Eight new species of Pseudophilautus(Amphibia: Anura: Rhacophoridae) from Sripada World Heritage Site (Peak Wilderness), a local amphibian hotspot in Sri Lanka

L.J. Mendis Wickramasinghe¹, Dulan Ranga Vidanapathirana ²,_, M.D. Gehan Rajeev ³, S. Chathuranga Ariyarathne ⁴, A.W. Amila Chanaka ⁵, L.L. Dharshana Priyantha ⁶,

Imesh Nuwan Bandara ⁷& Nethu Wickramasinghe ⁸

^{1, 2, 3, 4, 5, 6, 7, 8} Herpetological Foundation of Sri Lanka, 31/5, Alwis Town, Hendala, Wattala, Sri Lanka

¹ boiga2000@gmail.com (corresponding author), ² dulanrangavp@gmail.com,³ gehanrajeev@gmail.com, ⁴ harithasihinaya@yahoo.com, ⁵aamilachanaka@gmail.com, ⁶ dharshana.dpl@gmail.com, ⁷ imeshnu1@gmail.com,⁸ nemzy821@gmail.com

Abstract: Eight new species of Pseudophilautus(Pseudophilautus bambaradeniyai, P. dayawansai, P. jagathgunawardanai, P. karunarathnai, P. newtonjayawardanei, P. puranappu, P. samarakoon, and P. sirilwijesundarai) were discovered as a result of a survey carried out to study the herpetofaunal diversity with the changes in elevation in the Sripada World Heritage Site (Peak Wilderness), Central Hills of Sri Lanka. Detailed descriptions of new species along with colour photographs and line drawings for each species are provided herein. The new species possess unique morphological characters and are well distinguishable from one another that could be easily identified in the field. The conservation status of all species described here, have been considered Critically Endangered, except for P. newtonjayawardanei, as all the new species are recorded from single locations, and their habitats are under severe threat.

Keywords: Amphibian, new species, Peak Wilderness, Pseudophilautus,Sri Lanka, Sripada.

DOI: http://dx.doi.org/10.11609/JoTT.o3099.3789-920 | ZooBank: urn:lsid:zoobank.org:pub:8E8AF0BF-38D8-4B31-A67F-5BFD3735F66D

Editor: Annemarie Ohler, Muséum national d’Histoire naturelle, Paris, France Date of publication: 13 March 2013 (online & print)

Manuscript details: Ms # o3099 | Received 14 February 2012 | Final received 11 January 2013 | Finally accepted 19 February 2013

Citation: Wickramasinghe, L.J.M., D.R. Vidanapathirana, M.D.G. Rajeev, S.C. Ariyarathne, A.W.A. Chanaka, L.L.D. Priyantha, I.N. Bandara & N. Wickramasinghe (2013). Eight new species of Pseudophilautus (Amphibia, Anura, Rhacophoridae) from Sripada World Heritage Site (Peak Wilderness), a local amphibian hotspot in Sri Lanka. Journal of Threatened Taxa 5(4): 3789–3920; doi:10.11609/JoTT.o3099.3789-920

Copyright: © Wickramasinghe et al. 2013. Creative Commons Attribution 3.0 Unported License. JoTT allows unrestricted use of this article in any medium, reproduction and distribution by providing adequate credit to the authors and the source of publication.

Funding: Biodiversity Secretariat of the Ministry of Environment, Nagao Natural Environment Foundation, and Dilmah Conservation.

Competing Interest: None.

Acknowledgments: The authors wish to acknowledge the Biodiversity Secretariat of the Ministry of Environment, Sri Lanka for funding and collaborating the project, Mr. R.H.S.S. Samarathunga (Secretary-Ministry of Environment), Mr. Gamini Gamage (Additional Secretary-Policy Planning), Ms. Padma Abayakoon (Director), Mr. N.K.G.K. Nannawaththa (Additional Secretary-Natural Resources), Mr. R.A.R. Roopasinghe (Aditional Secretary- Administrations), Mr. Ajith de Silva (Former Director), Mrs. Dakshini Perera, and Mrs. Hasula Wickramasinghe for all the support renderd. To the Nagao Natural Environment Foundation for part funding, to the Department of Wildlife Conservation for permission granted (Permit no. WL/3/3/354), the Director General Mr. R.M. Rathnayake, and the Deputy Director Mr. S.R.B. Dissanayake (Research and Training) for all the suppport. Staff of the Nallathanniya and Palabaddala Wildlife beats. Mr. Saman Gamage, Mr. Chaminda Pushpakumara, Mr. Dilshan Maduranga de Silva, Mr. E.A. Thusitha Jayanath, Mr. Lankaputhra Wimaladharma, Mr. L.P.D. Wasantha Kumara, Mr. Kovida Herath, Mr. Jagath Krishantha, Mr. Ruwan Chinthaka are thanked for their immense help in the field, and late Dr. Amith Munindradasa fondly remembered for his support and encouragements. The authors wish to acknowledge the contributions of Mr. Prasanna Samarawickrama, Mr. Chamara Hettiarachchi, in preparing the location illustrations, to Mr. Charith Pubudu Lakmal and Mr. Lalith Senanayake for preparing images. The authors wish to thank the Director National Museum of Sri Lanka, Dr. Nanda Wickramasinghe and staff members (Assistant Director Mrs. Manori Nandasena and Mrs. Manaram de Silva), are gratefully acknowledged for their assistance in museum reference work. Dr. David Gower, Mr. Patrick Campbell, Dr. Barry Clarke, and Dr. Mark Wilkinson of the Natural History Museum, London are gratefully acknowledged for assisting the first author during his visit to the BMNH, also to Mr. Roshan Rodrigo, Mr. Nalaka Aththanayaka and Mr. Naalin Perera for their hospitality extended. Mr. Uditha Hettige, Mr. Sameera Suranjan Karunarathna, Mr. Gayan Chathuranga, Mr. Sanjiv de Silva provided valuable literature relevant, and Dr. Neelesh Dahanukar for doing the statistical analysis for this work are sincerely acknowledged. We wish to thank Mr. Bhathiya Kekulandala for his valuable comments. The authors also like to thank their colleagues at Herpetological Foundation of Sri Lanka (HFS), for various courtesies. Thanks goes to Dr. Sanjay Molur for giving vital suggestions to improve the manuscript, to Ms. Chamila Weerathunga and Mr. Ranil Nanayakkara for proof reading the final manuscript, to Mr. Asanka Abayakoon for the encouragements and to Dr. Annemarie Ohler whose invaluable comments undoubtedly improved the quality of the paper. Finally to our Principal Sponsors Dilmah Conservation, for funding project activities.

Author Details:

L.J. Mendis Wickramasinghe, is the founder, President of the Herpetological Foundation of Sri Lanka, and is the Principal Investigator in the current project. He has close to two decades of field herpetological (reptile and amphibian) experience in Sri Lanka with a focus on taxonomic identifications, and also providing education and awareness to the general public on Venomous Snakes in the island. He has contributed his expertise towards national projects on identification of threatened species in Sri Lanka, and has facilitated the declaration of several protected areas in Sri Lanka. A member of the Experts Committee on herpetofauna in Sri Lanka, under the National Species Conservation Advisory Group (NSCAG), and in several international bodies including several Species Survival Commission groups of the International Union for Conservation of Nature (IUCN/SSC).

Dulan Ranga Vidanapathirana, is the Vice-President, and a founder member of the Herpetological Foundation of Sri Lanka, with over 10 years of field herpetological, and birding experience in Sri Lanka, where he is also a Natural History Tour Leader at the Bird and Wildlife Team. Currently working in several projects on herpetology in the country at the HFS.

M. D. Gehan Rajeev, is working as a Medical Doctor at Base Hospital, Kanthale. Is a naturalist with an interest in writing and photography currently involved in herpetofaunal research at the Herpetological Foundation Sri Lanka, involved in field data gathering. Member of the Young Zoologist’s Association of Sri Lanka, and working to conserve the wildlife in the Island.

S. Chathuranga Ariyarathne, is a member of the Herpetological Foundation of Sri Lanka. A B.Sc graduate from the University of Colombo, also shows keen interest on Chiropterans, in Sri Lanka.

A.W. Amila Chanaka, is the Secretary at the Herpetological Foundation of Sri Lanka. A naturalist, researcher and member of the Young Zoologists’ Association (YZA). He currently works as a Program officer in Wildlife Trust Sri Lanka and Following a BSc. degree at the Open University of Sri Lanka. He has had a Special interest in Amphibians, Reptiles and Land Snails, also having an extensive talent for conducting lectures and presentation to increase knowledge of wildlife and environment conservation.

L.L. Dharshana Priyantha, is a Management Accountant by profession doing Management Consultancy for Small & Medium Entrepreneurs of Sri Lanka. He is working as a researcher cum treasurer of Herpetological Foundation of Sri Lanka. He has also been a member of Young Zoologists’ Association of Sri Lanka, working to conserve the herpetofauna in the Island.

Imesh Nuwan Bandara, is pursuing his Master Degree from the Heller School for Social Policy and Management at Brandeis university, after having obtained the Bachelor of Science (B.Sc) Degree having specialized in Zoology at the University of Peradeniya - Sri Lanka. He has been a member of the Youth Exploration society, Herpetological Foundation of Sri Lanka and has a keen interest in sustainable development, community based conservation, traditional agricultural practices, ethnobotany, local biodiversity and behavioral ecology of herpetofauna and other wild fauna.

Nethu Wickramasinghe, is the Projects Co-ordinator, at the Herpetological Foundation of Sri Lanka, completed the basic degree in chemistry at the University of Delhi. Currently a freelance Science journalist, contributing to the dissemination of the conservation aspects of herpetofauna to the general public.

Author Contributions: LJMW was involved in designing the study, conducting the field survey, conducting museum reference work, gathering, and analyzing the data, and compiling the MS. DRVP was involved in conducting the field survey, conducting museum reference work, gathering, analyzing the data, and writing the MS. MDGR was involved in conducting the field survey, conducting museum reference work, gathering, analyzing the data, and writing the MS. SCA was involved in field research work. AWAC was involved in field research work. LLDP was involved in field research work. INB was involved in field research work, writing and data compiling. NW was involved in the writing of the MS.

