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Two new endemic genera and a new species of toad (Anura: Bufonidae) from the Western Ghats of India

Two new endemic genera and a new species of toad (Anura: Bufonidae) from the Western Ghats of India Background: Bufonidae are a large family of toads with a subcosmopolitan distribution. Recent molecular phylogenetic analyses have revealed a radiation of toads (Adenominae) with distinct adult and larval ecomorphs on the Southern parts of the Indian subcontinent. The Indian torrential species "Ansonia" ornata has a basal position in this clade and does not group with South-East Asian Ansonia. Additionally, the nested position of "Bufo" koynayensis and an undescribed sister species, and their distinct ecologies including a non-typical egg-laying strategy within bufonids, support the recognition of a second distinct genus. In this paper we describe two new genera and one new species from the Adenominae clade. Findings: Ansonia ornata Günther, 1876 "1875" is transferred to Ghatophryne gen. nov., a genus of torrentially adapted toads that are endemic to the Western Ghats of India. On the basis of close morphological resemblance and distribution, Ansonia rubigina Pillai and Pattabiraman, 1981 is provisionally transferred to this new genus. The Western Ghats endemic toad Bufo koynayensis Soman, 1963 is transferred to a new genus Xanthophryne gen. nov. Based on molecular and morphological evidence, we additionally describe a new species, Xanthophryne tigerinus sp. nov., from Amboli in the Western Ghats. Conclusion: The descriptions and subsequent taxonomic changes we propose result in three genera of bufonids recognised as being endemic to the Western Ghats (Ghatophryne gen. nov., Xanthophryne gen. nov. and Pedostibes), and one to Sri Lanka (Adenomus). The spatial distribution, and arrangement of these lineages at the base of Adenominae diversification, reflects their Early Neogene isolation in the Western Ghats-Sri Lanka hotspot. Page 1 of 6 (page number not for citation purposes) BMC Research Notes 2009, 2:241 http://www.biomedcentral.com/1756-0500/2/241 Introduction Bufonidae are a family of toads with over 500 extant spe- cies distributed on most continents. Current opinions on bufonid taxonomy are very divergent, and range from the recognition of multiple genera [1], to favouring a subcos- mopolitan genus Bufo with plenty of subgenera [2]. Although the phylogenetic relationships of bufonids have been studied intensively [1,3,4], the evolutionary position of species on the Indian subcontinent had remained unclear. However, recent molecular phylogenetic analyses revealed that these toads belong to a radiation containing distinct ecomorphs in adult and/or larval forms [5]. The early diversification of this clade, for which the name Ade- nominae is available [6], has been reconstructed as occur- Ghatophryne orna mm, collected fr nad, Kerala Figure 1 om Kurichiy ta specimen armala, 11°3 SDB 6361, a5 d'N, ult ma 75°58'E, le, SVL Waya- 29.2 ring on the Southern parts of the Indian subcontinent. Ghatophryne ornata specimen SDB 6361, adult male, Early diversification of Adenominae, timed at the early SVL 29.2 mm, collected from Kurichiyarmala, Miocene, led to several endemic lineages ("Ansonia" 11°35'N, 75°58'E, Wayanad, Kerala. ornata, Pedostibes tuberculosus, the "Bufo" koynayensis group and Adenomus) [5] and to the origin of the more wide- spread Duttaphrynus clade. The unexpected evolutionary The original description of this species was based on nine position of "Ansonia" ornata and "Bufo" koynayensis (see syntypes (BMNH 1947.2.20.65-1947.2.20.73) collected abstract) requires recognition of two new genera to in "Bramagherries" (= Brahmagiri Hills, Coorg) by R. H. accommodate these Western Ghats endemic species and Beddome. We hereby designate one of them, BMNH their relatives. 1947.2.20.66 (ex BMNH 74.4.29.945), an adult male, SVL 29.0, as lectotype of this nominal taxon. Methods Collection of amphibian specimens was made during Diagnosis and comparison fieldtrips in the Western Ghats between 1997 and 2002. Ghatophryne can be distinguished from other bufonid Specimens were preserved in 5% formaldehyde for 2 days, genera by the combination of the following characters: and subsequently transferred to 70% ethanol. Measure- small-sized adults (male SVL 26.9-30.1, N = 5; female ments were taken to the nearest 0.1 mm, using a digital 35.0, N = 1); dorsum reddish brown, ventrum dark slide-caliper or a binocular microscope with a micrometer brownish black with prominent yellowish-orange spots; ocular. The description (all measurements in mm) of the head without cranial ridge, lack of parotoid glands; fingers types follows terminology used elsewhere [7]. To comply free, toes medium webbed; skin on dorsum with sparsely with regulations of the International Code of Zoological granular projections especially on anterior half of the Nomenclature we have deposited copies of this article at body; eggs non-pigmented, tadpoles with a suctorial disk, the following publicly accessible libraries: Bombay Natu- adapted to mountain streams throughout the life cycle ral History Society, Mumbai, India (BNHS); Koninklijk (SD Biju, personal observations). Ghatophryne can be Belgisch Instituut voor Natuurwetenschappen, Brussels, characterized in a phylogenetic framework as the most Belgium (KBIN); Natural History Museum, London, UK inclusive clade that contains Ansonia ornata Günther, (BMNH); American Museum of Natural History, New "1876" 1875, but not Bufo melanostictus Schneider, 1799 York, USA (AMNH); Museum National d'Histoire and Ansonia hanitschi Inger, 1960 [5]. Naturelle, Paris, France (MNHN); Russian Academy of Sciences, Moscow, Russia (RAS). Ghatophryne is morphologically very similar to species of Ansonia, but is endemic to the southern part of the West- Descriptions ern Ghats, while Ansonia is known only from South-East 1. Ghatophryne gen. nov. (Figure 1) Asia (Sumatra, Borneo and Philippines). Etymology The generic epithet is derived from the words 'Ghats' and Contents 'phryne'. The former is the Sanskrit word for 'steps' and Ghatophryne ornata (Günther, "1876" 1875) comb. refers to the Western Ghats mountain range, where this nov., Ghatophryne rubigina (Pillai and Pattabiraman, genus is endemic, while 'phryne' means toad in Greek. 