Fejervarya neilcoxi, 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 : 482-485

Fejervarya neilcoxi sp. nov.

http://zoobank.org/urn:lsid:zoobank.org:act:E3B863F6-7443-495C-A698-8543FF6C4197 Neil Cox’s Burrowing Frog

( Tables 1–7; Figs. 1–6 , 12 )

Etymology. This species is named for Dr Neil Cox, Manager of the IUCN-Conservation International Biodiversity Assessment Unit. Neil has been associated with the IUCN Red List in a variety of capacities including species assessment and management, and the new species is named particularly in appreciation of his contribution towards the Global Amphibian Assessment. The species epithet neilcoxi is treated as a noun in the genitive case.

Holotype. ZSI/ WGRC /V/A/951, an adult male, from Parambikulam (10°24’36.5” N 76°46’04.6” E, 650 m asl), Palakkad district, Kerala state, India, collected by SDB and SG on 0 8 July 2015.

Paratypes. ZSI/ WGRC /V/A/952–953, two adult males, and ZSI/ WGRC /V/A/954, an adult female collected along with the holotype; ZSI/ WGRC /V/A/955, an adult female, from the same locality as holotype, collected by SDB and team on 12 August 2012.

Genetic relationship. Phylogenetically, Fejervarya neilcoxi sp. nov. is nested in the Fejervarya rufescens group ( Figs. 1–2 ) of the Western Ghats. The average uncorrected pairwise genetic divergence with F. rufescens is 3.9% (range 3.7–4.1%, N = 14) for 16S, 11.2% (range 10.7–11.5%, N = 10) for COI, and 9.7% (range 8.7–10.8%, N = 10) for Cytb; 4.9% (range 4.9–5.0%, N = 8) for 16S, 11.0% (range 10.9–11.1%, N = 6) for COI, and 12.8% (range 12.7–12.9%, N = 6) for Cytb with F. cepfi sp. nov.; 4.7% (N = 2) for 16S, 11.1% (N = 2) for COI, and 10.8% (N = 2) for Cytb with F. kadar sp. nov.; and 3.5% (N = 8) for 16S, 8.9% (N = 6) for COI, and 10.4% (range 10.3–10.4%, N = 6) for Cytb with F. manoharani sp. nov. (Table 2).

Diagnosis. Fejervarya neilcoxi sp. nov. can be distinguished from known congeners by the following combination of morphological characters: (1) medium male adult size (SVL 32.0– 33.3 mm, N = 3); (2) stout body; (3) dorsal skin prominently granular with spinular projections; (4) snout subovoid in dorsal view and obtuse in lateral view; (5) presence of rictal gland at labial commissure of the mouth; (6) eye length nearly equal to snout length (male EL/SL ratio 97.6–97.7%, N = 3); (7) tympanum diameter less than half of eye length, (male TYD/EL ratio 38.1–39.0%, N = 3); (8) inter upper eyelid width nearly three-fourth of upper eyelid width (male IUE/UEW ratio 72.0–75.0%, N = 3) and internarial distance (male IUE/IN ratio 70.0–75.0%, 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 neilcoxi sp. nov. could be confused with the known species F. rufescens and three new species, F. cepfi sp. nov., F. kadar sp. nov. and F. manoharani sp. nov.

However, Fejervarya neilcoxi differs from F. rufescens by its dorsal skin prominently granular with spinular projections (vs. shagreened to granular without prominent warts); snout obtuse in lateral view (vs. rounded); tympanum to eye distance relatively longer or nearly equal to tympanum diameter, male TYE 1.4–1.6 mm, TYD 1.6 mm, TYE/TYD ratio 87.5–100%, N = 3 (vs. shorter, 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 relatively wider than upper eyelid width, male IUE 1.8–2.1 mm, UEW 2.5–2.9 mm, IUE/UEW ratio 72.0–75.0%, N = 3 (vs. relatively narrower, male IUE 1.4–1.8 mm, UEW 2.9–3.5 mm, IUE/UEW ratio 42.9–58.6%, N = 6); thigh shorter than shank length, male TL 14.0– 14.5 mm, SHL 15.3–15.8, TL/SHL ratio 91.5–92.2%, N = 3 (vs. equal or longer, male TL 14.7–15.8, SHL 14.5–15.8, TL/SHL ratio 100–102.1%, N = 6); thigh length shorter than foot length, male TL 14.0– 14.5 mm, FOL 16.3–16.9 mm, TL/ FOL ratio 85.4–87.1%, 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 less webbing between toes, male I2 – –2 II2 – –3– III2 1/2–3 1/2 IV3 1/2– 2V, specifically the third toe webbing below the first subarticular tubercle on the outside and fourth toe webbing below the second subarticular tubercle on either side (vs. more, male I2 – –2 II2 – – 3–III 2– 3IV 3– 2V).

