An integrative approach to infer systematic relationships and define species groups in the shrub frog genus Raorchestes, with description of five new species from the Western Ghats, India

The genus Raorchestes is a large radiation of Old World tree frogs for which the Western Ghats in Peninsular India is the major center for origin and diversification. Extensive studies on this group during the past two decades have resolved long-standing taxonomic confusions and uncovered several new species, resulting in a four-fold increase in the number of known Raorchestes frogs from this region. Our ongoing research has revealed another five new species in the genus, formally described as Raorchestes drutaahu sp. nov., Raorchestes kakkayamensis sp. nov., Raorchestes keirasabinae sp. nov., Raorchestes sanjappai sp. nov., and Raorchestes vellikkannan sp. nov., all from the State of Kerala in southern Western Ghats. Based on new collections, we also provide insights on the taxonomic identity of three previously known taxa. Furthermore, since attempts for an up-to-date comprehensive study of this taxonomically challenging genus using multiple integrative taxonomic approaches have been lacking, here we review the systematic affinities of all known Raorchestes species and define 16 species groups based on evidence from multi-gene (2,327 bp) phylogenetic analyses, several morphological characters (including eye colouration and pattern), and acoustic parameters (temporal and spectral properties, as well as calling height). The results of our study present novel insights to facilitate a better working taxonomy for this rather speciose and morphologically conserved radiation of shrub frogs. This will further enable proper field identification, provide momentum for multi-disciplinary studies, as well as assist conservation of one of the most colourful and acoustically diverse frog groups of the Western Ghats biodiversity hotspot

Seven new species of Night Frogs (Anura, Nyctibatrachidae) from the Western Ghats Biodiversity Hotspot of India, with remarkably high diversity of diminutive forms

The Night Frog genus Nyctibatrachus (Family Nyctibatrachidae) represents an endemic anuran lineage of the Western Ghats Biodiversity Hotspot, India. Until now, it included 28 recognised species, of which more than half were described recently over the last five years. Our amphibian explorations have further revealed the presence of undescribed species of Nights Frogs in the southern Western Ghats. Based on integrated molecular, morphological and bioacoustic evidence, seven new species are formally described here as Nyctibatrachus athirappillyensis sp. nov., Nyctibatrachus manalari sp. nov., Nyctibatrachus pulivijayani sp. nov., Nyctibatrachus radcliffei sp. nov., Nyctibatrachus robinmoorei sp. nov., Nyctibatrachus sabarimalai sp. nov. and Nyctibatrachus webilla sp. nov., thereby bringing the total number of valid Nyctibatrachus species to 35 and increasing the former diversity estimates by a quarter. Detailed morphological descriptions, comparisons with other members of the genus, natural history notes, and genetic relationships inferred from phylogenetic analyses of a mitochondrial dataset are presented for all the new species. Additionally, characteristics of male advertisement calls are described for four new and three previously known species. Among the new species, six are currently known to be geographically restricted to low and mid elevation regions south of Palghat gap in the states of Kerala and Tamil Nadu, and one is probably endemic to high-elevation mountain streams slightly northward of the gap in Tamil Nadu. Interestingly, four new species are also among the smallest known Indian frogs. Hence, our discovery of several new species, particularly of easily overlooked miniaturized forms, reiterates that the known amphibian diversity of the Western Ghats of India still remains underestimated

The vocal repertoire of <i>Pseudophilautus kani</i>, a shrub frog (Anura: Rhacophoridae) from the Western Ghats of India

<div><p>Taxonomic descriptions of new anuran species are published with high frequency in the recent systematics literature. In contrast, there are fewer detailed quantitative descriptions of the vocalizations of these new species. Here, we describe the vocalizations of a recently described shrub frog, <i>Pseudophilautus kani</i> (Anura: Rhacophoridae; Biju and Bossuyt 2009) endemic to the Western Ghats region of India. We recorded two distinct, pulsatile call types that could be distinguished by their temporal and spectral properties as well as their mode of production. Type 1 calls (<i>creek</i>) were short ( ≈ 85 ms), consisted of about six or seven pulses (92 pulses/s) and had a spectrum with a single frequency peak at about 3.5 kHz. By comparison, Type 2 calls (<i>ta-ta-ta-ta</i>) were typically longer ( ≈ 320 ms), contained fewer pulses (4–5) produced at much lower rates (12 pulses/s) and had broadband spectra with multiple frequency peaks, two of which were consistently present at about 2.3 and 4.6 kHz. Video analyses indicated that males produced pulses in Type 1 calls using a single, continuous contraction of the trunk musculature, whereas pulses of Type 2 calls were produced with separate muscle contractions. In both call types, spectral properties were significantly negatively related with body length and body mass. The pulse rate and pulse period of Type 2 calls were also related to body size. No call properties were correlated with body condition. Patterns of individual variation in call properties were generally consistent with those described for other anurans.</p> </div