1981) comb. nov. The latter is provisionally included based on distributional data and morphological resem- Type species blances [8]. Ansonia ornata Günther, "1876" 1875, by original designa- tion. Page 2 of 6 (page number not for citation purposes) BMC Research Notes 2009, 2:241 http://www.biomedcentral.com/1756-0500/2/241 Distribution Diagnosis and comparison Ghatophryne has only been reported from Kerala and Kar- Xanthophryne can be distinguished from other bufonid nataka in the Western Ghats of India. genera by the combination of the following characters: small-sized adults (male SVL 26.5-32.9, N = 12; female Description of lectotype of Ansonia ornata Günther, "1876" 1875 SVL 33.3-35.3, N = 3) having light brown dorsum with a Medium sized frog (SVL 29.0), body rather elongate. suffusion of dull chrome-yellow; head with discontinu- Head slightly longer than wide (HL 8.9; HW 8.5; MN 7.2; ous and weak canthal and preorbital ridges on the ante- MFE 5.4; MBE 1.3); outline of snout in dorsal view nearly rior part, flanks and sides of the abdomen have chrome- oval, its length (SL 4.5) larger than the horizontal diame- yellow patches, or sometimes 2-4 continuous bands; tym- ter of the eye (EL 3.7); loreal region vertically acute; dis- panum indistinct, rather weak parotoid glands; toes and tance between anterior margins of eyes (IFE 4.6) 1.8 times fingers without webbing, tips rounded; eggs in clutches. in distance between posterior margins of eyes (IBE 8.4); Xanthophryne can be characterized in a phylogenetic interorbital area (IUE 2.8) broader than upper eyelid framework as the most inclusive clade that contains Bufo width (UEW 2.3); tympanum (TYD 2.7) rather indistinct; koynayensis Soman, 1963 but not Bufo melanostictus supratympanic fold rather indistinct; parotoid glands Schneider, 1799 and Bufo kelaartii Günther, 1858. absent; parietal ridges absent; tongue elongate. Forearm (FLL 7.5) shorter than hand (HAL 8.8; TFL 4.0), strong; Contents relative length of fingers, shortest to longest: I < II < IV < Xanthophryne koynayensis (Soman, 1963) comb. nov. (Fig- III, tip of fingers enlarged, rounded, without distinct ure 2) and Xanthophryne tigerinus sp. nov. (Figure 3A) grooves, without lateral dermal fringe, webbing absent; subarticular tubercles weakly developed; supernumerary Distribution tubercles weakly developed; palmar tubercle absent, nup- Xanthophryne has only been reported from the northern tial pads present. Hind limbs moderately long, shank part of the Western Ghats of India. (ShL 14.1) longer than thigh (TL 13.3), longer than dis- tance from base of inner metatarsal tubercle to tip of toe 3. Xanthophryne tigerinus sp. nov.(Figure 3A-E) IV (FOL 12.6), shorter than distance from heel to tip of toe Etymology The species epithet tigerinus is attributed to this species IV (TFOL 19.2); relative length of toes, shortest to longest: I < II < V < III < IV, tips of toes enlarged, rounded, webbing due to similarity in the species colour pattern to the prom- reduced, reaching up to second subarticular tubercle on inent lateral stripes in Tigers (Panthera tigris). both sides of toe IV; dermal fringe along toe V absent; sub- Type material articular tubercles distinct; supernumerary tubercles weakly developed; inner metatarsal tubercle elongate Holotype, BNHS 5175, an adult male, SVL 30.5, collected (IMT 1.5), outer metatarsal tubercle smaller but distinctly by SDB and VG on 23 August 2002 from Amboli, raised. Skin of snout, between eyes, and upper eyelids sha- 15°56'N, 74°00'E, about 720 m asl, Sindhudurg, Mahar- greened to sparsely granular, side of head, back, flanks ashtra, India; paratypes, BNHS 5177, an adult male, and dorsal part of limbs sparsely granular. BNHS 5176, an adult female, collected along with the hol- Colour of lectotype: in preservation dorsally uniform light greyish brown with a dorsal light yellowish brown patch; ventral side light brown with white irregular spots. Other specimens studied Ovary of single female specimen (SDB 022) had 35 non- pigmented, creamy white eggs, 1.3 ± 0.4 mm in diameter. Dorsum can have continuous (SDB 6362) or discontinu- ous (SDB 6361) yellowish bands. 2. Xanthophryne gen. nov Etymology derived from two Greek words, 'xanthos' meaning yellow, and 'phryne' meaning toad. Type species Xanthophryne koynayensis type locality Figure 2 , a male adult (SDB 6040) from the Bufo koynayensis Soman, 1963, by original designation, Xanthophryne koynayensis, a male adult (SDB 6040) from the type locality. ZSIC A1784, Shivaji Sagar lake at Koyna (= Humbarli Vil- lage), Maharashtra, collected by P. W. Soman. Page 3 of 6 (page number not for citation purposes) BMC Research Notes 2009, 2:241 http://www.biomedcentral.com/1756-0500/2/241 prominent canthal and preorbital ridges; snout longer than eye length (SL 4.0 ± 0.3 mm, EL 3.4 ± 0.5 mm, N = 5, male) vs. snout shorter than eye length (SL 3.3 ± 0.3 mm, EL 4.3 ± 0.2 mm, N = 7, male); shank longer than thigh (ShL 11.5 ± 0.6 mm, TL 9.9 ± 0.4 mm, N = 5, males) vs. shank about equal to thigh (ShL 11.1 ± 0.7 mm, TL 11.1 ± 0.8 mm, N = 7, male); foot length longer than shank and thigh (FOL 12.8 ± 0.6, N = 5, male) vs. equal to shank and thigh (FOL 11.2 ± 0.7, N = 7, male). Description of the holotype Medium sized frog (SVL 30.5), body rather elongate (Fig- ure 3A). Head length subequal (HL 10.1) to head width (HW 10.3; MN 8.8; MFE 7.2; MBE 4.2) (Figure 3B, C); out- line