Raorchestes kempiae ( Boulenger, 1919 )

Boruah, Bitupan, Deepak, V. & Das, Abhijit, 2025, Revision of bush frogs, Raorchestes and Philautus (Amphibia: Anura: Rhacophoridae) from the northeast Indian biodiversity hotspot with description of thirteen new species, Vertebrate Zoology 75, pp. 517-625 : 517-625

Raorchestes kempiae ( Boulenger, 1919)

Figures 21 , 22; Tables 1, 2, S 6, S 12

Synonymy and chresonymy.

Ixalus kempiae Boulenger, 1919: 208 .

Rhacophorus ( Philautus) kempiae — Ahl (1931): 53, 69.

Philautus kempiae — Bourret (1942): 450 –451; Inger (1985): 91; Ahmed et al. 2009: 16.

Philautus ( Philautus) kempiae — Bossuyt and Dubois (2001): 41.

Raorchestes asakgrensis Naveen et al., 2024: 362 , 365–370.

Raorchestes kempiae — Naveen et al. (2024): 365 –367.

Comments on taxonomic status.

Boulenger (1919) described Raorchestes kempiae based on a single specimen collected by A. Kemp from “ above Tura ”, Garo Hills. The type locality refers to the ridge / peak immediately above Tura (see Paiva 1919). It is possible that this specimen was collected in 1917, which is evident from Paiva (1919) and Brunetti (1918) that A. Kemp was part of their expedition. However, there was no allocated voucher number available for the type in the original description. Chanda (1994) mentioned a possible location of the type specimen as the “ British Museum, (Natural History), London, U. K. ” which was followed by Bossuyt and Dubois (2001) and repeated in Frost (2025). Chanda et al. (2000) did not provide the presence of the type specimen of “ Ixalus kempiae ” or “ Philautus kempiae ” in the collection of the National Zoological Collection of Zoological Survey of India ( ZSI), Kolkata (Calcutta). Recently Naveen et al. (2024) provided a photograph of a specimen in ZSI as the “ holotype ” ( ZSI 18859 ) of R. kempiae . As there was no published literature that mentions the presence of a holotype with the voucher number “ ZSI 18859 ” prior to Naveen et al. (2024), we were unaware of this specimen during our examination of type specimens of Raorchestes / Philautus housed in ZSI. It was lost and found recently in the ZSI collections ( https://zsicollections.in/specimen/ZSI0000002582). We later examined this specimen.

On 27 May 2022, we encountered a few individuals of bush frogs (Fig. 21 ) in Tura Peak ( 25.502825°N; 90.23921°E; elevation 1060 m a. s. l.), West Garo Hills District of Meghalaya syntopic with R. garo but having a distinctive call pattern. These individuals resemble the original description of Boulenger (1919) by the following morphological characters: 1) dark bar on interorbital space, 2) crossbars on limbs, vertical bars on upper jaw, tubercles on dorsal surface of head and dorsum, 3) barely visible tympanum (although mentioned as “ tympanum hidden ” in original description), 4) indistinct canthus rostralis, and 5) brown marbling on throat and abdomen. Following exceptions were observed in the newly collected materials: 1) snout shape rounded or sub-oval in dorsal view (rounded in holotype), 2) nostril equidistant between eye and snout tip or closer to snout tip than eye (nostril equidistant between eye and snout tip in holotype), 3) concave stripes on dorsum diffused or not visible. For taxonomic stability, we provided an expanded description and determined the phylogenetic position based on specimens collected from the type locality. Thus, based on morphological similarity with the original description of R. kempiae , we refer to this population from “ above Tura ” as topotypic material of the species.

Material examined.

Holotype. ZSI 18859 (Fide: Naveen et al. (2024)), collected by A. Kemp from “ above Tura ” .

Newly collected material.

