neck, cross bars on dorsum narrower, irregular and posteriorly with white (Fig. S2); WII-ADR3060 has six dark brown cross bands on back between the level of axilla and groin, consists of irregular shaped enlarged individual spots, mid dorsally these spots are remarkably disjunct,. Precloacal pores in females are smaller than that of males.

Cyrtodactylus namdaphaensissp. nov. differs from C. mombergi by fewer dorsal tubercle rows, DTR 17–19 (vs. DTR 23–27), by fewer precloacal pores, PcP 7–9 (vs. PcP 10–11). Morphological differences with other members of khasiensis group is presented in Table 2.

Cyrtodactylus namdaphaensissp. nov. has a moderate genetic divergence of 6.2–7.9% from the closely related C. mombergi. The intraspecific divergence among the thirteen samples of C. namdaphaensissp. nov. ranges between 0.1–5%.

The specific epithet is a toponym named after its type locality Namdapha Tiger Reserve in Arunachal Pradesh, India.

Namdapha bent-toed gecko.

So far, Cyrtodactylus namdaphaensissp. nov. has been recorded within an elevational range of 400–650 m a.s.l. inside Namdapha Tiger Reserve. All the localities are south of Noa-Dihing River in Changlang District, Arunachal Pradesh, India. We recorded this species between May and September 2022. Individuals were recorded on tree bark, ferns, and riparian vegetation and along forest trails between 18:00–23:00 hrs. The forest type can be classified as Assam Valley Tropical Evergreen Forest. The area had a distinct understory with a thick covering of leaf-litter. Currently the species has been recorded from 25 Mile, Burma Nullah (40 Mile), Gibbons Land, Motijheel trail and at Hornbill camp located at the north bank of Noa-Dihing River within the Namdapha Tiger Reserve. We also recorded this species on rocks and vegetation near Kamlang River at Sinabrai, near the Kamlang Tiger Reserve, at an elevation of 470 m a.s.l. We observed juveniles of this species on the forest floor in the leaf litter during May, 2022. During 2023, adult individuals were seen on the ferns overhanging first order streams. At the slightest disturbance, the lizards would drop into the thick vegetation below. Other arboreal reptile taxa from the area included Ptyctolaemus sp., Japalura sp. and Pareas sp.

Adult female (WII-ADR1177; Fig. 9A, B, E–I), from Kalek stream (28.1711°N; 95.2420°E, elevation 210 m a.s.l.; Fig. 3A); 1.8 km (aerial distance) northwest from Bodak village, East Siang District, Arunachal Pradesh, India collected by Bitupan Boruah on 23 October 2021.

Two adult females (WII-ADR1581 and WII-ADR1582; Fig. 9C, D) from a hill slope near Kalek stream, East Siang District, Arunachal Pradesh, India collected by Bitupan Boruah on 15 June 2022.

Medium-sized gecko (SVL 70.1–72.1 mm in females); males unknown; supralabials 8–12 and infralabials 9–12; tubercles on dorsum weakly keeled and bluntly conical, 26–32 paravertebral tubercles between the level of axilla and level of groin; 15 or 16 dorsal tubercle rows at mid body; 40–45 mid ventral scale rows; 8–10 small precloacal pores in a continuous series; 13–17 subdigital lamellae on finger IV and 14–19 subdigital lamellae on toe IV; six or seven irregular and broken dark-brown bands on dorsum between the level of axilla and level of groin, or irregular dark-brown reticulation on dorsum.