of snout in dorsal view nearly oval, in ventral view oval to pointed, its length (SL 4.1) longer than the hori- zontal diameter of the eye (EL 3.1); loreal region vertically acute; distance between anterior margins of eyes (IFE 4.6) 1.9 times in distance between posterior margins of eyes (IBE 8.7); interorbital area (IUE 2.3) less than upper eye- lid (UEW 3.1); tympanum rather indistinct; supratym- panic fold absent; parotoid glands present, rounded; canthal and preorbital ridges well developed, continuous; tongue entire, oval. Forearm (FLL 6.1) shorter than hand (HAL 7.0; TFL 3.5); relative length of fingers, shortest to longest: I < II < IV < III (Figure 3D), tip of fingers enlarged, rounded, without distinct grooves, without lateral dermal fringe, webbing absent; subarticular tubercles weakly developed; prepollex rounded, prominent, palmar tuber- Xa Figure 3 nthophryne tigerinus sp. nov (holotype, BNHS 5175) cle one, rounded, prominent; supernumerary tubercles Xanthophryne tigerinus sp. nov (holotype, BNHS 5175). A, holotype in life; B, dorsal view of head showing weakly developed; nuptial pads present. Hindlimbs mod- prominent granular projections with horny spinules; C, lat- erately long, shank (ShL 11.8) longer than thigh (TL eral view of head; D, ventral view of hand; E, ventral view of 10.1), shorter than distance from base of inner metatarsal foot; F. Xanthophryne koynayensis, dorsal view of the head, tubercle to tip of toe IV (FOL 13.1); distance from heel to showing scattered granular projections with horny spinules. tip of toe IV (TFOL18.1); relative length of toes, shorter to longest: I < II < III < V < IV (Figure 3E), tips of toes enlarged, rounded, webbing absent; subarticular tubercles otype; BNHS 4064, an adult male, BNHS 4063, an adult rather indistinct; supernumerary tubercles weakly devel- female, collected by VG & Hegde on 9 September 2000, oped; inner metatarsal tubercle elongated (IMT 1.2). Skin BNHS 4208-4209, two males, BNHS 4207, an adult of snout, between eyes, upper eyelids, side of head, and female, collected by Kehimkar and Agarwal on 16 August back have granular projections with horny spinules; dor- 2003. sal part of limbs granular with horny spinules; throat sha- greened to granular; chest, belly, and posterior surface of Diagnosis and comparison thighs granular. Xanthophryne tigerinus can be distinguished by the follow- ing combination of characters: (1) medium size, male Color: in preservation: dorsum light-brown, loreal and tym- adult SVL 27.8-32.9, female adult SVL 33.3-35.3; (2) body panic region light brown, lateral region dark grey with rather elongate; (3) presence of discontinuous canthal light colour stripes; fingers I and II, and toes I and II and preorbital ridges; (4) stripes on lateral and dorsal creamy white; ventral side light brownish grey with dark side; (5) absence of webbing between fingers and toes. grey irregular spots; in life: dorsum golden yellow, lateral region brown with light yellow stripes extending from Xanthophryne tigerinus sp. nov. (Figure 3A-E) differs from dorsum; fingers I and II, and toes I and II whitish; ventral X. koynayensis (Figure 2, 3F) by the presence of a denser side light grey with dark greyish brown irregular spots, arrangement of granular projections with horny spinules lower jaw margins white. on dorsal and lateral parts of head, back and flank; more Page 4 of 6 (page number not for citation purposes) BMC Research Notes 2009, 2:241 http://www.biomedcentral.com/1756-0500/2/241 Secondary sexual characteristics sonal observation). The eggs are laid in temporary pud- Male (BNHS 5175) has nuptial spines on the prepollex; dles on laterite rocks (Figure 4). Ovary of single female specimen ((BNHS 4207) had 59 pigmented eggs, 1.2 ± 0.3 mm in diameter. Notes Bufo koynayensis Soman, 1963 was described twice based Variation on the same type series but bearing different numbers. Measurements of eight type specimens are given in Table Soman first described Bufo koynayensis, without designat- 1. During the breeding season, both male and female ing any type other than mentioning, "was collected in toads have a bright yellow dorsum with light yellow and good numbers" [9]. Almost 40 years later, specimen ZSI light blue lateral stripes. Outside the breeding season, A.1784 was recognized as the holotype of Bufo koynayensis they have a light brownish grey dorsum with light yellow Soman, 1963 [10]. This specimen was examined and is lateral stripes. badly damaged (SDB, personal observation). The samples collected by Soman were utilized to describe Bufo sul- Distribution and natural history phureus (holotype, BNHS 377 and 26 paratypes including Xanthophryne tigerinus sp. nov. is known only from the 5 males, 11 females and 10 juveniles) [11], which as a type locality Amboli. The type series was collected from consequence is a junior synonym of Bufo koynayensis. As the ground near disturbed evergreen forest patches during described in the original description, the type series of a rainy evening after 19:00 h. Amplexus is axillary. Egg Bufo sulphureus were deposited in the British Museum of clutches contain 30-35 eggs per clutch (N = 4; VG, per- Natural History, the Indian Museum and the Bombay Table 1: Measurements of the two Xanthophryne gen. nov. species Xanthophryne tigerinus sp. nov. (type series) Male Female BNHS BNHS BNHS BNHS BNHS Mean STDV BNHS BNHS BNHS Mean STDV 5175 5177 4064 4208 4209 5176 4063 4207 SVL 30.5 31.0 32.9 32.0 27.8 30.8 1.9 33.9 33.3 35.3 34.2 1.0 HW 10.3 10.1 11.0 10.2 10.1 10.3 0.4 12.2 11.5 13.9 12.5 1.2 HL 10.1 10.0 10.7 10.1 9.0 10.0 0.6 11.2 10.9 12.2 11.4 0.7 IUE 2.3 2.1 2.3 2.1 2.5 2.3 0.2 2.3 2.5 3.0 2.6 0.4 UEW 3.1 2.9 3.6 3.1 2.9 3.1 0.3 3.4 3.4 3.2 3.3 0.1 SL 4.1 3.6 4.5 4.1 3.8 4.0 0.3 4.5 4.8 4.6 4.6 0.2 EL 3.1 3.0 4.0 4.0 3.0 3.4 0.5 4.4 4.6 4.5 4.5 0.1 FLL 6.1 6.8 5.4 6.5 5.8 6.1 0.6 7.4 6.1 6.1 6.5 0.8 