Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920
Eight new species of Pseudophilautus (Amphibia: Anura: Monograph Rhacophoridae) from Sripada World Heritage Site (Peak
Wilderness), a local amphibian hotspot in Sri Lanka ISSN Online 0974-7907 1 2 3 Print 0974-7893 L.J. Mendis Wickramasinghe , Dulan Ranga Vidanapathirana ,, M.D. Gehan Rajeev , 4 5 6 S. Chathuranga Ariyarathne , A.W. Amila Chanaka , L.L. Dharshana Priyantha , OPEN ACCESS 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 [email protected] (corresponding author), 2 [email protected], 3 [email protected], 4 [email protected], 5 [email protected], 6 [email protected], 7 [email protected], 8 [email protected]
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 forP. 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.
3789 Eight new species of Pseudophilautus Wickramasinghe et al.
INTRODUCTION trails, heavy rainfall, and steep geographic variations, proper field surveys in this region were lacking. Due The genus Pseudophilautus consists of 65 known to this reason inspite of the geographical and climatic species currently, all of which are endemic to Sri variations the region was considered to have a lower Lanka (Manamendra-Arachchi & Pethiyagoda 2005; faunal diversity. Hence, we carried out the survey with Meegaskumbura & Manamendra-Arachchi 2005, 2011; the intention of setting a basis for the actual diversity Meegaskumbura et al. 2007, 2009). These direct of herpetofauna of Peak Wilderness. The novel species developing rhacophorids (Bossuyt & Dubois 2001; described here from the Peak Wilderness, Central Hills, Meegaskumbura et al. 2002; Bahir et al. 2005; Grosjean of Sri Lanka, as results of the preliminary two phases of et al. 2008; Li et al. 2008, 2009; Biju & Bossuyt 2009; Yu an extensive field survey of the region which lasted for et al. 2009) previously belonged to the genus Philautus a period of two years (end of 2009-end of 2011), were which were widely distributed throughout tropical Asia based on morphological and morphometric characters. (Bossuyt & Dubois 2001; Bossuyt et al. 2004; Frost et al. Sripada Peak is the highest point in this mountain range, 2006; Yu et al. 2009). But later, considering molecular and because of its religious importance, and aesthetic phylogenetics, the populations in Sri Lanka and India beauty, pilgrims and travellers have identified six major were found to be a separate radiation belonging to a trails leading to the top. For the 1st phase, sampling was separate genus known now as Pseudophilautus (Li et al. carried out in the trail leading from Palabaddala to the 2009; Yu et al. 2010; Meegaskumbura & Manamendra- Sripada peak, and in the 2nd phase we chose Kuruvita to Arachchi 2011). The most recent publication of this genus the Sripada peak. Our work here describes a collection by Meegaskumbura & Manamendra-Arachchi (2011) in of eight new species, with each species having a detailed the country described two species which brought the description based on strong morphological characters, total number to 65 species. But sadly, a considerable accompanied by colour photographs. portion of the total number of identified species, have already vanished (17 species) (Manamendra-Arachchi & Pethiyagoda 2005; Meegaskumbura et al. 2007) where MATERIALS AND METHODS most have not even been sighted after their initial descriptions, showing an urgency to conserve the few The approximate total distances of the 1st trail (from remaining habitats. Palabaddala to the Sripada peak) was 10km, and the 2nd Peak Wilderness as part of the Central Hills was recently trail (from Kuruvita to the Sripada peak) was 10km. The designated as a World Heritage Site. Geographical area field survey commenced at the end of 2009, and the of the sanctuary is about 240sq.km, of which 211.75sq. Phases I and II were completed in December 2011. The km comprises natural or semi natural vegetation; the sampling sites were selected considering the accessibility, rest is no longer forest and includes tea