Fejervarya cepfi, 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 : 472-475

Fejervarya cepfi sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:6A7B7C4D-31C3-4C43-B91B-EE78532847DA CEPF Burrowing Frog

( Tables 1–7; Figs. 1–6 , 9 )

Etymology. The species is named after the Critical Ecosystem Partnership Fund http://www.cepf.net (CEPF) for its effort to protect global biodiversity hotspots by providing grants in general, and specifically for a grant supporting research and conservation planning in the Western Ghats biodiversity hotspot through the Project Western Ghats Network of Protected Areas for Threatened Amphibians http://www.wnpata.org (WNPATA) to SDB (University of Delhi). The specific epithet cepfi is treated as a noun in the genitive case.

Holotype. ZSI/ WGRC /V/A/937, an adult male, from Amboli (15°57’16.61” N 73°59’54.97” E, 750 m asl), Sindhudurg district, Maharashtra state, India, collected by SDB and SG on 22 July 2013.

Paratypes. ZSI/WGRC/V/A/938, an adult male, and ZSI/WGRC/V/A/939, an adult female, collected along with the holotype.

Other referred material. SDBDU 2012.1429 , from Phansad WLS (18°27’18.5’’N 72°55’34.3’’E, 221 m asl), Raigad district , Maharashtra state, India, collected by SDB and SG on 0 3 December 2012 ; SDBDU 2007.1561 and SDBDU 2007.1569 (only tissue sample, voucher not collected), from Koyna (17°24’18.24’’N 73°44’17.93’’E, 820 m asl), Satara district , Maharashtra state, India, collected by SDB on 25 August 2007. These samples were only used for genetic identification.

Genetic relationship. Phylogenetically, Fejervarya cepfi sp. nov. is nested in the Fejervarya rufescens group ( Figs. 1–2 ) of the Western Ghats. The average uncorrected pairwise genetic divergences with F. rufescens are: 4.5% (range 3.9–5.2%, N = 28) for 16S, 11.3% (range 10.7–11.9%, N = 15) for COI, and 13.1% (range 12.7–13.9%, N = 15) for Cytb (Table 2). For comparison with F. kadar sp. nov., F. manoharani sp. nov., and F. neilcoxi sp. nov., see ‘Genetic relationship’ section of those species.

Diagnosis. Fejervarya cepfi sp. nov. can be distinguished from known congeners by the following combination of morphological characters: (1) medium male adult size (SVL 29.9–33.1 mm, N = 2); (2) stout body; (3) snout subovoid 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 72.9–73.9%, N = 2); (6) tympanum diameter nearly half of eye length, (male TYD/EL ratio 51.4–52.9%, N = 2); (7) inter upper eyelid width nearly equal to the upper eyelid width (male IUE/UEW ratio 96.0–96.2%, N = 2) and internarial distance (male IUE/IN ratio 96–100%, N = 2); (8) thigh length shorter than shank length (male TL/SHL ratio 93.9–94.3%, N = 2) and foot length (male TL/FOL ratio 89.7–90.1%, N = 2); (9) prominent shovel-shaped 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 cepfi sp. nov. could be confused with the known species F. rufescens and three new species F. kadar sp. nov., F. manoharani sp. nov., and F. neilcoxi sp. nov.

However, Fejervarya cepfi differs from F. rufescens by its snout obtuse in lateral view (vs. rounded); eye length relatively shorter compared to snout length, male EL 3.4–3.5 mm, SL 4.6–4.8, EL/SL ratio 72.9–73.9%, N = 2 (vs. relatively longer, male EL 3.9–4.9 mm, SL 4.2–5.0 mm, EL/SL ratio 88.6–98.0%, N = 6); tympanum diameter relatively larger compared to eye length, male TYD 1.8 mm, SL 4.6–4.8, TYD/EL ratio 51.4–52.9%, N = 2 (vs. relatively smaller, male TYD 1.6–2.2 mm, SL 4.2–5.0 mm, TYD/EL ratio 39.5–46.2%, N = 6); tympanum to eye distance relatively larger compared to tympanum diameter, male TYE 1.5–1.6, TYD 1.8 mm, TYE/TYD ratio 83.3–88.9%, N = 2 (vs. relatively smaller, male TYE 1.1–1.3 mm, TYD 1.6–2.2 mm, TYE/TYD ratio 50.0–76.5%, N = 6); inter upper eyelid width nearly equal to the upper eyelid width, male IUE 2.4–2.5 mm, UEW 2.5–2.6 mm, IUE/UEW ratio 96.0–96.2%, N = 2 (vs. narrower, male IUE 1.4–1.8 mm, UEW 2.9–3.5 mm, IUE/UEW ratio 42.9–58.6%, N = 6); inter upper eyelid width nearly equal to internarial distance, male IUE 2.4–2.5 mm, IN 2.5 mm, IUE/IN ratio 96–100%, N = 2 (vs. narrower, male IUE 1.4–1.8 mm, IN 2.4–3.2 mm, IUE/IN ratio 46.9–72.0%, N = 6); thigh length shorter than shank length, male TL 14.8–15.4 mm, SHL 15.7–16.4 mm, TL/SHL ratio 93.9–94.3%, N = 2 (vs. nearly equal, male TL 14.7–15.8 mm, SHL 14.5–15.8 mm, TL/SHL 100–102.1%, N = 6); thigh length shorter than foot length, male TL 14.8–15.4 mm, FOL 16.5–17.1 mm, TL/FOL ratio 89.7–90.1%, N = 2 (vs. nearly equal, male TL 14.7–15.8 mm, FOL 14.8–15.9 mm, TL/FOL 98.0–99.4%, N = 6); and relatively more webbing between toes, male I1 +–2– II1 +–3–III2– 3IV 3–1 1/ 2V (vs. less, male I2 – – 2II 2– –3–III2– 3IV 3– 2V).

