Taxonomic review of the tree frog genus Rhacophorus from the Western Ghats, India (Anura: Rhacophoridae), with description of ontogenetic colour changes and reproductive behaviour Author Biju, S. D. Author Kamei, Rachunliu G. Author Mahony, Stephen Author Thomas, Ashish Author Garg, Sonali Author Sircar, Gargi Author Suyesh, Robin text Zootaxa 2013 3636 2 257 289 journal article 10.11646/zootaxa.3636.2.3 494fb970-c19c-4fae-9a0b-ae3f4f5aa0e3 1175-5326 218525 1BD9544C-0D09-4F36-A3EF-07F53E129D39 Rhacophorus calcadensis Ahl, 1927 Kalakkad Tree Frog (Das and Dutta 1998) ( Figures 1B , 2A , 3, 3A–B , 4A–F , 6A ; Tables 1–3 ) Original name and description. Rhacophorus calcadensis Ahl, 1927 , Zur Systematik der asiatischen Arten der Froschgattung Rhacophorus . Sitzungsberichte der Gesellschaft Naturforschender Freunde zu Berlin , 1927:46. Replacement name for Rhacophorus beddomi Boulenger, 1882 . Name-bearing type . Holotype , NHM 1947.2.8.43 (ex BMNH 82.2.10.24), by monotypy, an adult male. Type locality. “Calcad” (Kalakkad), Tirunelveli district, Tamil Nadu, India . Junior secondary homonym of Ixalus beddomi Günther, 1876 “1875”. Current status of specific name. Va li d n a m e, a s Rhacophorus calcadensis Ahl, 1927 . Synonym. Rhacophorus beddomii Boulenger, G.A. 1882 , Catalogue of the Batrachia Salientia s. Ecaudata in the Collection of the British Museum , Ed. 2:469. FIGURE 3. A–H. Rhacophorus of the Western Ghats. A–B. R. calcadensis ; C–E. R. lateralis ; F–G. R. malabaricus ; H. R. pseudomalabaricus . Other material studied/observed. KERALA: Kollam district , Pandimotta, SDBDU 2011.291 (adult male), collected by SDB on 13 September 2011 ; Thiruvananthapuram district , Athirimala, SDBDU 569 (adult male), collected by SDB on 11 June 2002 ; Palakkad district , Kaikatti, SDBDU 2010.258–2010.259 (two adult males), collected by SDB on 29 May 2010 ; Idukki district , Kadalar, SDBDU 2012.864 (adult male), collected by SDB on 12 July 2011 . TAMIL NADU: Tirunelveli district , Ooth, SDBDU 2042 and SDBDU 2043 (two adult males), collected by SDB on 20 June 2002 ; Upper Kodayar (Kalakkad Mundanthurai Tiger Reserve), SDBDU 2005–2007 (three adult males), collected by SDB on 15 November 2008 ; Coimbatore district , Andiparai Shola, Valparai, SDBDU 1156–1157 (two adult males), collected by SDB on 8 June 2010 . Diagnosis. Rhacophorus calcadensis can be distinguished from known congeners by the following combination of characters: (1) medium male adult size (SVL 33.2–47.0 mm); (2) brown dorsal colour with grey to yellowish-brown webbing between fingers and toes in life ( Figure 3A–B ); (3) finger and toe discs well developed (FD III 2.1 ± 0.4 mm , FW III 0.9 ± 0.3 mm , N = 20; TD IV 1.8 ± 0.3 mm , TW IV 0.7 ± 0.3 mm , N = 20); (4) webbing extensive, reaching the base of the disc in inner and outer side of finger III; (5) thick dermal fringe from base of pad of toe V extends along lateral edge of tarsus to heel where it develops into broad flange with lateral projecting calcar on lateral edge of heel; (6) toe webbing extensive, reaching beyond distal subarticular tubercle on either side of toe IV ( Figure 4E, F ); (7) dorsal skin with prominent granular projections ( Figure 4A ). FIGURE 4. A–F. Holotype of Rhacophorus calcadensis , NHM 1947.2.8.43 (ex BMNH 82.2.10.24), from "Calcad” (Kalakkad): A. dorsal view; B. lateral view of head; C. ventral view of hand; D. schematic illustration of webbing on hand; E. ventral view of foot; F. schematic illustration of webbing on foot. Comparison. Rhacophorus calcadensis can not be confused with