estates and representative habitats and spatial distribution in the village settlements. The sanctuary lies in the center National Park, through an initial reconnaissance survey. of the western ridge of the Central Highlands, north- Fifteen field visits were made, each sampling session east of Ratnapura and spanning the border of Central spanning eight continuous days. Sampling was done and Sabaragamuwa provinces. Its eastern boundary mostly nocturnally, and photographs of most species is contiguous with Horton Plains National Park. Peak were done in the wild to avoid any confusion of change Wilderness occupies the escarpment that rises steeply in colour after captivity and to avoid over collection of from the lowlands to the south and west. Much of the the same species. Specimens collected in the field were terrain is very rugged, with altitudes ranging from 600m first fixed in 90% ethanol for two hours and stored in near Ratnapura to 2,238m at Sripada Peak, Sri Lanka’s 70% ethanol. fourth highest peak. It is an important watershed for Sex and maturity were determined by examining three major river systems, the Kelani Ganga, the Kalu secondary sexual characters, or when absent, by Ganga, and the Walawe Ganga (DWC 2007). examining the gonads through a small lateral incision The last vertebrate species discovered from in the specimen. The following morphological the studied area were Cnemaspis samanalensis charactetristics were common to all eight species hence (Wickramasinghe & Munindradasa 2007); Lankascincus they have not been mentioned under each species; munindradasai and Lankascincus sripadensis nostrils oval, without flap of skin laterally, small, close (Wickramasinghe et al. 2007), depicting its diversity. to tip of snout; latero-dorsal folds, lateral line system, Because of relatively high altitudes, difficult Fejervarya line, and macroglands were absent. We here
3790 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. 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 © L.J. Mendis Wickramasinghe (Appendix 4). The specimens that formerly belonged to Image 1. Weakly granular (Chest of Pseudophilautus the Wildlife Heritage Trust (WHT) bearing WHT numbers jagathgunawardanai sp. nov. holotype, NMSL 2013.03.01 NH). 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 © L.J. Mendis Wickramasinghe 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 Pseudophilautus species. 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 Image 2. Granular (Belly of Pseudophilautus newtonjayawardanei of eyes (DBE), measured between posterior edge of sp. nov. holotype NMSL 2013.05.01 NH) 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 (EN), measured between anterior most point of eye and of the third finger (DW), measured across the inner and middle of nostril; eye to snout distance (ES), measured outer edges of circum-marginal groove; eye diameter between anterior most point of eye and tip of snout; (ED), horizontal diameter of eye; eye to nostril distance thigh (femur) length (FEL), distance between vent and
Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3791 Eight new species of Pseudophilautus Wickramasinghe et al.
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), © L.J. Mendis Wickramasinghe 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 Image 3. Coarsely granular (Belly of Pseudophilautus sirilwijesundarai sp. nov. holotype, NMSL 2013.08.01 NH) 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 knee with both thigh and shank flexed; first finger margin of the most proximal subarticular tubercle or length (FL-1), measured between posterior margin of crease of articulation and the tip of third toe; fourth the most proximal subarticular tubercle or crease of toe length (TL-4), measured between posterior margin articulation and the tip of first finger; second finger of the most proximal subarticular tubercle or crease of length (FL-2), measured between posterior margin of articulation and the tip of fourth toe; fifth toe length the most proximal