Fejervarya kadar, Garg & Biju, 2017

Garg, Sonali & Biju, S. D., 2017, Description of four new species of Burrowing Frogs in the Fejervarya rufescens complex (Dicroglossidae) with notes on morphological affinities of Fejervarya species in the Western Ghats, Zootaxa 4277 (4), pp. 451-490 : 475-478

Fejervarya kadar sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:A9F6F16C-839B-4B13-95CF-A5DCA9165D68 Kadar Burrowing Frog

( Tables 1–7; Figs. 1–6 , 10 )

Etymology. The species is named after the Kadar tribe of Kerala, who live in the Vazhachal forest where the type series was collected. We enjoyed their support and hospitality during amphibian field studies in the region. The specific epithet kadar is treated as an invariable noun in apposition to the generic name.

Holotype. ZSI/ WGRC /V/A/940, an adult male, from Thavalakuzhipara , (10°17’01.7” N 76°41’06.4” E, 577 m asl), Vazhachal forest division, Thrissur district, Kerala state, India, collected by SDB and SG on 12 September 2015.

Paratypes. ZSI/WGRC/V/A/941–942, two adult males and ZSI/WGRC/V/A/943–944, two adult females, collected along with the holotype.

Genetic relationship. Phylogenetically, Fejervarya kadar sp. nov. is nested in the Fejervarya rufescens group ( Figs. 1–2 ) of the Western Ghats. The average uncorrected pairwise genetic divergences with other members of the group are: 3.2% (range 2.8–3.9%, N = 7) for 16S, 9.5% (range 9.3–9.7%, N = 5) for COI, and 8.2% (range 8.0–8.4%, N = 5) for Cytb with F. rufescens ; and 6.2% (6.2–6.3%, N = 4) for 16S, 12.1% (range 12.0–12.3%, N = 3) for COI, and 13.7% (range 13.6–13.7%, N = 3) for Cytb with F. cepfi sp. nov. (Table 2). For comparison with F. manoharani sp. nov. and F. neilcoxi sp. nov., see ‘Genetic relationship’ section of those species.

Diagnosis. Fejervarya kadar sp. nov. can be distinguished from known congeners by the following combination of morphological characters: (1) medium male adult size (SVL 31.0– 33.5 mm, N = 3); (2) stout body; (3) snout rounded in dorsal view and obtuse in lateral view; (4) presence of rictal gland at labial commissure of the mouth; (5) eye length shorter than snout length (male EL/SL ratio 88.9–93.3%, N = 3); (6) tympanum diameter less than half of eye length, (male TYD/EL ratio 37.5–40.5%, N = 3); (7) inter upper eyelid width narrower than upper eyelid width (male IUE/UEW ratio 53.1–60.0%, N = 3) and internarial distance (male IUE/IN ratio 63.0–69.2%, N = 3); (8) thigh length nearly equal to shank length (male TL/SHL ratio 99.3–99.4%, N = 3); (9) prominent shovelshaped inner metatarsal tubercle prominent and small outer metatarsal tubercle; (10) webbing between toes small.

Morphological comparison. Based on the overall morphology and comparable body size, Fejervarya kadar sp. nov. could be confused with the known species F. rufescens and three new species F. cepfi sp. nov., F. manoharani sp. nov. and F. neilcoxi sp. nov.

However, Fejervarya kadar differs from F. rufescens by its dorsal skin with scattered glandular warts (vs. shagreened to granular); snout obtuse in lateral view (vs. rounded) and rounded in dorsal view (vs. subovoid); tympanum to eye distance relatively longer or nearly equal to tympanum diameter, male TYE 1.2–1.5 mm, TYD 1.5–1.7 mm, TYE/TYD ratio 80.0–93.3%, N = 3 (vs. relatively shorter, male TYE 1.1–1.3 mm, TYD 1.6–2.2 mm, TYE/TYD ratio 50.0–76.5%, N = 6); thigh length shorter than foot length, male TL 14.5–15.4 mm, FOL 15.7–16.1 mm, TL/FOL ratio 92.4–95.7%, N = 3 (vs. nearly equal, male TL 14.7–15.8 mm, FOL 14.8–15.9 mm, TL/FOL ratio 98.0–99.4%, N = 6); and relatively more webbing between toes, male I2 – –2– II2 – – 3–III 2–3 2/3 IV3 2/3– 2V, specifically the fourth toe webbing slightly above the third subarticular tubercle on either side (vs. less, male I2 – –2 II2 – – 3–III 2– 3IV 3– 2V, specifically the fourth toe webbing extending up to the second subarticular tubercle on either side).