For figures, images, tables -- click here

INTRODUCTION

The genus Pseudophilautus consists of 65 known species currently, all of which are endemic to Sri Lanka (Manamendra-Arachchi & Pethiyagoda 2005; Meegaskumbura & Manamendra-Arachchi 2005, 2011; Meegaskumbura et al. 2007, 2009). These direct developing rhacophorids (Bossuyt & Dubois 2001; Meegaskumbura et al. 2002; Bahir et al. 2005; Grosjean et al. 2008; Li et al. 2008, 2009; Biju & Bossuyt 2009; Yu et al. 2009) previously belonged to the genus Philautuswhich were widely distributed throughout tropical Asia (Bossuyt & Dubois 2001; Bossuyt et al. 2004; Frost et al. 2006; Yu et al. 2009). But later, considering molecular phylogenetics, the populations in Sri Lanka and India were found to be a separate radiation belonging to a separate genus known now as Pseudophilautus(Li et al. 2009; Yu et al. 2010; Meegaskumbura & Manamendra-Arachchi 2011). The most recent publication of this genus by Meegaskumbura & Manamendra-Arachchi (2011) in the country described two species which brought the total number to 65 species. But sadly, a considerable portion of the total number of identified species, have already vanished (17 species) (Manamendra-Arachchi & Pethiyagoda 2005; Meegaskumbura et al. 2007) where most have not even been sighted after their initial descriptions, showing an urgency to conserve the few remaining habitats.

Peak Wilderness as part of the Central Hills was recently designated as a World Heritage Site. Geographical area of the sanctuary is about 240sq.km, of which 211.75sq.km comprises natural or semi natural vegetation; the rest is no longer forest and includes tea estates and village settlements. The sanctuary lies in the center of the western ridge of the Central Highlands, north-east of Ratnapura and spanning the border of Central and Sabaragamuwa provinces. Its eastern boundary is contiguous with Horton Plains National Park. Peak Wilderness occupies the escarpment that rises steeply from the lowlands to the south and west. Much of the terrain is very rugged, with altitudes ranging from 600m near Ratnapura to 2,238m at Sripada Peak, Sri Lanka’s fourth highest peak. It is an important watershed for three major river systems, the Kelani Ganga, the Kalu Ganga, and the Walawe Ganga (DWC 2007).

The last vertebrate species discovered from the studied area were Cnemaspis samanalensis (Wickramasinghe & Munindradasa 2007); Lankascincus munindradasai and Lankascincus sripadensis(Wickramasinghe et al. 2007), depicting its diversity.

Because of relatively high altitudes, difficult trails, heavy rainfall, and steep geographic variations, proper field surveys in this region were lacking. Due to this reason inspite of the geographical and climatic variations the region was considered to have a lower faunal diversity. Hence, we carried out the survey with the intention of setting a basis for the actual diversity of herpetofauna of Peak Wilderness. The novel species described here from the Peak Wilderness, Central Hills, of Sri Lanka, as results of the preliminary two phases of an extensive field survey of the region which lasted for a period of two years (end of 2009-end of 2011), were based on morphological and morphometric characters. Sripada Peak is the highest point in this mountain range, and because of its religious importance, and aesthetic beauty, pilgrims and travellers have identified six major trails leading to the top. For the 1^st phase, sampling was carried out in the trail leading from Palabaddala to the Sripada peak, and in the 2^nd phase we chose Kuruvita to the Sripada peak. Our work here describes a collection of eight new species, with each species having a detailed description based on strong morphological characters, accompanied by colour photographs.

MATERIALS AND METHODS

The approximate total distances of the 1^st trail (from Palabaddala to the Sripada peak) was 10km, and the 2^nd trail (from Kuruvita to the Sripada peak) was 10km. The field survey commenced at the end of 2009, and the Phases I and II were completed in December 2011. The sampling sites were selected considering the accessibility, representative habitats and spatial distribution in the National Park, through an initial reconnaissance survey. Fifteen field visits were made, each sampling session spanning eight continuous days. Sampling was done mostly nocturnally, and photographs of most species were done in the wild to avoid any confusion of change in colour after captivity and to avoid over collection of the same species. Specimens collected in the field were first fixed in 90% ethanol for two hours and stored in 70% ethanol.

Sex and maturity were determined by examining secondary sexual characters, or when absent, by examining the gonads through a small lateral incision in the specimen. The following morphological charactetristics were common to all eight species hence they have not been mentioned under each species; nostrils oval, without flap of skin laterally, small, close to tip of snout; latero-dorsal folds, lateral line system, Fejervaryaline, and macroglands were absent. We here have explained the terminology used to explain the skin textures, specially the variations in granular skin on the ventral side of body: weakly granular (Image 1); granular (Image 2); coarsely granular (Image 3). The new type material discussed in this paper is also deposited in NMSL and the Department of Wildlife Conservation (DWC), National Wildlife Research and Training Center, Girithale, Sri Lanka.

The new species were compared with all types from Sri Lanka, deposited in the NMSL and specimens deposited in the Natural History Museum, London (Appendix 4). The specimens that formerly belonged to the Wildlife Heritage Trust (WHT) bearing WHT numbers are currently deposited in the NMSL, catalogued under the same numbers. For the description section, all known species of Sri Lanka were grouped into four categories based on their snout to vent length: very small (<15mm), small (16–30 mm), moderate (31–45 mm) and large (46–60 mm). Comparison for each species was carried out amongst all species belonging to the genus Pseudophilautus, by cancelling common characters and taking the best combination of characters that best distinguishes. In the comparison tables for each species, for convenience of distinguishing the species, an empty cell in the table denotes either the same character as that of the species considered, or for few extinct species when there was no data available. Hence in order to minimise this error we included all characteristics and all species in one table provided in Appendix 1 and 2, by selecting the best combination of characters where Appendix 1, provides morphological characters that differentiate Pseudophilautus species and Appendix 2 provides characters in the skin structure that differentiate Pseudophilautusspecies. We paid special attention to extinct species and their characters in the comparison (Wickramasinghe et al. 2013) of each species.

Forty four external measurements of specimens were taken with a Mitutoyo digital vernier calliper to the nearest 0.1mm. Nomenclature of external anatomy abbreviated in the text and external measurements taken are listed alphabetically herein: Breadth of disk of the third finger (DB), taken from anterior to posterior edge of circum-marginal groove; distance between back of eyes (DBE), measured between posterior edge of eyes; distance between front of eyes (DFE), measured between anterior edges of eyes; length of disk of the third finger (DL), taken from anterior edge of circum-marginal groove to posterior edge of disk; width of disk of the third finger (DW), measured across the inner and outer edges of circum-marginal groove; eye diameter (ED), horizontal diameter of eye; eye to nostril distance (EN), measured between anterior most point of eye and middle of nostril; eye to snout distance (ES), measured between anterior most point of eye and tip of snout; thigh (femur) length (FEL), distance between vent and knee with both thigh and shank flexed; first finger length (FL-1), measured between posterior margin of the most proximal subarticular tubercle or crease of articulation and the tip of first finger; second finger length (FL-2), measured between posterior margin of the most proximal subarticular tubercle or crease of articulation and the tip of second finger; third finger length (FL-3), measured between posterior margin of the most proximal subarticular tubercle or crease of articulation and the tip of third finger; fourth finger length (FL-4), measured between posterior margin of the most proximal subarticular tubercle or crease of articulation and the tip of fourth finger; foot length (FOL), distance between heel and tip of fourth toe with both foot and shank flexed; groin to knee distance (GK), measured between groin and the tip of knee; hand length (HNL), taken from posterior-most margin of inner palmar tubercle to tip of disk of third finger; head depth (HD), distance between the apex of eye and the bottom of jaw; head length (HL), distance between angle of jaws and snout tip; head width (HW), measured across angle of jaws; inner metatarsal tubercle length (IML), distance between the anterior and posterior edges of the inner metatarsal tubercle; internarial distance (IN), least distance between the inner margin of nares; interorbital width (IO), least distance between the upper margins of orbits; knee-angle length (KT), distance from knee-angle to tibio-tarsal articulation; forearm length (LAL), taken from elbow to posterior-most margin of inner palmar tubercle; mandible-back of eye distance (MBE), distance between angle of jaws and posterior-most point of eye; mandible-front of eye distance (MFE), taken as distance between angle of jaws and anterior-most point of eye; mandible-nostril distance (MN), taken as distance between angle of jaws and middle of nostril; snout-nostril distance (SN), taken as distances between middle of nostril and tip of snout; snout-vent length (SVL), measured from tip of snout to vent; tympanum-back of eye distance (TAD), distance between anterior-most point of inner margin of inner rim of tympanum and posterior most point of eye; tarsal length (TAS), measured between the tibio-tarsal articulation and anterior edge of inner metatarsal tubercle; tibia length (TBL), distance between knee and heel with both shank and foot flexed; first toe length (TL-1), measured between posterior margin of the most proximal subarticular tubercle or crease of articulation and the tip of first toe; second toe length (TL-2), measured between posterior margin of the most proximal subarticular tubercle or crease of articulation and the tip of second toe; third toe length (TL-3), measured between posterior margin of the most proximal subarticular tubercle or crease of articulation and the tip of third toe; fourth toe length (TL-4), measured between posterior margin of the most proximal subarticular tubercle or crease of articulation and the tip of fourth toe; fifth toe length (TL-5), measured between posterior margin of the most proximal subarticular tubercle or crease of articulation and the tip of fifth toe; tympanum-nostril distance (TND), distance between anterior-most point of inner margin of inner rim of tympanum and middle point of nostril; tympanum-front of eye distance (TPD), taken as distance between anterior-most point of inner margin of inner rim of tympanum and anterior most point of eye; tympanum height (TYH), vertical diameter of the inner rim; tympanum width (TYW), horizontal diameter of the inner rim; upper arm length (UAW), distance between axilla and elbow; upper eyelid width (UEW), measured from bony edge of supraorbital to outer edge of upper eyelid; vent to knee-angle length (VKL), distance measured from knee-angle to vent.