Holotype well preserved except an incision below left axilla ventrolaterally. Snout-vent length 72.1 mm. Head moderately large (HL/SVL = 0.26), dorsoventrally depressed, longer than width (HW/HL = 0.72), distinct from neck, broader at occipital region; snout rounded in both dorsal and lateral view; loreal region convex; canthus rostralis rounded, indistinct; interorbital space flat, a longitudinal furrow on dorsal surface of the snout, snout short (SO/HL = 0.39), longer than orbit (OD/SO = 0.73); nostril nearly rounded, opening directed posterolaterally; ear opening oval and oblique; scales on head heterogeneous, largest on snout and loreal region, posteriorly smaller in upper eyelid, interorbital space and occipital region, granular juxtaposed; scales on upper eyelids heterogeneous; supraciliaries outwardly sharp giving serrated appearance in dorsal view, size anteriorly and posteriorly decreases, largest at approximately anterior one third of it; rostral wide, a short groove at the middle on top, rostral connected with nasals, supranasals, an internasal and first supralabials; a single scale between the supranasals, larger than the rest of the granular snout scales; granular scales at parietal region and occipital region intermixed with slightly large rounded granular tubercles, dense in occipital and temporal region and size increases towards nape; supralabial eight on right and 11 on left side, supralabials upto midorbit seven on right and eight on left side, size decreases towards angle of jaw; a series of scales nearly rounded, granular and slightly larger than the loreal scales present above the supralabials; mental as wide as rostral, triangular, connected with first infralabials, inner postmentals; 10 infralabials on right and nine on left side, size decreases towards angle of jaw, first infralabials connected with mental, second infralabial, inner and outer postmentals; inner pair of postmentals are larger than the outer postmentals, posterior margin of the inner postmentals bordered by eight granular scales of different size; one or two rows of enlarged scales along the infralabials starting below the outer postmentals, posteriorly size of those decreases, elongated and narrow; rest of the gular scales are small, granular juxtaposed, homogeneous, size increases towards throat where they become imbricate.

Habitus slender (BW/SVL = 0.18, TRL/SVL = 0.45), dorsoventrally depressed; dorsal scales granular, rounded, heterogeneous, intermixed with rounded, weakly keeled and bluntly conical tubercles irregularly arranged, starting from occipital region to fourth segment of the tail, size increases towards posterior body, pronounced at sacrum and base of the tail; 15 dorsal tubercles across mid dorsum; 26 paravertebral tubercles; ventrolateral fold weak; ventral scales larger than those of dorsal, flat, smooth, cycloid subimbricate to imbricate, largest on belly; 40 mid-ventral scales between ventrolateral fold; very small, 10 precloacal pores arranged in an inverted “V” shaped continuous series, followed by 7 unpored, large scales below it, largest at the apex.

Forelimbs and hindlimbs slender (FL/SVL = 0.14, CL/SVL = 0.17); digits strongly inflected at the joints, all bearing large recurved claw, enlarged subdigital lamellae, lamellae beneath digit IV of right and left manus (given as basal + distal) is 6+11 and 6+10 respectively, the second most lower lamellae of the basal series of digit IV of both sides are divided (not included in lamellar count); lamellae beneath digit IV of right and left pes (given as basal + distal) is 7+11 and 6+10 respectively; dorsal scales on forelimbs heterogeneous, granular juxtaposed, smooth and subimbricate at distal end of forearm; forearm intermixed with a few rounded, large tubercles; dorsal scales of hindlimbs granular intermixed with large, rounded, bluntly conical tubercles; scales on thigh towards knee smooth subimbricate; ventral scales of forelimbs granular, juxtaposed, mostly homogeneous; scales on palm heterogeneous, granular juxtaposed; scales on hindlimbs smaller than those of belly, smooth, cycloid and subimbricate; scales on the knee, above cloaca and on thigh below the level of precloacal pores are smaller and granular; scales on soles heterogeneous, granular, juxtaposed to subimbricate.

Tail complete, regenerated (TL = 80 mm), slender, gradually tapering, segments indistinct; dorsal scales small, granular, juxtaposed at the base, posteriorly size increases, flat, smooth, subimbricate, heterogeneous in shape and size; large feebly keeled scales up to fourth segment of the tail at the distal end of each segment, those on basal segment are pronounced; subcaudal scales smooth, subimbricate, wider than that of dorsal, heterogeneous in shape and size; below cloaca scales are granular, smaller than the rest of the tail scales; no enlarged plate like series of subcaudal scales; three bluntly conical spurs on each side of the tail base present.

