Systematic status of Fejervarya (( Amphibia, Anura, Dicroglossidae) from South and SE Asia with the description of a new species from the Western Ghats of Peninsular India Author Dinesh, K. P. Author Vijayakumar, S. P. Author Channakeshavamurthy, B. H. Author Torsekar, Varun R. Author Kulkarni, Nirmal U. Author Shanker, Kartik text Zootaxa 2015 3999 1 79 94 journal article 10.11646/zootaxa.3999.1.5 3ef5a58e-b058-4c33-a79d-f768fdc73427 1175-5326 289212 4DB46BCD-D37E-4B27-8C15-6B43057E7FFD Fejervarya gomantaki sp. nov. (Table 1; Figure 1,2,3,4,5,6) Holotype : ZSI/ WGRC /V/A/865 ( CESF 2861), an adult male (SVL 19.3 mm ) collected by K.P. Dinesh, B.H. Channakeshavamurthy and Nirmal U. Kulkarni in July 2014 from Chigule village ( N 15.741 ; E 74.211 , 780m amsl), Belgaum, Karnataka. Paratypes : ZSI/ WGRC /V/A/866 ( CESF 2871), an adult male (SVL 19.4 mm ) collected by K.P. Dinesh, B.H. Channakeshavamurthy and Nirmal U. Kulkarni in July 2014 from Honda village ( N 15.547 ; E 74.046 , 50m amsl), Goa; ZSI/ WGRC /V/A/867 ( CESF 2294), an adult male (SVL 19.2 mm ) collected by Varun R. Torsekar, S.R. Chandramouli and Saunak P. Pal in August 2012 from Codal village ( N 15.594 ; E 74.207 , 80m amsl), Goa; ZSI/ WGRC /V/A/868 ( CESF 2859), an adult male (SVL 18.4 mm ) collected by K.P. Dinesh, B.H. Channakeshavamurthy and Nirmal U. Kulkarni in July 2014 from Chigule village ( N 15.741 ; E 74.211 , 780m amsl), Belgaum, Karnataka. Lineage diagnosis. Fejervarya gomantaki sp. nov. can be diagnosed phylogenetically as a member of the fejervaryan clade ( Fig 2 ), showing a sister relationship to Fejervarya sahyadris and exhibiting high genetic divergence of 4% for 16S rRNA and 1% for tyrosinase (to the GenBank accession number AB489009.1, taxa labeled as Fejervarya syhadrensis which is a population of Fejervarya sahyadris having 0.9% genetic distance for 16S sRNA of the new species). Field diagnosis. Morphology: In the field, Fejervarya gomantaki sp. nov. does not have any morphologically similar congeneric sympatric species, and it can be identified on the basis of its small size, presence of white horizontal band on the upper lip and presence of rictal glands. For comparisons of the new species Fejervarya gomantaki sp. nov. with its sister lineage Fejervarya sahyadris and the morphologically similar looking species F. chilapata , see section below. Geography: This species is restricted in distribution to low and medium elevations ( 80 – 780 m ) with a small range around the Goa region in the northern Western Ghats ( Fig 1 ). It is geographically disjunct (southern range limit 15.44 degrees North) from its sister species, Fejervarya sahyadris , whose known northern range limit is around 13.98 degrees North. Considering that there are no known biogeographic barriers in the zone of absence, additional sampling is required to establish the range limits of the sister lineages. Description