subarticular tubercle or crease of (TL-5), measured between posterior margin of the most articulation and the tip of second finger; third finger proximal subarticular tubercle or crease of articulation length (FL-3), measured between posterior margin of and the tip of fifth toe; tympanum-nostril distance the most proximal subarticular tubercle or crease of (TND), distance between anterior-most point of inner articulation and the tip of third finger; fourth finger margin of inner rim of tympanum and middle point of length (FL-4), measured between posterior margin of nostril; tympanum-front of eye distance (TPD), taken as the most proximal subarticular tubercle or crease of distance between anterior-most point of inner margin of articulation and the tip of fourth finger; foot length inner rim of tympanum and anterior most point of eye; (FOL), distance between heel and tip of fourth toe with tympanum height (TYH), vertical diameter of the inner both foot and shank flexed; groin to knee distance (GK), rim; tympanum width (TYW), horizontal diameter of the measured between groin and the tip of knee; hand inner rim; upper arm length (UAW), distance between length (HNL), taken from posterior-most margin of inner axilla and elbow; upper eyelid width (UEW), measured palmar tubercle to tip of disk of third finger; head depth from bony edge of supraorbital to outer edge of upper (HD), distance between the apex of eye and the bottom eyelid; vent to knee-angle length (VKL), distance of jaw; head length (HL), distance between angle of measured from knee-angle to vent. jaws and snout tip; head width (HW), measured across Morphometric data of both males and females was angle of jaws; inner metatarsal tubercle length (IML), considered for statistical analysis to understand the distance between the anterior and posterior edges morphometric difference among the individuals of eight of the inner metatarsal tubercle; internarial distance new species of Pseudophilautus. To account for the (IN), least distance between the inner margin of nares; size difference among individuals, all the morphometric
3792 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. characters were taken as %SVL. Principle Component than horizontal diameter of eye (ES/ED 1.1); internasal Analysis (PCA) was performed on the correlation matrix space flat; canthus rostralis rounded, loreal region between 25 variables using the software PAST (Hammer concave; interorbital space convex, larger than upper et al. 2001). Dendrogram showing the relationship eyelid (IO/UEW 1.1), and equal to internasal distance between different species was plotted using mean values (IO/IN 1.0); distance between front of eyes 2/3rd the of morphometric data for each species using Euclidian distance between back of eyes (DBE/DFE 1.6); nostrils distances and Ward’s method for clustering in PAST. oval, without flap of skin laterally, closer to tip of snout Geographical coordinates were determined from than to eye (SN/EN 0.8); pupil horizontally elliptical; GPS readings (Gamin eTrex Gista) with WGS84 (World tympanum distinct, oval (TYH/TYW 0.8), smaller than Geodetic System) datum at the locality. Additional data the eye diameter (TYH/ED 0.4), tympanum-eye distance for comparison were collected from Pethiyagoda & about 2/5th of tympanum width (TAD/TYW 0.4); pineal Manamendra-Arachchi (2005) and WGS 84 world grid ocellus, vomerine teeth and lingual papilla absent, but system. a depression with a conical tubercle present on tongue; Collectors are indicated by their initials as follows: tongue small, lanceolate. L.J. Mendis Wickramasinghe (L.J.M.W), Dulan Ranga Arm short, thin (LAL/FEL 0.5, UAL/FEL 0.3); forearm