HAL 7.0 6.9 6.7 6.8 6.7 6.8 0.1 6.7 6.9 7.4 7.0 0.4 ShL 11.8 11.9 11.8 11.7 10.4 11.5 0.6 11.6 11.7 12.2 11.8 0.3 TL 10.1 10.2 9.3 10.3 9.7 9.9 0.4 10.7 8.8 11.6 10.4 1.4 FOL 13.1 12.8 13.4 12.6 11.9 12.8 0.6 12.6 13.2 13.9 13.2 0.7 Xanthophryne koynayensis Males BNHS* BNHS** BNHS** BNHS** BNHS** BNHS BNHS Mean STDV 377 357 358 372 365 5174 5188 SVL 29.6 29.8 31.8 31.1 31.2 27.1 26.5 29.6 2.1 HW 11.1 10.3 11.0 11.0 10.9 10.6 10.5 10.8 0.3 HL 10.6 10.0 11.4 10.2 10.0 8.4 9.1 10.0 1.0 IUE 2.4 2.0 2.4 2.4 2.2 2.5 2.7 2.4 0.2 UEW 3.3 2.8 3.1 3.1 3.1 3.1 3.2 3.1 0.2 SL 3.2 3.1 3.2 3.7 3.8 3.2 3.2 3.3 0.3 EL 4.1 4.2 4.2 4.7 4.5 4.2 4.2 4.3 0.2 FLL 6.2 5.3 5.8 5.8 6.2 5.2 6.0 5.8 0.4 HAL 6.3 6.0 7.1 7.6 7.3 6.0 7.1 6.8 0.7 ShL 11.8 10.1 11.5 11.3 11.5 10.2 11.2 11.1 0.7 TL 11.9 10.0 11.6 11.2 11.6 10.1 11.3 11.1 0.8 FOL 11.9 10.2 11.5 11.4 11.5 10.3 11.5 11.2 0.7 * holotype and ** paratype of Bufo sulphureus Grandison and Daniel, 1964. Page 5 of 6 (page number not for citation purposes) BMC Research Notes 2009, 2:241 http://www.biomedcentral.com/1756-0500/2/241 and Forest (MoEF) and University of Delhi (Scheme to strengthen R&D and Doctoral Research Program) for funds for the research. IVB is supported by a research grant from the Institute for the Promotion of Innovation through Science and Technology in Flanders (IWT-Vlaanderen). FB receives a postdoctoral fellowship from the Fonds voor Wetenschappelijk Onder- zoek-Vlaanderen (FWO). Financial support was provided FWO-Vlaan- deren (grants FWO 1.5.039.03N, FWO G.0056.03, FWO G.0307.04) and the Vrije Universiteit Brussel (grant VUB OZR834). References 1. Frost DR, Grant T, Faivovich J, Bain RH, Haas A, Haddad CFBdS RO, Channing A, Wilkinson M, Donnellan SC, Raxworthy CJ, et al.: The amphibian tree of life. Bull Am Mus Nat Hist 2006, 297:1-370. 2. Smith HM, Chiszar D: Dilemma of name-recognition: Why and when to use new combinations of scientific names. Herpeto- logical Conservation and Biology 2006, 1(1):6-8. 3. Pramuk JB, Robertson T, Sites JW, Noonan BP: Around the world in 10 million years: Biogeography of the nearly cosmopolitan true toads (Anura: Bufonidae). Global Ecol Biogeogr 2007, 17:72-83. Xa pu Figure 4 d nd tho les phryne tigerinus eggs are laid in clutches in temporary 4. Pauly GB, Hillis DM, Cannatella DC: The history of a Nearctic col- Xanthophryne tigerinus eggs are laid in clutches in onization: Molecular phylogenetics and biogeography of the temporary puddles. neartic toads (Bufo). Evolution 2004, 58(11):2517-2535. 5. Van Bocxlaer I, Biju S, Loader S, Bossuyt F: Toad radiation reveals into-India dispersal as a source of endemism in the Western Ghats-Sri Lanka biodiversity hotspot. BMC Evolutionary Biology 2009, 9(131):1-10. Natural History Society. In this study, we examined the 6. Dubois A: La nomenclature supragénérique des Amphibiens Anoures. Mémoires du Muséum national d'histoire naturelle 1984, holotype and 18 paratypes (14 adult males - SVL 24.0- série A, Zoologie, 131:1-64. 31.8 mm and 4 subadults - SVL 20.9-25.0 mm) deposited 7. Biju SD, Bossuyt F: Systematics and phylogeny of Philautus gis- in BNHS and five paratypes in BMNH under the nomen tel, 1848 (Anura, rhacophoridae) in the Western Ghats of India, with descriptions of 12 new species. Zool J Linn Soc 2009, 'Bufo sulphureus'. In addition, recent collections of 'Bufo' 155(2):374-444. koynayensis from the type locality were compared. All 8. Pillai RS, Pattabiraman R: A new species of torrent toad (genus: available material of this species is consistent with the Ansonia) from silent valley, s. India. Proc Indian Acad Sci (Anim Sci) 1981, 90(2):203-208. original description. 9. Soman PW: A new Bufo from maharashtra. Journal of Biological Science Bombay 1963, 6:74. 10. Chanda KS, Das I, Dubois A: Catalogue of amphibian types in Conclusion the collection of the zoological survey of India. Hamadryad Ghatophryne gen. nov. and Xanthophryne gen. nov. are both 2000, 25(2):100-128. endemic to the Western Ghats of India. Together with the 11. Grandison AG, Daniel JC: Description of a new species of toad from surat district, maharashtra, India. Journal Bombay Natural arboreal toad genus Pedostibes (Western Ghats endemic; History Society 1964, 61:192-194. species from South-East Asia belong to another clade) and the genus Adenomus (Sri Lanka endemic), they form the oldest lineages in the Adenominae clade. The recognition of each of these lineages as distinct genera highlights their early diversification on the southern parts of the Indian subcontinent [5]. Competing interests The authors declare that they have no competing interests. Publish with Bio Med Central and every Authors' contributions scientist can read your work free of charge All authors contributed equally, and read and approved "BioMed Central will be the most significant development for the final manuscript. disseminating the results of biomedical researc h in our lifetime." Sir Paul Nurse, Cancer Research UK Acknowledgements Your research papers will be: We thank G. Ramakrishna and Kausik Deuti (ZSI), Asad Rahmani, J.C. Dan- available free of charge to the entire biomedical community iel (BNHS) and B. Clarke (NHM) for their support in our studies. Anil Zach- arya, Kehimkar, I. Agarwal and V. Patil provided valuable support in the peer reviewed and published immediately upon acceptance field; I. Das provided valuable literature; I. Agarwal and Hemant Ogale made cited in PubMed and archived on PubMed Central photographs of Xanthophryne available. The State Forest Departments of yours — you keep the copyright Kerala and Maharashtra kindly provided study permits to SDB and VG, and BioMedcentral Submit your manuscript here: are greatly acknowledged. SDB is grateful to the Ministry of Environment http://www.biomedcentral.com/info/publishing_adv.asp Page 6 of 6 (page number not for citation purposes) http://www.deepdyve.com/assets/images/DeepDyve-Logo-lg.png BMC Research Notes Springer Journals

