taxonID	type	description	language	source
039F6F025479D33DA485B43BFB92F846.taxon	materials_examined	Type species: Polypedates bijui Zachariah et al., 2011	en	Abraham, Robin Kurian, Pyron, R. Alexander, Zachariah, Arun, Zachariah, Anil (2013): Two novel genera and one new species of treefrog (Anura: Rhacophoridae) highlight cryptic diversity in the Western Ghats of India. Zootaxa 3640 (2): 177-189, DOI: 10.11646/zootaxa.3640.2.3
039F6F025479D33DA485B43BFB92F846.taxon	etymology	Etymology: The generic epithet is derived from a combination of the cognomen of Colonel Richard Henry Beddome, in honor of his pioneering and extensive fieldwork, which contributed to a fundamental understanding of amphibian diversity of the Western Ghats during the colonial period, and Ixalus (Dumeril & Bibron, 1839), often used as a suffix for names of rhacophorid genera. Gender is masculine.	en	Abraham, Robin Kurian, Pyron, R. Alexander, Zachariah, Arun, Zachariah, Anil (2013): Two novel genera and one new species of treefrog (Anura: Rhacophoridae) highlight cryptic diversity in the Western Ghats of India. Zootaxa 3640 (2): 177-189, DOI: 10.11646/zootaxa.3640.2.3
039F6F025479D33DA485B43BFB92F846.taxon	materials_examined	Material examined: ZSI / WGRC / V / A / 842, 843, 844, adult males, Kadalar; ZSI / WGRC / V / A / 849, adult female, Kadalar; TNHM (H) 12.6.18 / 54, adult male, Kadalar; TNHM (H) 12.6.18 / 73, tadpole, Kadalar. Measurements of specimens provided in Table 1.	en	Abraham, Robin Kurian, Pyron, R. Alexander, Zachariah, Arun, Zachariah, Anil (2013): Two novel genera and one new species of treefrog (Anura: Rhacophoridae) highlight cryptic diversity in the Western Ghats of India. Zootaxa 3640 (2): 177-189, DOI: 10.11646/zootaxa.3640.2.3
039F6F025479D33DA485B43BFB92F846.taxon	diagnosis	Diagnosis: Beddomixalus differs from all other rhacophorid genera by a combination of the following characters: a slender, elongated medium-sized frog (average male SVL 40.1, N = 3; female SVL 60.8 mm, N = 1); dorsum yellowish-buff or reddish-brown in colour with two distinct yellowish-cream coloured parallel longitudinal stripes, extending from the supratympanic fold to the vent along the dorsolateral margin; vomerine teeth and lingual papilla absent; supratympanic fold and tympanum distinct; canthus rostralis rounded, loreal region obtusely concave (the canthus rostralis and the loreal region had been erroneously interpreted in the type description as being concave and angular, respectively); Wolffian duct simple, tubular (Fig. 6). Additionally, early development of non-pigmented eggs occur exposed on moist swamp beds, without any form of protection, neither in the form of foam nests or parental care; has free-living aquatic tadpoles adapted to lentic conditions; inhabits mid- to highelevation forests. Beddomixalus currently contains one species and this diagnosis is valid for the genus and species. Colour in life: Male: dorsum buff-brown with a dark stripe between the eyes in the inter-orbital region; a dark stripe extending medially from inter-orbital region to vent; two prominent light-coloured, yellowish-cream longitudinal stripes extending along the dorsolateral region, from the outer border of upper eyelid to the groin, on either side of the body, bordered on the lower side by a series of black dots, which appear as a line; a broad dark band extending from below the snout along the tympanum up to the middle of the flank, where it transforms into a pale patch covered in dark-brown to black vermiculations. Female: reddish- to orange-brown dorsum; similar dark and light bands and vermiculations as in the male; ventral surface white. Geographic distribution: B. bijui is found in mid- to high-elevation (1100 – 1600 m ASL) evergreen forests along the western slopes of the Eravikulam plateau (Fig. 7). We recorded the species in the Mankulam forest division (10.1344 N, 76.9947 E; 1431 m ASL) of Idukki district and the higher parts of the Malayattoor forest division (10.1869 N, 76.9928 E; 1584 m ASL) of Ernakulam district in Kerala, and also on the Valparai plateau