Top of head, limbs and tail pale-brown; dorsum pale-greyish-brown; a slightly dark-brown stripe radiating from posterior orbital border to occipital region, medially “W” shaped; an another short dark-brown stripe at the middle of the head; a short, posteriorly diffused dark-brown stripe starting from posterior corner of the upper eyelids; a faint brown stripe on the loreal region; rostral and mental scale, labial scales dark-greyish-brown with white patch posteriorly; short and irregular shaped, longitudinal dark-brown stripe on dorsal side of neck; six irregular shaped, zig-zagged dark-brown bands on dorsum; irregular shaped, large dark-brown spots on sacrum and tail base; zig-zagged dark-brown bands on anterior part of the tail, posterior half of the tail (regenerated part) with brown speculation; indistinct brown reticulation on dorsal side of the limbs; ventrally head, body and limbs creamy-white with brown speculation; chin scales and scales below infralabials heavily speckled; lamellae, palm and feet heavily speckled with brown; indistinct brown patches on ventral side of the tail.

Detailed morphological variations among the collected individuals of Cyrtodactylus siangensis are provided in Table S2. Dorsal dark brown markings in the two paratypes are variable in the form of irregular zig-zagged bands to reticulation; reticulation on limbs of the two paratypes distinct than the holotype; cross bands on tail of WII-ADR1581 darker posteriorly (Fig. 9).

Morphologically Cyrtodactylus siangensissp. nov. is close to C. cayuensis, however, phylogenetically it is distinct from the later (Fig. 2; Table S3), also the type locality of the new species is separated by Siang River from the C. cayuensis localities recorded in this study (Fig. 3B). Morphological differences with other members of khasiensis group is presented in Table 2.

Cyrtodactylus siangensissp. nov. has a high genetic divergence of 11.7–17.0% from its closely related and the only other species in the clade, C. cayuensis.

The specific epithet is a toponym derived from the name of the river “Siang”. The type locality of this species lies in the Siang valley of Arunachal Pradesh.

Siang Valley bent-toed gecko.

Cyrtodactylus siangensissp. nov. is currently only known from the type locality, Bodak, East Siang District. It appears to be uncommon, and likely a habitat generalist. The holotype was encountered on a shrub at the edge of Kalek stream (a tributary of Siang River) at a height of approximately one meter above the ground, at 18:15 hrs. The stream at the time of survey was flowing at a moderately speed, was approximately 10 m in width and contained boulders (sedimentary rock) and fallen logs. The two paratypes were recorded on a small tree at a height of 2.0 m on the slope of a hill close to the collection site for the holotype. The type locality is ca. 300 m from Siang River and approximately 150 m from the road connecting Pasighat and Yinkiong.

Cyrtodactylus khasiensis cayuensis – Li (2007)

Cyrtodactylus cayuensis – Agarwal et al. (2018: 337)

Cyrtodactylus arunachalensis – Mirza et al. (2021)

(Fig. 10). Arunachal Pradesh: one male (WII-ADR1219) and one female (WII-ADR1218) collected from Balek village (28.0624°N; 95.2721°E; elevation 450 m a.s.l.), East Siang District on 29 October 2021 by Bitupan Boruah; one male (WII-ADR1213) collected from Ramsing (28.6563°N; 94.9795°E; elevation 600 m a.s.l.), Mouling National Park, Upper Siang District on 27 October 2021 by Bitupan Boruah; one female (WII-ADR1199) collected from Syrnyup stream (28.5340°N; 95.0305°E; elevation 890 m a.s.l.), Jengging, Mouling National Park, Upper Siang District on 26 October 2021 by Bitupan Boruah; two males (WII-ADR695 and WII-ADR696) and two females (WII-ADR697 and WII-ADR698) collected from Potin (27.3478°N, 93.8497°E, elevation 900 m a.s.l.), Lower Subansiri District on 5 October 2019 by Bitupan Boruah; one female (WII-ADR3017) collected near Glaw lake (27.6960°N; 96.4456°E; elevation 1200 m a.s.l.), Kamlang Tiger Reserve, Lohit District on 2 September 2022 by Abhijit Das, Bitupan Boruah and Naitik G. Patel; one male (WII-ADR1682) and one female (WII-ADR1681) collected from Ezengo (28.1565°N; 95.8638°E; elevation 560 m a.s.l.), Lower Dibang Valley District on 3 August 2022 By Bitupan Boruah; three females (WII-ADR453, WII-ADR454 and WII-ADR459) collected from Jengging (28.5499°N; 95.0537°E; elevation 760 m a.s.l.) by Abhijit Das on 5 October 2018; one female (WII-ADR478) and one male (WII-ADR473) collected from 6 km northwest to Pasighat (28.0945°N; 95.2682°E; elevation 410 m a.s.l.) by Abhijit Das on 2 October 2018.