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

Description of holotype (measurements in mm). Adult male (SVL 29.9), rather stout; head nearly as long as wide (HW 11.2, HL 11.1); snout subovoid in dorsal view, obtuse in lateral view, snout length (SL 4.6) longer than horizontal diameter of eye (EL 3.4); loreal region acute, rounded canthus rostralis; interorbital space flat, nearly as wide (IUE 2.4) as upper eyelid (UEW 2.5) and internarial distance (IN 2.5); nostril oval, as close to snout (NS 1.6) as to eye (EN 1.6); tympanum (TYD 1.8) 52.9% of eye diameter (EL 3.4); tympanum-eye distance (TYE 1.5), 83.3% of the tympanum diameter (TYD 1.8); 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 then the hand (HAL 7.2); relative length of fingers IV<II<I<III (FL I 3.6, FL II 2.6, FL III 4.3, FL IV 2.2); 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 14.8) shorter than shank (SHL 15.7) and foot (FOL 16.5); distance from the base of tarsus to the tip of toe IV (TFOL 23.5); toes long, relative length of toes I<II<III<V<IV; toe tips rounded, slightly enlarged without discs, toes without fringes, webbing between toes small: I1 +–2– II1 +–3–III2– 3IV 3–1 1/ 2V; inner toe length (ITL 2.9); minute spinular projections on the outer margins of toe V; subarticular tubercles prominent, all present, circular; inner metatarsal tubercle prominent (IMT 2.0), shovel-shaped; outer metatarsal tubercle, small (OMT 0.5), rounded; supernumerary tubercles absent ( Fig. 9 ).

Skin of snout shagreened with scattered granular 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 larger glandular warts; 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. 9 ).

Colour of holotype. In life. Dorsum, upper eyelids and lateral side of snout brick red with prominent greenishbrown markings ( Fig. 9 A); lateral surfaces of head light yellowish-brown; upper and lower lip with prominent greenish-brown cross bands; tympanum light yellowish-brown with a greenish-brown patch; forelimbs and hind limbs (including toes) light brick red with greenish-brown transverse bands; anterior parts of flank light greenishbrown and posterior parts light greenish-brown to yellow; groin light yellow; webbing light brown; anterior parts of thigh light yellowish with faint grey reticulation. Ventral surface of throat light flesh colour with minute dark brown speckles and 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 greyish-brown with blackish-brown patches, forelimbs and hind limbs light brown with dark greyish-brown transverse bands, fingers yellowish-white with minute dark brown speckles on the third and fourth fingers. Ventral surface of throat light grey with dark grey speckles and two lateral black calling patches; belly off-white; margins of the limbs light grey with greyish-brown mottling ( Fig. 9 ).

Variations. Morphometric data from two adult males and an adult female, including the holotype, is given in Table 7. Colour in preservation. ZSI/WGRC/V/A/938 and ZSI/WGRC/V/A/939: dorsum greyish-brown with more prominent and larger blackish-brown blotches.

Secondary sexual characters. Male: nuptial pad on finger I present; distinct calling patches on either side of the throat. Female (ZSI/WGRC/V/A/939): pigmented eggs present (diameter 1.9 ± 0.4 mm, N = 15).

Distribution and natural history. Fejervarya cepfi sp. nov. is currently known only from the northern Western Ghats state of Maharashtra ( Fig. 3 ). Apart from the type locality Amboli, we also genetically confirmed the presence of this species in Phansad WLS (Raigad district) and Koyna (Satara district). During the monsoon season (June–July), large congregations of males and females were observed on emerging laterite rock surfaces inside open grassland areas at Amboli. These sites were usually adjacent to shallow streams or temporary water collections. Calling males were extremely sensitive to nearby movements, and would immediately stop calling and try to hide upon being approached. This species was observed to be locally abundant during the breeding season but only for a very short duration lasting a couple of weeks. It was difficult to locate individuals outside the breeding season.

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 9. Holotype of Fejervarya cepfi sp. nov. (ZSI / WGRC / V / A / 937): 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.