Morphometric data of both males and females was considered for statistical analysis to understand the morphometric difference among the individuals of eight new species of Pseudophilautus. To account for the size difference among individuals, all the morphometric characters were taken as %SVL. Principle Component Analysis (PCA) was performed on the correlation matrix between 25 variables using the software PAST (Hammer et al. 2001). Dendrogram showing the relationship between different species was plotted using mean values of morphometric data for each species using Euclidian distances and Ward’s method for clustering in PAST.

Geographical coordinates were determined from GPS readings (Gamin eTrex Gista) with WGS84 (World Geodetic System) datum at the locality. Additional data for comparison were collected from Pethiyagoda & Manamendra-Arachchi (2005) and WGS 84 world grid system.

Collectors are indicated by their initials as follows: L.J. Mendis Wickramasinghe (L.J.M.W), Dulan Ranga Vidanapathirana (D.R.V)_, M.D. Gehan Rajeev (M.D.G.R), S. Chathuranga Ariyarathne (S.C.A.), A. W. Amila Chanaka (A.W.A.C.), L.L. Dharshana Priyantha (L.L.D.P.), Imesh Nuwan Bandara (I.N.B.).

RESULTS

Pseudophilautus bambaradeniyai sp. nov.

(Figs. 1–4; Images 4–7; Tables 1–3)

urn:lsid:zoobank.org:act:FB630D8A-7354-4307-A343-5B5343921C73

Holotype: NMSL 2013.01.01 NH, 05.iv.2010, adult male SVL 17.3mm (Images 4, 5A,D), Sripada (Peak Wilderness), Ratnapura District, Sabaragamuwa Province, Sri Lanka (06⁰48’31”N & 80⁰8’14”E; 1334m) (Fig. 1), coll. L.J.M.W, D.R.V., S.C.A. & A.W.A.C.

Paratypes: DWC 2013.01.001, adult male, SVL 19.9mm (Images 5B,E, 6); DWC 2013.01.002, adult male, SVL 20.2mm (Images 5C,F, 7); same data as holotype.

Diagnosis: Pseudophilautus bambaradeniyai sp. nov., can be distinguished from known congeners by the following combination of characters: small size frog (SVL 17.3–20.2 mm); head dorsally convex; canthus rostralis rounded; interorbital space convex; internarial space flat; vomerine teeth, lingual papilla, fringe on fingers, calcar and nuptial pad absent; snout, interorbital area, side of head, upper and lower flank smooth; anterior dorsum prominent tubercles and horny spinules, posterior dorsum horny spinules. Distinguishing characters provided in comparison tables 2 and 3.

Description of holotype: Small sized frog (SVL 17.3mm), elongate (SVL/HW 2.4); head large (HL/SVL 0.5), convex above; snout truncate in lateral aspect, mucronate in dorsal aspect, and sub-elliptical in ventral aspects (Fig. 2 A–C) (ES/DFE 0.8, SN/IN 0.7), larger than horizontal diameter of eye (ES/ED 1.1); internasal space flat; canthus rostralis rounded, loreal region concave; interorbital space convex, larger than upper eyelid (IO/UEW 1.1), and equal to internasal distance (IO/IN 1.0); distance between front of eyes 2/3^rd the distance between back of eyes (DBE/DFE 1.6); nostrils oval, without flap of skin laterally, closer to tip of snout than to eye (SN/EN 0.8); pupil horizontally elliptical; tympanum distinct, oval (TYH/TYW 0.8), smaller than the eye diameter (TYH/ED 0.4), tympanum-eye distance about 2/5^th of tympanum width (TAD/TYW 0.4); pineal ocellus, vomerine teeth and lingual papilla absent, but a depression with a conical tubercle present on tongue; tongue small, lanceolate.

Arm short, thin (LAL/FEL 0.5, UAL/FEL 0.3); forearm shorter than hand length (LAL/HNL 0.9), longer than upper arm (LAL/UAL 1.4); fingers thin, (Fig. 3), relative length of fingers I < II < IV < III (FL-1/FL-3 0.5, FL-2/FL-3 0.7, FL-4/FL-3 0.9) (Table 1), and webbing formula I2½—2 2/3II2^-— 3½III2½—2^-IV; tips of fingers rounded enlarged, discs present on all fingers, with distinct basal and circum marginal grooves; dermal fringes absent; rudimentary webbing present on all fingers; subarticular tubercles prominent, oval, single, all present, III2 and IV2 relatively smaller, supernumerary tubercles present on finger III and IV; inner palmar tubercle indistinct, single, oval, larger than outer palmar tubercle; outer palmar tubercle indistinct, single, oval, smaller than the subarticular tubercles; nuptial pads absent; prepollex absent; femur 2 2/5 times longer than fourth toe length (FEL/TL-4 2.4); foot length longer than thigh (FOL/FEL 1.3); toes thin, (Fig. 4), relative length of toes I < II < III < V < IV (TL-1/TL-4 0.3, TL-2/TL-4 0.4, TL-3/TL-4 0.7, TL-5/TL-4 0.7); webbing formula I2⁺—2^-II2⁺—3^-III2⁺—3^-IV3—2V; tips of toes rounded, enlarged, discs present on all toes with distinct basal and circum marginal grooves; thin dermal fringe present on all toes, thin fringe along postaxial edge of toe V, small tubercles present on outer edge of tarsal fold; subarticular tubercles prominent, oval, single, all present, IV2, IV3, and V2 relatively smaller; supernumerary tubercles present, indistinct, on toe III and IV; inner metatarsal tubercle oval prominent and large, its length 3/4 times in length of toe I (IML/TL-1 0.6); outer metatarsal tubercle indistinct.

Skin of snout dorsally, laterally and interorbital space smooth; upper eyelid tubercular; head laterally smooth; dorsum anterior with prominent tubercles and horny spinules, posterior with prominent horny spinules; upper and lower part of flank smooth; supratympanic fold distinct; upper arm, forearm, and hand smooth; thigh inner and outer smooth, dorsally with tubercles, macroglands absent; leg smooth with few tubercles; calcar absent, small tubercles present at heel; vent dorsally shagreen, either side shagreen.

Ventral side of body: Throat and chest weakly granular; belly granular; upper arm weakly granular to smooth; forearm smooth; thigh, leg and tarsus smooth.

Colour in life: Dorsum dark brown with blackish blotches, large dark brown marking covering much of the anterior, blackish cross band between eyes, a prominent off-white vertebral stripe from the tip of snout to anus, and continuing down the hind limbs symmetrically, lateral body lighter; limbs dark brown, fore limbs, hind limbs, fingers and toes with blackish-brown cross bands; ventral side off white with light brown blotches, hands, feet and webbing lighter (Image 4).

Colour in alcohol: Colour pattern remains with a little fading, off-white vertebral stripe prominent on dark brown, ventral side off-white all over (Images 5 A,D).

Paratype and variation: Dorsum colour varies ranging from orange to a lighter brownish tinge and the thickness of the vertebral line varies; in some this line discontinues towards limbs (Images 5 B,E, 6).

Etymology: The species epithet bambaradeniyai is named after Channa Bambaradeniya, a leading wetland scientist who has been working immensely to protect the wetlands and its ecosystems in Sri Lanka, in great appreciation of his dedicated efforts towards the conservation of the biodiversity at large in the country. The species name bambaradeniyai is a noun in the genitive case.

Suggested vernacular names: Bambaradeniyage panduru madiya, and Bambaradeniya’s Shrub Frog in Sinhala and English, respectively.

Natural history: The species was found in elevations of 750–1400 m from the forest floor to about 1.5m high shrubs in lowland rain forests to lower montane rain forests.

Pseudophilautus dayawansai sp. nov.

(Figs. 5–7; Images 8–12; Tables 4–6)

urn:lsid:zoobank.org:act:3D56DDDE-770F-4D0E-BBE3-04C357922CDE

Holotype: NMSL 2013.02.01 NH, 23.x.2011, adult male SVL 24.5mm (Images 8, 9A,D). Sripada (Peak Wilderness), Ratnapura District, Sabaragamuwa Province, Sri Lanka (06⁰48’30”N & 80⁰29’19”E; altitude 1679m) (Fig. 1), coll. L.J.M.W, D.R.V., M.D.G.R., S.C.A., A.W.A.C. & L.L.D.P..

Paratypes: DWC 2013.01.003, 14.xi.2011, adult female, SVL 30.1mm (Image 10); DWC 2013.01.004, 14.xi.2011, adult male, SVL 26.2mm (Image 11); information same as holotype.

Diagnosis: Small sized frog (adult male SVL 24.5–26.2 mm, adult female SVL 30.1mm); head dorsally convex; snout laterally truncate, skin dorsally tuberculated, a single prominent tubercle medially placed; canthus rostralis rounded; loreal region, interorbital space and internarial space concave; vomerine teeth absent; lingual papilla, fringe on fingers and blunt small calcar present; interorbital area smooth, with a ridge across braking medially; side of head tubercular; anterior dorsum smooth; posterior dorsum weakly tubercular; median dermal ridge absent; upper and lower flank granular; throat shagreen.

Description of holotype: Small sized frog (SVL 24.5mm), elongate (SVL/HW 2.3); head large (HL/SVL 0.4), about as wide as long (HW/HL 1.0), dorsally convex; snout truncate in lateral aspect, mucronate in dorsal aspect, and sub-elliptical in ventral aspect (Fig. 5 A) (ES/DFE 0.8, SN/IN 0.5); internasal space concae; canthus rostralis rounded; loreal region concave; interorbital space mid dorsally concave, upper eyelid smaller than interorbital distance (IO/UEW 1.2), smaller than internasal distance (IN/UEW 1.2); distance between front of eyes half its distance between back of eyes (DBE/DFE 1.9); nostrils oval without flap of skin laterally, closer to tip of snout than to eye (SN/EN 0.7); pupil horizontally elliptical; tympanum distinct, oval and vertical (TYH/TYW 1.2), tympanum smaller than the eye diameter (TYH/ED 0.3), tympanum-eye distance 1/3 of tympanum diameter (TAD/TYW 0.8); pineal ocellus absent; vomerine ridge absent; tongue moderate, lanceolate; lingual papilla present, blunt, depressed, rounded, and flattened, with conical tubercles present on tongue.