Topotype: three adult males ( WII-ADA 1497 , WII-ADA 1498 and WII-ADA 1500 ) collected by BB, AD and VJ on 27 May 2022 from Tura peak ( 25.502825°N, 90.23921°E, elevation 1060 m a. s. l.), West Garo Hills District, Meghalaya . Referred materials: two adult males ( WII-ADA 1951 , WII-ADA 1952 ) and one adult female ( WII-ADA 1953 ) collected by BB, AD and VD on 1 July 2016 from Nongkhyllem Wildlife Sanctuary ( 25.90282°N, 91.80384°E, elevation 370 m a. s. l.), Ri-Bhoi District, Meghalaya ; one adult male ( WII-ADA 933 ) collected by BB and AD on 14 August 2021 from Athibung ( 25.541758°N, 93.626190°E, elevation 770 m a. s. l.), Peren District, Nagaland ; three adult males ( WII-ADA 1464 – WII-ADA 1466 ) collected by BB, AD and VJ on 24 May 2022 from Tura Forest IB campus ( 25.51596°N, 90.17733°E, elevation 230 m a. s. l.), West Garo Hills District, Meghalaya ; three adult males ( WII-ADA 1476 – WII-ADA 1478 ) and one female ( WII-ADA 1480 ) collected by BB, AD and VJ on 24 May 2022 from Daribokgre ( 25.48974°N, 90.32374°E, elevation 1140 m a. s. l.), West Garo Hills District, Meghalaya .

Diagnosis.

Medium sized Raorchestes, SVL 17.8–22.9 mm in adult males and SVL 23.0– 23.3 mm in adult females; head width equal to or greater than length ( HL / HW = 0.91–1.01); snout rounded to sub-oval in dorsal view; snout longer than inter-upper eyelid width; canthus rostralis indistinct; numerous tubercles on dorsal surface of head, dorsum, and limbs; numerous spinules on dorsum; pale brown dorsally; pair faint concave brown stripes (“) (”) on dorsum; faint crossbars on limbs; short brown streak present on groin.

Redescription of holotype ( ZSI 18859 ).

Holotype is in bad condition and completely dehydrated (Fig. 21 ). Lower jaw broken; forelimbs broken at elbow; outer two fingers of the right hand damaged; hindlimbs broken at tibio-tarsal articulation; left foot broken at the base; except first toe, all other toes damaged. Small sized Raorchestes, SVL 16.5 mm; head slightly wider than its length ( HL / HW = 0.93); snout rounded in dorsal view and lateral view; snout length equal to eye length; nostril equidistant between eye and snout tip; canthus rostralis indistinct; loreal slightly concave; eyes moderate sized ( EL / HL = 0.46); internarial distance smaller than inter-upper eyelid width ( IN / IUE = 0.88) and greater than upper eyelid width ( UEW / IN = 0.62); tympanum and supratympanic fold invisible; vomerine teeth absent; tongue notch at posterior end; trunk nearly half of snout-vent length ( AG / SVL = 0.48); tibia slightly longer than thigh ( TL / TBL = 0.93); subarticular tubercles on fingers and toes distinct, rounded; inner metatarsal tubercle present; circum-marginal groove present on finger and toe disc.

Skin on dorsal aspect of head and dorsum shagreened; flat tubercles visible on upper eyelid visible; skin on dorsal aspect of forelimb and hindlimb smooth; skin on throat smooth; belly and ventral side of thigh granular; tibia smooth.

Colouration of holotype ( ZSI 18859 ) in preservative.

Dorsal aspect of head, dorsum and limbs brown; upper eyelids pale greyish brown; slightly dark broad patch below eye present; pair of slightly dark concave stripes on dorsum barely visible; a broad and very faint cross bar visible on thigh and tibia; throat, belly and ventral aspect of limbs brown, paler than dorsal colour.

Expanded description of the newly collected topotype ( WII-ADA 1497 : Fig. 22 A – H).

Specimen well preserved except for an incision on the ventral aspect of the right thigh. Snout-vent length 22 mm; head as long as wide; snout sub oval in dorsal view, nearly vertical in lateral view, slightly protruding beyond lower jaw in ventral aspect; snout length slightly smaller than eye length ( SL / EL = 0.94); snout depressed dorsally at internarial region; narial region pronounced; nostrils oval, laterally positioned and obliquely oriented; nostril closer to snout tip than eye; internarial distance slightly smaller than inter-upper eyelid width ( IN / IUE = 0.86) and greater than upper eyelid width (UE/ IN = 0.90); canthus rostralis indistinct, oblique; loreal region concave; eye protruding, moderate in size, less than half of eye length ( EL / HL = 0.41); interorbital space flat; tympanum barely visible, round, upper part concealed by supratympanic fold; supratympanic fold distinct; vomerine teeth absent, choanae round; tongue posteriorly notched; pair of internal vocal sac opening on lower jaw towards angle of jaw; habitus dorso-ventrally flattened, length less than half of snout-vent length ( AG / SVL = 0.46).