Vidanapathirana (D.R.V), M.D. Gehan Rajeev (M.D.G.R), shorter than hand length (LAL/HNL 0.9), longer than S. Chathuranga Ariyarathne (S.C.A.), A. W. Amila Chanaka upper arm (LAL/UAL 1.4); fingers thin, (Fig. 3), relative (A.W.A.C.), L.L. Dharshana Priyantha (L.L.D.P.), Imesh length of fingers I < II < IV < III (FL-1/FL-3 0.5, FL-2/ Nuwan Bandara (I.N.B.). 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 RESULTS and circum marginal grooves; dermal fringes absent; rudimentary webbing present on all fingers; subarticular Pseudophilautus bambaradeniyai sp. nov. tubercles prominent, oval, single, all present, III2 and (Figs. 1–4; Images 4–7; Tables 1–3) IV2 relatively smaller, supernumerary tubercles present urn:lsid:zoobank.org:act:FB630D8A-7354-4307-A343-5B5343921C73 on finger III and IV; inner palmar tubercle indistinct, single, oval, larger than outer palmar tubercle; outer Holotype: NMSL 2013.01.01 NH, 05.iv.2010, adult palmar tubercle indistinct, single, oval, smaller than the male SVL 17.3mm (Images 4, 5A,D), Sripada (Peak subarticular tubercles; nuptial pads absent; prepollex Wilderness), Ratnapura District, Sabaragamuwa absent; femur 2 2/5 times longer than fourth toe length Province, Sri Lanka (06048’31”N & 8008’14”E; 1334m) (FEL/TL-4 2.4); foot length longer than thigh (FOL/FEL (Fig. 1), coll. L.J.M.W, D.R.V., S.C.A. & A.W.A.C. 1.3); toes thin, (Fig. 4), relative length of toes I < II < III < Paratypes: DWC 2013.01.001, adult male, SVL V < IV (TL-1/TL-4 0.3, TL-2/TL-4 0.4, TL-3/TL-4 0.7, TL-5/ 19.9mm (Images 5B,E, 6); DWC 2013.01.002, adult male, TL-4 0.7); webbing formula I2+—2-II2+—3-III2+—3-IV3— SVL 20.2mm (Images 5C,F, 7); same data as holotype. 2V; tips of toes rounded, enlarged, discs present on all Diagnosis: Pseudophilautus bambaradeniyai sp. toes with distinct basal and circum marginal grooves; nov., can be distinguished from known congeners by the thin dermal fringe present on all toes, thin fringe along following combination of characters: small size frog (SVL postaxial edge of toe V, small tubercles present on outer 17.3–20.2 mm); head dorsally convex; canthus rostralis edge of tarsal fold; subarticular tubercles prominent, rounded; interorbital space convex; internarial space flat; oval, single, all present, IV2, IV3, and V2 relatively vomerine teeth, lingual papilla, fringe on fingers, calcar smaller; supernumerary tubercles present, indistinct, on and nuptial pad absent; snout, interorbital area, side of toe III and IV; inner metatarsal tubercle oval prominent head, upper and lower flank smooth; anterior dorsum and large, its length 3/4 times in length of toe I (IML/TL-1 prominent tubercles and horny spinules, posterior 0.6); outer metatarsal tubercle indistinct. dorsum horny spinules. Distinguishing characters Skin of snout dorsally, laterally and interorbital space provided in comparison tables 2 and 3. smooth; upper eyelid tubercular; head laterally smooth; Description of holotype: Small sized frog (SVL dorsum anterior with prominent tubercles and horny 17.3mm), elongate (SVL/HW 2.4); head large (HL/SVL spinules, posterior with prominent horny spinules; 0.5), convex above; snout truncate in lateral aspect, upper and lower part of flank smooth; supratympanic mucronate in dorsal aspect, and sub-elliptical in ventral fold distinct; upper arm, forearm, and hand smooth; aspects (Fig. 2 A–C) (ES/DFE 0.8, SN/IN 0.7), larger thigh inner and outer smooth, dorsally with tubercles,
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Eight new species of Pseudophilautus Wickramasinghe et al. Figure 1. Distribution of the type localities▲ within the Peak Wilderness: “●” Pseudophilautus bambaradeniyai sp. nov., P. karunarathnai sp. nov., and P. samarakoon sp. nov.; “ ” P. dayawansai sp. nov., and P. sirilwijesundarai sp. nov.; “♦” P. jagathgunawardanai sp. nov.; “” P. newtonjayawardanei sp. nov.; “■” P. puranappu sp. nov.
3794 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al.
Figure 2. Pseudophilautus bambaradeniyai sp. nov. holotype (NMSL 2013.01.01 NH), head; (A) lateral aspect (B) dorsal aspect (C) ventral aspect. Scale bar: 2mm. (© Dulan Ranga Vidanapathirana).
Figure 4. Pseudophilautus bambaradeniyai sp. nov. holotype (NMSL 2013.01.01 NH), ventral aspect of left foot. Scale bar: 5mm. (© L.J. Mendis Wickramasinghe) © L.J. Mendis Wickramasinghe