Medium-sized gecko, SVL 61.2–83.5 mm in adult males and SVL 59.4–83.6 mm in adult females; 8–13 supralabials and 8–12 infralabials; 18–26 rows of bluntly conical and feebly keeled enlarged tubercles across mid-dorsum; 27–38 paravertebral tubercles between the level of axilla and groin; 28–44 mid-ventral scales; 13–21 subdigital lamellae beneath the fourth digit of manus; 11–23 subdigital lamellae beneath the fourth digit of pes; colouration and marking pattern variable; dorsally greyish-brown, pale-brown or pale-yellowish-brown; five to eight dark-brown bands consisting of irregular shaped and sized enlarged spots on back between axilla and groin, or dorsum with dark-brown reticulation; tail with 9–13 dark and 9–12 light bands alternatively arranged on top.

Medium-sized gecko, SVL 61.2–75.1 mm in males (n = 6) and SVL 59.4–83.6 mm in females (n = 11); head moderately large (HL/SVL = 0.25–0.28), dorsoventrally depressed, longer than width (HW/HL = 0.67–0.75), distinct from neck, broader at occipital region; snout rounded in both dorsal and lateral view; loreal region convex; canthus rostralis rounded, indistinct; interorbital space flat, a longitudinal furrow on dorsal surface of the snout, snout short (SO/HL = 0.38–0.43), longer than orbit (OD/SO = 0.49–0.78); nostril opening directed posterolaterally; ear opening oblique; scales on head heterogeneous, largest on snout and loreal region, posteriorly smaller in upper eyelid, interorbital space and occipital region, granular juxtaposed; scales on upper eyelids heterogeneous; supraciliaries outwardly sharp giving serrated appearance in dorsal view, size anteriorly and posteriorly decreases; rostral wide, a short groove at the middle on top; granular scales at parietal region and occipital region intermixed with slightly large rounded granular tubercles, dense in occipital and temporal region and size increases towards nape; 9–13 supralabials present, size decreases towards angle of jaw; a series of scales, slightly larger than the loreal scales present above the supralabials; 8–12 infralabials, size decreases towards angle of jaw; two or three rows of slightly enlarged and narrow scales along the infralabials, posteriorly size of those decreases; rest of the gular scales are small, granular juxtaposed, nearly homogeneous, scales on throat are imbricate.

Habitus slender (BW/SVL = 0.16–0.22, TRL/SVL = 0.41–0.47), dorsoventrally depressed; dorsal scales granular, heterogeneous, intermixed with rounded, weakly keeled and bluntly conical tubercles irregularly arranged, continuing up to third or fifth segment of the tail, size of the tubercles increases towards posterior body, pronounced at sacrum and base of the tail, posteriormost rows indistinctly keeled; 15–21 rows of dorsal tubercles across mid dorsum; 27–38 paravertebral tubercles; ventrolateral fold present; ventral scales on chest and belly larger than those of dorsal, flat, smooth, cycloid, subimbricate to imbricate; 34–44 mid ventral scales; seven or nine distinct precloacal pores in male and 7–10 small precloacal pores in female; PcP followed by three to six unpored enlarged scales below.