Forearm shorter than hand length ( FAL / HAL = 0.95); relative length of fingers = I <II <IV <III; each finger with rounded discs; circum-marginal groove present on finger discs; discs on third and fourth finger wider than tympanic diameter; palmar tubercles present; subarticular tubercles distinct, rounded; proximal subarticular tubercles on third and fourth fingers smaller than distal ones; fine granular nuptial pad on first finger; no webbing on fingers.

Hindlimbs slender; thigh length slightly longer than half of snout-vent length ( TL / SVL = 0.52); tibia length nearly equal to thigh length ( TBL / TL = 0.98); relative length of toes = I <II <III <V <IV; rounded disc on toes; discs slightly smaller than that of fingers; circum-marginal groove present; subarticular tubercles rounded; proximal subarticular tubercle on fourth and fifth toes indistinct compared to that of toe III, IV and V; a distinct inner metatarsal tubercle present, its length smaller than fourth and fifth toe disc widths; outer metatarsal tubercle absent.

Skin on dorsal aspect of snout and head shagreen with irregular sized tubercles; upper eyelids shagreen with indistinct tubercles; prominent tubercles on mandibular region behind the angle of jaw; indistinct tubercles on lateral side of head below loreal region; numerous spinules on dorsum, denser towards dorsolateral side, posteriorly density of spinules decrease and skin nearly smooth in preserved condition; prominent bluntly conical tubercles visible above supratympanic folds in life; indistinct blunt tubercles scattered on middle of dorsum; on dorsal aspect of forelimbs smooth but tubercles visible in life; thighs smooth on top indistinct tubercles on top of the tibia (distinct in life); tubercles on tarsus indistinct; scattered spinules on dorsal part of flank, ventral part of flank smooth; on ventrum, chin smooth; gular skin granular; chest smooth; abdomen coarsely granular; granules on ventral aspect of thighs barely visible unlike in life; tibia and tarsus smooth on ventral aspect; flat granular tubercles sparsely present on ventral aspect of the foot.

Colouration in life.

Dorsal aspect and lateral side of head, dorsum, and dorsal aspect of limbs brown with slightly darker brown spots; dark brown bar on inter-upper eyelid space; pair of diffused concave dark brown stripes on dorsum, posterior ends of it extended toward groin; broad dark brown crossbar on each tibia and thigh; cross bar barely visible on forearms; on ventral side, head, abdomen and limbs flesh-coloured with brown mottling that is absent on posterior part of abdomen; pale creamy white blotches on granules of abdomen, these blotches decrease posteriorly; similar flecks on ventral aspect of thigh, tibia, and tarsus; disc on fingers and toes yellowish on ventral side, becoming brighter on inner two fingers and inner three toes.

Colouration in preservative.

Dorsal aspect of head and dorsum greyish brown; upper eyelids dark grey; grey patch at the middle of posterior part of head; slightly darker streaks present on upper jaw below eyes; dorsal colour on dorsal aspect of forelimbs and hindlimbs slightly paler than that of dorsum except thighs, which is pale yellowish brown; enlarged dark brown patch on vent area; dark brown concave stripes on dorsum visible; dark brown crossbar on dorsal aspect of forearm distinct; similar patches on hand and outer two fingers; crossbar on thigh faint which is distinct on tibia; similar patches on tibia, foot and toes; ventral side of head, trunk, limbs creamy-white with brown mottling; mottling denser along lower jaw, gradually decreasing towards abdomen and completely absent on posterior part of abdomen; brown mottling dense on outer ventrolateral side of forearm, tarsus, and towards knee; mottling absent on inner ventrolateral side of forearm and middle of tibia.

Sexual dimorphism and morphological variation.