Rhacophorus malabaricus Jerdon 1870

Rhacophorus malabaricus Jerdon, 1870 Malabar Flying Frog (Daniel and Sekar 1989) (Figures 1 F–G, 2 C, 3 F–G, 6 C, 8 A–G; Tables 1–3) Original name and description. Rhacophorus malabaricus Jerdon, 1870, Notes on Indian herpetology. Proceedings of the Asiatic Society of Bengal, 1870: 84. Name-bearing type. Holotype, NHM 72.4. 17.106, by monotypy, an adult female, collected by Beddome, presented to NHM by Jerdon. Type locality. “Malabar”. Current status of specific name. Valid name, as Rhacophorus malabaricus Jerdon, 1870. Other material studied/observed. KERALA: Idukki district, Kattapana, AT personal observation (not collected) on 23 June 2011; Kollam district, Kattalappara, SDBDU 2138 (adult male), collected by RGK, SG & RS on 12 September 2011; Kozhikodu district, Kakkayam, SDBDU 431 (adult male), collected by SDB on 11 August 2003; Pathanamthitta district, Moozhiyar, SDB personal observation (not collected) on 23 July 2001; Thiruvananthapuram district, Athirimala, SDBDU 200 (adult female), SDBDU 550 (adult male), SDBDU 6311, 1615 (two adult females), SDBDU 6312–6314 (three adult males), collected by SDB on 6 July 2004; Chathankod, SDBDU 2074 (adult male), collected by SDB on 6 September 2002, SDBDU 4075 (adult male), collected by SDB on 23 November 2002, SDBDU 40117 (an adult male), collected by SDB on 7 July 2003; Palode, SDBDU 2314 (adult male), collected by SDB on 23 June 2000; Ponmudi, SDBDU 1276 (adult male), collected by SDB on 2 August 2001; Thrissur district, Athirapplly, SDBDU 4045 (adult male), collected by SDB on 8 June 2003; Peechi, SDB personal observation, (not collected) on 4 August 2001; Wayanad district, Kalpetta, SDBDU 6019 (adult male), collected by SDB on 11 August 2007; Sulthanbathery, SDBDU 201 (adult male), collected by SDB on 12 August 2007, SDBDU 202 (adult female), collected by SDB on 11 August 2007; Kurichiyarmala, SDB personal observation, (not collected) on 14 August 2007. TAMIL NADU: Coimbatore district, Valparai, SDBDU 20056 (TBGRI 2001.0010) (adult female), collected by SDB on 19 July 2001; KARNATAKA: Chikmagalur district, Bygoor Estate, SDBDU 2010.329 (adult male), collected by RS on 26 June 2010; Merthkarv Estate, BNHS 1800 (female), collected by K.R. Sethna in June 1959; Kudremukh, SDB personal observation (not collected); Bhadra Wildlife Sanctuary, SDB personal observation, (not collected); Mysore district, near Sagarmalnad forest, ZSI 17710, collected by M.O. Parthasarathy Ayyangar; Uttara Kannada district, Castle Rock, SDB personal observation, (not collected) on 4 June 2010; Anshi Wildlife Sanctuary, SDBDU 12321 (adult male), collected by SDB on 11 June 2000; Dandeli, SDBDU 2345 (adult male), collected by SDB on 20 October 2011; Anmod, RS personal observation, (not collected) on 19 July 2011; Kodagu district, Honey Valley Estate, SDBDU 2011.75 (adult male), collected by SDB on 13 June 2011; Shimoga district, Kachigebailu, SDBDU 2011.35 (adult male), collected by SDB on 10 June 2011; Jog, SDBDU 40182, (adult female), SDBDU 40183 (adult male), mating pair collected by SDB on 26 June 2003; Sagar, SDBDU 23142, (adult male), collected by SDB on 10 August 2004; Hassan district, Kempholey, SDBDU 43211 (adult male), collected by SDB on 11 June 2009. GOA: North Goa district, Volpoi, Kodal Forest, BNHS 2749 (female), BNHS 2748 (adult male), collected by AG Sekar & Vithoba; Volpoi, Kodal Village, BNHS 2612 (adult male), collected by AG Sekar; Volpoi, BNHS 2600 (adult male), collected by Abdulah & AG Sekar; Bondla Sanctuary, BNHS 2829 (adult male), collected by Sarkar & Vithoba. MAHARASHTRA: Sindhudurg district, Amboli, BNHS 3045 (adult male), collected by AG Sekar & VM Hegde, BNHS 4066 (adult male), collected AG Sekar & VM Hegde. Malabar: NHM 74.4. 