in the Anamalai Hills of Tamil Nadu (10.2756 N, 77.0067 E; 1246 m ASL). Breeding behaviour and reproductive mode: The species is primarily arboreal and inhabits trees in and around seasonal highland swamps or marshes (Fig. 2 B). We carried out observations on breeding and development from April to July 2012. During the early part of the breeding season (late April to early June), hundreds of males aggregated around optimal breeding locations in moist swamp beds that were not yet filled with water. Males begin migrating towards breeding sites (swamp beds) (Fig. 2 B) during the onset of pre-monsoon showers in late April from surrounding forest (Fig. 2 A), where they congregate. These highland swamps are few in number and form part of the headwaters of regional streams and the substrate is mostly moist clay in April, when the pre-monsoon showers begin. Most males perch on shrubs and bushes (at up to 2 m height) along the periphery of the dry swamp and in unison make a pulsating breeding chorus (audio file of call deposited in the Dryad Repository as Supplementary file 1; http: // dx. doi. org / 10.5061 / dryad. pm 166). But, individual males that occupy territories on shrubs within the swamp bed make a different call that could not be recorded (Fig. 2 C), akin to that of some Polypedates spp. from Southern India. The air was dominated by a strong odour, which was reminiscent of the ‘ burnt rubber’ smell characteristic of aggregating males of Hylarana malabarica (Daniel, 1976). Close examination of individual males suggested this odour to be produced by glands on their dorsolateral margin. Such an ambient odour and loud pulsating call could be reason for prompting males to form mass aggregations and attracting females, but further observations are needed to validate this. Males descend from their calling perches as the night progresses and occupy positions closer to grass patches in the muddy swamp bed. In spite of several nights (n = 7) of observation, only two individual females could be located and followed to make observations of reproductive behaviour. Amplexus is pectoral (Fig. 2 D) and an amplectant pair was observed moving within the swamp. The pair initiated oviposition on wet grass within the swamp basin. Non-pigmented, semiterrestrial eggs (terminology of Altig and McDiarmid 2007) (average diameter of an egg when freshly laid = 3.2 mm) were scattered on moist soil or a grassy substrate, either as scattered loose arrays or in clumps (terminology of Altig and McDiarmid 2007; Fig. 2 E). Total clutch size was 175 eggs. Clutches of eggs deposited by other individuals were observed on wet clay. Eggs scattered on exposed rocks were seen to have desiccated the next day. Subsequent rains over the next few days raised the water level of the swamp, after which free-swimming tadpoles could be observed. Tadpole morphology: Five tadpoles at various stages were collected within or close to grassy agglomerations in a swamp pool on 22 nd June, 2012. Sample tissue from a tadpole was DNA sequenced and analyzed to determine its species status. Description is based on a single tadpole [TNHM (H) 12.6.22 / 73] in Stage 38 (Fig. 3 A-C). The small exotrophic, nektonic tadpole is oval and depressed (Fig. 2 F) and is consistently dark pigmented from the snout to the tip of the tail, including fins. The ventral and ventrolateral body sides are pale pink and less pigmented. Eyes are reddish and of moderate size, positioned dorsolaterally and directed more laterally than anteriorly; not visible in ventral view. In lateral view, the body is slightly depressed, and snout slightly rounded. The musculus interhyoideus and intestine are visible through the ventral parts of the body. Naris equidistant from both snout and eye. Spiracle sinistral, ventrolaterally positioned at midbody. Myotomes of the tail musculature of moderate development. Tail fin moderate, rounded at the end. Upper fin smaller than lower fin. Oral disc anteroventrally positioned, of triangular shape in relaxed state, of oval shape in expanded state, and slightly emarginated laterally. Marginal