Arm short, thin (LAL/FEL 0.5, UAL/FEL 0.4); lower-arm shorter than hand length (LAL/HNL 0.8), longer than upper arm (UAL/LAL 0.8); fore-arm distinctly enlarged; fingers thin, (Fig. 6), relative length of fingers I < II < IV < III (FL-1/FL-3 0.4, FL-2/FL-3 0.5, FL-4/FL-3 0.8) (Table 4); tips of fingers rounded enlarged; discs present on all fingers, with distinct basal and circum marginal grooves; dermal fringe present on inside of all fingers; fringe on external edge of finger IV and palm; ulnar fold indistinct but few small undulating tubercles present; webbing formula I2^-—2 ½II2^-—3^-III3—2 ½IV; subarticular tubercles prominent, oval, single, all present, III2 and IV2 relatively smaller; inner palmar tubercle single, oval, larger than outer palmer tubercle; outer palmar tubercle single, indistinct; nuptial pads absent; supernumerary tubercles present on all fingers, and a few indistinct ones on palm; femur twice as long as the fourth toe length (FEL/TL-4 2.1), foot length longer than thigh (FOL/FEL 1.5), toes long, thin (Fig. 7); relative length of toes I <II <III <V <IV (TL-1/TL-4 0.4, TL-2/TL-4 0.4, TL-3/TL-4 0.6, TL-5/TL-4 0.8); tips of toes rounded, discs present on all toes, with distinct basal and circum marginal grooves; webbing formula I2—2^-II2⁺—3⁺III1 ½—2 ½IV2^-—1⁺V; lateral dermal fringe present on both sides of all toes, fringe along exterior edge of toe V and metatarsal fold; fringe absent on tarsus but few small undulating tubercles present; subarticular tubercles prominent, oval, single, all present, III3 relatively smaller; supernumerary tubercles present on toe III, IV and V, becomes indistinct after preservation; inner metatarsal tubercle oval prominent and large, its length 3/5 times the length of toe I (IML/TL-1 0.6); outer metatarsal tubercle indistinct; tarsal with a single blunt small calcar and a few blunt small tubercles present at heel.

Skin between eyes smooth with two symmetrically placed ridges that do not meet mid dorsally; dorsal and lateral snout tubercular, a prominent tubercle present dorsally on middle of snout; upper eyelid bearing few horny tubercles with one being more prominent; lateral head tubercular with prominent tubercle present near the gape of mouth; a median dermal ridge absent; anterior part of dorsum smooth, posterior part weakly tubercular; upper and lower part of flank granular; supratympanic fold prominent; upper arm shagreen; forearm weakly tubercular; hand shagreen; inner, outer, and dorsal thigh smooth; dorsally leg weakly tubercular; tarsus weakly tubercular; foot smooth.

Ventral side of body: Throat shagreen; chest weakly granular; belly granular; upper arm weakly granular; forearm smooth; thigh weakly granular; leg smooth.

Colour in life: Dorsally brown with dark brown blotches; much of the dorsum covered with a dark brown marking, forming a clear semicircular blackish margin anteriorly, its center with a large light brown triangle, prominent, symmetrically placed on dorsum with the base on sacrul hump pointing towards head, its margins blackish-brown and outer area dark brown, three prominent blackish-brown circular spots, two between eyes, and right in front another circular dark brown spot, posterior part from sacrum light brown; lateral body, and behind eyes dark brown with an olive green tinge; limbs dark brown, forelimbs, hind limbs, fingers and toes with maroon cross bands, hind limbs laterally olive green tinge; ventrally light brown with dark brown blotching, hands, feet and webbing darker.

Colour in alcohol: Colour pattern remains with a little fading, maroon to dark brown, venter light brown to off-white with darker marbling, green tinge absent.

Paratype and variation: Dorsum colour varies ranging from a reddish-brown (Image 10) to a lighter brown shades (Image 12); in the lighter coloured specimens the dorsal colour of foot and hand light yellowish, and limbs were cream with brown cross bands.

Etymology: The species epithet dayawansai is named after Nihal Dayawansa, Senior Lecturer of the Department of Zoology, University of Colombo in recognition of his dedicated service and commitment towards amphibian research work in Sri Lanka. The species name dayawansai is a noun in the genitive case.

Suggested vernacular names: Dayawansage panduru madiya in Sinhala and Dayawansa’s Shrub Frog in English.

Natural History: The species was found in cloud forests of elevation 1550–1900 m above sea level from forest floor to about 2m high shrubs.

Remarks: Pseudophilautus dayawansai sp. nov., possesses a unique colour pattern which is not possessed by any of its congeners, hence can easily be identified. In Dutta & Manamendra-Arachchi 1996: “Fig. 232. Rhacophorus macropus MCZ 20873 Queenwood, Dimbula, Sri Lanka” in page 194, closely ressembles Pseudophilautus dayawansai from its unique mark on head; dark brown marking, forming a clear semicircular blackish margin. Rhacophorus macropus (Pseudophilautus macropus) is restricted to the Knuckles Range (Manamendra-Arachchi & Pethiyagoda 2005), but the specimen’s locality Dimbula is closer to the Peak Wilderness, hence given the geographical similarity and the unique marking we feel that this specimen could be Pseudophilautus dayawansai.

Pseudophilautus jagathgunawardanai sp. nov.

(Figs. 8–10; Images 13–20; Tables 7–9)

urn:lsid:zoobank.org:act:A5A913CB-9D12-4024-9776-F8E904F3887E

Holotype: NMSL 2013.03.01 NH, 05.ii.2009, adult female 35.8mm SVL (Image 13, 14A,D). Sripada (Peak Wilderness), Ratnapura District, Sabaragamuwa Province, Sri Lanka (06⁰48’31”N & 80⁰29’10”E; 1642m) (Fig. 1), coll. L.J.M.W, D.R.V., M.D.G.R., S.C.A., A.W.A.C., L.L.D.P. & I.N.B.

Paratypes: DWC 2013.01.005, adult female, SVL 40.4mm (Image 14 B&E); DWC 2013.01.006, adult female, SVL 36.3 mm (Image 14 C&F, 15); data same as holotype.

Diagnosis: Pseudophilautus jagathgunawardanai sp. nov., can be distinguished from known congeners by the following combination of characters: Body moderate size (SVL adult female 35.8–40.4 mm); head dorsally, loreal region, and interorbital space convex; lateral snout rounded; canthus rostralis sharp; internarial space concave; vomerine teeth, fringe on fingers, rudimentary webbing on fingers, calcar, and nuptial pad absent; lingual papilla present; snout, side of head, anterior dorsum, posterior dorsum, upper flank smooth; interorbital area smooth, with two cross dermal fringes; throat and chest weakly granular; underside of thigh granular; supernumerary tubercles on palm and foot present. Comparative details between congeners in tables 8 and 9.

Description of holotype: Body moderate size (SVL 35.8), elongate (SVL/HW 2.2); head large (HL/SVL 0.5), as wide as long (HW/HL 1.0), convex above; snout rounded in lateral and ventral aspects, truncate in dorsal aspect (Fig. 8 A–C) (ES/DFE 0.8, SN/IN 0.6) its length slightly longer than the horizontal diameter of eye (ES/ED 1.1); internasal space concave; canthus rostralis sharp, loreal region and interorbital space convex, upper eyelid smaller than interorbital distance (IO/UEW 1.2), internasal distance equal to upper eyelid (IN/UEW 1.0); distance between front of eyes almost half the distance between back of eyes (DBE/DFE 1.9); nostrils oval without flap of skin laterally, closer to tip of snout than to eye (SN/EN 0.6); pupil horizontally elliptical; tympanum distinct, oval (TYH/TYW 1.4), one third the eye diameter (TYH/ED 0.3), tympanum-eye distance equal to tympanum width (TAD/TYW 1.0); pineal ocellus and vomerine teeth absent; tongue large, lanceolate, lingual papilla present, rounded and blunt with several conical tubercles present evenly distributed on tongue.

Arm short, thin (LAL/FEL 0.5, UAL/FEL 0.4); forearm shorter than hand length (LAL/HNL 0.8); forearm longer than upper arm (LAL/UAL 1.4); fore-arm distinctly enlarged; fingers thin, (Fig. 9), relative length of fingers I < II < IV < III (FL-1/FL-3 0.5, FL-2/FL-3 0.6, FL-4/FL-3 0.7); tips of fingers rounded enlarged, discs present on all fingers, with distinct basal and circum marginal grooves; no dermal fringe on fingers, but indistinct dermal fringe present on exterior edge of finger IV; webbing formula I3⁺—3⁺II2—3^-III3⁺—3⁺IV; subarticular tubercles prominent, rounded or oval, single, IV2 very small and indistinct; inner palmar tubercle single, oval, prominent, larger than outer palmer tubercle; nuptial pads absent; outer palmar tubercle single, oval, indistinct, smaller than the distal subarticular tubercles; supernumerary tubercles present on all fingers, and palm; prepollex absent; femur twice as long as the fourth toe length (FEL/TL-4 2.1); foot length longer than thigh (FOL/FEL 1.5); toes thin, (Fig. 10), relative length of toes I < II < V < III <IV (TL-1/TL-4 0.4, TL-2/TL-4 0.4, TL-3/TL-4 0.7, TL-5/TL-4 0.5) (Tables 7); tips of toes rounded, enlarged, discs present on all toes with distinct basal and circum marginal grooves; webbing formula I2—2^-II2⁺—3III2⁺—3^-IV2^—2⁺V; dermal fringe on all toes and fringe along exterior edge of toe V present; fringe absent on metatarsus and tarsus but tubercles present; subarticular tubercles prominent, rounded, single, all present, IV3, and V2 relatively smaller; supernumerary tubercles concentrated on foot, present on all toes; inner metatarsal tubercle oval prominent and large, its length 2/3 times the length of toe I (IML/TL-1 0.6); outer metatarsal tubercle absent.