Fejervarya neilcoxi differs from F. cepfi by its dorsal skin with relatively more prominent glandular warts (vs. less); horizontal diameter of eye nearly equal to snout length, EL 4.1–4.2 mm, SL 4.2–4.3 mm, EL/SL ratio 97.6–97.7%, N = 3 (vs. smaller, male EL 3.4–3.5 mm, SL 4.6–4.8 mm, EL/SL ratio 72.9–73.9%, N = 2); tympanum diameter less than half of horizontal diameter of eye, male TYD 1.6 mm, EL 4.1–4.2 mm, TYD/EL ratio 38.1–39.0%, 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); inter upper eyelid width narrower than upper eyelid width, male IUE 1.8–2.1 mm, UEW 2.5–2.9 mm, IUE/UEW ratio 72.0–75.0%, 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 1.8–2.1 mm, IN 2.5–3.0 mm, IUE/IN ratio 70.0–75.0%, N = 3 (vs. nearly equal, male IUE 2.4–2.5 mm, IN 2.5 mm, IUE/IN ratio 96–100%, N = 2); and relatively less webbing between toes, male I2 – –2 II2 – –3– III2 1/2–3 1/2 IV3 1/2– 2V (vs. more, male I1 +–2– II1 +– 3–III 2– 3IV 3–1 1/ 2V).

Fejervarya neilcoxi differs from F. kadar by its dorsal skin with prominent granular projections (vs. with scattered glandular warts); snout subovoid in dorsal view (vs. rounded); eye length nearly equal to snout length, male EL 4.1–4.2 mm, SL 4.2–4.3 mm, EL/SL ratio 97.6–97.7%, N = 3 (vs. relatively shorter, male EL 4.0– 4.2 mm, SL 4.4–4.5 mm, EL/SL ratio 88.9–93.3%, N = 3); inter upper eyelid width relatively wider than upper eyelid width, male IUE 1.8–2.1 mm, UEW 2.5–2.9 mm, IUE/UEW ratio 72.0–75.0%, N = 3 (vs. narrower, male IUE 1.7–1.8 mm, UEW 3.0– 3.3 mm, IUE/UEW ratio 53.1–60.0%, N = 3); thigh length shorter than shank length, male TL 14.0– 14.5 mm, SHL 15.3–15.8 mm, TL/SHL ratio 91.5–92.2%, N = 3 (vs. nearly equal, male TL 14.5–15.4 mm, SHL 14.6–15.5 mm, TL/SHL ratio 99.3–99.4%, N = 3); and relatively more webbing between toes, male I2 – –2 II2 – –3– III2 1/2–3 1/2 IV3 1/2– 2V (vs. less, male I2 – –2– II2 – – 3–III 2–3 2/3 IV3 2/3– 2V).

Fejervarya neilcoxi differs from F. manoharani by its prominently granular dorsal skin with spinular projections (vs. glandular with interrupted linear warts); snout subovoid in dorsal view (vs. rounded) and obtuse in lateral view (vs. vertical); relatively larger snout-vent size, male SVL 32.0– 33.3 mm, N = 3 (vs. smaller, male SVL 28.1–30.0 mm, N = 4); inter upper eyelid width relatively wider than upper eyelid width, male IUE 1.8–2.1 mm, UEW 2.5–2.9 mm, IUE/UEW ratio 72.0–75.0%, N = 3 (vs. relatively narrower, male IUE 1.7–2.0 mm, UEW 3.0 mm, IUE/UEW ratio 56.7–66.7%, N = 4); forearm length relatively longer than hand length, male FAL 6.1–6.2 mm, HAL 7.1–7.4 mm, FAL/HAL ratio 83.8–86.1%, N = 3 (vs. relatively shorter, male FAL 5.1–5.4 mm, HAL 6.4–6.9 mm, FAL/HAL ratio 76.8–79.7%, N = 4); thigh length relatively shorter than shank length, male TL 14.0– 14.5 mm, SHL 15.3–15.8 mm, TL/SHL ratio 91.5–92.2%, N = 3 (vs. relatively longer or nearly equal, male TL 12.9–13.5 mm, SHL 13.2–13.8 mm, TL/SHL ratio 96.3 – 99.2%, N = 4); and relatively less webbing between toes, male I2 – –2 II2 – –3– III2 1/2–3 1/2 IV3 1/2– 2V (vs. more, male I2–2 II2 – – 3–III 2–3 1/3 IV3 1/3– 2V, specifically the third toe webbing upto the first subarticular tubercle on the outside and fourth toe webbing extending closer to the second subarticular tubercle on either side).