29.1167 (female) collected by Beddome, now damaged (SDB, personal observation); Travancore, NHM 1903.9.26.24– 28 [ex BMNH 74.4.29.1108– 1112] (two adult males, two adult females and one juvenile), collected by Beddome. Diagnosis. Rhacophorus malabaricus can be distinguished from known congeners by the following combination of characters: (1) large male adult size (SVL 49.5–68.2 mm); (2) dark green dorsal colour with bright red webbing between fingers and toes in life (Figure 3 F–G); (3) finger and toe discs well developed (FD III 4.2 ± 0.7 mm, FW III 1.5 ± 0.4 mm, N = 13, male; TD IV 3.2 ± 0.5 mm, TW IV 1.3 ± 0.2 mm, N = 13, male); (4) dorsal skin with prominent granular projections (Figure 3 F); (5) webbing extensive, reaching the base of disc on either side of finger III; (6) 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 outer edge of heel; and (7) webbing extensive, reaching the base of disc on either side of toe IV (Figure 8 F–G). Comparison. Rhacophorus malabaricus is unique by its dorsal dark green skin coloration and extensive red webbing on both fingers and toes in life. However, this species could be confused with Rhacophorus calcadensis and R. pseudomalabaricus due to some similarity in preservation. Rhacophorus malabaricus differs from R. calcadensis by its larger adult male size, SVL 49.5–68.2 mm, N = 13 (vs. SVL 33.2 –47.0 mm, N = 20), outline of snout in dorsal view oval (vs. dorsal view pointed); differs from R. pseudomalabaricus by its larger adult male size, SVL 49.5–68.2 mm, N = 13 (vs. SVL 47.2–53.6 mm, N = 6), canthus rostralis rounded (vs. indistinct), webbing between fingers and toes bright red (vs. yellowish-orange), webbing on fingers extensive, reaching the base of disc in outer side of finger III (vs. webbing moderate, reaching the distal subarticular tubercle on inner side and just above distal subarticular tubercle on outer side of finger III), webbing on toes extensive, reaching the base of disc on either side of toe IV (vs. reaching the distal subarticular tubercle on either side of toe IV), usually green dorsum or green with minute black spots in life (vs. faint black lines radiating on dorsum and limbs, prominent in juveniles). Description of holotype (all measurements in mm): Large sized female frog (SVL 71.2); head wider than long (HL 21.6; HW 22.5; MN 19.0; MFE 3.9; MBE 8.0); outline of snout in dorsal and ventral view oval, snout (SL 9.8) longer than horizontal diameter of eye (EL 6.0); loreal region obtuse, canthus rostralis rounded; nostril oval and closer to tip of snout than eye (NS 3.1; EN 6.0); interorbital area (IUE 8.1) wider than upper eyelid (UEW 5.2); distance between anterior margins of eyes (IFE 19.5) 1.6 times the distance between posterior margins of eyes (IBE 11.9); tympanum (TYD 4.2) distinct, rounded, its distance to eye (TYE 1.2) ca. 30 % of tympanum diameter, supratympanic fold indistinct, from posterior corner of upper eyelid to near the shoulder; pineal ocellus present; vomerine ridge horizontal in position, near anterior corner of choanae; space between the two ridges is longer than the length of the ridge; tongue notched; median lingual process absent. Forearm (FAL 13.8), shorter than hand (HAL 21.6); fingers with dermal fringes, presence of dermal fringes on outer border of forelimbs; finger discs with distinct circummarginal grooves, third finger disc 3.4 times wider than finger (FD III 5.2; FW III 1.5); webbing complete, reaching the base of disc on either side of all fingers; relative length of fingers: (I <II <IV <III); subarticular tubercles prominent, oval, all present; prepollex distinct, oval. Hind limbs moderately long, shank length (SHL 43.6) longer to thigh (TL 32.7), and foot length (FOL 35.0); distance from heel to tip of toe IV (TFOL 50.2); thick dermal fringe from base of pad on toe V extends along lateral edge of tarsus to heel where it develops into broad flange with lateral projecting calcar on outer edge of heel; webbing complete, reaching the base of disc on either side of all toes; relative length of toe: I <II <III <V <IV; toe discs with distinct circummarginal grooves, fourth toe disc 2.9 times wider than toe (TD IV 3.8; TW IV 1.3); subarticular tubercles prominent, oval, all present; inner metatarsal tubercle distinct, oval; outer metatarsal tubercle, supernumerary tubercles and tarsal tubercle absent. Skin of snout and between eyes prominently granular; anterior parts of back shagreened, dorsal parts of forelimb, thigh and shank are shagreened (Figure 8 A); chest shagreened, belly granular (tree frog belly skin), thigh and anal region coarsely granular; and limbs shagreened (Figure 8 B); a conspicuous squarish dermal fold above the anal opening. Colour of holotype. In preservation. Dorsum uniform brownish-grey (Figure 8 A); ventral side greyish-brown, hand, feet and webbing dark grey (Figure 9 B). In life (SDBDU 2001.0011). Dorsum uniform green, lateral side of abdomen light yellow, limbs dorsally dark green; ventral side uniform yellow, feet and webbing dark red. Species Sex Locality Museum Number SVL HW HL SL MN EN NS EL FAL HAL FDIII FWIII TL SHL FOL TDIV TWIV Rhacophoru M “Colcat hills, NHM 1947.2.8.43) 46.5 14.6 16.2 7.3 12.5 3.5 3.0 5.2 7.9 12.0 1.9 0.9 22.2 21.3 16.4 1.8 1.0 calcadensis Tinievelly ” (ex BMNH 82.2. 