papillae frame the oral disc ventrally and laterally, but absent on the upper labium. Small submarginal papillae cover the entire inner rim of oral disc. Upper jaw sheath narrow and stretched into a wide U-shaped arch. Lower jaw sheath V-shaped. Labial tooth row formula (LTRF) is 5 (2 - 5) / 3 (see Fig. 3 B). Measurements (mm) of the representative tadpole were as follows: 29.1 total length from tip of snout to tip of tail, 9.9 body length as the distance from the tip of the snout to the junction of the body and tail, 20.3 tail length from the bodytail junction to the tip of the tail, 5.6 body width at the plane of the spiracle, 4.1 body height at the plane of the eyes, 2.7 tail muscle height at the base of the tail, 2.4 tail muscle width at the base of the tail, 1.2 maximum upper fin height, 1.3 maximum lower fin height, 4.9 maximum tail height including the fins, 1.1 eyenaris distance between the centres of the nares to the anterior edge of the eyes, 1.8 naris-snout distance between the centres of the narial apertures to tip of snout, 7.0 snout-spiracle distance from the tip of the snout and the terminal end of the spiracle, 1.7 internarial distance between the centres of the narial apertures, 1.5 interorbital distance between the medial borders of the eyes, and 1.5 eye diameter. Larval metamorphosis: We observed tadpoles with completely developed hindlimbs at Stage 40 (Fig. 3 D) in the first week of July. A pale dorsolateral band appears at Stage 42 (Fig. 3 E). Eyes are bright red and overall body pigmentation changes to greenish-yellow by Stage 44 (Fig. 3 F), when metamorphs begin emerging on land. At Stage 46, froglets can be found among vegetation surrounding the swamp (Fig. 3 G).	en	Abraham, Robin Kurian, Pyron, R. Alexander, Zachariah, Arun, Zachariah, Anil (2013): Two novel genera and one new species of treefrog (Anura: Rhacophoridae) highlight cryptic diversity in the Western Ghats of India. Zootaxa 3640 (2): 177-189, DOI: 10.11646/zootaxa.3640.2.3
039F6F02547CD333A485B038FE68FA8D.taxon	materials_examined	Holotype: TNHM (H) 12.6.18 / 69, an adult male (SVL 35.4 mm) (Fig. 5), collected by Ansil B. R. on 18 th May, 2012, from bushes at the periphery of a temple compound, Arippa (8.8392 N, 77.0314 E, altitude approximately 184 m ASL), near Kulathupuzha Reserve Forest, Kollam district, Kerala, India. Paratypes: TNHM (H) 12.6.18 / 70 and TNHM (H) 12.6.18 / 74, two adult males collected by Robin K. Abraham on 12 th July, 2012 from the type locality. Measurements of all specimens provided in Table 1.	en	Abraham, Robin Kurian, Pyron, R. Alexander, Zachariah, Arun, Zachariah, Anil (2013): Two novel genera and one new species of treefrog (Anura: Rhacophoridae) highlight cryptic diversity in the Western Ghats of India. Zootaxa 3640 (2): 177-189, DOI: 10.11646/zootaxa.3640.2.3
039F6F02547CD333A485B038FE68FA8D.taxon	etymology	Etymology: The generic epithet is derived from ‘ Mercury’ as a tribute to Freddie Mercury, late lead singer of the British rock band Queen, whose vibrant music inspires the authors, in combination with Rana (Linnaeus, 1758), a suffix commonly used for many frog taxa. Gender is masculine. The specific epithet, a combination of the words ‘ myristica’ for the nutmeg family, and ‘ palustris’, which is latin for swampy, emphasizes the Myristica swamp forest habitat of this frog, which is a fragile and threatened habitat type of the Western Ghats.	en	Abraham, Robin Kurian, Pyron, R. Alexander, Zachariah, Arun, Zachariah, Anil (2013): Two novel genera and one new species of treefrog (Anura: Rhacophoridae) highlight cryptic diversity in the Western Ghats of India. Zootaxa 3640 (2): 177-189, DOI: 10.11646/zootaxa.3640.2.3