Fejervarya kadar differs from F. cepfi by its dorsal skin with relatively more prominent glandular warts (vs. less prominent); snout rounded in dorsal view (vs. subovoid); inter upper eyelid width narrower than upper eyelid width, male IUE 1.7–1.8 mm, UEW 3.0– 3.3 mm, IUE/UEW ratio 53.1–60.0%, N = 3 (vs. nearly equal, male IUE 2.4–2.5 mm, UEW 2.5–2.6 mm, IUE/UEW ratio 96.0–96.2%, N = 2); inter upper eyelid width narrower than internarial distance, male IUE 1.7–1.8 mm, IN 2.6–2.8 mm, IUE/IN ratio 63.0–69.2%, N = 3 (vs. nearly equal, male IUE 2.4–2.5 mm, IN 2.5 mm, IUE/IN ratio 96–100%, N = 2); tympanum diameter less than half of horizontal diameter of eye, male TYD 1.5–1.7 mm, EL 4.0– 4.2 mm, TYD/EL ratio 37.5–40.5%, N = 3 (vs. nearly half, male TYD 1.8 mm, EL 3.4–3.5 mm, TYD/EL ratio 51.4–52.9%, N = 2); horizontal diameter of eye relatively larger compared to snout length, male EL 4.0– 4.2 mm, SL 4.4–4.5 mm, EL/SL ratio 88.9–93.3%, N = 3 (vs. relatively smaller, male EL 3.4–3.5 mm, SL 4.6–4.8 mm, EL/SL ratio 72.9–73.9%, N = 2); thigh length nearly equal to shank length, male TL 14.5–15.4 mm, SHL 14.6–15.5 mm, TL/SHL ratio 99.3–99.4%, N = 3 (vs. shorter, male TL 14.8–15.4 mm, SHL 15.7–16.4 mm, TL/SHL ratio 93.9–94.3%, N = 2); and relatively less webbing between toes, male I2 – –2– II2 – – 3–III 2–3 2/3 IV3 2/3– 2V (vs. more, male I1 +–2– II1 +– 3–III 2– 3IV 3–1 1/ 2V).

For differences with Fejervarya manoharani and F. neilcoxi , see ‘morphological comparison’ of those species.

Description of holotype (measurements in mm). Adult male (SVL 33.5), rather stout; head wider than long (HW 12.7, HL 12.1); snout rounded in dorsal view, obtuse in lateral view, snout length (SL 4.5) longer than horizontal diameter of eye (EL 4.2); loreal region acute, rounded canthus rostralis; interorbital space flat, narrower than (IUE 1.8) upper eyelid width (UEW 3.3) and internarial distance (IN 2.8); nostril oval, nearly as close to the tip of snout (NS 2.0) as to the eye (EN 2.1); tympanum (TYD 1.7) 40.5% of eye diameter (EL 4.2); tympanum-eye distance (TYE 1.5) 88.2% of the tympanum diameter (TYD 1.7); supratympanic fold well developed, extends from posterior corner of eye to near the shoulder; vomerine ridge present, bearing small teeth, at an angle of 45° to the body axis, as close to choanae as to each other; tongue moderately large, emarginated, bearing no median lingual process; rictal gland present at labial commissure of the mouth. Arms short, forearm length (FAL 6.2) shorter than hand length (HAL 7.1); relative length of fingers IV<II<I<III (FL I 4.0, FL II 3.4, FL III 4.4, FL IV 2.5); finger tips rounded, slightly enlarged without discs, fingers without fringes, webbing between fingers absent; subarticular tubercles prominent, circular; one distinct palmar tubercle, oval, bifid; supernumerary tubercles absent. Hind limbs short, thigh (TL 15.4) subequal to shank (SHL 15.5) and shorter than foot (FOL 16.1), distance from the base of tarsus to the tip of toe IV (TFOL 21.7); toes long, relative length of toes I<II<V<III<IV; toe tips rounded, slightly enlarged without discs, toes without fringes, webbing between toes small: I2 – –2– II2 – – 3–III 2–3 2/3 IV3 2/3– 2V; inner toe length (ITL 2.7); subarticular tubercles prominent, all present, circular; inner metatarsal tubercle prominent, shovel-shaped (IMT 2.1); outer metatarsal small (OMT 0.7), rounded; supernumerary tubercles absent.