10.24 (HT) M Athirimala SDBDU 569 41.3 13.4 14.1 6.3 12.6 2.8 2.8 4.6 7.4 11.2 2.2 1.2 20.1 20.7 15.1 1.9 0.6 M Kadalar SDBDU 2012.863 44.9 13.7 15.4 7.0 12.0 3.9 3.0 5.2 5.6 12.5 2.4 1.1 19.3 20.5 18.2 2.2 0.9 M Kadalar SDBDU 2012.864 42.4 12.9 14.0 6.5 12.0 3.7 2.8 4.8 7.4 11.5 2.3 0.9 19.1 20.0 16.4 1.8 0.7 M Kadalar SDBDU 2012.811 45.5 13.5 13.3 7.0 12.9 4.0 3.9 5.4 7.7 13.4 2.8 1.5 21.0 21.3 17.4 2.5 1.2 M Kadalar SDBDU 2011.304 45.8 13.6 13.0 7.1 13.0 3.9 4.0 5.4 7.8 13.3 2.9 1.4 21.2 21.4 17.5 2.4 1.2 M Kaikatti SDBDU 2010.258 33.2 9.8 12.3 6.0 9.6 3.1 3.0 3.8 5.4 9.2 1.7 0.7 16.6 16.4 12.6 1.4 0.8 M Kaikatti SDBDU 2010.261 37.0 11.0 12.1 6.2 8.8 3.4 2.6 5.0 6.2 10.0 2.2 0.9 17.2 17.3 13.1 1.6 0.6 M Kaikatti SDBDU 2010.260 36.1 9.6 12.6 6.5 9.8 3.5 2.3 3.8 5.9 9.9 1.6 0.6 16.2 16.5 13.2 1.4 0.4 M Kaikatti SDBDU 2010.259 40.0 12.5 13.8 7.0 13.9 4.0 3.1 4.9 6.6 11.6 2.1 0.6 20.0 20.0 16.1 1.7 0.3 M Kaikatti SDBDU 2010.257 33.7 9.8 12.6 5.5 9.2 2.5 2.2 3.6 6.7 9.0 1.5 0.8 15.5 16.5 13.1 1.3 0.5 M Ooth SDBDU 2042 42.9 12.5 14.4 6.8 11.6 3.1 3.2 5.3 6.9 11.6 2.1 0.8 19.1 18.6 15.6 1.5 0.6 M Ooth SDBDU 2043 36.9 11.2 12.4 5.4 10.6 2.8 3.0 4.2 6.6 10.0 2.2 1.0 18.3 17.5 14.9 1.8 0.5 M Upper SDBDU 2005 47.0 14.0 16.8 7.8 13.5 3.3 3.3 5.1 7.1 12.1 2.8 1.0 23.3 22.0 18.2 2.0 1.1 Kodayar M Upper SDBDU 2006 43.5 11.7 13.9 6.6 12.0 2.7 2.8 4.5 6.4 11.5 2.0 1.0 19.4 18.3 15.4 1.7 0.5 Kodayar M Upper SDBDU 2007 39.8 11.9 14.1 6.5 11.4 2.7 2.8 4.7 6.3 11.8 1.7 0.6 19.2 18.7 15.2 1.8 0.5 Kodayar M Valparai SDBDU 1156 39.0 11.8 12.5 5.2 10.0 2.5 2.0 3.6 5.7 10.9 1.8 0.6 18.6 19.0 14.6 1.5 0.5 M Valparai SDBDU 1157 41.1 11.0 11.8 5.8 9.4 3.5 2.5 4.5 7.5 10.7 2.1 0.6 17.9 17.7 15.9 1.4 0.5 M Valparai SDBDU 1159 41.2 12.4 14.9 7.5 12.1 3.4 3.0 4.9 7.2 11.9 2.3 1.2 19.7 20.5 16.5 2.2 1.6 M Valparai SDBDU 40245 41.9 13.0 15.0 6.7 11.9 3.0 2.9 5.7 7.1 11.7 2.3 1.2 20.0 19.9 16.5 1.9 0.9 Average 41.0 12.2 13.8 6.5 11.4 3.3 2.9 4.7 6.8 11.3 2.1 0.9 19.2 19.2 15.6 1.8 0.7 Standarddeviation 4.1 1.5 1.4 0.7 1.5 0.5 0.5 0.6 0.8 1.2 0.4 0.3 1.9 1.8 1.7 0.3 0.3 Rhacophor M “Malabar” NHM 1947.7. 2.8. 38 35.0 9.8 9.7 4.2 9.3 2.5 1.1 3.1 5.8 9.3 1.2 0.5 15.0 16.3 11.9 1.2 0.5 us lateralis (ex BMNH 1882.2.10.75) (HT) M Bygoor SDBDU 2010.331 32.0 10.3 10.4 4.2 9.5 2.7 1.0 4.5 6.1 9.9 1.8 0.6 16.8 17.5 13.7 1.3 0.6 Estate M Bygoor SDB DU 2010.330 32.4 10.4 10.8 3.9 9.3 2.6 1.0 4.1 6.2 8.9 1.5 0.7 16.2 17.7 13.8 1.5 0.6 Estate M Honey SDBDU 2011.84 31.1 9.9 10.0 3.9 9.4 2.4 1.1 3.5 5.6 9.9 1.9 0.5 15.1 16.9 13.8 1.5 0.6 Valley Estate M Kalpetta SDBDU 395 29.5 10.3 10.6 3.8 9.6 2.2 1.1 3.3 6.3 8.6 1.9 0.7 15.2 16.3 12.6 1.3 0.6 M Kalpetta SDBDU 396 29.0 9.5 11.1 3.8 9.1 2.1 1.0 3.6 5.9 8.4 1.7 0.6 14.9 15.7 13.0 1.4 0.5 …… continued on the next page Species Sex Locality Museum Number SVL HW HL SL MN EN NS EL FAL HAL FDIII FWIII TL SHL FOL TDIV TWIV M Kalpetta SDBDU 397 30.6 10.2 10.7 3.7 9.3 2.5 0.9 3.2 5.6 9.2 1.6 0.5 16.0 17.2 13.3 1.5 0.6 M Kalpetta SDBDU 398 29.7 10.2 10.4 3.6 9.1 2.2 1.0 3.3 5.9 9.8 1.6 0.6 14.9 15.9 13.5 1.7 0.6 M Kalpetta SDBDU 711 30.6 9.3 9.8 3.5 9.2 2.5 1.1 3.6 5.7 9.3 1.8 0.5 15.1 15.9 14.0 1.6 0.6 M Kalpetta SDBDU 712 31.3 9.9 10.1 3.7 9.2 2.6 1.0 3.8 6.2 9.2 1.5 0.6 15.1 16.1 13.7 1.4 0.7 M Kudremukh SDBDU011.1421 28.2 9.1 9.2 3.7 8.8 2.3 0.9 3.6 5.6 8.1 1.3 0.5 13.8 15.6 12.3 1.3 0.4 M Mananthav SDBDU 6302 28.3 9.9 10.1 3.9 9.3 2.2 1.0 3.6 4.8 8.3 1.5 0.6 14.2 15.5 12.6 1.5 0.5 ady M Mananthav SDBDU 6303 28.7 10.1 10.4 3.3 9.6 2.4 1.0 4.1 5.6 8.5 1.7 0.6 13.8 15.6 12.9 1.5 0.4 ady M Sulthanbath SDBDU 204 29.0 10.2 10.4 3.9 9.1 2.6 1.1 3.2 5.0 9.2 1.5 0.5 14.5 16.5 13.6 1.4 0.6 ery Average 30.4 9.9 10.3 3.8 9.3 2.4 1.0 3.6 5.7 9.0 1.6 0.6 15.0 16.3 13.2 1.4 0.6 Standarddeviation 1.9 0.4 0.5 0.2 0.2 0.2 0.1 0.4 0.4 0.6 0.2 0.1 0.9 0.7 0.6 0.1 0.1 F Bhadra SDBDU 36.0 10.5 11.8 5.1 10.4 3.3 1.6 4.9 6.9 10.9 2.1 0.7 18.4 19.8 15.8 1.8 0.7 WLS 2011.1420 F Kalpetta SDBDU 40114 33.5 10.7 10.8 4.8 10.1 2.9 1.3 3.9 6.5 10.5 2.1 0.8 17.1 18.6 14.0 1.6 0.6 Average 34.8 10.6 11.3 5.0 10.3 3.1 1.5 4.4 6.7 10.7 2.1 0.8 17.8 19.2 14.9 1.7 0.7 Standarddeviation 1.8 0.1 0.7 0.2 0.2 0.3 0.2 0.7 0.3 0.3 0.0 0.1 0.9 0.8 1.3 0.1 0.1 Rhacophorus malabaricus M Malabar, NHM 74.4. 