Males have a pair of internal vocal sac openings on the lower jaw, an external subgular vocal sac, and a nuptial pad on the first finger. Supratympanic fold may be distinct or indistinct; concave stripes on dorsum, dark bar on interorbital space, and crossbars on limbs may be diffused or not visible in some individuals; a mid-dorsal line on dorsum starting from snout extending to above vent and a similar line on hind limb radiating from the mid-dorsal line above vent extending to heel present in WII-ADA 1478 and WII-ADA 933 ; slightly darker brown streak may be present on groin ( WII-ADA 933 , WII-ADA 1478 ) or this may be continuous with posterior ends of concave stripes on dorsum ( WII-ADA 1497 ). According to Boulenger (1919) the tympanum is hidden in the type. However, our newly collected material reveals that the tympanum is barely visible in most individuals but hidden in others ( WII-ADA 1466 , WII-ADA 1497 , WII-ADA 1951 and WII-ADA 1952 ). Details of morphometric variation are given in Table S 12.

Morphological comparison.

Raorchestes kempiae differs from R. annandalii in having numerous tubercles and spinules on the dorsum (vs. dorsum smooth); it differs from R. andersoni by presence of a faint short brown streak on groin (vs. enlarge black irregular spot with two yellow spots); it differs from R. cinerascens nov. comb. by its snout being longer than inter-upper eyelid width (vs. snout length equal to inter-upper eyelid width); differs from R. dulongensis , R. hillisi , and R. menglaensis by its head length being less than or equal to width (vs. head longer than width); it differs from R. garo by presence of a faint short brown streak on groin (vs. enlarged dark brown patches with or without yellow or pale white spots present on groin), canthus rostralis indistinct (vs. canthus rostralis distinct); it differs from R. hekouensis by presence of nuptial pad only on first finger (vs. nuptial pad present on first and second fingers); it differs from R. huanglianshan and R. tytthus nov. comb. by its snout length being smaller than or equal to eye length (vs. snout length longer than eye length); it differs from R. jadoh by larger body size in adult males, SVL 17.8–22.9 mm (vs. SVL 13.6–14.0 mm in adult males); it differs from R. jadoh and R. jakoid by its snout length being less than or equal to eye length (vs. snout length greater than eye length), and thigh length being greater than or equal to tibia length (vs. thigh length less than tibia length); it differs from R. leiktho by larger body size adult males, SVL 17.8–22.9 mm (vs. SVL 15.7–15.8 mm); it differs from R. longchuanensis and R. yadongensis by its inter upper eyelid width being smaller than eye length (vs. inter upper eyelid width greater than eye length); differs from R. malipoensis by its snout being longer than inter upper eyelid width (vs. snout length less than or equal to inter-upper eyelid width); it differs from R. menglaensis by its head length being less than or equal to its width (vs. head longer than its width); it differs from R. mindat by presence of short streak on the groin (vs. enlarged black and white patches on groin); it differs from R. parvulus by presence of a faint short brown streak (vs. dark brown marbling enclosing a whitish blotch present on groin); it differs from R. rezakhani by presence of spinules intermixed with tubercle on dorsum (vs. scattered tubercles on dorsum). A detailed morphological comparison with other congeners included in this study is provided in Table 1.

Boulenger (1919) noted that R. garo has truncated snout, whereas R. kempiae has a rounded snout, distinguishing the two species despite the limited descriptions. However, our examination of the holotype of R. garo revealed that its snout is damaged (squeezed; see Fig. 15 ). Further assessment of newly collected specimens of both species in this study showed that their snouts are actually similar in shape (rounded to sub-ovoid in dorsal view). Therefore, we conclude that snout shape is not a diagnostic character for distinguishing these two species. Similarly, a light-coloured line on mid-dorsum and dorsal aspect of thigh and tibia may be present in some individuals of both R. kempiae and R. garo (Figs 16 N, O, U and 22 I, L ).

Acoustics.