other currently known members of the Western Ghats due to its dark brown or yellowish-brown dorsal skin colouration and extensive webbing on both fingers and toes. The original description by Boulenger (1882) and subsequent description by Ahl (1927) were rather brief. For a more comprehensive understanding of this species morphology we provide a redescription of the holotype . FIGURE 5. Ordination plot of Principal Component Analysis (PCA) for morphometric variables studied in four populations of Rhacophorus calcadensis . Percent variance explained by each PC axis is given in parentheses. Redescription of holotype ( all measurements in mm ): Medium sized frog (SVL 46.5); head longer than wide (HL 16.2; HW 14.6; MN 12.5; MFE 9.8; MBE 5.9); outline of snout in dorsal and ventral view pointed, snout (SL 7.3) longer than horizontal diameter of eye (EL 5.2); loreal region obtuse, canthus rostralis indistinct; nostril oval and closer to tip of snout than eye (NS 3.0; EN 3.5); interorbital area (IUE 4.0) wider than upper eyelid (UEW 3.5); distance between posterior margins of eyes (IBE 11.8) 1.4 times the distance between anterior margins of eyes (IFE 8.4); tympanum ( TYD 2.4) distinct, rounded, its distance to eye (TYE 0.9) less than half of tympanum diameter, supratympanic fold distinct, from posterior corner of upper eyelid to near the shoulder; pineal ocellus absent; vomerine ridge horizontal in position, near anterior corner of choanae; space between the two ridges is less than the length of the ridge; tongue notched; median lingual process absent. Forearm (FAL 7.9), shorter than hand (HAL 12.0); fingers with dermal fringes, presence of dermal flaps on border of forelimbs; finger discs with distinct circummarginal grooves, third finger disc 2 times wider than finger (FD III 1.9; FW III 0.9); webbing extensive, on finger I, II, III and IV to base of disc ( Figure 4C, D ); relative length of fingers: (I <II <IV <III); subarticular tubercles prominent, oval, all present; prepollex distinct, oval; one palmar tubercle, oval, indistinct; nuptial pad well developed. Hind limbs moderately long, shank length (SHL 21.3) shorter than thigh (TL 22.2), and longer than the foot length (FOL 16.4); distance from heel to tip of toe IV (TFOL 25.9); thick dermal fringe from base of pad of toe V extends along lateral edge of tarsus to heel where it develops into broad flange with lateral projecting calcar on lateral edge of heel; webbing extensive, on toe I, to base of disc; toe II, to base of disc (inner and outer); toe III, to base of disc (inner and outer); toe IV, to above distal subarticular tubercle (both outer and inner); and toe V, to base of disc; relative length of toe: I <II <III <V <IV; finger discs with distinct circummarginal grooves, fourth toe disc 1.8 times wider than toe (TD IV 1.8, TW IV 1.0); subarticular tubercles prominent, oval, all present; inner metatarsal tubercle distinct, oval; outer metatarsal tubercle, supernumerary tubercles and tarsal tubercle absent. FIGURE 6. A–D. Distribution maps of the four Rhacophorus species in the Western Ghats. A. R. calcadensis ; B. R. lateralis ; C. R. malabaricus ; and D. R. pseudomalabaricus . Positioning of symbols is approximate; see Table 1 for coordinates. Skin of snout and between eyes prominently granular with spinular tips, upper eyelids tuberculate; anterior parts of back coarsely wrinkled with keratinised spines throughout the body, posterior parts of back with minute granules and skin folds ( Figure 4A ); ventral side, throat, chest, belly prominently granular (tree frog belly skin), thigh granular, forelimbs smooth; a conspicuous squarish dermal fold above the anal opening. Colour of holotype . In preservation. Dorsally dark grey with irregular dark patches; ventral side light grey, thigh darker than belly and chest, hand, feet and webbing dark grey ( Figure 4 ). In life (SDBDU 2005). Dorsally dark grey, brownish-grey, with or without yellow tinge ( Figure 3A, B ) and scattered inconspicuous patches; lateral side light yellowish-grey; limbs dorsally light brown, forelimbs, hind limbs and fingers with faint brown cross bands; flanks light grey with dark spots; ventral side light grey, webbing dark grey. Secondary sexual characters. Male , nuptial pads present. Variation. Measurements of 20 male specimens, including holotype , are given in Table 2 . Rhacophorus calcadensis shows considerable variation in morphometric characters, especially snout to vent length (33.2–47.0 mm) in adult males. Though PCA of the ordination plot ( Figure 5 ) indicates no clear demarcation between the populations based on their morphometric characters, population from Kadalar (Idukki, Kerala) shows relatively less overlapping characters. However, the combination of low genetic distance and lack of distinct morphological differences do not warrant this population to be named as a new species. Distribution and natural history. This species is known from Athirimala, Pandimotta, Kadalar, Kaikatti, Nelliyampathy, in Kerala and Kakachi, Kalakkad, Ooth, Upper Kodayar, Andiparai Shola, Valparai in Tamil Nadu ( Figure 6A , Table 1 ). The species occurs at elevations between ca. 200 to 1400 m asl. Specimens from Upper Kodayar, Kaikatti, Nelliyampathy and Athirimala were found in evergreen forest patches at height of about six to seven meters above ground level. The population from Ooth was collected from disturbed forest fringes close to tea plantations at about three meters height. Specimens (SDBDU 2012.863, SDBDU 2012.811) from Kadalar were collected from leaves and stems of wild cardamom plants and shrubs on the roadside ca. one meter above ground level between 19:30 to 21:00 hours. During the breeding period (June–August) animals aggregated near marshy areas covered with vegetation. TABLE 1. Distribution and collection localities of Rhacophorus that are discussed in the text. Distribution is given state wise in the following sequence: district followed by specific locality. Altitude is in meters. * locality requires confirmation, see ‘Discussion’ for explanation. Distribution Coordinates Altitude Reference Latitude (ºN) Longitude (ºE) Rhacophorus calcadensis KERALA Kollam dist. Pandimotta 8.8269 77.2164 1222 Present study Palakkad dist. Kaikatti, Nelliyampathy 10.5372 76.6764 965 Present Study Idukki dist. Kadalar 10.1311 76.9986 1429 Present study Thiruvananthapuram dist Athirimala 8.6178 77.2297 993 Present study TAMIL NADU Coimbatore dist. Valparai 10.3669 76.9697 600 Present study Tirunelveli dist. Kakachi 8.5442 77.4278 1200 Vasudevan et al . 2001 “Calcad” (= Kalakkad) 8.7050 77.3136 198 Boulenger 1882; Ahl 1927 Ooth, Tirunelveli 8.5917 77.5031 1100 Present study Upper Kodayar, Tirunelveli 8.5292 77.3656 1346 Present study ......continued on the next page TABLE 1 . (Continued)

Distribution	Coordinates	Altitude	Reference
Latitude (ºN) Longitude (ºE)
Rhacophorus lateralis
KERALA
Idukki dist. Periyar Tiger Reserve Eravikulam National Park Thekkady	9.4903 77.2083 10.2000 77.0333 9.5833 77.2000	1171 1477 1162	*Daniels 2005; Ravichandran & Pillai 1990 *Dinesh et al . 2010 *Easa 2003
Wayanad dist.