Two new endemic genera and a new species of toad (Anura: Bufonidae) from the Western Ghats of India

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Abstract

Background: Bufonidae are a large family of toads with a subcosmopolitan distribution. Recent molecular phylogenetic analyses have revealed a radiation of toads (Adenominae) with distinct adult and larval ecomorphs on the Southern parts of the Indian subcontinent. The Indian torrential species "Ansonia" ornata has a basal position in this clade and does not group with South-East Asian Ansonia. Additionally, the nested position of "Bufo" koynayensis and an undescribed sister species, and their distinct ecologies including a non-typical egg-laying strategy within bufonids, support the recognition of a second distinct genus. In this paper we describe two new genera and one new species from the Adenominae clade. Findings: Ansonia ornata Günther, 1876 "1875" is transferred to Ghatophryne gen. nov., a genus of torrentially adapted toads that are endemic to the Western Ghats of India. On the basis of close morphological resemblance and distribution, Ansonia rubigina Pillai and Pattabiraman, 1981 is provisionally transferred to this new genus. The Western Ghats endemic toad Bufo koynayensis Soman, 1963 is transferred to a new genus Xanthophryne gen. nov. Based on molecular and morphological evidence, we additionally describe a new species, Xanthophryne tigerinus sp. nov., from Amboli in the Western Ghats. Conclusion: The descriptions and subsequent taxonomic changes we propose result in three genera of bufonids recognised as being endemic to the Western Ghats (Ghatophryne gen. nov., Xanthophryne gen. nov. and Pedostibes), and one to Sri Lanka (Adenomus). The spatial distribution, and arrangement of these lineages at the base of Adenominae diversification, reflects their Early Neogene isolation in the Western Ghats-Sri Lanka hotspot. Page 1 of 6 (page number not for citation purposes) BMC Research Notes 2009, 2:241 http://www.biomedcentral.com/1756-0500/2/241 Introduction Bufonidae are a family of toads with over 500 extant spe- cies distributed on most continents. Current opinions on bufonid taxonomy are very divergent, and range from the recognition of multiple genera [1], to favouring a subcos- mopolitan genus Bufo with plenty of subgenera [2]. Although the phylogenetic relationships of bufonids have been studied intensively [1,3,4], the evolutionary position of species on the Indian subcontinent had remained unclear. However, recent molecular phylogenetic analyses revealed that these toads belong to a radiation containing distinct ecomorphs in adult and/or larval forms [5]. The early diversification of this clade, for which the name Ade- nominae is available [6], has been reconstructed as occur- Ghatophryne orna mm, collected fr nad, Kerala Figure 1 om Kurichiy ta specimen armala, 11°3 SDB 6361, a5 d'N, ult ma 75°58'E, le, SVL Waya- 29.2 ring on the Southern parts of the Indian subcontinent. Ghatophryne ornata specimen SDB 6361, adult male, Early diversification of Adenominae, timed at the early SVL 29.2 mm, collected from Kurichiyarmala, Miocene, led to several endemic lineages ("Ansonia" 11°35'N, 75°58'E, Wayanad, Kerala. ornata, Pedostibes tuberculosus, the "Bufo" koynayensis group and Adenomus) [5] and to the origin of the more wide- spread Duttaphrynus clade. The unexpected evolutionary The original description of this species was based on nine position of "Ansonia" ornata and "Bufo" koynayensis (see syntypes (BMNH 1947.2.20.65-1947.2.20.73) collected abstract) requires recognition of two new genera to in "Bramagherries" (= Brahmagiri Hills, Coorg) by R. H. accommodate these Western Ghats endemic species and Beddome. We hereby designate one of them, BMNH their relatives. 1947.2.20.66 (ex BMNH 74.4.29.945), an adult male, SVL 29.0, as lectotype of this nominal taxon. Methods Collection of amphibian specimens was made during Diagnosis and comparison fieldtrips in the Western Ghats between 1997 and 2002. Ghatophryne can be distinguished from other bufonid Specimens were preserved in 5% formaldehyde for 2 days, genera by the combination of the following characters: and subsequently transferred to 70% ethanol. Measure- small-sized adults (male SVL 26.9-30.1, N = 5; female ments were taken to the nearest 0.1 mm, using a digital 35.0, N = 1); dorsum reddish brown, ventrum dark slide-caliper or a binocular microscope with a micrometer brownish black with prominent yellowish-orange spots; ocular. The description (all measurements in mm) of the head without cranial ridge, lack of parotoid glands; fingers types follows terminology used elsewhere [7]. To comply free, toes medium webbed; skin on dorsum with sparsely with regulations of the International Code of Zoological granular projections especially on anterior half of the Nomenclature we have deposited copies of this article at body; eggs non-pigmented, tadpoles with a suctorial disk, the following publicly accessible libraries: Bombay Natu- adapted to mountain streams throughout the life cycle ral History Society, Mumbai, India (BNHS); Koninklijk (SD Biju, personal observations). Ghatophryne can be Belgisch Instituut voor Natuurwetenschappen, Brussels, characterized in a phylogenetic framework as the most Belgium (KBIN); Natural History Museum, London, UK inclusive clade that contains Ansonia ornata Günther, (BMNH); American Museum of Natural History, New "1876" 1875, but not Bufo melanostictus Schneider, 1799 York, USA (AMNH); Museum National d'Histoire and Ansonia hanitschi Inger, 1960 [5]. Naturelle, Paris, France (MNHN); Russian Academy of Sciences, Moscow, Russia (RAS). Ghatophryne is morphologically very similar to species of Ansonia, but is endemic to the southern part of the West- Descriptions ern Ghats, while Ansonia is known only from South-East 1. Ghatophryne gen. nov. (Figure 1) Asia (Sumatra, Borneo and Philippines). Etymology The generic epithet is derived from the words 'Ghats' and Contents 'phryne'. The former is the Sanskrit word for 'steps' and Ghatophryne ornata (Günther, "1876" 1875) comb. refers to the Western Ghats mountain range, where this nov., Ghatophryne rubigina (Pillai and Pattabiraman, genus is endemic, while 'phryne' means toad in Greek. 