The calls of R. kempiae were recorded from three localities: Tura Peak on 27 May 2022 at an ambient temperature of 28.3 ° C; in the Forest Rest House Campus, Tura on 24 May 2022 at an ambient temperature of 28.0 ° C and in Daribokgre on 25 May, 2022 at an ambient temperature of 28.0 ° C. All calls were recorded between 18: 30 hrs and 19: 30 hrs. the description of the calls is based on analyses of 40 calls from three different individuals ( WII-ADA 1464 , WII-ADA 1476 and WII-ADA 1497 ). The calls of R. kempiae are single type, pulsatile and emitted mostly at regular intervals and not in groups (Fig. 18 ). The mean call duration is 149.29 ± 43.92 ms (67–200 ms) with 3.43 ± 0.59 pulses per call (2–4 pulses per call) and at a pulse rate of 19.21 ± 2.92 pulses / sec (18.52–21.74 pulses / sec). The call rise time is 1 ms and the fall time is 146 ± 46.88 ms (64–199 ms). The pulses are not closely packed; the mean pulse duration is 21.52 ± 8.09 ms (9–37 ms) and the mean pulse period is 54.55 ± 5.06 ms (45–63 ms). The mean inter-call interval is 833.97 ± 298.61 ms (398–1695 ms). The mean dominant frequency is 3306.41 ± 101.26 Hz (3057.7–3488.4 Hz). A detailed comparison of acoustic characters with that of its congeners is given in Table 2.

Phylogenetic relationship and genetic divergence.

Raorchestes kempiae is sister to an undescribed lineage from eastern Arunachal Pradesh ( UFB 100 and PP 1.0; Figs 2 , 3 ) with genetic divergences of 2.6–3.3 % in the 16 S, 9.0–10.4 % in the cyt b and 5.0–5.8 % in the COI genes. The genetic divergences of R. kempiae with other congeners included in this study is 4.4–9.4 % in the 16 S, 9.0–20.4 % in the cyt b and 5.0–16.4 % in the COI genes (Table S 7 A – C).

Distribution and natural history.

From our current sampling it appears that R. kempiae has a patchy distribution encompassing Tura peak, Nokrek Biosphere Reserve in Garo Hills and Nongkhyllem Wildlife Sanctuary of Meghalaya State and Athibung in Nagaland State within an elevation range of 230–1140 m a. s. l. (Fig. 19 B ). Mathew and Sen (2009) reported this species from Sohra, East Khasi Hills, Meghalaya. Naveen et al. (2024) reported this species from Nongkhyllem as “ R. asakgrensis ”. However, we recover the population from Nongkhyllem Wildlife Sanctuary as R. kempiae sensu stricto (see remark below).

We recorded R. kempiae on shrubs at a perch height of approximately one metre, inside semi-evergreen forest as well as near settlements (Fig. 20 A, B ). We observed calling individuals of R. kempiae and R. garo in the same habitat type at Tura peak and Daribokgre.

Remarks.

Naveen et al. (2024) provided a phylogenetic status of “ R. kempiae ” based on a specimen ( SACON VA 806 ) collected from Mikadogre Community Reserve ( 25.433660°N, 90.398981°E,), South Garo Hills District, Meghalaya at an elevation of 174 m. This collection locality is approximately 18 km southeast of the original type locality of “ above Tura ” which is located at an elevation of 1000–1200 m a. s. l (see Paiva 1919). We argue that Naveen et al. (2024) overlooked the original description by Boulenger (1919) and did not provide a justification for their treatment of the material from Mikadogre as topotypes of R. kempiae .

In our phylogenetic analyses, the “ R. kempiae ” of Naveen et al. (2024) did not cluster with the topotypical R. kempiae collected from Tura peak, instead it is nested within the topotypical samples of R. garo (Figs 2 , 3 ) we collected from the same locality. Thus, our robust phylogenetic analyses suggest that the “ R. kempiae ” of Naveen et al. (2024) should be referred as R. garo . The photograph of an uncollected male with white bar on head provided on page 366 of Naveen et al. (2024) as “ R. kempiae ” falls within the morphological variation in R. garo across its range (see morphological variation section under R. garo and Figs 16 , 17 ). In the present study, we found that the topotypical morph of R. kempiae corresponds to the original publication of Boulenger (1919) in having a blackish bar on the inter-upper eyelid space which is not found in R. garo (see variation in R. garo , Figs 16 , 17 ). Although the type locality of R. kempiae and R. garo are the same and the two species are found in sympatry, they can be clearly distinguished based on their acoustic characteristics (Fig. 18 ). Furthermore, in our PCA the type specimens of R. garo and R. kempiae clustered with its respective topotypes collected in this study (Fig. 6 E ).