Kalpetta Kurichiyarmala	11.6300 76.0825 11.7367 76.3753	796 1200	Biju 2000; Dinesh et al. 2010 Present study
Mananthavady	11.8067 76.0094	750	Present study
Sulthanbathery	11.6642 76.2653	908	Present study
KARNATAKA
Chikmagalur dist. Bhadra Wildlife Sanctuary Bygoor Estate	13.4350 75.6430 13.3114 75.6119	1409 1024	Dinesh et al. 2010 Present study
Kudremukh National Park	13.3006 75.3528	1222	Dinesh et al . 2010
Kodagu dist. Mojo Plantation	12.4666 75.7000	1140	Goel & Goel 2010
Shanthi Estate	12.4000 75.7333	932	Molur & Molur 2010
Lakunda Estate, Virajpet Virajpet Honey Valley Estate "Malabar"	12.0500 76.0400 12.1904 75.8056 12.2200 75.6556	900 911 1153	Bennet et al. 2000; Das 2000 Present Study Present Study
Rhacophorus malabaricus
KERALA
Thiruvananthapuram dist
Athirimala	8.6178 77.2297	993	Present study
Chathankod	8.6500 77.1750	180	Present study
Palode	8.7600 77.0233	355	Present study
Ponmudi	8.7608 77.1178	852	Inger et al. 1984; Dutta 1997; Chandramouli & Ganesh 2010
Trivandrum	8.4825 76.9267	43	ZSI/Calcutta specimen by HS Ferguson

Kollam dist. Kattalappara 8.9178 77.0953 130 Present Study Pandimotta 8.8264 77.2122 1221 Present Study Pathanamthitta dist. Moozhiyar 9.3161 77.0683 566 Present Study ......continued on the next page ......continued on the next page Rhacophorus pseudomalabaricus TABLE 1. (Continued)

Distribution	Coordinates	Altitude	Reference
Latitude (ºN) Longitude (ºE)
Rhacophorus malabaricus
Thrissur dist. Athirapplly Peechi	10.2953 76.5483 10.5378 76.3728	250 86	Present Study Present Study
Idukki dist.
Vagamon Kattapana	9.5789 77.0917 9.7511 76.9978	854 836	Present Study Present Study
Kozhikodu dist.
Kakkayam	11.5542 75.9194	752	Present study
Wayanad dist.
Kalpetta	11.6078 76.0867	770	Abraham et al. 2001
Sulthanbathery	11.6642 76.2653	908	Abraham et al. 2001
Kurichiyarmala	11.5936 75.9742	1287	Abraham et al. 2001
Palakkad dist.
Silent Valley NP	11.0953 76.4458	887	Andrews et al. 2005; Pillai 1986
Mukkali	11.0642 76.5372	530	Present study
TAMIL NADU
Coimbatore dist.
Valparai	10.3669 76.9697	600	Present study
KARNATAKA
Chikmagalur dist.
Bygoor Estate	13.3114 75.6119	1024	Present study
Merthkarv estate	- -	-	BNHS 1800, Collected by Sethna
Kudremukh	13.3006 75.3528	1894	Krishnamurthy 2003; Dinesh et al. 2010
Bhadra WLS	13.4494 75.6431	768	Present study
Mysore dist.
Sagarmalnad forest	12.2765 76.6738	748	ZSI 17710, collected by MO Parthasarthy A
Uttara Kannada dist.
Castle Rock	15.4011 74.3253	597	Present study
Anshi WLS	15.0078 74.3883	530	Present study
Dandeli	15.1667 74.6322	494	Present study
Anmod	15.5643 74.5317	657	Present study
Kodagu dist.
Honey Valley Estate	12.2200 75.6556	1153	Present study
Shimoga dist.
Kachigebailu	13.9833 75.1097	640	Present study
Jog	14.2247 75.4067	600	Present study

TABLE 1. (Continued)

Distribution	Coordinates	Altitude	Reference
Latitude (ºN) Longitude (ºE)
Rhacophorus malabaricus
Sagar	14.1639 75.0350	588	Present study
Hassan dist.
Kempholay	12.8567 75.6956	880	Kadadevaru & Kanamadi 2000
GOA
North Goa dist.
Volpoi, Kodal Forest	15.5281 74.1425	153	BNHS 2749 and BNHS 2748, collected by AG Sekar & Vithoba
Volpoi, Kodal Village	- -	-	BNHS 2612, collected by AG Sekar
Volpoi	15.5306 74.1373	55	BNHS 2600, collected by Abdulah & Sekar
Bondla WLS	15.4292 74.1078	195	BNHS 2829, collected by Sarkar & Vithoba
MAHARASHTRA
Sindhudurg dist.
Amboli	15.9599 73.9973	702	BNHS 3042 and BNHS 3046, collected by Sekar & Hegde

KERALA Palakkad dist.