29.1108 58.0 19.6 19.2 9.2 16.2 4.8 2.6 6.0 10.2 17.9 3.4 1.1 30.1 32.5 28.9 2.0 1.1 Beddome (ex BMNH 1903.9.26.24) M Malabar, NHM 74.4. 29.1109 56.4 17.6 18.0 8.2 13.9 4.3 3.5 6.0 9.0 14.3 3.6 1.5 25.8 27.1 25.3 2.8 0.8 Beddome (ex BMNH 1903.9.26.25) M Amboli BNHS 3045 60.1 19.0 21.4 10.1 16.9 5.2 2.9 6.0 10.3 19.7 4.1 1.3 29.2 30.9 27.7 3.1 1.3 M Amboli BNHS 4066 62.5 19.1 20.9 10.2 17.3 5.6 3.2 6.9 10.9 19.1 4.2 1.3 28.1 29.6 26.8 3.2 1.3 M Bygoor SDBDU 2010.329 60.5 17.1 19.9 9.9 16.5 5.2 3.4 6.2 9.9 18.1 4.5 1.3 27.1 28.9 25.8 3.3 1.5 Estate M Chathankod SDBDU 2074 68.2 21.1 22.9 10.8 18.3 6.6 3.5 8.0 11.8 20.5 6.0 2.7 33.9 35.9 30.1 3.7 1.6 M Honey SDBDU 2011.75 56.5 16.5 19.0 9.2 15.2 5.5 3.3 5.8 8.4 16.6 4.6 1.6 24.1 26.2 23.1 3.3 1.1 Valley Estate M Jog SDBDU 40183 60.9 18.5 19.8 10.3 16.7 5.4 3.0 5.9 9.8 18.1 3.6 1.2 29.8 31.7 28.4 3.0 1.5 M Kachigebailu SDBDU 2011.35 53.4 15.1 16.9 8.9 13.8 4.1 3.1 5.1 7.6 15.4 4.0 1.6 24.9 26.5 23.5 3.0 13 M Kalpetta SDBDU 66.4 18.2 21.8 10.3 17.5 6.9 3.6 7.6 11.2 18.3 4.3 1.9 29.4 31.6 28.6 3.9 1.3 2001.0011 M Kalpetta SDBDU 6019 54.5 17.0 18.1 8.9 14.1 4.3 3.2 5.9 8.5 15.5 3.5 1.5 28.6 28.9 26.9 3.5 1.3 M Volpoi, BNHS 2617 62.5 19.1 20.5 10.1 17.1 5.1 2.8 6.6 10.1 18.0 4.4 1.8 29.8 31.8 28.2 3.8 1.2 Kodal Forest …… continued on the next page Species Sex Locality Museum Number SVL HW HL SL MN EN NS EL FAL HAL FDIII FWIII TL SHL FOL TDIV TWIV M Volpoi, BNHS 2747 49.5 15.1 16.3 8.1 13.5 5.0 2.8 4.5 7.5 14.9 3.9 1.1 24.6 26.2 23.6 3.0 1.1 Kodal Forest Average 59.2 17.9 19.6 9.6 15.9 5.2 3.1 6.2 9.6 17.4 4.2 1.5 28.1 29.8 26.7 3.2 1.3 Standard deviation 5.2 1.8 1.9 0.9 1.6 0.8 0.3 0.9 1.3 1.9 0.7 0.4 2.8 2.9 2.3 0.5 0.2 F “Malabar” NHM 72.4. 17.106 71.2 22.5 21.6 9.8 19.0 6.0 3.1 6.0 13.8 21.6 5.2 1.5 32.7 43.6 35.0 3.8 1.3 (HT) F “Malabar” NHM 74.4. 29.1167 88.8 27.6 26.5 12.0 22.3 7.7 3.9 8.3 13.2 25.3 5.4 1.9 39.4 40.6 38.9 4.3 1.1 F Chathankod SDBDU 2001.008 87.7 29.0 27.5 13.0 24.1 8.1 3.9 7.5 15.4 26.9 6.7 2.6 40.7 41.5 39.5 4.2 1.1 F Jog SDBDU 40182 78.8 25.8 26.7 13.0 22.0 8.4 3.8 6.3 14.1 25.6 7.3 1.6 41.1 42.9 40.3 4.0 1.2 F Valparai SDBDU 200 90.3 31.1 32.2 13.5 27.4 9.3 4.0 8.2 17.7 28.0 7.7 2.9 49.0 51.1 48.1 5.6 1.9 (TBGRI 2001.0010) Average 82.2 27.2 26.9 12.3 23.0 7.9 3.7 7.3 14.8 25.5 6.5 2.1 40.6 43.9 40.4 4.4 1.3 Standarddeviation 7.6 3.3 3.8 1.5 3.1 1.2 0.4 1.1 1.8 2.4 1.1 0.6 5.8 4.2 4.8 0.7 0.3 Rhacophorus pseudomal abaricus M Kadalar SDBDU 53.6 17.9 18.1 8.0 15.5 3.7 2.8 5.5 8.0 15.2 3.3 2.2 24.8 25.0 21.0 2.2 1.2 2011.1010 M Poopara SDBDU 47.2 15.0 16.5 6.8 13.5 3.9 3.2 6.0 7.9 13.1 3.0 1.1 23.5 23.6 17.8 2.2 1.6 2011.1422 M Valparai SDBDU 1162 48.6 14.9 17.4 7.2 14.9 3.1 2.8 5.1 9.1 14.1 2.7 1.0 23.7 23.7 20.9 2.7 1.5 M Valparai SDBDU 1163 52.2 15.6 18.5 8.5 18.2 3.7 2.5 6.6 9.2 15.5 3.2 1.1 26.3 25.5 21.5 2.3 1.2 M Valparai SDBDU 1164 51.1 15.1 16.5 7.2 14.1 4.6 3.7 5.5 7.5 15.1 3.5 1.7 24.1 24.5 23.2 2.2 1.3 M Valparai SDBDU 1165 47.5 14.1 16.2 6.6 13.9 5.2 3.8 5.5 8.5 14.8 2.6 1.3 22.2 22.3 20.5 3.3 1.8 Average 50.0 15.4 17.2 7.4 15.0 4.0 3.1 5.7 8.4 14.6 3.1 1.4 24.1 24.1 20.8 2.5 1.4 Standarddeviation 2.6 1.3 1.0 0.7 1.7 0.7 0.5 0.5 1.7 0.9 0.4 0.5 1.4 1.1 1.8 0.4 0.2 F Andiparai BNHS 3095 (HT) 66.0 21.2 22.1 9.4 19.2 4.7 3.8 6.5 11.6 18.5 3.1 2.0 32.6 32.4 2.8 3.0 1.7 Shola F Valparai SDBDU 1160 67.6 22.1 23.5 10.3 20.4 6.1 3.5 7.9 11.6 21.4 4.0 2.5 33.2 33.4 31.8 3.9 1.9 F Valparai SDBDU 1166 71.5 24.0 25.4 10.4 24.4 5.6 4.0 6.1 12.0 20.7 4.1 1.8 36.4 36.1 32.5 4.3 2.1 Average 68.4 22.4 23.7 10.0 21.3 5.5 3.8 6.8 11.7 20.2 3.7 2.1 34.1 34.0 30.7 3.7 1.9 Standarddeviation 2.8 1.4 1.7 0.6 2.7 0.7 0.3 0.9 0.2 1.5 0.6 0.4 2.0 1.9 2.5 0.7 0.2 Secondary sexual characters. Male, nuptial pads present; female, eggs large (diameter 2.4 ± 0.3 mm, N = 30), unpigmented. Variation. Measurements of 13 male and five female specimens, including holotype, are given in Table 2. Predominantly green dorsally with limited variation in dorsal colour. SDBDU 