Skin on dorsal and lateral part of snout smooth; interorbital space smooth, with two cross dermal fringes one connecting the front of eyes and the other placed behind it, a faint median dermal ridge from tip of snout to vent; upper eyelid bearing a prominent tubercle and a few intermediate sized tubercles on top; head laterally smooth a few prominent tubercle cluster behind gape of mouth, dorsal body smooth, except behind eyes which is shagreen; upper part of flank smooth; lower part of flank weakly granular; supratympanic fold prominent; upper arm smooth; forearm weakly tubercular; few tubercles present on hand; inner and outer thigh smooth, dorsally weakly tubercular; leg dorsally with tubercles, a small calcar like tubercle at heel; tarsus bearing indistinct tubercles; foot weakly tubercular.

Ventral side of body: Throat and chest weakly granular; belly granular; forearm and upper arm weakly granular; thigh granular; leg smooth; tarsus with prominent tubercles.

Colour in life: Dorsum light brown with a green tinge, a dark brown circle on snout and a broad dark brown band connecting upper eyelids, dark brown triangular patch pointed towards head on anterior dorsum, another broad dark brown cross band on sacrum, side olive green; limbs light brown with a green tinge, fore limbs, hind limbs, fingers and toes with dark brown cross bands, finger tips off white; ventral side off white, throat and chest marbled with brown blotches, belly, hands, feet and webbing darker (Image 13).

Colour in alcohol: Colour darkens, pattern remains, light brown with green tinge changes to dark brown and dark brown patches to maroon, and ventrally off white colour changes to a light brown (Image 14 A&D).

Paratype and variation: The species showed quite a variation of dorsal colour depending on its surrounding environment, ranging from maroonish tinges to light brown to cream colour, where they lacked the prominent greenish tinge found on holotype. Markings and patterns remained the same even with change in colour (Image 16–20).

Etymology: The species epithet jagathgunawardanai is named after Jagath Gunawardana, a leading attorney at law and naturalist, in great appreciation of his services towards the conservation of Sri Lanka’s biodiversity and for his efforts on environmental policy making in the island. The species name jagathgunawardanaiis a noun in the genitive case.

Suggested vernacular names: Jagath Gunawardanage panduru madiya and Jagath Gunawardana’s Shrub Frog in Sinhala and English, respectively.

Natural History: The species is found mainly in cloud forests of elevation 1600–1750 m from forest floor to about 6m high trees. They are commonly observed on tree trunks whith lichens, or were hidden on crevises of barks, or on shrubs.

Remarks: Pseudophilautus jagathgunawardanai sp. nov. is not confused with its congeners, due to its unique patterns and markings on dorsum.

Pseudophilautus karunarathnaisp. nov.

(Figs. 11–13; Images 21–25; Tables 10–12)

urn:lsid:zoobank.org:act:0B077FB6-CC9D-48AA-8AC8-D5822D4BC0CA

Holotype: NMSL 2013.04.01 NH, 03.iv.2010, adult male 16.2mm SVL (Images 21, 22A,D). Sripada (Peak Wilderness), Ratnapura District, Sabaragamuwa Province, Sri Lanka (06⁰48’31”N & 80⁰28’14”E; 1334m) (Fig. 1), coll. L.J.M.W, D.R.V., M.D.G.R., S.C.A. & A.W.A.C..

Paratypes: DWC 2013.01.007, adult female, 19.2mm SVL (Images 22 B,E, 23); DWC 2013.01.008, adult male, 16.3mm (Images 22 C,F, 24); the same date, locality and collectors.

Diagnosis: Pseudophilautus karunarathnai sp. nov. can be distinguished from known congeners by the following combination of characters: body rather small (males SVL 16.2–16.3 mm, females 19.2mm); dorsal head and Interorbital space convex; lateral snout, canthus rostralis rounded; loreal region and internarial space concave; lingual papilla and calcar absent; vomerine teeth and nuptial pad present; snout smooth, a prominent blunt tubercle present; interorbital area weakly tubercular; anterior dorsum and dorsal thigh smooth; posterior dorsum spinulate; upper flank weakly granular; lower flank, throat and chest granular; supernumerary tubercles on palm and foot absent. Comparative characters that distinguish the species from its congeners are provided in tables 11 and 12.

Description of holotype: Body rather small (SVL 16.2), moderately elongate (SVL/HW 2.1); head large (HL/SVL 0.5), wider than long (HW/HL 0.9), convex above; snout rounded in lateral aspect, mucronate in dorsal and sub-elliptical in ventral aspects (Fig. 11 A–C) (ES/DFE 0.8, SN/IN 0.6), longer than the horizontal diameter of eye (ES/ED 1.1); internasal space concave; canthus rostralis rounded, loreal region concave; interorbital space convex, larger than upper eyelid (IO/UEW 1.1), equal to internasal distance (IN/UEW 1.1); distance between front of eyes 3/5^th the distance between back of eyes (DBE/DFE 1.7); nostrils oval, without flap of skin laterally, closer to tip of snout than to eye (SN/EN 0.7); pupil horizontally elliptical; tympanum distinct, oval (TYH/TYW 1.4), smaller than the eye diameter (TYH/ED 0.4), tympanum-eye distance 2/3^rd tympanum width (TAD/TYW 0.7); pineal ocellus absent; vomerine teeth present, small, few, odontophores oblique and widely separated, between choanae with an angle of 60° relative to body axis; tongue large, lanceolate; lingual papilla absent, depression with conical tubercles present on tongue.

Arm short, thin (LAL/FEL 0.49, UAL/FEL 0.3); forearm shorter than hand length (LAL/HNL 0.9); forearm longer than upper arm (LAL/UAL 1.5); fingers thin, (Fig. 12), relative length of fingers I < II < IV < III (FL-1/FL-3 0.5, FL-2/FL-3 0.6, FL-4/FL-3 0.9) (Table 10); tips of fingers rounded enlarged, discs present on all fingers, with distinct basal and circum marginal grooves; dermal fringe present on fingers; webbing formula I2—2 1/3 II2^—3 2/3III3⁺—2 ½IV; subarticular tubercles prominent, rounded, single, all present, IV2 and III2 relatively smaller; inner palmar tubercle indistinct, single, oval, larger than outer palmer tubercle; nuptial pads present; outer palmar tubercle indistinct, single, oval, smaller than the distal subarticular tubercles; supernumerary tubercles absent on fingers indistinct on palm; prepollex absent; femur 1 1/3 times longer than fourth toe length (FEL/TL-4 2.3); foot length longer than thigh (FOL/FEL 1.2); toes thin, (Fig. 13), relative length of toes I < II < III < V < IV (TL-1/TL-4 0.4, TL-2/TL-4 0.5, TL-3/TL-4 0.7, TL-5/TL-4 0.7); tips of toes rounded, enlarged, discs present on all toes with distinct basal and circum marginal grooves; webbing formula I2—2^-II2⁺—3⁺III2⁺—3^-IV2 ½—2⁺V; dermal fringe present on inside of toes III, IV and V, thin fringe along exterior edge of toe V, and absent on metatarsus and tarsal fold; subarticular tubercles prominent, rounded, single, all present, V2 and IV3 relatively smaller; supernumerary tubercles absent; inner metatarsal tubercle oval prominent and large, its length half the length of toe I (IML/TL-1 0.5); outer metatarsal tubercle present, smaller than penultimate subarticular tubercles.

Skin of snout dorsally smooth, a large, prominent, blunt tubercle present, snout laterally weakly tubercular; interorbital space and upper eyelid weakly tubercular; head laterally weakly tuberculated, a faint median dermal ridge intermittent with tubercles extends from tip of snout to back of the head; dorsum anteriorly smooth, posteriorly spinulate; upper part of flank weakly granular; lower part of flank granular; supratympanic fold distinct; upper arm, forearm and hand smooth; inner, outer and dorsal thigh smooth; leg and tarsus smooth; small calcar like tubercle present at heel; foot smooth.

Ventral side of body: Throat, chest and belly granular; upper arm proximally granular, distally smooth; forearm smooth; thigh granular; leg and tarsus smooth.

Colour in life: Dorsum uniform cream, with dark brown blotches, dark brown cross band between eyes, and another faint cross band on snout, broad dark brown cross band on sacral hump; laterally head dark brown with a light brown unprominent spot, dark brown band below supratympanic fold; limbs cream colour, fore limbs, hind limbs, fingers and toes with dark brown cross bands; ventral side off-white, hands blotching all over, belly, and hands lighter, feet and webbing dark brown (Image 21).

Colour in alcohol: Colour pattern remains with a little fading. All dark brown patches more prominent in off white, ventral side off-white all over (Image 22 A & D).

Paratype and variation: Apart from the slightly dark brownish dorsum colouration all specimens observed had similar colour not much variation seen (Image 23–25).

Etymology: The species name karunarathnai is named after Y.G.P. Karunarathna (retired Assistant Director, Department of Wildlife Conservation, Sri Lanka), in great appreciation for his commendable services to wildlife management which has resulted in the minimizing of the human-wildlife conflict on the island. The species name is a noun in the genitive case.

Suggested vernacular names: Karunarathnage panduru madiya and Karunarathna’s Shrub Frog in Sinhala and English, respectively.

Natural history: The species is found in elevations of 750–1400 m, from the forest floor to about 1.5m high shrubs in lowland rain forests and lower montane rain forests. Apart from forests they are also observed in home gardens without much canopy cover.

Remarks: Pseudophilautus karunarathnai sp. nov. possesses a unique colour pattern which is not possessed by any of its congeners, hence can easily be identified.

Pseudophilautus newtonjayawardanei sp. nov.

(Figs. 14–16; Images 26–28; Tables 13–15)

urn:lsid:zoobank.org:act:54DAD829-148A-47C4-BCD4-92B994D8E581

Holotype: NMSL 2013.05.01 NH, 05.ii.2010, adult female 38.4mm SVL (Images 26–27). Sripada (Peak Wilderness), Ratnapura District, Sabaragamuwa Province, Sri Lanka (06⁰48’36”N & 80⁰28’59”E; 1559m) (Fig. 1), coll. L.J.M.W, D.R.V., M.D.G.R., S.C.A. & A.W.A.C.