of Holotype (CESF 2861) ( Fig 4 ). A small sized dicroglossid frog (SVL = 19.3 mm ) with slender body; head length sub equal to head width (HL = 7.3 mm ; HW = 7.1 mm ); snout pointed (SL = 2.9 mm ) and sub equal to eye diameter (EL = 2.3 mm ); canthus rostralis angled, loreal region concave, inter orbital space flat (IUE = 1.8 mm ) sub equal to upper lid (UEW = 1.3 mm ) and internarial distance (IN = 1.8 mm ); distance between back of eyes 1.3 times more than front of eyes (IFE = 3.5 mm ; IBE = 4.8 mm ); nostrils oval, nearer to tip of snout; symphysial knob moderate; tympanum indistinct barely visible below the supratympanic fold ( TYD = 1.1 mm ); vomerine ridges absent; tongue bifid without a papilla. Fore arm slender and short (FLL = 3.6 mm ) slightly longer than hand (HAL = 4.6 mm ); finger short and thin without any dermal fringes (TFL = 3.3 mm ), tips blunt without any enlarged discs, webbing between fingers absent; subarticular tubercles distinct, rounded and pre-pollex tubercle distinct, supernumerary tubercles absent. Hind limbs short, do not touch when folded at right angles to the body and tibio-tarsal articulation reaches back of eyes; femur length sub equal to tibia length (FL = 8.4 mm ; TiL = 8.0 mm); foot length is 2.4 times tarsus length (FOL = 9.6 mm , TAL = 3.9 mm ), relative toe length I<II<III<V<IV (FTL = 6.0 mm); webbing rudimentary ( I 1-2 II 2-2 ½ III 2 ½-3 IV 3-2 V); inner (IMTL = 0.7 mm ) and outer metatarsal tubercle, supernumerary tubercles and tarsal tubercle present. Skin on the dorsum, flanks and belly smooth; few faint longitudinal glandular folds on the dorsum; ventrally smooth on throat and belly but granular at the region of thighs. Colour on the dorsum light brown, black streak from tip of snout to groin passing canthus rostralis, supratympanic fold and flanks on either side of the body; lower lip and region of rictal gland yellowish; fore and hind limbs barred; ventrally throat and thighs translucent, belly creamish yellow with fejervaryan lines on either side of the body. Iris copper brown with fine blackish reticulations ( Fig 3 ). FIGURE 3. Fejervarya gomantaki sp. nov. in life Secondary sexual characters: Adult males have an external vocal sac at the region of throat which is mostly light black in colour, faint nuptial pad present on the first finger. Additional information from paratypes and variations: Morphological data are given in Table 1. Paratypes range from 18.4 to 19.4 mm in SVL; in all the external morphological characters, they are similar to the holotype and reference collections. All the 16S rRNA sequences generated for ZSI/WGRC/V/A/867 (CESF 2294) ( paratype ), CESF 2289, CESF 2295 and CESF 2306 (reference collections) are homogenous. Etymology. The specific epithet is derived from the term 'Gomantak', the cultural and historical name of the state of Goa, India . Suggested common name: 'Goan fejervarya'. FIGURE 