40183 light yellowish-green with minute black spots; SDBDU 2011.75, dark green with scattered white spots; SDBDU 6019, light green dorsum, laterally with grey reticulation on a yellow background. The red webbing colour is constant. Distribution and natural history. Rhacophorus malabaricus is a widely distributed species in the Western Ghats occurring on both sides of the Palghat Gap from Tamil Nadu through Kerala, Karnataka, Maharashtra and Goa. The southernmost known range is in Palode, Thiruvananthapuram district, Kerala and the northernmost in Amboli, Sindhudurg district of Maharashtra (Figure 6 C). For complete list of specific localities refer to Table 1. According to published work this species has a wide altitudinal range (43 to 1894 m asl) and is reported from a range of habitats- from primary forests, secondary forests, plantations and urban areas. During the breeding season individuals aggregate on vegetation overhanging natural or artificial pools of water (Vasudevan & Dutta 2000; present study). Remarks. Jerdon described Rhacophorus malabaricus from “Malabar” based on a specimen presented to him by Beddome, simply providing a vague comparison with Rhacophorus bipunctatus (as R. reinwardtii), as follows “The head and body of the Malabar are indistinctly though finely tuberculated; the habit is more slender, and there is a distinct fold of skin over the eye in Reinwardtii, absent in this. The head too is perhaps a trifle longer… It has the spots on the sides of the body, so conspicuous in Khasi specimens of Reinwardtii ” (Jerdon 1870: 84). The original name-bearing type of this nominal species was never clarified, therefore the interpretation of the species that applies to this name relied entirely on the original description. It is not understood what Jerdon meant by skin folds over the eyes which are not present on R. bipunctatus (or R. reinwardtii), and spots on the flanks, which are not present on R. malabaricus. However, there has never been any doubts about the species that applies to this name. According to Frost (2013) the type was presumably originally deposited in ZSI, and recommended evaluating the type status of a specimen from Malabar in BMNH (=NHM) purchased from Jerdon. Dutta (1997) regarded this BMNH specimen as the type, but did not provide any justification for his decision, thus further investigation was necessary. Though Jerdon supplied both the ZSI and BMNH with the type specimens of Indian species which he had named, many of his type specimens are considered “lost” (Chanda et al. 2000), or likely incorrectly identified. In order to determine whether the holotype of R. malabaricus is indeed lost, it is necessary to identify where Jerdon deposited the specimen. The assumption that he may have deposited the specimen in the ZSI, can be discounted through studying the historical literature. Anderson (1871), provided a list of the amphibian accessions at the ZSI between 1865–1870, on which this species is not included.Sclater (1892 a) published a list of type specimens in the ZSI collection which also did not mention this species. Further Sclater (1892 b) produced a list of all amphibian specimens in the ZSI which explicitly mentioned only a single specimen of R. malabaricus was available in the museum at that time (ZSI 13581; from Trivandrum, presented by H. Ferguson), therefore it is safe to say that Jerdon had never deposited the type specimen in the ZSI. Upon Jerdon’s return to England, he deposited a sizable amphibian collection in the BMNH in 1872, which included the single Malabar specimen, and a number of his other type specimens (Günther 1912). This specimen was later listed in Boulenger (1882, 1890) without the designation of type specimen. However, considering Jerdon based the description of R. malabaricus on the single specimen available to him, and soon after deposited only a single specimen in BMNH (NHM 72.4.17.106), we follow Dutta (1997) to regard this specimen as the holotype of this species.Published as part of Biju, S. D., Kamei, Rachunliu G., Mahony, Stephen, Thomas, Ashish, Garg, Sonali, Sircar, Gargi & Suyesh, Robin, 2013, Taxonomic review of the tree frog genus Rhacophorus from the Western Ghats, India (Anura: Rhacophoridae), with description of ontogenetic colour changes and reproductive beha