Figure 3. Pseudophilautus bambaradeniyai sp. nov. holotype (NMSL 2013.01.01 NH), ventral aspect of left hand. Scale bar: 5mm. (© L.J. Mendis Wickramasinghe) macroglands absent; leg smooth with few tubercles; Image 4. Pseudophilautus bambaradeniyai sp. nov. holotype in life calcar absent, small tubercles present at heel; vent (NMSL 2013.01.01 NH). dorsally shagreen, either side shagreen. Ventral side of body: Throat and chest weakly granular; belly granular; upper arm weakly granular to
Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3795 Eight new species of Pseudophilautus Wickramasinghe et al. © L.J. Mendis Wickramasinghe
Image 5. Type series of Pseudophilautus bambaradeniyai sp. nov. Left to right, NMSL 2013.01.01 NH, DWC 2013.01.001, DWC 2013.01.002. A-C dorsal aspect and D-F ventral aspect. Scale bar: 5mm. © L.J. Mendis Wickramasinghe © L.J. Mendis Wickramasinghe Image 6. Pseudophilautus bambaradeniyai sp. nov., paratype in life (DWC 2013.01.001). Image 7. Pseudophilautus bambaradeniyai sp. nov. paratype in life (DWC 2013.01.002). smooth; forearm smooth; thigh, leg and tarsus smooth. feet and webbing lighter (Image 4). Colour in life: Dorsum dark brown with blackish Colour in alcohol: Colour pattern remains with a blotches, large dark brown marking covering much of the little fading, off-white vertebral stripe prominent on dark anterior, blackish cross band between eyes, a prominent brown, ventral side off-white all over (Images 5 A,D). off-white vertebral stripe from the tip of snout to anus, Paratype and variation: Dorsum colour varies and continuing down the hind limbs symmetrically, ranging from orange to a lighter brownish tinge and the lateral body lighter; limbs dark brown, fore limbs, hind thickness of the vertebral line varies; in some this line limbs, fingers and toes with blackish-brown cross bands; discontinues towards limbs (Images 5 B,E, 6). ventral side off white with light brown blotches, hands, Etymology: The species epithet bambaradeniyai
3796 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Range 3.1—3.3 5.3—5.6 3.6—3.7 4.2—4.7 2.9—3.0 1.5—1.7 1.4—1.5 6.2—6.8 1.5—1.5 8.7—9.1 0.6—0.6 4.6—4.8 3.0—3.5 1.3—1.5 1.3—1.5 3.4—3.5 1.9—2.2 9.4—9.6 6.0—6.3 7.5—8.0 10.5—10.9 17.3—20.2 SD 0.2 0.1 0.1 0.2 0.0 0.1 0.1 0.3 0.2 0.0 0.2 0.0 0.2 1.5 0.1 0.1 0.1 0.1 0.2 0.1 0.3 0.1 5.5 3.2 3.7 4.5 2.9 1.7 1.4 6.4 1.5 8.9 0.6 3.2 4.7 1.5 1.4 3.5 2.1 6.2 7.8 9.5 10.7 19.0 Mean 5.6 3.2 3.7 4.6 3.0 1.7 1.4 6.8 1.5 8.7 0.6 3.5 4.8 1.3 1.5 3.5 2.1 6.0 7.8 9.6 10.9 20.2 Paratype 2 Paratype 2013.01.002 5.5 3.3 3.7 4.7 2.9 1.7 1.5 6.3 1.5 8.9 0.6 3.0 4.8 1.5 1.5 3.5 2.2 6.3 8.0 9.6 10.8 19.6 Paratype 1 Paratype 2013.01.001 5.3 3.1 3.6 4.2 2.9 1.5 1.4 6.2 9.1 1.5 0.6 3.2 4.6 1.5 3.4 1.9 1.3 6.1 7.5 9.4 10.5 17.3 Holotype 2013.01.01. NH KT SN LAL TBL SVL TAS MN VKL TYH TPD TAD UAL TL-5 TL-4 TL-3 TL-2 TL-1 TND MFE TYW MBE DWC UEW NMSL/ sp. nov. 14.2 Range 13.3— 5.3—6.4 4.2—4.6 8.7—9.2 8.4—8.6 2.8—3.0 3.2—3.7 1.5—1.7 2.1—2.3 3.2—3.4 4.3—4.3 1.9—2.1 2.2—2.7 7.4—8.3 0.8—0.9 6.8—7.6 0.7—0.8 0.8—0.9 3.2—3.4 2.0—2.4 0.4—0.5 9.6—10.2 SD 0.6 0.2 0.3 0.1 0.5 0.1 0.3 0.1 0.1 0.1 0.0 0.1 0.3 0.5 0.1 0.4 0.4 0.0 0.0 0.1 0.2 0.0 of Pseudophilautus bambaradeniyai 5.9 4.4 9.0 8.5 2.9 3.5 1.6 2.2 3.3 4.3 2.0 2.5 8.0 0.9 7.2 0.7 0.9 3.3 2.2 0.5 13.9 10.0 Mean 6.1 4.6 9.2 8.5 3.0 3.5 1.7 2.1 3.2 4.3 2.1 2.6 8.2 0.9 7.3 0.8 0.9 3.2 2.0 0.5 14.2 10.2 Paratype 2 Paratype 2013.01.002 6.4 4.4 8.7 8.6 2.9 3.7 1.7 2.3 3.3 4.3 2.1 2.7 8.3 0.8 9.6 7.6 0.7 0.8 3.4 2.4 0.5 14.1 Paratype 1 Paratype 2013.01.001 5.3 4.2 9.2 8.4 2.8 3.2 1.5 2.1 3.4 4.3 1.9 2.2 7.4 0.9 0.7 0.8 3.2 6.8 2.1 0.4 13.3 10.1 Holotype 2013.01.01. NH 2013.01.01. IN IO ES HL DL ED EN GK DB HD FEL DW HW FOL IML DFE DBE FL-4 FL-3 FL-1 FL-2 HNL DWC NMSL/ Table 1. The morphometric measurements (mm) of the type series measurements 1. The morphometric Table
Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3797 Eight new species of Pseudophilautus Wickramasinghe et al. Present Present Present Nuptial Nuptial pad Absent Present Present Present Present Present Present Present Present Calcar Absent Present Present Present Present Present Present Present Present Present Present Present Present Present Present Fringe on Fringe fingers Absent Present Present Present Present Present Present Present Present Present Present Present Present Present Present Absent Lingual papilla Present Present Present Present Present Present Present Present Present Absent Vomerine teeth Vomerine Present Present Present Present Present Concave Concave Concave Concave Concave Concave Concave Concave Concave Flat Internarial space Internarial Concave or flat Concave Concave Concave Concave Concave Concave Concave Flat Flat Flat Flat Concave Concave Convex Interorbital Interorbital space Concave Flat Flat Concave Flat Flat Flat Concave or flat Concave Flat Flat Sharp Sharp Sharp Sharp Rounded Canthus Canthus rostralis Sharp Sharp Sharp Sharp Sharp Sharp Sharp Pseudophilautus of the genus among congeners sp. nov. Rounded pointed Obtusely Rounded Rounded or oval Pointed pointed Obtusely Blunt pointed Obtusely Oval Rounded Rounded Truncate Snout lateral Sharply Pointed pointed Obtusely Rounded Blunt or truncate Rounded Rounded Rounded Rounded Rounded Concave Concave Concave Flat Flat Flat Concave Convex Head dorsally Flat Flat Flat Flat Flat Flat Concave or Concave flat Pseudophilautus bambaradeniyai 20.4–24.6 22.3–32.5 18.9–27.3 17.6–28.9 38.4 25.4–37.0 33.3–41.7 17.0–27.5 21.7–26.4 16.3–19.0 38.4–48.9 33.6–51.1 16.2–19.2 35.8-40.4 24.5–30.1 17.3–20.2 SVL 18.3–23.9 44.8 35.3 43.5 23.4–28.8 23.7–29.4 26.7–29.3 26.9–44.5 33.4–46.8 32.9–42.9 sp. nov. sp. nov. sp. nov. sp. nov. frankenbergi hallidayi P. samarakoon samarakoon P. sirilwijesundarai P. sp. nov. asankai P. cuspis P. P. newtonjayawardanei newtonjayawardanei P. sp. nov. puranappu P. abundus P. adspersus P. alto P. auratus P. caeruleus P. cavirostris P. P. jagathgunawardanai P. sp. nov. P. karunarathnai karunarathnai P. P. bambaradeniyai P. sp. nov. dayawansai P. P. decoris P. Species name P. dimbullae P. P. eximius P. extirpo P. femoralis P. folicola P. P. P. fergusonianus P. fulvus P. P. Table 2. Characters that differentiate differentiate that 2. Characters Table
3798 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Nuptial Nuptial pad Present Present Present Present Present Present Present Present Calcar Present Present Present Present Present Present Present. Present Fringe on Fringe fingers Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Lingual papilla Present Present Present Present Present Present Vomerine teeth Vomerine Absent or present Absent Present Present Present Present Present Present Present Present Present Present Internarial space Internarial Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave or Convex concave Concave or flat Concave Concave Concave Concave Concave Concave Concave Concave Interorbital Interorbital space Flat Flat Flat Concave Concave Concave Flat Flat Flat Flat Flat Flat Concave Concave Flat Concave Flat Concave Flat Flat Flat Flat Concave Canthus Canthus rostralis Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Snout lateral Rounded Pointed Rounded Blunt Acuminate Oval Obtusely pointed Obtusely Rounded Oval Pointed or oval Pointed Rounded Rounded Rounded Obtusely pointed Obtusely Blunt Pointed Oval Pointed Oval Pointed Rounded Oval Oval Pointed Head dorsally Uneven Flat Flat Concave Flat flat or Convex Concave Flat Flat Convex or Convex concave Concave Flat Flat SVL 27.9 18.3–21.9 22.0–28.6 11.2–22.4 19.8 21.2–23.4 25.7 17.5 18.3–21.7 42.5–61.1 25.6–30.3 25.1–26.8 32.1 21.3–26.0 17.7–24.7 23.1–33.4 18.6 40 27.4–42.7 46.6 14.9 40.9 17.4 20.7 29.4–35.0 34.8 23.9–49.0 16.3–18.4 hoffmanni P. halyi P. P. hankeni P. P. hoipolloi P. hypomelas P. leucorhinus P. P. limbus P. P. rugatus P. P. regius P. reticulatus P. P. pleurotaenia P. procax P. P. pardus P. poppiae P. popularis P. P. ocularis P. oxyrhynchus P. papillosus P. Species name P. macropus P. P. maia P. malcolmsmithi P. P. lunatus P. P. nasutus P. nemus P. P. mooreorum P. nanus P. P. microtympanum P. mittermeieri P.
Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3799 Eight new species of Pseudophilautus Wickramasinghe et al. Present Present Present Nuptial Nuptial pad Present Present Present Present Calcar Present Present Present Present Present Present Present Present Present Fringe on Fringe fingers Present Present Present Present Present Lingual papilla Present Present Present Present Present Present Present Present Present Vomerine teeth Vomerine Present Concave Concave Concave Concave Concave Concave Concave Concave Internarial space Internarial Concave Flat Flat Concave Flat Flat Concave Concave Flat Flat Flat Flat Flat Interorbital Interorbital space Flat Flat Sharp Sharp Sharp Sharp Sharp Sharp Canthus Canthus rostralis Sharp Sharp Rounded Obtusely pointed Obtusely Rounded pointed Obtusely or rounded Blunt Rounded Obtusely pointed Obtusely Pointed Oval Oval Oval Rounded Blunt Rounded Rounded Oval Rounded Snout lateral Blunts Pointed Flat Flat Flat Concave Flat Flat Head dorsally Concave 35.7 28.4–31.0 23.2–36.0 13.5–13.9 27.4–36.3 20.0–32.5 24.2–32.4 17.7–30.0 19.9–22.8 13.4 35.4–51.8 24.3–31.3 12.6–15.6 16.1–16.6 22.2–39.4 39.6–55.3 30.4–41.6 20.6–24.1 22.6–38.1 SVL 31.6 22.9–30.1 stuarti steineri P. variabilis P. P. temporalis P. P. stictomerus P. P. tanu P. viridis P. zal P. P. schmarda P. schneideri P. semiruber P. silus P. silvaticus P. simba P. singu P. sordidus P. P. stellatus P. P. rus P. sarasinorum P. Species name zimmeri P. P. zorro P.
3800 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Smooth Smooth Smooth Smooth Very fine, horny fine, horny Very spinules Tuberculated Shagreen Glandular warts Smooth Weakly shagreen Weakly Smooth Dorsal thigh Dorsal Smooth Smooth Smooth or with glandular warts Smooth Smooth Granular Granular in males Granular Granular Granular Very fine, Very horny spinules Smooth Weakly Weakly granular Glandular warts Granular Granular Granular Granular Lower Lower flank Granular Granular Weakly Weakly granular Granular Granular Glandular warts and Glandular warts spinules horny Granular or smooth Granular Granular Shagreen Horny spinules Horny Smooth Shagreen Glandular warts Granular Weakly tubercular Weakly Tubercular spinules in Horny males Upper flank Weakly granular Weakly Areolate
Heavily tuberculated, tuberculated, Heavily glandular warts Glandular warts and Glandular warts spinules (females horny spinules) lack horny Shagreen Weakly tubercular Weakly Shagreen Very fine, horny spinules fine, horny Very Horny spinules Horny Shagreen Glandular warts Smooth Smooth spinules, horny Few tubercles Prominent Tubercular spinules in males, Horny in females warty Posterior dorsum Posterior Spinulate Shagreen
Heavily tuberculated, tuberculated, Heavily glandular warts Glandular warts and horny and horny Glandular warts lack horny spinules (females spinules) Shagreen Smooth Shagreen Very fine, horny spinules fine, horny Very Prominent tubercles and tubercles Prominent spinules horny Shagreen Glandular warts Smooth except for a few a few for Smooth except warts glandular scattered Smooth shagreen Weakly Tubercular in spinules (absent Horny females) spinules in males, Horny in females warty spinules with a “) Horny of tubercles pattern (”shaped Anterior dorsum Anterior Smooth tubercles with few Shagreen sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners sp. nov. Glandular warts Glandular warts Glandular warts spinules and horny Shagreen Tubercular Shagreen Very fine, horny fine, horny Very spinules Smooth Shagreen Glandular warts Smooth except for for Smooth except scattered a few glandular warts Side of head Weakly tubercular Weakly shagreen Weakly
Pseudophilautus bambaradeniyai Glandular warts Glandular warts and Glandular warts spinules horny Shagreen Smooth, with a ridge Smooth, with a ridge breaking across medially Shagreen Horny spinules in Horny males Tubercular fine, horny Very spinules Smooth Shagreen, with a Shagreen, ridge prominent Glandular warts Smooth except for for Smooth except scattered a few glandular warts spinules Horny Interorbital area Interorbital Weakly tubercular Weakly tubercle A prominent present Smooth,with two Smooth,with two dermal fringes cross Glandular warts Glandular warts and Glandular warts spinules horny Shagreen Tuberculated, a single Tuberculated, tubercle prominent medially Smooth, with a blunt Smooth, with a blunt tubercle Shagreen Horny spinules in males in spinules Horny Prominent tubercle tubercle