1981) comb. nov. The latter is provisionally included based on distributional data and morphological resem- Type species blances [8]. Ansonia ornata Günther, "1876" 1875, by original designa- tion. Page 2 of 6 (page number not for citation purposes) BMC Research Notes 2009, 2:241 http://www.biomedcentral.com/1756-0500/2/241 Distribution Diagnosis and comparison Ghatophryne has only been reported from Kerala and Kar- Xanthophryne can be distinguished from other bufonid nataka in the Western Ghats of India. genera by the combination of the following characters: small-sized adults (male SVL 26.5-32.9, N = 12; female Description of lectotype of Ansonia ornata Günther, "1876" 1875 SVL 33.3-35.3, N = 3) having light brown dorsum with a Medium sized frog (SVL 29.0), body rather elongate. suffusion of dull chrome-yellow; head with discontinu- Head slightly longer than wide (HL 8.9; HW 8.5; MN 7.2; ous and weak canthal and preorbital ridges on the ante- MFE 5.4; MBE 1.3); outline of snout in dorsal view nearly rior part, flanks and sides of the abdomen have chrome- oval, its length (SL 4.5) larger than the horizontal diame- yellow patches, or sometimes 2-4 continuous bands; tym- ter of the eye (EL 3.7); loreal region vertically acute; dis- panum indistinct, rather weak parotoid glands; toes and tance between anterior margins of eyes (IFE 4.6) 1.8 times fingers without webbing, tips rounded; eggs in clutches. in distance between posterior margins of eyes (IBE 8.4); Xanthophryne can be characterized in a phylogenetic interorbital area (IUE 2.8) broader than upper eyelid framework as the most inclusive clade that contains Bufo width (UEW 2.3); tympanum (TYD 2.7) rather indistinct; koynayensis Soman, 1963 but not Bufo melanostictus supratympanic fold rather indistinct; parotoid glands Schneider, 1799 and Bufo kelaartii Günther, 1858. absent; parietal ridges absent; tongue elongate. Forearm (FLL 7.5) shorter than hand (HAL 8.8; TFL 4.0), strong; Contents relative length of fingers, shortest to longest: I < II < IV < Xanthophryne koynayensis (Soman, 1963) comb. nov. (Fig- III, tip of fingers enlarged, rounded, without distinct ure 2) and Xanthophryne tigerinus sp. nov. (Figure 3A) grooves, without lateral dermal fringe, webbing absent; subarticular tubercles weakly developed; supernumerary Distribution tubercles weakly developed; palmar tubercle absent, nup- Xanthophryne has only been reported from the northern tial pads present. Hind limbs moderately long, shank part of the Western Ghats of India. (ShL 14.1) longer than thigh (TL 13.3), longer than dis- tance from base of inner metatarsal tubercle to tip of toe 3. Xanthophryne tigerinus sp. nov.(Figure 3A-E) IV (FOL 12.6), shorter than distance from heel to tip of toe Etymology The species epithet tigerinus is attributed to this species IV (TFOL 19.2); relative length of toes, shortest to longest: I < II < V < III < IV, tips of toes enlarged, rounded, webbing due to similarity in the species colour pattern to the prom- reduced, reaching up to second subarticular tubercle on inent lateral stripes in Tigers (Panthera tigris). both sides of toe IV; dermal fringe along toe V absent; sub- Type material articular tubercles distinct; supernumerary tubercles weakly developed; inner metatarsal tubercle elongate Holotype, BNHS 5175, an adult male, SVL 30.5, collected (IMT 1.5), outer metatarsal tubercle smaller but distinctly by SDB and VG on 23 August 2002 from Amboli, raised. Skin of snout, between eyes, and upper eyelids sha- 15°56'N, 74°00'E, about 720 m asl, Sindhudurg, Mahar- greened to sparsely granular, side of head, back, flanks ashtra, India; paratypes, BNHS 5177, an adult male, and dorsal part of limbs sparsely granular. BNHS 5176, an adult female, collected along with the hol- Colour of lectotype: in preservation dorsally uniform light greyish brown with a dorsal light yellowish brown patch; ventral side light brown with white irregular spots. Other specimens studied Ovary of single female specimen (SDB 022) had 35 non- pigmented, creamy white eggs, 1.3 ± 0.4 mm in diameter. Dorsum can have continuous (SDB 6362) or discontinu- ous (SDB 6361) yellowish bands. 2. Xanthophryne gen. nov Etymology derived from two Greek words, 'xanthos' meaning yellow, and 'phryne' meaning toad. Type species Xanthophryne koynayensis type locality Figure 2 , a male adult (SDB 6040) from the Bufo koynayensis Soman, 1963, by original designation, Xanthophryne koynayensis, a male adult (SDB 6040) from the type locality. ZSIC A1784, Shivaji Sagar lake at Koyna (= Humbarli Vil- lage), Maharashtra, collected by P. W. Soman. Page 3 of 6 (page number not for citation purposes) BMC Research Notes 2009, 2:241 http://www.biomedcentral.com/1756-0500/2/241 prominent canthal and preorbital ridges; snout longer than eye length (SL 4.0 ± 0.3 mm, EL 3.4 ± 0.5 mm, N = 5, male) vs. snout shorter than eye length (SL 3.3 ± 0.3 mm, EL 4.3 ± 0.2 mm, N = 7, male); shank longer than thigh (ShL 11.5 ± 0.6 mm, TL 9.9 ± 0.4 mm, N = 5, males) vs. shank about equal to thigh (ShL 11.1 ± 0.7 mm, TL 11.1 ± 0.8 mm, N = 7, male); foot length longer than shank and thigh (FOL 12.8 ± 0.6, N = 5, male) vs. equal to shank and thigh (FOL 11.2 ± 0.7, N = 7, male). Description of the holotype Medium sized frog (SVL 30.5), body rather elongate (Fig- ure 3A). Head length subequal (HL 10.1) to head width (HW 10.3; MN 8.8; MFE 7.2; MBE 4.2) (Figure 3B, C); out- line of snout in dorsal view