4. Holotype of Fejervarya gomantaki sp. nov. (a) dorsal view. (b) ventral view FIGURE 5. Fejervarya sahyadris and Fejervarya chilapata in life. Distribution and Natural history. Fejervarya gomantaki sp. nov. is known from Virdi (Maharashtra), Chigule (Karnataka), Chorla Ghat, Bondla, Surla, Keri, Honda and Vagheri (Goa) in the Western Ghats. We found this species to be locally abundant and to occur sympatrically with Fejervarya syhadrensis . An amplected pair laid 45 eggs under captive conditions in the month of July 2014 . This species was observed calling in stagnant mud pools, artificial ponds and from grasses or herbs. Comparisons. Fejervarya gomantaki sp. nov. can be distinguished from its sympatric congeners by its small size, presence of white horizontal band on the upper lip and presence of rictal glands. However, this trait is shared with the other clade member Fejervarya sahyadris and the morphologically similar species Fejervarya chilapata . Fejervarya gomantaki sp. nov. shows a disjunct distribution with regard to its morphologically similar congeners, Fejervarya sahyadris (separated aerially by at least 170 km ) and Fejervarya chilapata (separated by at least 2000 km ). TABLE 1. Morphometric data (in mm) for the type series of Fejervarya gomantaki sp. nov. Prefix CESF for other referred specimens. ZSI/WGRC/ ZSI/WGRC/ ZSI/WGRC/ ZSI/WGRC/ 2852& 2863& 2306& Average ± SD (Range) 2295& 2869& 2860& Average ± SD (Range). No. V/A/865 V/A/866 V/A/867 V/A/868 (CESF 2861)# (CESF 2871)$ (CESF 2294)$ (CESF 2859)$ Male Male Male Male Male Male Male Female Female Female

SVL	19.3	19.4	19.2	18.4	18.5	20.0	20.2	19.3 ± 0.68 (18.4-20.2) 23.0	23.8	21.2	22.7 ± 1.33 (21.2–23.8)
HW*	7.3	7.8	7.6	7.4	6.7	6.8	7.9	7.4 ± 0.46 (6.7–7.9) 8.1	8.6	7.4	8.0 ± 0.60 (7.4–8.6)
HL*	7.1	7.4	8.0	7.2	5.2	6.4	8.1	7.1 ± 1.00 (6.4–8.1) 7.8	8.2	7.2	7.7 ± 0.50 (7.2–8.2)
IN*	1.8	2.1	2.0	1.9	1.9	2.0	2.1	2.0 ± 0.11 (1.8–2.1) 2.3	2.2	2.3	2.3 ± 0.06 (2.2–2.3)
NE*	1.4	1.4	1.4	1.6	1.1	1.5	1.5	1.4 ± 0.16 (1.1–1.6) 1.7	1.7	1.5	1.6 ± 0.12 (1.5–1.7)
MN*	5.2	5.5	5.0	5.7	4.1	5.0	6.2	5.2 ± 0.66 (4.1–6.2) 5.5	4.8	4.8	5.0 ± 0.40 (4.8–5.5)
MFE*	3.9	4.2	4.1	3.6	3.2	3.5	4.8	3.9 ± 0.53 (3.2–4.8) 4.2	3.4	3.8	3.8 ± 0.40 (3.4–4.2)
MBE*	2.1	2.4	2.3	2.3	1.9	2.0	2.6	2.2 ± 0.24 (1.9–2.6) 2.3	2.0	2.0	2.1 ± 0.17 (2.0–2.3)
SL	2.9	2.5	3.2	2.6	2.2	3.1	2.8	2.8 ± 0.35 (2.2–3.2) 2.8	2.9	2.3	2.7 ± 0.32 (2.3–2.9)
EL*	2.3	2.3	2.9	2.6	2.2	2.5	2.7	2.5 ± 0.25 (2.2–2.9) 2.8	2.4	2.3	2.5 ± 0.26 (2.3–2.8)
IUE	1.8	1.9	1.9	2.3	1.5	1.9	2.1	1.9 ± 0.25 (1.5–2.3) 2.2	2.2	2.3	2.2 ± 0.06 (2.2–2.3)
UEW	1.3	1.5	1.8	1.5	1.2	1.3	1.6	1.5 ± 0.21 (1.2–1.8) 2.1	1.4	1.5	1.7 ± 0.38 (1.2–2.1)
IFE* IBE*	3.5 4.8	3.8 5.4	4.0 5.5	3.8 5.2	3.3 4.9	4.4 6.3	4.1 5.8	3.8 ± 0.37 (3.3–4.4) 4.3 5.4 ± 0.52 (4.8–6.3) 6.5	4.0 5.9	3.7 5.4	4.0 ± 0.30 (3.7–4.3) 5.9 ± 0.55 (5.4–6.5)
TYD*	1.1	1.2	1.0	1.2	0.8	1.5	1.3	1.2 ± 0.22 (0.8–1.5) 1.4	1.3	1.0	1.2 ± 0.21 (1.0–1.4)