039F6F02547CD333A485B038FE68FA8D.taxon	diagnosis	Diagnosis: The diagnosis is valid for the genus and species. Mercurana myristicapalustris can be distinguished from other rhacophorid genera by the following combination of characters: medium sized frog (average male SVL 36.1, N = 3; female SVL 65.1 mm, N = 1); weakly-developed vomerine teeth present; simple tubular Wolffian duct; eggs non-pigmented; early embryonic development occurs in wet mud after pre-monsoon showers; free living aquatic tadpoles; arboreal adults inhabit low-elevation swamp forests. Mercurana, which could potentially be confused with Beddomixalus, differs from the latter in the following characters: presence of vomerine teeth and lingual papilla (vs. absence); symphysial knob not sharply pointed (vs. sharply pointed); no stripe on the dorsolateral margin (vs. pale distinct stripe on the dorsolateral margin); canthus rostralis indistinct (vs. rounded); toes 3 / 4 th webbed (vs. half / moderately webbed); finger discs large (vs. moderate); subarticular tubercle on finger IV bifid (vs. rounded); oviposited eggs mixed with mud in shallow pit (vs. eggs openly scattered on ground substrates) and; inhabits low-elevation forests (vs. mid- to high- elevation forest).	en	Abraham, Robin Kurian, Pyron, R. Alexander, Zachariah, Arun, Zachariah, Anil (2013): Two novel genera and one new species of treefrog (Anura: Rhacophoridae) highlight cryptic diversity in the Western Ghats of India. Zootaxa 3640 (2): 177-189, DOI: 10.11646/zootaxa.3640.2.3
039F6F02547CD333A485B038FE68FA8D.taxon	description	Description of Holotype: A slender, medium sized frog (average male SVL 35.4 mm); dorsal skin texture shagreened; ventral surface including chest, belly and posterior surface of thighs coarsely glandular; throat slightly glandular; lateral profile of snout rounded and protruding; canthus rostralis indistinct, loreal region obtusely concave; pupil oval and horizontal; tympanum rounded and faintly distinct, faint supratympanic fold from back of eye to shoulder; weakly-developed vomerine teeth oblique and anterior to choanae; tongue lanceolate; lingual papilla absent; small vocal sac; fingers free, toes 3 / 4 th webbed (from inner aspect of disc on Toe V reaching up to the distal subarticular tubercle of Toe IV, resuming from penultimate subarticular tubercle of Toe IV to distal subarticular tubercle of Toe III); subarticular tubercles prominent, rounded, except on finger IV, which is strongly bifid; supernumerary tubercles well developed; discs well developed and distinct; Wolffian duct simple, tubular (not illustrated, but similar to that of Beddomixalus; Fig. 6). Colour in life: Male: Dorsal surface of forearm, hindlimbs and entire body rusty-brown with small black speckles scattered throughout; handful of tiny, sparsely spaced yellow blotches on dorsum; axillary region on the underside of arms white, suffused with yellow splotches; upper-arm from shoulder to elbow and hand yellowish; flank white with diffused yellow on groin; upper lip whitish; ventral surface uniformly off-white; forelimbs and hindlimbs without any crossband pattern; anterior and posterior border and ventral surface of thigh whitish; vent fold absent; iris silvery-brown with a yellowish tinge. In non-breeding condition, the dorsum of the male is pale yellowish-brown, but with the black speckles. Female: pale greenish-yellow dorsum; ventral surface white. Geographic distribution and habitat: The species Mercurana myristicapalustris was documented only in lowland swampy areas of the western foothills of the Agasthyamalai Hill Range in Kerala (see Fig. 7). These areas are dominated by Myristica swamp formations (Fig. 4 A), a unique biotope of the Western Ghats. The type locality is outside the protected area network, but at the periphery of the Kulathupuzha Reserve Forest. We have also recorded the species in lowland areas of the Shendurney Wildlife Sanctuary in Kollam district (8.8678 N, 77.0939 E; 188 m ASL) and further south in Peppara Wildlife Sanctuary, Thiruvananthapuram district (8.6697 N, 77.1197 E; 208 m ASL). Breeding behaviour: The frog Mercurana myristicapalustris is an arboreal species whose breeding behaviour is remarkably similar to that of Beddomixalus bijui, in terms of large male aggregations at breeding sites during the pre-monsoons and oviposition on the swamp floor. Vocalizing males (Fig. 4 C) (audio file of call deposited in the Dryad Repository as Supplementary file 2; http: // dx. doi. org / 10.5061 / dryad. pm 166) call from atop perches inside the swamp forest with standing water or intermittent streams, at 2 - 5 m height from the forest floor. As the night progresses, the males make their way slowly towards the forest floor, where competition for space is greater and on one occasion, we witnessed physical combat between males. We were able to make only a brief observation of amplectic behaviour in M. myristicapalustris. A male and female in amplexus were seen descending to the swamp floor, from the base of a shrub. On reaching the ground, the pair moved into the leaf litter where both individuals slowly changed colour, becoming almost inconspicuous on the forest floor. The female was seen digging into the slushy soil and ovipositing into the resulting shallow burrow in the mud, following which she mixed the semiterrestrial eggs with muddy soil. Digging and mixing was aided by employing her well-developed hindlimb webbing. Ensuing oviposition, the pair moved away, after which they could not be relocated. Photographic documentation of breeding behaviour was not possible because of failure of camera equipment at the time. However, clutches of developing eggs washed loosely in the swamp floor along a rivulet, and a single early stage tadpole (Fig. 4 D), which did not yield an accurate LTRF, were photographed after a few days of pre-monsoon showers. The tadpole was sequenced to determine species identity, and confirmed to be that of Mercurana. Museum Sex SVL HW HL IUE UEW SL EL FLL HaL TL ShL TaL FoL Vo u c h e r. N o. Beddomixalus bijui ZSI / WGRC / V / Male 40.1 14.1 12.0 4.7 3.6 5.7 5.0 9.0 11.6 20.0 20.0 11.0 17.1 A / 842 ZSI / WGRC / V / Male 42.0 14.4 12.0 4.7 3.9 5.6 5.0 8.9 12.0 20.0 21.4 11.0 17.3 A / 843 ZSI / WGRC / V / Male 40.3 14.3 11.9 4.4 3.8 5.7 5.0 8.9 12.1 20.0 20.1 11.0 17.2 A / 844 ZSI / WGRC / V / Female 60.8 24.4 21.6 7.3 5.3 8.1 6.5 13.8 18.8 32.1 32.3 17.7 28.5 A / 849 Mercurana myristicapalustris	en	Abraham, Robin Kurian, Pyron, R. Alexander, Zachariah, Arun, Zachariah, Anil (2013): Two novel genera and one new species of treefrog (Anura: Rhacophoridae) highlight cryptic diversity in the Western Ghats of India. Zootaxa 3640 (2): 177-189, DOI: 10.11646/zootaxa.3640.2.3
039F6F02547CD333A485B038FE68FA8D.taxon	materials_examined	Holotype Male 35.4 11.9 9.8 4.1 3.0 5.2 5.1 8.4 8.2 17.8 17.7 9.8 13.0 TNHM (H) 12.6.18 / 69 TNHM (H) Male 35.1 12.9 12.0 4.3 2.8 5.0 5.0 8.5 8.0 21.1 18.2 11.7 14.5 12.6.18 / 70 TNHM (H) Male 37.8 14.1 15.7 4.8 4.2 5.6 5.5 9.9 10.6 18.7 20.5 12.0 13.4 12.6.18 / 74 Mean 36.1 13.0 12.5 4.4 3.4 5.2 5.2 8.9 8.9 19.2 18.8 11.2 13.6 Beddomixalus bijui Tadpole Gosner Stage 38 TL BL TAL BW BH NSD END TMH TMW SSD IND IOD ED TNHM (H) 12.6.22 / 73 29.1 9.9 20.3 5.6 4.1 1.8 1.1 2.7 2.4 7.0 1.7 1.5 1.5 Abbreviations for adult measurements: SVL = snout-vent length; HW = head width, at angle of jaw; HL = head length, from rear of mandible to tip of snout; IUE = inter upper eyelid width, the shortest distance between the upper eyelids; UEW = maximum upper eyelid width; SL = snout length, from tip of snout to anterior orbital border of eye; EL = eye length; FLL = forelimb length, from elbow to base of outer palmar tubercle; HaL = hand length, from base of outer palmar tubercle to tip of third finger; TL = thigh length; ShL = shank length; Tal = tarsal length; FoL = foot length, from base of inner metatarsal tubercle to tip of fourth toe. Abbreviations for tadpole measurements: TL = total length; BL = body length; TAL = tail length; BW = body width; BH = body height; NSD = naris-snout distance; END = eye-naris distance; TMH = tail muscle height at the base of the tail; TMW = tail muscle width; SSD = snout-spiracle distance; IND = internarial distance; IOD = interorbital distance; ED = eye diameter.	en	Abraham, Robin Kurian, Pyron, R. Alexander, Zachariah, Arun, Zachariah, Anil (2013): Two novel genera and one new species of treefrog (Anura: Rhacophoridae) highlight cryptic diversity in the Western Ghats of India. Zootaxa 3640 (2): 177-189, DOI: 10.11646/zootaxa.3640.2.3