Mistaken identity of R. kempiae by Naveen et al. (2024) was subsequently repeated in Naveen et al. (in press) and Warjri et al. (2025). In Naveen et al. (in press) another “ topotype ” of R. kempiae was reported to be collected near Tura ( 25.522648°N, 90.185480°E) at an elevation of 210 m a. s. l. Interestingly, Naveen et al. (in press) did not include their previous sequences of “ R. kempiae ” provided in Naveen et al. (2024) without any explanation. Additionally, Naveen et al. (in press) provided three specimens ( RSNFM 13–15) under a single GenBank accession number ( PV 061647) in the phylogenetic figure (“ Figure 1 ”) and “ Table 1 ”. Furthermore, the locations of these specimens in the text and their reference in the phylogenetic tree do not correspond. However, in our phylogenetic analyses all these sequences of the claimed “ topotype ” of R. kempiae and sequences from Mizoram previously referred as “ R. cangyuanensis ” were clustered with topotype of R. garo sensu stricto.

Naveen et al. (in press) synonymised R. manipurensis based on specimens ( MZMU 2179 and MZMU 2029 ) collected from Ralruwng (Larong) ca. 50 km southeast of the actual type locality, which they claimed to be the topotype. Similarly, they synonymised “ P. namdaphaensis ” based on a specimen ( V/ APRC /A-317 ) collected from Gibbons Land, Namdapha Tiger Reserve which is located on the southern slope of the Noa-Dihing river. Another specimen ( V/ APRC /A-563 ) collected from Magoring village in the Itanagar Wildlife Sanctuary in Arunachal Pradesh on the northern slope of the Brahmaputra River referred to as “ near topotype ” of “ P. namdaphaensis ” without any justification or molecular evidence. In contrast, Naveen et al. (in press) used “ P. namdaphaensis ” sample ( ON 493540 ) available in GenBank with the voucher number MZMU-V 1277 but did not use either of the two specimens in their phylogenetic analysis.

However, in our study, sequences of “ R. manipurensis ” and “ P. namdaphaensis ” of Naveen et al. (in press) and actual topotype of these two species collected in this study were clustered with topotype of R. garo sensu stricto. Thus, we suggest “ R. manipurensis ” and “ P. namdaphaensis ” are junior synonym of R. garo and not a synonym of R. kempiae .

Naveen et al. (2024) described a new species, “ Raorchestes asakgrensis ” from Eman Asakgre Community Reserve ( 25.36788°N; 90.54344°E; 174 m) of Meghalaya State. In our phylogenetic analyses, the specimen of “ R. asakgrensis ” ( SACON VA 805 ) is nested with R. kempiae (but not the “ R. kempiae ” of Naveen et al. (2024)) (Figs 2 , 3 ). The genetic divergence of “ R. asakgrensis ” and specimens of R. kempiae collected from Tura Peak is 0.7–1.7 % in the 16 S gene (Table S 7 A – C). Thus, we consider “ R. asakgrensis ” as a junior subjective synonym of R. kempiae . We consider this taxonomic chaos created by Naveen et al. (2024) is an artefact of i) sampling gaps involving topotypic material of R. garo and R. kempiae , ii) not assigning materials according to the original morphological description of Boulenger (1919), and iii) providing single DNA sequence data each from the materials collected far away from the original type locality.

Figures

Figure 1. Map of northeast India showing distribution. A Type localities of previously described bush frog species from northeast India, Indochina and Indo-Burma; B sampling localities for bush frogs during this study between 2016 and 2024 (see Table S 1 for locality details). Political boundaries of neighbouring countries are marked with black lines and state boundaries within the political boundary of India are marked with grey lines.

Figure 2. A, B Maximum likelihood tree inferred from the concatenated data set (16 S, cyt b, COI and RHO) for Raorchestes species from northeast India, Indo-Burma and Indo-China. Outgroups are not shown in this tree. Numbers at the internal branches represent ultrafast bootstrap values (UFB). Values below 50 are not shown. Scale bar: substitutions per site.

Figure 3. A, B Bayesian tree inferred from the concatenated data set (16 S, cyt b, COI and RHO) for Raorchestes species from northeast India, Indo-Burma and Indochina. Outgroups are not shown in the tree. Numbers at the nodes represent posterior probability (PP) value. Values <0.5 are not shown. Scale bar: substitutions per site.