Diagnosis: Pseudophilautus newtonjayawardanei sp. nov. can be distinguished from known congeners by the following combination of characters: moderate size (SVL 38.4); dorsal head, interorbital space and internarial space concave; lateral snout, canthus rostralis and tips of toes rounded; vomerine teeth, lingual papilla, calcar, nuptial pad, and tympanum absent; fringe on fingers present; tarsal tubercle present; snout, side of head, anterior and posterior dorsum, upper flank and dorsal thigh shagreen; interorbital area shagreen, with a prominent ridge; lower flank weakly granular; throat and underside of thigh granular; supernumerary tubercles present on palm. Comparative characters that distinguish the species from its congeners are provided in tables 14 and 15.

Description of holotype: Body moderate size (SVL 38.4), elongate (SVL/HW 2.3); head large (HL/SVL 0.4), about as wide as long (HW/HL 1.0), concave above; snout rounded in lateral, dorsal and ventral aspects (Fig. 14 A–C) (ES/DFE 0.7, SN/IN 0.6), its length equal to horizontal diameter of eye (ES/ED 1.0); internasal space concave; canthus rostralis rounded, loreal region concave; interorbital space concave, upper eyelid smaller than interorbital distance (IO/UEW 1.3); internasal distance less than upper eye width (IN/UEW 0.9); distance between front of eyes half the distance between back of eyes (DBE/DFE 2.0); nostrils oval without flap of skin laterally, closer to tip of snout; pupil horizontally elliptical; tympanum absent; pineal ocellus and vomerine teeth absent; tongue large, lanceolate, lingual papilla absent, but conical tubercles present on tongue.

Arm short, robust (LAL/FEL 0.6, UAL/FEL 0.4); forearm shorter than hand length (LAL/HNL 0.9), longer than upper arm (LAL/UAL 1.4); fore arm distinctly enlarged.; fingers robust, (Fig. 15), relative length of fingers I < II < IV < III (FL-1/FL-3 0.5, FL-2/FL-3 0.6, FL-4/FL-3 0.9) (Table 13); tips of finger disks semi circular, enlarged, discs present on all fingers, with distinct basal and circum marginal grooves; dermal fringe present on inside of all fingers, but indistinct dermal fringe present on exterior edge of finger IV and hand, absent on ulnar fold; webbing formula I2^-—2II2^-—3⁺III3—3IV; subarticular tubercles prominent, rounded, single, all present, IV2 and III2 relatively smaller; inner palmar tubercle, large, single, oval, prominent; outer palmar tubercle single, oval, distinct, larger than the distal subarticular tubercles; supernumerary tubercles prominent and widespread on palm and on all fingers; prepollex absent; femur 2 1/4 times longer than fourth toe length (FEL/TL-4 2.3); foot length longer than thigh (FOL/FEL 1.5); toes thin, (Fig. 16), relative length of toes I < II < III <V <IV (TL-1/TL-4 0.4, TL-2/TL-4 0.5, TL-3/TL-4 0.7, TL-5/TL-4 0.8), tips of toes rounded, enlarged, discs present on all toes with distinct basal and circum marginal grooves; webbing formula I2—2II2⁺—3^-III2⁺—3^-IV2—1⁺V; lateral dermal fringe on inside of all toes and undulating fringe along exterior edge of toe V and metatarsal fold, fringe absent on tarsus but tubercles present; subarticular tubercles prominent, rounded and single, IV3 relatively small; supernumerary tubercles prominent, wide spread and concentrated on foot and all toes; inner metatarsal tubercle oval prominent and large, its length 2/3^rd the length of toe I (IML/TL-1 0.6); outer metatarsal tubercle absent.

Skin on dorsal and lateral parts of snout shagreen, with four tubercles; interorbital space shagreen with a prominent ridge; upper eyelid shagreen with prominent tubercles; head laterally shagreen, a prominent tubercle present near the gape of mouth; a median dermal ridge extends from tip of snout to vent (more prominent till back of head); dorsum shagreen; upper part of flank shagreen; lower part of flank weakly granular; supratympanic fold prominent; upper arm, forearm and hand shagreen; inner thigh smooth, dorsally shagreen, outer thigh weakly granular; leg shagreen; many prominent tubercles at heel; tarsal fold absent but tarsus shagreen with prominent tubercles; foot shagreen.

Ventral side of body: Throat, chest and belly granular; upper arm granular; forearm weakly granular; thigh granular, leg smooth; tarsus weakly granular.

Colour in life: Dorsum bronze with dark brown patches, vertebral stripe off white, a pair of broad dark brown longitudinal dorsal bands extends from the back of the eye to the groin, dark brown cross band between eyes, small bronze blotch on snout, dark brown band on canthal edge, below eye bronze patch, lateral body lighter with a green tinge dark brown spots marbled with off white spots; limbs dorsally bronze, fore limbs, hind limbs, fingers and toes with brown cross bands, finger tips off white; ventral side off white all over, hands feet and webbing darker (Images 26 & 28).

Colour in alcohol: All bronze in life turns to dark brown, and the colour pattern remains with a little fading (Image 27).

Etymology: The species epithet newtonjayawardanei named after Newton Jayawardane, a leading surgeon by profession and is an ardent naturalist, in great appreciation of his dedicated service to the country, both as a surgeon and a wildlife conservationist.

Suggested vernacular names: Newton Jayawardanege panduru madiya and Newton Jayawardane’s Shrub Frog in Sinhala and English, respectively.

Natural history: The specimen was observed in the forest canopy at very high altitudes with elevations ranging fron 1800–2000 m and was very rare. Only two specimens have been observed so far, from about 8m high trees. They perch on leaves and are very hard to sight.

Pseudophilautus puranappu sp. nov.

(Figs. 17–19; Images 29–34; Tables 16–18)

urn:lsid:zoobank.org:act:2961FA3C-4696-4A1D-B38C-1DE1A86D741D

Holotype: NMSL 2013.06.01 NH, 21.xi.2009, adult male 34.2mm SVL (Images 29, 30A,D). Sripada (Peak Wilderness), Nuwara Eliya District, Central Province, Sri Lanka (06⁰ 48’26”N & 80⁰29’58”E; 2100m) (Fig. 1), coll. L.J.M.W, D.R.V. & S.C.A.

Paratypes: DWC 2013.01.009, 14.xi.2011, adult male, 33.6mm SVL (Images 30 B,E, 31); DWC 2013.01.010, adult female, 51.1mm (Images 30C,F, 32); the same locality, coll. D.R.V. & S.C.A.

Diagnosis: Pseudophilautus puranappu sp. nov. can be distinguished from known congeners by the following combination of characters: Moderate size (males SVL 33.6–34.2 mm; female 51.1mm); dorsal head, loreal region and internarial space concave; lateral snout and canthus rostralis rounded; interorbital space flat; vomerine teeth, lingual papilla and nuptial pad present; fringe on fingers and calcar absent; snout smooth, with a blunt tubercle; interorbital area and side of head smooth; anterior dorsum and dorsal thigh weakly shagreen; posterior dorsum few horny spinules, prominent tubercles; upper flank weakly tubercular; lower flank granular; throat, chest and underside of thigh weakly granular; supernumerary tubercles on foot indistinct. Comparative characters that distinguish the species from its congeners are provided in tables 17 and 18.

Description of holotype: moderate size (SVL 34.2), body moderately elongate (SVL/HW 2.6); head large (HL/SVL 0.4), concave above; snout rounded in lateral, truncate in dorsal and nearly rounded in ventral aspects (Fig. 17 A–C) (ES/DFE 0.7, SN/IN 0.7), its length shorter than horizontal diameter of eye(ES/ED 0.9); canthus rostralis rounded; loreal region concave; interorbital space flat, less than upper eyelid (IO/UEW 0.9), larger than internasal distance (IO/IN 1.2), internarial space concave; distance between front of eyes 3/5^th the distance between back of eyes (DBE/DFE 1.7); pupil horizontally elliptical; tympanum prominent, oval (TYH/TYW 1.2), vertical, smaller than the eye (TYH/ED 0.3); tympanum-eye distance 4/5^th tympanum width (TAD/TYW 0.8); pineal ocellus absent; vomerine teeth present, (N=7) prominent, odontophores oblique and widely separated, between choanae with an angle of 65° relative to body axis; tongue large, tip bifurcate, lanceolate, lingual papilla and conical tubercles present on tongue.

Arm short (HNL/LAL 1.1); lower-arm longer than upper arm (UAL/LAL 0.7); fore-arm distinctly enlarged; fingers long and thin (Fig. 18), relative length of fingers I < II <IV <III (FL-1/FL-3 0.4, FL-2/FL-3 0.7, FL-4/FL-3 0.9) (Table 16), tips of fingers rounded, enlarged; discs present on fingers I – II – III – IV, with distinct basal and circum marginal grooves; dermal fringe absent on inside of fingers, but indistinct on exterior edge of finger IV and hand; ulnar fold absent; webbing formula I3⁺–2 ½II3⁺–2 2/3^rd III2 2/3 – 2 ½IV; subarticular tubercles prominent, rounded, single, all present, IV2 & III2 relatively smaller; inner palmar tubercle single, oval, prominent, equal to outer palmer tubercle; nuptial pads present; outer palmar tubercle single, oval, and prominent; supernumerary tubercle present on all fingers; prepollex absent; femur about twice as long as the fourth toe length (FEL/TL-4 2.2); foot length longer than thigh (FOL/FEL 1.5); toes long (Fig. 19), thin; relative length of toes I <II <III <V <IV (TL-1/TL-4 0.4, TL-2/TL-4 0.5, TL-3/TL-4 0.7, TL-5/TL-4 0.8); tips of toes rounded, discs present on all toes, with distinct basal and circum marginal grooves; webbing formula I2 – 2^-II2+ –2 ½III2⁺–3^-IV3⁺ – 2^-V; lateral dermal fringe on inside of all toes; fringe along exterior edge of toe V and metatarsal fold; fringe absent on tarsus but few small undulating tubercles present; subarticular tubercles prominent, rounded, single, all present, V2, IV3, and III2 relatively smaller; supernumerary tubercles indistinct; inner metatarsal tubercle oval prominent and large, its length about 1/2 times in length to that of toe I (IML/TL-1 0.50), outer metatarsal tubercle absent.