Vocal Behavior of the Elusive Purple Frog of India (<i>Nasikabatrachus sahyadrensis</i>), a Fossorial Species Endemic to the Western Ghats

<div><p>Quantitative descriptions of animal vocalizations can inform an understanding of their evolutionary functions, the mechanisms for their production and perception, and their potential utility in taxonomy, population monitoring, and conservation. The goal of this study was to provide the first acoustical and statistical analysis of the advertisement calls of <i>Nasikabatrachus sahyadrensis</i>. Commonly known as the Indian purple frog, <i>N. sahyadrensis</i> is an endangered species endemic to the Western Ghats of India. As the only known species in its family (Nasikabatrachidae), it has ancient evolutionary ties to frogs restricted to the Seychelles archipelago (Sooglossidae). The role of vocalizations in the behavior of this unique species poses interesting questions, as the animal is fossorial and potentially earless and it breeds explosively above the soil for only about two weeks a year. In this study, we quantified 19 acoustic properties of 208 calls recorded from 10 males. Vocalizations were organized into distinct call groups typically composed of two to six short (59 ms), pulsatile calls, each consisting of about five to seven pulses produced at a rate of about 106 pulses/s. The frequency content of the call consisted of a single dominant peak between 1200–1300 Hz and there was no frequency modulation. The patterns of variation within and among individuals were typical of those seen in other frogs. Few of the properties we measured were related to temperature, body size, or condition, though there was little variation in temperature. Field observations and recordings of captive individuals indicated that males engaged in both antiphonal calling and call overlap with nearby calling neighbors. We discuss our findings in relation to previous work on vocal behavior in other fossorial frogs and in sooglossid frogs.</p></div