Since Fejervarya neilcoxi is found in the same geographical region as Fejervarya parambikulamana ( Rao 1937) , for which the type specimen is now lost ( Dubois 1984), we also compared this new species with the original description of Rana (Tomopterna) parambikulamana Rao 1937 . Fejervarya neilcoxi differs from F. parambikulamana by its relatively smaller snout-vent size, male SVL 32.0– 33.3 mm, N = 3 (vs. larger, male “From snout to vent 39.00 mm”, N = 1); prominently granular dorsal skin with spinular projections (vs. “skin smooth above” and “no granulation on any part of the body”); head wider than long, male HW 11.9–12.1 mm, HL 11.4–11.7 mm, N = 3 (vs. head “distinctly longer than broad”, HW “11.50” mm, HL “15.00” mm, N = 1); snout nearly equal to the diameter of the eye, male SL 4.2–4.3 mm, EL 4.1–4.2 mm, N = 3 (vs. “longer than the eye”, SL “6.50” mm, EL “5.00” mm, N = 1); first finger longer than the second, FL I 3.6 mm, FL II 2.9 mm, N = 1 (vs. “first nearly equal to the 2nd”, FL I “6.00 mm”, FL II “ 5.75 mm ”, N = 1); and fourth finger length considerably shorter than the diameter of the eye, FL IV 2.6 mm, EL 4.2 mm, N = 1 (vs. “fourth digit equals the diameter of the eye”, FL IV 5.0 mm, EL 5.0 mm, N = 1). Based on the original description and the accompanying illustration ( Rao 1937, Pl. XXI, Fig. 1 ), F. parambikulamana is likely to be a member of the Fejervarya nilagirica group.

Description of holotype (measurements in mm). Adult male (SVL 33.3), rather stout; head wider than long (HW 12.1, HL 11.7); snout subovoid in dorsal view and obtuse in lateral view, its length (SL 4.3) nearly equal to horizontal diameter of eye (EL 4.2); loreal region acute with rounded canthus rostralis; interorbital space flat, narrower (IUE 2.1) than upper eyelid (UEW 2.9) and internarial distance (IN 3.0); nostril nearly as close to eye (EN 2.1) as to tip of snout (NS 2.0); tympanum (TYD 1.6) 38.1% of eye diameter (EL 4.2); tympanum-eye distance (TYE 1.4), 87.5% of the tympanum diameter (TYD 1.6); 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 hand length (HAL 7.4); relative length of fingers IV<II<I<III (FL I 3.6, FL II 2.9, FL III 4.5, FL IV 2.6); 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 14.5) shorter than shank (SHL 15.8) and foot (FOL 16.9), distance from the base of tarsus to the tip of toe IV (TFOL 22.7); 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– III2 1/2–3 1/2 IV3 1/ 2– 2V; inner toe length (ITL 2.8); subarticular tubercles prominent, all present, circular; inner metatarsal tubercle prominent, shovel-shaped (IMT 1.9); outer metatarsal tubercle small (OMT 0.7), rounded; supernumerary tubercles absent.

Skin of snout shagreened to prominently granular, upper eyelids prominently tuberculate, anterior and posterior parts of back, and upper and lower parts of flank shagreened to prominently granular with spinular projections; interrupted inverse V-shaped ridge on center of dorsum; dorsal surfaces of forelimb, thigh and shank shagreened with scattered spinular projections. Ventral surface of throat, chest, belly and limbs shagreened, anterior part of thigh sparsely granular; fejervaryan line present on both sides of the belly ( Fig. 12 ).

Colour of holotype. In life. Snout, upper eyelids and anterior and posterior parts of back greyish-brown with prominent blackish-brown blotches ( Fig. 12 A), upper and lower lip with faint alternate brown and light grey cross bands; tympanum light grey; flanks light grey with scattered dark grey mottling; forelimbs and hind limbs light brown with dark brown transverse bands; groin off-white with faint reticulations; anterior part of thigh light grey with greyish-brown reticulations; webbing light brown. Ventral surface of throat light flesh red with two lateral black calling patches; belly white; forearm and foreleg light flesh red. In preservation. Dorsum dark grey with blackish-grey blotches, forelimbs and hind limbs light greyish-brown dark greyish-brown transverse bands, posterior part of thigh greyish-brown with faint dark brown reticulations. Ventral surface of throat light grey with two lateral black calling patches on either side; belly off-white ( Fig. 12 ).

Variations. Morphometric data from three adult males and two adult females, including the holotype, is given in Table 7. Colour in preservation. ZSI/WGRC/V/A/953: dorsum light grey with dark grey blotches; ZSI/WGRC/ V/A/954 and ZSI/WGRC/V/A/955: ventral surface of throat with grey speckles.

Secondary sexual characters. Male: calling patches on either side of the throat. Female (ZSI/WGRC/V/A/ 955): pigmented eggs present (diameter 1.5 ± 0.3 mm, N = 15).

Distribution and natural history. Fejervarya neilcoxi sp. nov. is currently known only from its type locality Parambikulam, south of Palghat gap in the Western Ghats state of Kerala ( Fig. 3 ). The type series was observed near roadside vegetation with temporary pools of water. The specific site was close to a large moss-covered rock cutting surrounded with primary forest. Calling males were found close to water puddles and they stopped calling with any slight movement around them. Collections were made between 20:00–21:00 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 12. Holotype of Fejervarya neilcoxi sp. nov. (ZSI / WGRC / V / A / 951): 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.