Dorsal snout smooth with a blunt tubercle, lateral snout smooth; between eyes smooth; upper eyelid tubercular; head laterally smooth with prominent tubercles present near the gape of mouth; supratympanic fold prominent; a median dermal ridge extends from tip of snout to back of the head, with tubercles; back of head with a large prominent tubercle, and a pair of prominent tubercles on either side of the ridge end; anterior part of back weakly shagreen, few horny spinules, posterior part of back with few horny spinules and prominent tubercles; upper part of flank weakly tubercular, lower part of flank granular; upper arm smooth; forearm weakly tubercular; hand smooth with few tubercles; inner thigh smooth, dorsal part of thigh weakly shagreen, outer thigh smooth; leg tuberculated; tarsal fold absent but tarsus weakly tuberculated; calcar absent but small tubercles present at heel; foot smooth with few tubercles present on fifth toe.

Ventral side of body: Throat and chest weakly granular, belly granular; upper arm and forearm weakly granular; thigh weakly granular, leg smooth, tarsus weakly granular.

Colour in life: Dorsum uniform light brown with a maroon longitudinal band covering snout to vent (hourglass-shaped), mid dorsally colour discontinues, and gets darker down to vent, a faint strip between eyes demarcating a fairly lighter triangular region on snout, canthal edge and tip of snout light brown, laterally lighter with a green tinge; limbs dark brown with a green tinge, forelimbs, hindlimbs, fingers and toes with blackish maroon bars; ventral side off-white marbled with dark brown, hands, feet and webbing lighter (Image 29).

Colour in alcohol: All maroon in life changed to a blackish tinge, the colour pattern remains with fading (Image 30 A & D).

Paratype and variation: Specimens showed dorsal colour variations. Mainly the hour glass maroonish broad patch remains the same in all and only in the holotype the patch discontinued, but the colouration outside this patch on either side of body changes ranging from pink, and some instances to a yellowish tinge (Image 31–34).

Etymology: The species epithet puranappu is named after Veera Puran Appu (Veerahennadige Francisco Fernando, 1812–1848) Puran Appu was a leader who stood up against the might of the British Empire, and died a courageous death, named for his sacrifice for freedom of Sri Lanka. The specific name is an invariable noun in apposition to the generic name.

Suggested vernacular names: Puran Appuge panduru madiya and Puran Appu’s Shrub Frog in Sinhala and English, respectively.

Natural history: The species was observed even in day time, commonly seen found perched on rocks in shady places during the day and nocturnally they may be observed perched on 1m high bushes. Mostly seen in places where there was no canopy cover, in disturbed areas even on the Peak top. The species was commonly recorded from the highest elevations of 2000–2100 m, but low populations were also observed from about 1800m.

Pseudophilautus samarakoon sp. nov.

(Figs. 20–22; Images 35–39; Tables 19–21)

urn:lsid:zoobank.org:act:DE23DB51-4DD4-47B0-A974-1F8D70F90E8A

Holotype: NMSL 2013.07.01 NH, 03.iv.2010, adult female 24.6mm SVL (Images 35, 36 A&D). Sripada (Peak Wilderness), Ratnapura District, Sabaragamuwa Province, Sri Lanka (06⁰48’31”N & 80⁰28’14”E; 1335m) (Fig. 1), coll. L.J.M.W., D.R.V., S.C.A. & A.W.A.C.

Paratypes: DWC 2013.01.011, adult male, 20.4mm SVL (Images 36B,E, 37); DWC 2013.01.012, adult female, 22.3mm (Images 36C,F, 38); the same date, locality and collectors.

Diagnosis: Pseudophilautus samarakoon sp. nov. can be distinguished from known congeners by the following combination of characters: Rather small size (male SVL 22.3mm; females SVL 22.4–24.6); dorsal head and interorbital space convex; snout laterally truncate; canthus rostralis rounded; loreal region concave; internarial space flat; vomerine teeth, lingual papilla and nuptial pad absent; fringe on fingers and calcar present; snout prominent tubercle present on centre of snout; interorbital area tubercular; side of head smooth; anterior and posterior dorsum, upper flank and dorsal thigh tubercular; lower flank granular; throat and chest weakly granular; supernumerary tubercles present on foot.

Description of holotype: Body rather small size (SVL 24.6), moderately robust (SVL/HW 2.9); head large size (HL/SVL 0.5), about as wide as long (HW/HL 1.0), convex above; snout truncate in lateral, mucronate in dorsal and rounded in ventral and aspects (Fig. 20 A–C), its length (ES/DFE 0.8, SN/IN 0.6) longer than horizontal diameter of eye (ES/ED 1.2); internasal space flat; canthus rostralis rounded, loreal region concave; interorbital space mid dorsally convex, larger than upper eyelid (IO/UEW 1.6), larger than internasal distance (IO/IN 1.1); distance between front of eyes 3/5 time the distance between back of eyes (DBE/DFE 1.7); nostrils oval, closer to tip of snout than to eye (SN/EN 0.6); pupil horizontally elliptical; tympanum distinct, elliptical (TYH/TYW 1.2), smaller than the eye (TYH/ED 0.40), tympanum-eye distance 4/5^thof tympanum width (TAD/TYW 0.8); pineal ocellus and vomerine teeth absent; tongue lanceolate; lingual papilla absent, with few conical tubercles on tongue.

Arm short (LAL/FEL 0.5, UAL/FEL 0.3) thin, lower-arm shorter than hand length (LAL/HNL 0.8), lower-arm longer than upper arm (LAL/UAL 1.4); fingers thin, (Fig. 21), relative length of fingers I < II < IV < III (FL-1/FL-3 0.5, FL-2/FL-3 0.6, FL-4/FL-3 0.8) (Table 19), tips of fingers rounded, enlarged, discs present on all fingers, with distinct basal and circum marginal grooves; dermal fringe on inside of all fingers, and fringe present on postaxial edge of finger IV, metacarpal fold weakly present; webbing formula I2 ½–2 ½II2^—3 2/3III3⁺–2 ½; subarticular tubercles prominent, rounded, single, all present, IV2 and III2 relatively smaller; inner palmar tubercle indistinct, single, oval, larger than outer palmer tubercle; nuptial pads absent; outer palmar tubercle indistinct, single, oval, smaller than the diatal subarticular tubercles; supernumerary tubercles present, indistinct; prepollex absent; femur twice as long as the fourth toe length (FEL/TL-4 2.2), foot length longer than thigh (FOL/FEL 1.5), toes thin, (Fig. 22), relative length of toes I < II< III < V < IV (TL-1/TL-4 0.4, TL-2/TL-4 0.4, TL-3/TL-4 0.7, TL-5/TL-4 0.7), tips of toes rounded enlarged, discs present on all toes, with distinct basal and circum marginal grooves, webbing formula I2—2^-II2⁺—3III2⁺—3^-IV2 ½—2⁺V; dermal fringe present on all toes, thin undulating fringe along exterior edge of toe V, and metatarsal fold; prominent, small tubercles present on outer edge of tarsal fold; subarticular tubercles prominent, rounded, single, all present, V2 and IV3, relatively smaller; supernumerary tubercles present, indistinct; inner metatarsal tubercle oval prominent and large, its length 3/4^th in length of toe I (IML/TL-1 0.5); outer metatarsal tubercle indistinct.

Skin between eyes tubercular, with a prominent tubercle in the center; a median dermal ridge from tip of snout to vent (more prominent till back of head) and a prominent tubercle present on centre of snout; lateral snout with few tubercles; above nostrils a pair of prominent tubercles present; on upper eyelid prominent tubercles; lateral head smooth, prominent tubercles present near the gape of mouth; supratympanic fold prominent; co-ossified skin on head and cephalic ridges absent; dorsum tubercular; upper part of flank tubercular; lower part of flank granular; upper arm smooth; forearm tubercular; hand smooth; inner and outer thigh smooth, dorsally tubercular, macroglands absent; leg dorsally tubercular, blunt calcar and small tubercles present at heel; tarsus and foot with a few tubercles. Comparative characters that distinguish the species from its congeners are provided in tables 20 and 21.

Ventral side of body: Throat and chest weakly granular; belly granular; thigh, leg and tarsus weakly granular; vent dorsally shagreen, either side prominent tubercles present.

Colour in life: Dorsum uniform creamy light brown with dark brown patches, a dark brown band between eyes and another in front, anteriorly a large dark brown ‘M’ shaped patch with a creamy light brown oval center, posteriorly two symmetric dark brown diagonal lines; laterally cream colour; limbs dorsally creamy light brown, forelimbs, hind limbs, fingers and toes with brown cross bands; ventral side cream, throat brown, feet and webbing light brown (Image 35).

Colour in alcohol: Colour pattern remains with a little fading. The colours change from dark brown to brown and the cream colour to off white (Images 36 A,D).

Paratype and variation: Dorsum colour varies ranging from a reddish brown to a lighter brown shade, to a cream colour. The markings and patterns are the same for all (Images 37–39). The ventral side of thigh granular in DWC 2013.01.012, adult female, SVL 22.3mm (Images 36 C,F, 38).

Etymology: The species name samarakoon is named after Ananda Vijith Samarakoon (Assistant Director, Department of Wildlife), in great appreciation of his efforts in educating the wildlife officers for the conservation of the country’s biodiversity at large. The species name samarakoon is used as a noun in apposition to the generic name.

Suggested vernacular names: Samarakoonge panduru madiya and Samrakoon’s Shrub Frog in Sinhala and English, respectively.

Natural history: The species was found in elevations of 1000–1400 m from the forest floor to about 2m high shrubs and in bamboo of the lower montane rain forests. Sympatric with P. bambaradeniyai sp. nov. and P. karunarathnei sp. nov.. Most abundantly found near the starting point of Kaluganga River. They were commonly found perched on bamboo plants close to the stream.

Pseudophilautus sirilwijesundarai sp. nov.