Skin of snout shagreened with scattered glandular projections, upper eyelids prominently tuberculate, anterior and posterior parts of back, and upper and lower parts of flank shagreened with prominent granular projections and scattered glandular warts; interrupted inverse V-shaped ridge on the center of dorsum; dorsal surfaces of forelimb, thigh and shank shagreened with prominent granular projections. Ventral surface of throat, chest, belly and limbs smooth; fejervaryan line present on both sides of the belly ( Fig. 10 ).

Colour of holotype. In life. Dorsum, upper eyelids, lateral side of snout brownish-grey with prominent dark brown mottling and blotches ( Fig. 10 A); upper and lower lip with prominent greyish-brown cross bands; tympanum light brown with a dark greyish-brown patch; anterior parts of flank light greyish-brown with dark grey reticulations, forelimbs and hind limbs (including toes) lighter brown compared to dorsum with greyish-brown transverse bands; groin off-white; webbing light grey; anterior part of thigh light yellowish with faint grey reticulation. Ventral surface of throat light flesh colour with two lateral black calling patches; belly white; forearm and foreleg light flesh red in colour with dark brown mottling on the margins. In preservation. Dorsum dark grey with blackish-brown patches, forelimbs and hind limbs lighter brown in colour than dorsum with dark grey transverse bands, posterior parts of thigh light brown with faint dark brown reticulations. Ventral surface of throat light greyish-brown with two lateral black calling patches on either side of the throat, belly off-white, and margins of the limbs light brown with dark greyish-brown mottling ( Fig. 10 ).

Variations. Morphometric data from three adult males and two adult females, including the holotype, is given in Table 7. In preservation. ZSI/WGRC/V/A/942: dorsum grey with scattered faint black spots; ZSI/WGRC/V/A/ 944: dorsum light grey with dark grey spots and more prominent glandular projections.

Secondary sexual characters. Male: nuptial pad on finger I present; distinct calling patches on either side of the throat. Female (ZSI/WGRC/V/A/944): mature ovum absent.

Distribution and natural history. Fejervarya kadar sp. nov. is currently known only from its type locality Thavalakuzhipara (Vazhachal forest), south of Palghat gap in the Western Ghats state of Kerala ( Fig. 3 ). The type series was collected from rocky surfaces of an open hill stream with small puddles or narrow water channels flowing down the slopes at the time of collection. The specific site is located inside an evergreen forest patch close to the Thavalakuzhipara tribal colony. Animals were sensitive to movement around them and tried to hide in nearby ground vegetation or grasses when approached. A few male individuals were found calling erratically but breeding activity was not observed. Collections were made in the late evening between 18:00–20:00 hours.

Figures

FIGURE 1. Maximum Likelihood phylogram based on the 16 S mitochondrial DNA dataset of 545 basepairs. Bayesian Posterior Probabilities (BPP) and RaxML bootstrap values of> 50 % are indicated above and below the branches, respectively. The uncorrected pairwise distances between species in the Fejervarya rufescens group are shown near the clade labels. Voucher number and locality details of the samples are referenced in Table 1.