Forelimbs and hindlimbs slender (FL/SVL = 0.13–0.15, CL/SVL = 0.16–0.18); digits strongly inflected at the joints, all bearing large recurved claw and enlarged subdigital lamellae, dorsal scales on forelimbs heterogeneous, granular juxtaposed, smooth and subimbricate at distal end of forearm; forearm intermixed with a few rounded, large tubercles; dorsal scales of hindlimbs granular intermixed with large, rounded, bluntly conical tubercles; thigh scales towards lateral and dorsolateral side are smooth and subimbricate; ventral scales of forelimbs granular, juxtaposed, mostly homogeneous; scales on palm heterogeneous, granular juxtaposed; scales on ventral side of hindlimbs slightly smaller than or nearly equal to those of belly, smooth, cycloid and subimbricate; scales on the knee, above cloaca and on thigh below the level of precloacal pores are smaller and granular; scales on soles heterogeneous, granular, juxtaposed to subimbricate.

Tail slender (TL = 58–97 mm), gradually tapering, segments indistinct; dorsal scales small, granular, juxtaposed at the base, posteriorly size increases, flat, smooth, subimbricate, heterogeneous in shape and size; large feebly keeled scales up to third to fifth segment of the tail, those on basal segment are pronounced and posterior rows with indistinct keel; subcaudal scales smooth, subimbricate, wider than that of dorsal, heterogeneous in shape and size; no enlarged plate like series of subcaudal scales; two to four bluntly conical spurs present at the base of the tail. Detailed morphological characteristics of the collected samples are provided in Table S4.

The intraspecific divergence within C. cayuensis ranges between 0.1–6.4%. Among the two clades identified by bPTP, both clades have an intraspecific divergence between 0.1–4.3% and 0.2–2.4%, respectively. The divergence between these two clades range between 3–6.4%.

Dorsal colour and marking pattern variable (Fig. S3). Head and body greyish-brown, pale-brown to pale-yellowish-brown on top; head on top irregularly mottled with dark-brown, slightly enlarged dark-brown irregular patches may be present on occipital area; dark-brown postorbital stripe and loreal stripe present; a light stripe above loreal stripe and postorbital stripe may be visible; a few pale-yellow spots may be present on temporal and occipital region; supraciliary yellowish-brown; neck with short dark-brown streak with irregular edge or with irregular shaped patches; trunk dorsally with five to eight dark-brown bands consisting of irregular shaped and sized enlarged spots and posteriorly pale-yellowish or light-edged, these bands mid dorsally interrupted, or dorsum with dark-brown reticulation; limbs dorsally greyish-brown or pale-yellowish-brown or pale-brown with distinct or indistinct dark-brown reticulation; a dark-brown band on sacrum, band mid dorsally intersected; tail dorsally with 9–13 dark and 9–12 light bands alternatively arranged, bands are with irregular edged, anterior one or two dark bands may be mid dorsally intersected, dark bands are broad and light bands are comparatively narrow, dark bands are posteriorly becomes black or more dark-brown, while light bands become whitish; light bands may have a few dark-brown spots.

The type locality, Xizang (Tibet), China by C. cayuensis also appears to be widely distributed in Arunachal Pradesh, India (Fig. 3B), within an elevational range of 410–1200 m a.s.l. Cyrtodactylus cayuensis is widespread in the foothills of Arunachal Pradesh from the western part of the state, at Seijusa (Pakke Tiger Reserve) to the eastern part (Kamlang Tiger Reserve) and it has successfully spread across major tributaries of the Brahmaputra River, i.e., the Subansiri, Siang, Dibang, Lohit up to at least north of Noa-Dihing. Populations of C. cayuensis have been recorded in degraded evergreen forest as well as in undisturbed semi-evergreen and evergreen forests. Cyrtodactylus cayuensis is particularly abundant in the vegetation close to the picturesque Glaw Lake in the Kamlang Tiger Reserve. We observed individuals at night on shrubs and bedrocks (sedimentary rock) along small forest streams. We also encountered them on small trees up to a 2.0 m inside the forest at Jengging, Kamlang Tiger Reserve and Mehao Wildlife Sanctuary.