nearly oval, in ventral view oval to pointed, its length (SL 4.1) longer than the hori- zontal diameter of the eye (EL 3.1); loreal region vertically acute; distance between anterior margins of eyes (IFE 4.6) 1.9 times in distance between posterior margins of eyes (IBE 8.7); interorbital area (IUE 2.3) less than upper eye- lid (UEW 3.1); tympanum rather indistinct; supratym- panic fold absent; parotoid glands present, rounded; canthal and preorbital ridges well developed, continuous; tongue entire, oval. Forearm (FLL 6.1) shorter than hand (HAL 7.0; TFL 3.5); relative length of fingers, shortest to longest: I < II < IV < III (Figure 3D), tip of fingers enlarged, rounded, without distinct grooves, without lateral dermal fringe, webbing absent; subarticular tubercles weakly developed; prepollex rounded, prominent, palmar tuber- Xa Figure 3 nthophryne tigerinus sp. nov (holotype, BNHS 5175) cle one, rounded, prominent; supernumerary tubercles Xanthophryne tigerinus sp. nov (holotype, BNHS 5175). A, holotype in life; B, dorsal view of head showing weakly developed; nuptial pads present. Hindlimbs mod- prominent granular projections with horny spinules; C, lat- erately long, shank (ShL 11.8) longer than thigh (TL eral view of head; D, ventral view of hand; E, ventral view of 10.1), shorter than distance from base of inner metatarsal foot; F. Xanthophryne koynayensis, dorsal view of the head, tubercle to tip of toe IV (FOL 13.1); distance from heel to showing scattered granular projections with horny spinules. tip of toe IV (TFOL18.1); relative length of toes, shorter to longest: I < II < III < V < IV (Figure 3E), tips of toes enlarged, rounded, webbing absent; subarticular tubercles otype; BNHS 4064, an adult male, BNHS 4063, an adult rather indistinct; supernumerary tubercles weakly devel- female, collected by VG & Hegde on 9 September 2000, oped; inner metatarsal tubercle elongated (IMT 1.2). Skin BNHS 4208-4209, two males, BNHS 4207, an adult of snout, between eyes, upper eyelids, side of head, and female, collected by Kehimkar and Agarwal on 16 August back have granular projections with horny spinules; dor- 2003. sal part of limbs granular with horny spinules; throat sha- greened to granular; chest, belly, and posterior surface of Diagnosis and comparison thighs granular. Xanthophryne tigerinus can be distinguished by the follow- ing combination of characters: (1) medium size, male Color: in preservation: dorsum light-brown, loreal and tym- adult SVL 27.8-32.9, female adult SVL 33.3-35.3; (2) body panic region light brown, lateral region dark grey with rather elongate; (3) presence of discontinuous canthal light colour stripes; fingers I and II, and toes I and II and preorbital ridges; (4) stripes on lateral and dorsal creamy white; ventral side light brownish grey with dark side; (5) absence of webbing between fingers and toes. grey irregular spots; in life: dorsum golden yellow, lateral region brown with light yellow stripes extending from Xanthophryne tigerinus sp. nov. (Figure 3A-E) differs from dorsum; fingers I and II, and toes I and II whitish; ventral X. koynayensis (Figure 2, 3F) by the presence of a denser side light grey with dark greyish brown irregular spots, arrangement of granular projections with horny spinules lower jaw margins white. on dorsal and lateral parts of head, back and flank; more Page 4 of 6 (page number not for citation purposes) BMC Research Notes 2009, 2:241 http://www.biomedcentral.com/1756-0500/2/241 Secondary sexual characteristics sonal observation). The eggs are laid in temporary pud- Male (BNHS 5175) has nuptial spines on the prepollex; dles on laterite rocks (Figure 4). Ovary of single female specimen ((BNHS 4207) had 59 pigmented eggs, 1.2 ± 0.3 mm in diameter. Notes Bufo koynayensis Soman, 1963 was described twice based Variation on the same type series but bearing different numbers. Measurements of eight type specimens are given in Table Soman first described Bufo koynayensis, without designat- 1. During the breeding season, both male and female ing any type other than mentioning, "was collected in toads have a bright yellow dorsum with light yellow and good numbers" [9]. Almost 40 years later, specimen ZSI light blue lateral stripes. Outside the breeding season, A.1784 was recognized as the holotype of Bufo koynayensis they have a light brownish grey dorsum with light yellow Soman, 1963 [10]. This specimen was examined and is lateral stripes. badly damaged (SDB, personal observation). The samples collected by Soman were utilized to describe Bufo sul- Distribution and natural history phureus (holotype, BNHS 377 and 26 paratypes including Xanthophryne tigerinus sp. nov. is known only from the 5 males, 11 females and 10 juveniles) [11], which as a type locality Amboli. The type series was collected from consequence is a junior synonym of Bufo koynayensis. As the ground near disturbed evergreen forest patches during described in the original description, the type series of a rainy evening after 19:00 h. Amplexus is axillary. Egg Bufo sulphureus were deposited in the British Museum of clutches contain 30-35 eggs per clutch (N = 4; VG, per- Natural History, the Indian Museum and the Bombay Table 1: Measurements of the two Xanthophryne gen. nov. species Xanthophryne tigerinus sp. nov. (type series) Male Female BNHS BNHS BNHS BNHS BNHS Mean STDV BNHS BNHS BNHS Mean STDV 5175 5177 4064 4208 4209 5176 4063 4207 SVL 30.5 31.0 32.9 32.0 27.8 30.8 1.9 33.9 33.3 35.3 34.2 1.0 HW 10.3 10.1 11.0 10.2 10.1 10.3 0.4 12.2 11.5 13.9 12.5 1.2 HL 10.1 10.0 10.7 10.1 9.0 10.0 0.6 11.2 10.9 12.2 11.4 0.7 IUE 2.3 2.1 2.3 2.1 2.5 2.3 0.2 2.3 2.5 3.0 2.6 0.4 UEW 3.1 2.9 3.6 3.1 2.9 3.1 0.3 3.4 3.4 3.2 3.3 0.1 SL 4.1 3.6 4.5 4.1 3.8 4.0 0.3 4.5 4.8 4.6 4.6 0.2 EL 3.1 3.0 4.0 4.0 3.0 3.4 0.5 4.4 4.6 4.5 4.5 0.1 FLL 6.1 6.8 5.4 6.5 5.8 6.1 0.6 7.4 6.1 6.1 6.5 0.8 HAL 7.0 6.9 6.7 6.8 6.7 