TE*	0.5	0.7	1.0	0.9	0.7	0.7	1.1	0.8 ± 0.21 (0.5–1.1) 0.8	1.0	1.0	0.9 ± 0.12 (0.8–1.0)
FLL*	3.7	4.8	3.3	4.7	3.4	3.5	3.3	3.8 ± 0.65 (3.3–4.8) 3.3	3.5	3.6	3.5 ± 0.15 (3.3–3.6)
HAL*	4.6	4.3	3.1	4.1	3.9	4.7	4.8	4.2 ± 0.59 (3.1–4.8) 4.9	5.4	5.0	5.1 ± 0.26 (4.9–5.1)
FL1	2.1	2.4	2.3	2.3	1.4	1.9	2.4	2.1 ± 0.36 (1.4–2.4) 2.4	2.5	2.5	2.5 ± 0.06 (2.4–2.5)
FL2	2.5	2.1	2.1	2.0	1.8	1.9	2.1	2.1 ± 0.22 (1.8–2.5) 2.4	2.4	2.3	2.4 ± 0.06 (2.3–2.4)
TFL*	3.3	3.3	3.0	2.7	2.7	2.8	3.0	3.0 ± 0.26 (2.7–3.3) 3.6	3.1	3.1	3.3 ± 0.29 (3.1–3.6)
AGL	7.8	8.2	7.0	7.0	7.1	7.8	7.5	7.5 ± 0.47 (7.0–8.2) 9.5	10.7	10.8	10.3 ± 0.72 (9.5–10.8)
WBS	6.6	7.1	7.5	6.2	6.7	7.7	8.3	7.2 ± 0.73 (6.2–8.3) 8.5	8.1	7.0	7.9 ± 0.78 (7.0–8.5)
WFG	4.8	4.1	6.5	4.6	4.4	5.7	5.4	5.1 ± 0.84 (4.1–5.7) 7.0	6.8	4.9	6.2 ± 1.16 (4.9–7.0)
ShL/ FL	8.4	9.0	8.8	8.0	7.7	9.1	9.1	8.6 ± 0.56 (7.7–9.1) 9.3	10.1	8.8	9.4 ± 0.66 (8.8–10.1)
TiL* Tal	8.0 3.9	9.2 5.0	8.3 4.1	7.8 4.2	7.6 4.2	9.1 4.5	9.8 4.0	8.5 ± 0.83 (7.6–9.8) 8.6 4.3 ± 0.37 (4.0–5.0) 4.7	9.6 4.7	8.6 3.8	8.9 ± 0.58 (8.6–9.6) 4.4 ± 0.52 (3.8–4.7)
FOL* FTL*	9.6 6.0	9.9 6.0	9.8 5.7	9.4 6.2	8.5 5.1	7.8 4.3	9.5 5.7	9.2 ± 0.77 (8.5–9.9) 11.4 5.6 ± 0.66 (4.3–6.2) 8.1	10.9 8.5	9.7 5.9	10.7 ± 0.87 (9.7–11.4) 7.5 ± 1.40 (5.9–8.5)
ITL*	1.6	1.6	1.7	1.5	1.5	1.7	1.7	1.6 ± 0.09 (1.5–1.7) 1.9	1.9	1.5	1.8 ± 0.23 (1.5–1.9)
IMT*	0.7	0.7	0.7	0.8	0.7	0.7	0.6	0.7 ± 0.06 (0.6–0.8) 0.8	0.8	0.8	0.8

* data used for PCA analysis; # holotype ; $ paratype ; & other referred specimens A multivariate Principal Component Analysis (PCA) was carried out for the morphologically similar and potential phylogenetic sister species Fejervarya sahyadris , F. chilapata and Fejervarya gomantaki sp. nov. ( Table 2 ). All the morphological variables were size corrected by SVL. The PCA results suggested three different clusters on the scatter plot representing Fejervarya sahyadris , F. chilapata and Fejervarya gomantaki sp. nov. (for both male and female morphometric analyses) ( Fig 6 ). For males, PC1 accounted for 32.4% variance (Eigen value of 6.4) and PC2 for 20.0% variance (Eigen value of 3.9), while for females PC1 accounted for 44.2% variance (Eigen value of 8.8) and PC2 for 25.9% variance (Eigen value of 5.1) ( Table 2 ). Although all three species cluster distinctly on the PCA scatter plot, the utility of these characters in the field identity are not realistic due to small adult size and range overlap in the morphological measurements and character crypticity. FIGURE 6. Multivariate Principal Component Analysis scatter plot (20 morphometric characters marked as * in Table 1 transformed to their ratio to SVL) for Fejervarya gomantaki sp. nov. , F. sahyadris and F. chilapata (a) for male individuals; (b) for female individuals. TABLE 2. Factor loadings of Principal Component Analysis (separately for males and females for the total of 20 morphometric characters marked as* in Table 1 transformed to their ratio to SVL).