A unique mating strategy without physical contact during fertilization in Bombay Night Frogs (Nyctibatrachus humayuni) with the description of a new form of amplexus and female call

Anurans show the highest diversity in reproductive modes of all vertebrate taxa, with a variety of associated breeding behaviours. One striking feature of anuran reproduction is amplexus. During this process, in which the male clasps the female, both individuals’ cloacae are juxtaposed to ensure successful external fertilization. Several types of amplexus have evolved with the diversification of anurans, and secondary loss of amplexus has been reported in a few distantly related taxa. Within Nyctibatrachus, a genus endemic to the Western Ghats of India, normal axillary amplexus, a complete loss of amplexus, and intermediate forms of amplexus have all been suggested to occur, but many species remain unstudied. Here, we describe the reproductive behaviour of N. humayuni, including a new type of amplexus. The dorsal straddle, here defined as a loose form of contact in which the male sits on the dorsum of the female prior to oviposition but without clasping her, is previously unreported for anurans. When compared to known amplexus types, it most closely resembles the form of amplexus observed in Mantellinae. Furthermore, we prove that, opposed to the situation in most anurans, male semen release happens before egg deposition. We hypothesize that the male ejaculates on the female’s dorsum and that sperm subsequently runs from her back and hind legs before fertilizing the eggs. A second feature characterizing anuran breeding is the advertisement call, mostly produced solely by males. Despite recent descriptions of several new Nyctibatrachus species, few studies have explored their vocal repertoire. We describe both the male advertisement call and a female call for N. humayuni. The presence of a female call has not been reported within Nyctibatrachidae, and has been reported in less than 0.5% of anuran species. Altogether, our results highlight a striking diversity and several unique aspects of Nyctibatrachus breeding behaviour

Descriptive statistics for call groups, calls, and pulses, including means (<i>X¯</i>), standard deviations (SDs), ranges, and coefficients of variation.

†<p>For calls per call group and pulses per call, the values reported in the columns headed <i>X¯</i> and SD, respectively, are the median and interquartile range. Note, however, that coefficients of variation for these two properties are computed from the <i>X¯</i> and SD.</p

Results of Spearman rank correlations between acoustic properties and temperature, body size, and condition (<i>N</i> = 10).

†<p>Scatterplots depicting correlations with <i>P</i>-values below the conventional (uncorrected) α level of 0.05 are included in <a href="http://www.plosone.org/article/info:doi/10.1371/journal.pone.0084809#pone.0084809.s004" target="_blank">Figure S1</a>.</p