6.8 0.1 6.7 6.9 7.4 7.0 0.4 ShL 11.8 11.9 11.8 11.7 10.4 11.5 0.6 11.6 11.7 12.2 11.8 0.3 TL 10.1 10.2 9.3 10.3 9.7 9.9 0.4 10.7 8.8 11.6 10.4 1.4 FOL 13.1 12.8 13.4 12.6 11.9 12.8 0.6 12.6 13.2 13.9 13.2 0.7 Xanthophryne koynayensis Males BNHS* BNHS** BNHS** BNHS** BNHS** BNHS BNHS Mean STDV 377 357 358 372 365 5174 5188 SVL 29.6 29.8 31.8 31.1 31.2 27.1 26.5 29.6 2.1 HW 11.1 10.3 11.0 11.0 10.9 10.6 10.5 10.8 0.3 HL 10.6 10.0 11.4 10.2 10.0 8.4 9.1 10.0 1.0 IUE 2.4 2.0 2.4 2.4 2.2 2.5 2.7 2.4 0.2 UEW 3.3 2.8 3.1 3.1 3.1 3.1 3.2 3.1 0.2 SL 3.2 3.1 3.2 3.7 3.8 3.2 3.2 3.3 0.3 EL 4.1 4.2 4.2 4.7 4.5 4.2 4.2 4.3 0.2 FLL 6.2 5.3 5.8 5.8 6.2 5.2 6.0 5.8 0.4 HAL 6.3 6.0 7.1 7.6 7.3 6.0 7.1 6.8 0.7 ShL 11.8 10.1 11.5 11.3 11.5 10.2 11.2 11.1 0.7 TL 11.9 10.0 11.6 11.2 11.6 10.1 11.3 11.1 0.8 FOL 11.9 10.2 11.5 11.4 11.5 10.3 11.5 11.2 0.7 * holotype and ** paratype of Bufo sulphureus Grandison and Daniel, 1964. Page 5 of 6 (page number not for citation purposes) BMC Research Notes 2009, 2:241 http://www.biomedcentral.com/1756-0500/2/241 and Forest (MoEF) and University of Delhi (Scheme to strengthen R&D and Doctoral Research Program) for funds for the research. IVB is supported by a research grant from the Institute for the Promotion of Innovation through Science and Technology in Flanders (IWT-Vlaanderen). FB receives a postdoctoral fellowship from the Fonds voor Wetenschappelijk Onder- zoek-Vlaanderen (FWO). Financial support was provided FWO-Vlaan- deren (grants FWO 1.5.039.03N, FWO G.0056.03, FWO G.0307.04) and the Vrije Universiteit Brussel (grant VUB OZR834). References 1. Frost DR, Grant T, Faivovich J, Bain RH, Haas A, Haddad CFBdS RO, Channing A, Wilkinson M, Donnellan SC, Raxworthy CJ, et al.: The amphibian tree of life. Bull Am Mus Nat Hist 2006, 297:1-370. 2. Smith HM, Chiszar D: Dilemma of name-recognition: Why and when to use new combinations of scientific names. Herpeto- logical Conservation and Biology 2006, 1(1):6-8. 3. Pramuk JB, Robertson T, Sites JW, Noonan BP: Around the world in 10 million years: Biogeography of the nearly cosmopolitan true toads (Anura: Bufonidae). Global Ecol Biogeogr 2007, 17:72-83. Xa pu Figure 4 d nd tho les phryne tigerinus eggs are laid in clutches in temporary 4. Pauly GB, Hillis DM, Cannatella DC: The history of a Nearctic col- Xanthophryne tigerinus eggs are laid in clutches in onization: Molecular phylogenetics and biogeography of the temporary puddles. neartic toads (Bufo). Evolution 2004, 58(11):2517-2535. 5. Van Bocxlaer I, Biju S, Loader S, Bossuyt F: Toad radiation reveals into-India dispersal as a source of endemism in the Western Ghats-Sri Lanka biodiversity hotspot. BMC Evolutionary Biology 2009, 9(131):1-10. Natural History Society. In this study, we examined the 6. Dubois A: La nomenclature supragénérique des Amphibiens Anoures. Mémoires du Muséum national d'histoire naturelle 1984, holotype and 18 paratypes (14 adult males - SVL 24.0- série A, Zoologie, 131:1-64. 31.8 mm and 4 subadults - SVL 20.9-25.0 mm) deposited 7. Biju SD, Bossuyt F: Systematics and phylogeny of Philautus gis- in BNHS and five paratypes in BMNH under the nomen tel, 1848 (Anura, rhacophoridae) in the Western Ghats of India, with descriptions of 12 new species. Zool J Linn Soc 2009, 'Bufo sulphureus'. In addition, recent collections of 'Bufo' 155(2):374-444. koynayensis from the type locality were compared. All 8. Pillai RS, Pattabiraman R: A new species of torrent toad (genus: available material of this species is consistent with the Ansonia) from silent valley, s. India. Proc Indian Acad Sci (Anim Sci) 1981, 90(2):203-208. original description. 9. Soman PW: A new Bufo from maharashtra. Journal of Biological Science Bombay 1963, 6:74. 10. Chanda KS, Das I, Dubois A: Catalogue of amphibian types in Conclusion the collection of the zoological survey of India. Hamadryad Ghatophryne gen. nov. and Xanthophryne gen. nov. are both 2000, 25(2):100-128. endemic to the Western Ghats of India. Together with the 11. Grandison AG, Daniel JC: Description of a new species of toad from surat district, maharashtra, India. Journal Bombay Natural arboreal toad genus Pedostibes (Western Ghats endemic; History Society 1964, 61:192-194. species from South-East Asia belong to another clade) and the genus Adenomus (Sri Lanka endemic), they form the oldest lineages in the Adenominae clade. The recognition of each of these lineages as distinct genera highlights their early diversification on the southern parts of the Indian subcontinent [5]. Competing interests The authors declare that they have no competing interests. Publish with Bio Med Central and every Authors' contributions scientist can read your work free of charge All authors contributed equally, and read and approved "BioMed Central will be the most significant development for the final manuscript. disseminating the results of biomedical researc h in our lifetime." Sir Paul Nurse, Cancer Research UK Acknowledgements Your research papers will be: We thank G. Ramakrishna and Kausik Deuti (ZSI), Asad Rahmani, J.C. Dan- available free of charge to the entire biomedical community iel (BNHS) and B. Clarke (NHM) for their support in our studies. Anil Zach- arya, Kehimkar, I. Agarwal and V. Patil provided valuable support in the peer reviewed and published immediately upon acceptance field; I. Das provided valuable literature; I. Agarwal and Hemant Ogale made cited in PubMed and archived on PubMed Central photographs of Xanthophryne available. The State Forest Departments of yours — you keep the copyright Kerala and Maharashtra kindly provided study permits to SDB and VG, and BioMedcentral Submit your manuscript here: are greatly acknowledged. SDB is grateful to the Ministry of Environment http://www.biomedcentral.com/info/publishing_adv.asp Page 6 of 6 (page number not for citation purposes)

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