Males	PC 1	PC 2	PC 3	PC 4	PC 5
HW	-0.29477	0.10633	0.18141	-0.25137	0.31006
HL	0.11123	0.33153	0.21077	-0.35456	-0.01153
MN	0.3381	0.21151	-0.02436	-0.06811	-0.09219
MFE	0.3159	0.24207	-0.06717	-0.04379	-0.08348
MBE	0.34021	0.21486	-0.04082	-0.00699	-0.02695
IFE	-0.19718	0.28918	0.23816	0.22868	0.20018
IBE	-0.19002	0.34459	0.13339	0.3231	0.095683
FLL	0.10961	0.14147	0.042123	0.58979	0.17017
HAL	0.13525	-0.22292	-0.17014	0.26061	0.52486
TFL	-0.12362	-0.29478	0.34586	-0.04655	0.21395
TiL	0.17888	-0.02917	0.37737	0.24136	-0.20624
FOL	0.2891	-0.09859	0.18839	-0.17694	0.32247
FTL	0.27996	-0.0929	0.12498	-0.2311	0.36559
IN	-0.18554	0.19924	0.25051	-0.05305	0.13151
NE	0.23916	0.08578	0.21674	-0.04372	-0.03592
EL	0.11002	0.27912	0.18809	-0.06222	0.032883
TYD	-0.17996	0.35689	-0.05376	-5.43E-05	-0.019
TE	-0.021	-0.20156	0.50256	0.003559	-0.10684
IMT	0.052829	0.22206	-0.28864	-0.09151	0.41999
ITL	0.34207	-0.09405	0.11679	0.26444	-0.00969
Explained Variance (%)	32.3	19.9	12.1	8.3	7.3
Eigenvalue	6.4	3.9	2.4	1.6	1.4
Females	PC 1	PC 2	PC 3	PC 4	PC 5
HW	-0.3032	0.029256	-0.13911	0.033742	-0.0104
HL	0.26111	-0.01929	0.11543	0.033648	-0.11311
MN	0.31375	0.13096	0.1016	-0.03255	0.006028
MFE	0.29811	0.18523	0.074262	-0.03014	0.10674
MBE	0.29922	0.18105	-0.0348	-0.02365	0.14113
IFE	-0.14712	0.30049	0.28945	0.091344	-0.18726
IBE	-0.1607	0.33215	0.054271	0.29442	0.17314
FLL	0.29237	0.13011	0.10615	-0.27259	0.023834
HAL	0.094086	-0.39535	0.1504	-0.09061	-0.0322
TFL	-0.13699	-0.10847	0.36724	0.46279	0.35834
TiL	0.29321	-0.10886	0.004552	-0.13279	-0.08912
FOL	0.26503	-0.16742	0.069156	0.32146	0.063505
FTL	0.076548	-0.20162	-0.24905	0.56342	-0.27879
IN	-0.15778	0.04958	0.57767	-0.14308	0.2166
NE	0.219	0.1123	0.27431	0.29977	-0.23542
EL	0.25031	0.24262	0.010387	0.13255	0.27281
TYD	-0.0428	0.37405	-0.325	0.14649	0.074424
TE	-0.00063	-0.41878	0.18837	0.049569	-0.02178
IMT	0.042731	-0.23892	-0.25711	-0.01583	0.6966
ITL	0.3182	-0.04561	-0.10818	0.11402	0.054537
Explained Variance (%)	44.2	25.8	9.9	8.2	5.5
Eigenvalue	8.8	5.1	1.9	1.6	1.1

Although none of these species are sympatric in distribution, for the sake of convention, detailed comparisons are presented for Fejervarya gomantaki sp. nov. with the morphologically similar species Fejervarya sahyadris (genetic sister) and Fejervarya chilapata in the following paragraphs. Comparisons are made only with male specimens unless specifically mentioned; PCA scatter plots for both male and female specimens of the three species of Fejervarya are presented in the Figure 6 . Fejervarya gomantaki sp. nov. can be distinguished from Fejervarya sahyadris in having a relatively larger adult male size of SVL 19.3 mm ±0.68, n=7 (vs relatively smaller adult male size SVL 18.4 mm ±0.67, n= 6 in F. sahyadris ); lower MN/SVL ratio of 0.272 (0.222–0.307, n=7) (vs higher MN/SVL ratio of 0.366 (0.331–0.428, n=6) in F. sahyadris ); lower MFE/SVL ratio of 0.202 (0.173–0.238, n=7) (vs higher MFE/SVL ratio of 0.298 (0.266–0.359, n=6) in F. sahyadris ); lower MBE/SVL ratio of 0.116 (0.100–0.129, n=7) (vs higher MBE/SVL ratio of 0.172 (0.147–0.206, n=6) in F. sahyadris ); finger I is shortest (vs finger IV is shortest in F. sahyadris ); vomerine ridge and palmar tubercle indistinct (vs distinct vomerine ridge and palmar tubercle in F. sahyadris ); brown dorsum, lateral sides of the body light creamish, black streak from the tip of snout passing through canthus rostralis and tympanic region tapering to the region of groin, hind limbs with dark cross bars (vs dorsum brick red to dark brown, lateral sides dark greyish, dark brown band on the canthus rostralis and at the region of tympanum below the supra tympanic fold, limbs with faint cross bars in F. sahyadris ); known distribution from Chigule village of Belgaum district in the North ( N 15.74 ) to Bondla Wildlife Sanctuary in the South ( N 15.44 ) (vs known distribution from Sagara of Shimoga district in the North ( N 13.98 ) to Mukkam of Calicut district in the South ( N 11.33 ) for F. sahyadris ). Fejervarya gomantaki sp. nov. can be distinguished from Fejervarya chilapata in having relatively small adult female size of SVL 22.7 mm ±1.3, n=3 (vs relatively large adult female size SVL 24.4 mm ±0.92, n= 2 in F. chilapata ); higher HW/SVL ratio of 0.382 (0.340–0.402, n=7) (vs lower HW/SVL ratio of 0.304 (0.291–0.312, n=6) in F. chilapata ); lower MN/SVL ratio of 0.272 (0.260–0.307, n=7) (vs higher MN/SVL ratio of 0.365 (0.357– 0.375, n=6) in F. chilapata ); lower MFE/SVL ratio of 0.202 (0.173–0.238, n=7) (vs higher MFE/SVL ratio of 0.277 (0.280–0.281, n=6) in F. chilapata ); lower MBE/SVL ratio of 0.116 (0.100–0.129, n=7) (vs higher MBE/ SVL ratio of 0.166 (0.160–0.174, n=6) in F. chilapata ); lower FoL/SVL ratio of 0.478 (0.390–0.511, n=7) (vs higher FoL/SVL ratio of 0.539 (0.517–0.570, n=6) in F. chilapata ); lower ITL/SVL ratio of 0.084 (0.081–0.089, n=7) (vs higher ITL/SVL ratio of 0.113 (0.105–0.121, n=6) in F. chilapata ); webbing rudimentary ( I 1-2 II 2-2 ½ III 2 ½-3 IV 3-2 V) (vs webbing small ( I 1-2 II 1-2 ½ III 1 ½-3 IV 3-1 V) in F. chilapata ); dorsum brown, lateral sides of the body light creamish, skin on the thigh smooth (vs dorsum dark brown, lateral sides of the body mottled with yellow, cream and black, skin on the thigh shagreened in F. chilapata ( Fig 5 )); allopatric in distribution with known distribution from Chigule village of Belgaum district in the North ( N 15.74 ) to Bondla Wildlife Sanctuary in the South ( N 15.44 ) (vs known distribution from Chilapata Reserve Forest ( N 26.58 ) in West Bengal for F. chilapata ).