Figure 6. Principal Component Analysis (PCA) of 176 specimens of Raorchestes using 38 morphometric characters (A – D). A Raorchestes garo group, B Raorchestes rezakhani group, C Raorchestes kempiae group + R. khonoma sp. nov. + R. eaglenestensis sp. nov., D Raorchestes annandalii group + R. mindat + R. arunachalensis sp. nov. + R. magnus sp. nov. Sister species plotted with same symbols but with different colours. E PCA of R. garo and R. kempiae specimens using eight morphometric characters.

Figure 15. Raorchestes garo. A – D Holotype of Raorchestes garo (ZSI 19187), A dorsal view, B ventral view, C, D lateral view of head; E – N “ Philautus namdaphaensis ”, E – G holotype (ZSIA 7177); H – J paratype (ZSIA 7179); K – N paratype (ZSIA 7178), N granular belly skin. Scale bar = 10 mm.

Figure 16. Raorchestes garo. A – H WII-ADA 1495 from the type locality, Tura Peak. A Dorsal view (hourglass shaped dark patch on dorsum is shown with a black arrow), B ventral view, C lateral view of head in preserved condition, D dorsal view, E dorsolateral view, F ventral view, G groin, H thighs. I – M WII-ADA 608 from Lakhicherra, Assam. I Dorsal view in preserved condition (Inset image showing spinules on dorsum), J dorsolateral view, K ventral view, L groin, M. thigh; N – Q WII-ADA 1499 from the type locality, N dorsolateral view, O dorsal view, P groin, Q thigh. R – T WII-ADA 1142 from Terei, Mizoram. R Dorsolateral view, S groin, T thigh; U WII-ADA 3217 from Haldibari, Arunachal Pradesh; V, W WII-ADA 607 and WII-ADA 603 respectively from Lakhicherra, Assam; X. WII-ADA 1035 from Ngengpui, Mizoram. Scale bar = 10 mm.

Figure 17. Different morphs of Raorchestes garo in their natural habitat. A – F Barail hills, Assam; G Ngengpui Wildlife Sanctuary, Mizoram; H Leimatak, Manipur (topotype of R. “ manipurensis ” WII-ADA 1645).

Figure 18. Advertisement calls of R. garo (WII-ADA 1495) on left and R. kempiae (WII-ADA 1497) on right. From top to bottom, A, B oscillograms, C spectrogram, D power spectrum. The call marked with an arrow on top in A is expanded in B.

Figure 19. Distribution of Raorchestes species in northeast India. A Raorchestes garo and three others closely related new species; B Raorchestes kempiae, R. shillongensis, R. mindat and three new species. Type localities are denoted with stars.

Figure 20. Habitat at the type localities of Raorchestes species of northeast India. A Forest trail at Tura Peak, the type locality of R. garo and R. kempiae; B forest cover and adjacent lowlands of Garo Hills, from Daribokgre; C a landscape view of forest covers and D a forest trail at Namdapha TR, type localities of R. orientalis sp. nov. and R. nasuta sp. nov.; E hills at Mawsynram, near the type locality of R. mawsynramensis sp. nov.; F Nohkalikai Waterfall and forest cover at the type locality of R. boulengeri sp. nov. in Cherrapunji; G dry stream bed and vegetation in Maruwacherra, near Barail WLS, the type locality of R. barakensis sp. nov. and H forest around Jaintia Hills in Narpuh WLS, at the type locality of R. narpuhensis sp. nov.

Figure 21. Holotype of R. kempiae (ZSI 18859). A Dorsal view, B ventral view, C, D lateral view of head.

Figure 22. Raorchestes kempiae. A – H Topotype (WII-ADA 1497), A dorsal view, B ventral view, C lateral view of head in preserved condition, D lateral view, E dorsal view, F ventral view, G dorsolateral view showing groin, H posterior view of thighs. I WII-ADA 1478 from Daribokgre near Nokrek Biosphere Reserve; J dorsal view and K ventral view of WII-ADA 1480 in life from Daribokgre; L WII-ADA 933 in life from Nagaland; M WII-ADA 1952 and WII-ADA 1953 in axillary amplexus, from Nongkhyllem WLS. Scale bar = 10 mm.