(Figs. 23–25; Images 40–42; Tables 22–24)

urn:lsid:zoobank.org:act:09A9CE69-1BC9-4577-8EFB-66597C984C52

Holotype: NMSL 2013.08.01 NH, 05.ii.2009, adult female 32.5mm SVL (Images 40, 41A,C). Sripada (Peak Wilderness), Ratnapura District, Sabaragamuwa Province, Sri Lanka (06⁰48’30”N 80⁰29’19”E; 1679m) (Fig. 1), coll. L.J.M.W, D.R.V., M.D.G.R., S.C.A., A.W.A.C. & I.N.B.

Paratypes: DWC 2013.01.013, 11.xi.2011, adult male, 22.3mm SVL (Images 41B,D, 42). The same locality, coll. D.R.V and S.C.A.

Diagnosis: Pseudophilautus sirilwijesundarai sp. nov., can be distinguished from known congeners by the following combination of characters: Moderate size (male SVL 22.3mm; female SVL 32.5mm); dorsal head and interorbital space convex; lateral snout and canthus rostralis rounded; loreal region concave; internarial space flat; fringe on fingers and vomerine ridge present; lingual papilla, nuptial pad and calcar absent; snout shagreen, with prominent dermal fold; interorbital area tubercular; side of head weakly shagreen; anterior dorsum shagreen with few tubercles; posterior dorsum and throat shagreen; upper flank areolate; dorsal thigh smooth; chest weakly granular; underside of thigh granular; supernumerary tubercles present on foot. Comparative characters that distinguish the species from its congeners are provided in tables 23 and 24.

Description of holotype: Body moderate size (SVL 32.5mm), and moderately elongate (SVL/HW 2.5); head moderate size (HL/SVL 0.4), convex above; snout truncate in dorsal, rounded in lateral, and ventral aspects (Fig. 23 A–C) (ES/DFE 0.8, SN/IN 0.4) its length equal to horizontal diameter of eye (ES/ED 1.0); internasal space flat; canthus rostralis rounded; loreal region concave; interorbital space convex, equal to upper eyelid (IO/UEW 1.0), smaller than internasal distance (IO/IN 0.9); distance between front of eyes is half times that of the distance between back of eyes (DBE/DFE 1.9); nostrils oval, small, closer to tip of snout than to eye (SN/EN 0.5); tympanum indistinct, oval (TYH/TYW 1.2), vertical, smaller than the eye (TYH/ED 0.3), tympanum-eye distance greater than tympanum width (TAD/TYW 1.1); pineal ocellus absent; vomerine teeth present, few, small, odontophores oblique and widely separated, between choanae with an angle of 70⁰ relative to body axis; tongue large, tip bifurcate, lanceolate, lingual papilla absent, but conical tubercles present on tongue.

Arm short (LAL/FEL 0.5, UAL/FEL 0.4) thin, lower-arm shorter than hand length (LAL/HNL 0.9), lower-arm longer than upper arm (LAL/UAL 1.4); fingers long and thin, (Fig. 24), relative length of fingers I < II < IV < III (FL-1/FL-3 0.4, FL-2/FL-3 0.6, FL-4/FL-3 0.6) (Tables 22), tips of fingers rounded, enlarged, discs present on fingers I – II – III – IV, with distinct basal and circum marginal grooves; dermal fringe on inside of all fingers, webbing formula I2 2/3 – 2 ½II2 – 3^-III2 2/3 – 2 2/3IV; prepollex absent; subarticular tubercles prominent, rounded, single, all present, IV2 and III2 relatively smaller; inner palmar tubercle single, oval, indistinct, larger than outer palmer tubercle; nuptial pads absent; outer palmar tubercle single, oval, and indistinct; supernumerary tubercles prominent on palm and all fingers; femur twice as long as the fourth toe length (FEL/TL-4 2.0); foot length longer than thigh (FOL/FEL 1.5); toes long, thin, (Fig. 25), relative length of toes I < II < V < III < IV (TL-1/TL-4 0.3, TL-2/TL-4 0.4, TL-3/TL-4 0.6, TL-5/TL-4 0.5), tips of toes rounded, discs present on all toes, with distinct basal and circum marginal grooves; webbing formula I2—2^-II1 2/3—3III1^-—3IV2—1⁺V; lateral dermal fringe on inside of all toes and fringe along exterior edge of toe V, on metatarsal and tarsal folds absent, but a few small tubercles present similar to an undulating fringe; subarticular tubercles prominent, rounded, single, all present, IV3, relatively smaller; supernumerary tubercles prominent, on foot and all toes; inner metatarsal tubercle oval, prominent and large, its length 2/3^rd in length of toe I (IML/TL-1 0.7); outer metatarsal tubercle absent.

Skin of snout shagreen with prominent dermal folds; upper eyelid shagreen, a few prominent tubercles present; Interorbital area with a prominent tubercle; head laterally weakly shagreen with a few tubercles near the gape of mouth; supratympanic fold prominent; a median dermal ridge extends from tip of snout to back of the head; anterior dorsum shagreen with few tubercles, posterior dorsum shagreen; upper part of flank areolate, lower part of flank granular; upper arm weakly shagreen; forearm shagreen with few tubercles; hand weakly shagreen; inner, dorsal and outer surface of thigh smooth; leg, tarsus and foot smooth; few tubercles present at heel.

Ventral side of body: Throat shagreen, chest weakly granular, belly coarsely granular; upper arm and forearm weakly granular; thigh granular, leg smooth, tarsus weakly shagreen.

Colour in life: Dorsum dark brown with a green tinge and maroon blotches fairly symmetrically placed, snout and interorbital area maroon demarcating a triangle; lateral head maroon patch below eye, flank yellowish white blotches on maroon; limbs dorsally olive green, fore limbs, hind limbs, fingers and toes with prominent maroon cross bands; ventral side off-white with brown blotches, throat area, hands and feet dark brown (Image 40).

Colour in alcohol: Colour pattern remains the same with a little fading, olive green colour changes to yellow (Images 41 A,C).

Etymology: The species epithet sirilwijesundarai named after Siril Wijesundara (Director, Royal Botanical gardens, Peradeniya, Sri Lanka), in great appreciation of his service to the country. The species name is a noun in the genitive case.

Suggested vernacular names: Siril Wijesundarage panduru madiya and Siril Wijesundara’s Shrub Frog in Sinhala and English, respectively.

Natural history: The species was found at high altitudes of elevations ranging from 1600–1700 m, montane cloud forests. They were found in the canopy on trees along side mosses, from about 10m tall tree branches.

DISCUSSION

PCA extracted 10 factors with eigenvalue more than unity. First three PCA axes explained total 61.29% of the total variation in the data with 34.92% being explained by the first component, 16.58% by the second and 9.78% by the third. PCA ordination on the first two axes (Fig. 26a) could virtually separate Pseudophilautus newtonjayawardanei, P. sirilwijesundarai, P. bambaradeniyai and P. puranappu, however the remaining species showed overlaps within the convex hulls. PCA ordination on the first three axes (Fig. 26b) could separate all the eight species. Correlation values for the variables along the first three PCA axes are provided in Fig. 27. These characters define the separation of different species from related species. For instance, variables with negative loadings on first and third principle axis and positive on the second principle component define the separation of P. newtonjayawardanei from other species as this species have high values for such variables.

Dendrogram depicting the similarity and differences among the species (Fig. 28) suggested that the eight species are divided into three morphometrical clusters. Pseudophilautus newtonjayawardanei was distinctly different from all other species and its separation was supported by high bootstrap value of 100. Out of the remaining species, the cluster comprising of Pseudophilautus bambaradeniyai, P. karunarathnai, P. samarakoon and P. sirilwijesundarai was separated from the cluster of P. dayawansai, P. jagathgunawardanai and P. puranappu with high value of bootstrap of 86.

Statistical analysis presented here has limited application as the number of individuals in each species is less. However, the analysis does reveal that proposed species are morphometrically distinct from each other. Most importantly, P. newtonjayawardanei, which is represented by a single individual, is distinctly different from all other species described in the current study. Further morphometric data of different species could be helpful in determining the morphological species boundaries among the proposed species (Appendix 3).

The conservation status of all species described here, except for P. newtonjayawardanei sp. nov., have been considered Critically Endangered, according to the IUCN Red List criteria, because for all species the extent of occurrence is <100km², are recorded from single locations, and their habitats are under severe threat. P. newtonjayawardanei sp. nov., will have to be treated as Data Deficient as it is described based on a single specimen.

The highest point of this mountain range the Sripada Peak, is a place of worship of all religions in the country and a place of aesthetic beauty. Millions visit this site every year during the pilgrim season which lasts for a period of six months, which is the time where a large amount of garbage gets collected, and the natural forest gets over exploited. Illegal gem mining on either sides of the river bank within the forest is another potential threat to the amphibian diversity. Tea cultivation in the surrounding areas are slowly expanding. Illegal tree felling to cultivate tea, has become a major threat in the area.

Amongst the most important findings in this region is the forest die back possibly due to pollution and/or climate change (Image 43). It has never been documented in this region before and a large area has now been effected with the natural forest cover slowly declining.

Most importantly, although these species have been described from a single locality, it must be emphasised that all these species possess quite unique morphological characters and are well distinguishable from one another. P. newtonjayawardanei sp. nov. as mentioned under its natural history notes is a very rare species and nothing is known about its behaviour as only two specimens have been observed by us till date. The high altitude, low temperatures, misty conditions, tall canopy dwelling habit and small size make it very difficult to locate the species. Hence, we preserved only a single specimen of the two. We have critically compared the novel species with the 17 extinct species of this genus, as several species morphologically similar have been recorded during these surveys, where further studies are ongoing to clarify their identity. The quite large number of species descriptions in this area in the recent past is due to the increased number of systematic surveys being carried out in the region, and many more species await their discoveries.

Although Peak Wilderness is part of the recently declared Central Hills World Heritage Site, and is one of the few remaining virgin forests with a continuous natural forest cover and altitudinal gradient forest influenced by different climatic zones (Image 44–45), the diversity of this area remains relatively unknown and hence under represented. The new species described here is just a glimpse of what remains to be uncovered. At the current rate of destruction of the natural habitats in the region the amphibians are at a higher risk of extinction, where they may be extripated without ever being known to science. The novel descriptions presented here hopefully will set a basis for future conservation efforts in the region.

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