FIGURE 2. Phylogenetic analyses. A. Maximum Likelihood phylogram based on mitochondrial (16 S, COI and Cytb) and nuclear (Rag 1) DNA of 2314 basepairs, showing phylogenetic relationships among members of the Fejervarya rufescens group. Bayesian Posterior Probabilities (BPP) and RaxML bootstrap values of> 50 % are indicated above and below the branches, respectively. Fejervarya granosa (2) was used as the outgroup taxon. Voucher number and locality details of the samples are referenced in Table 1; B. Median Joining haplotype network inferred from 554 basepair fragment of the nuclear Rag 1 gene. Circle sizes are proportional to the number of haplotypes, branch lengths are proportional to the number of mutational steps, and black bars represent mutational steps. Number of individuals used for each species: Fejervarya cepfi (3), Fejervarya kadar (1), Fejervarya manoharani (3), Fejervarya neilcoxi (2), and Fejervarya rufescens (5).

FIGURE 3. Geographical distribution of five species in Fejervarya rufescens group of the Western Ghats, Peninsular India. The Western Ghats biodiversity hotspot region is shown in orange colour. Locality details are referenced in Table 3.

FIGURE 4. Four morphologically recognized Fejervarya groups in the Western Ghats. A – B. Fejervarya nilagirica group: A. Fejervarya nilagirica in life (male, SDBDU 2014.2772); B. Fejervarya keralensis in life (male, SDBDU 2015.3139); C. Fejervarya rufescens group, Fejervarya rufescens in life (male, SDBDU 2015.3070); D. Fejervarya sahyadris group, Fejervarya sahyadris in life (male, SDBDU 2015.3046); E – F. Fejervarya syhadrensis group: E. Fejervarya syhadrensis in life (male, SDBDU 2015.2934); F. Fejervarya granosa in life (male, SDBDU 2014.2541). Numbers on the panels represents: 1. dorsal view, 2. ventral view of foot showing the inner metatarsal tubercle, 3. schematic illustration of foot webbing, 4. thighs showing the marking and colour pattern, 5. lateral view showing the groin markings.

FIGURE 5. Morphological comparison between members of the genus Sphaerotheca and Fejervarya rufescens group of the Western Ghats. A – H. Genus Sphaerotheca: A – E. Sphaerotheca cf. breviceps, in life (SDBDU 2014.2822): A. dorsolateral view, B. ventral view, C. ventral skin texture, D. lateral view showing the groin markings, E. thighs showing the marking and colour pattern; F – H. Inner metatarsal tubercle: F. Sphaerotheca breviceps (syntype ZMB 3351), G. S. dobsonii (Holotype BMNH 1947.2.28.45), H. S. rolandae (Holotype BMNH 1973.3024). I – R. Fejervarya rufescens group: I – K. F. rufescens in life (SDBDU 2015.3070): I. dorsolateral view, J. ventral view, K. ventral skin texture; L – M. F. kadar sp. nov. (Holotype ZSI / WGRC / V / A / 940): L. lateral view showing the groin markings, M. thighs showing the marking and colour pattern; N – R. Inner metatarsal tubercle: N. F. cepfi sp. nov. (Holotype ZSI / WGRC / V / A / 937), O. F. kadar sp. nov. (Holotype ZSI / WGRC / V / A / 940), P. F. manoharani sp. nov. (Holotype ZSI / WGRC / V / A / 945), Q. F. neilcoxi sp. nov. (Holotype ZSI / WGRC / V / A / 951), R. F. rufescens (SDBDU 2015.3070).

FIGURE 6. PCA plot for the first two principal component axes showing morphometric differentiation among adult males of five species in the Fejervarya rufescens groups of the Western Ghats. Factor loadings for each component are provided in Table 4.

FIGURE 10. Holotype of Fejervarya kadar sp. nov. (ZSI / WGRC / V / A / 940): A. dorsolateral view, in life; B. dorsal view, in preservation; C. ventral view, in preservation; D. lateral view of head, in preservation; E. ventral view of hand, in preservation; F. ventral view of foot, in preservation; G. schematic illustration of foot webbing.