Prominent of on centre present snout fine, horny Very spinules Smooth Shagreen Glandular warts Smooth except for a for Smooth except glandular scattered few warts spinules Horny Snout Smooth, a prominent Smooth, a prominent tubercle blunt with Shagreen, dermal fold prominent sp. nov. sp. nov. sp. nov. sp. nov. P. cavirostris cavirostris P. P. caeruleus P. P. jagathgunawardanai P. sp. nov. P. auratus P. P. dayawansai P. P. puranappu P. P. dimbullae P. P. alto P. P. samarakoon samarakoon P. asankai P. P. bambaradeniyai P. sp. nov. P. newtonjayawardanei newtonjayawardanei P. sp. nov. abundus P. P. decoris P. P. adspersus P. cuspis P. Species name karunarathnai P. sirilwijesundarai P. sp. nov. Table 3. Characters in skin structure that differentiate differentiate that in skin structure 3. Characters Table
Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3801 Eight new species of Pseudophilautus Wickramasinghe et al. Glandular warts Glandular warts with horny spinules Glandular warts Glandular warts Smooth with glandular warts, Shagreen Smooth Granular, not Granular, rough Smooth Shagreen, few few Shagreen, glandular warts Smooth Smooth Dorsal thigh Dorsal Smooth Smooth with a scattered few glandular warts Smooth Smooth Smooth Granular Granular Granular Granular Granular Granular Granular Granular Granular Granular Granular with glandular warts Granular Lower Lower flank Granular Granular Weakly Weakly granular Glandular Glandular warts Glandular warts bearing horny spinules Glandular warts Glandular warts, Glandular warts, spinules with horny in males Shagreen, males Shagreen, with glandular warts spinules and horny Shagreen Finely granular Granular Shagreen Granular Smooth with glandular warts Glandular warts Upper flank Granular Granular with a few with a few Granular glandular scattered warts
Glandular warts bearing Glandular warts spinules horny Shagreen with glandular Shagreen warts Glandular warts, with Glandular warts, spinules in males horny Few glandular warts, glandular warts, Few and males glandular warts spinules horny Shagreen Finely granular Weakly granular Weakly Shagreen, horny spinules horny Shagreen, scattered Shagreen, with a few with a few Shagreen, glandular warts Smooth Smooth Smooth or shagreen Smooth Smooth with a few warts glandular scattered Smooth Posterior dorsum Posterior Smooth Shagreen or smooth, with Shagreen glandular scattered a few warts Glandular warts bearing Glandular warts spinules horny Shagreen with glandular Shagreen warts Glandular warts, with horny with horny Glandular warts, spinules in males Shagreen, males glandular Shagreen, spinules and horny warts Shagreen Finely granular Weakly granular Weakly Shagreen, horny spinules horny Shagreen, scattered Shagreen, with a few with a few Shagreen, glandular warts Smooth Smooth Smooth or shagreen Smooth scattered Smooth with a few glandular warts Smooth Anterior dorsum Anterior Smooth or granular Shagreen or smooth, with a Shagreen warts glandular scattered few Glandular warts Glandular warts bearing horny spinules Shagreen with Shagreen glandular warts Glandular warts Few glandular Few warts Shagreen Finely granular Weakly granular Weakly Glandular warts Shagreen Side of head smooth Shagreen, or with a few glandular scattered warts Glandular warts Glandular warts Glandular warts bearing horny spinules Shagreen with Shagreen glandular warts Glandular warts, Glandular warts, spinules with horny in males Shagreen, males Shagreen, and glandular warts spinules horny Shagreen Finely granular Weakly granular Weakly Shagreen Shagreen Interorbital area Interorbital Smooth with a few glandular scattered warts Glandular warts Glandular warts spinules bearing horny Shagreen with Shagreen glandular warts Glandular warts, with Glandular warts, spinules in males horny Shagreen, males Shagreen, and glandular warts spinules horny Shagreen Finely granular Weakly granular Weakly Shagreen Shagreen, with a few with a few Shagreen, glandular warts Snout hoffmanni hallidayi frankenbergi frankenbergi P. P. hankeni P. P. halyi P. P. folicola P. P. P. fergusonianus fergusonianus P. P. lunatus P. P. fulvus P. P. femoralis femoralis P. P. limbus P. macropus P. P. eximius eximius P. P. P. hypomelas hypomelas P. P. extirpo P. hoipolloi P. Species name leucorhinus P.
3802 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Smooth Shagreen Smooth Smooth Glandular warts Smooth Smooth, few scattered glandular warts Smooth Dorsal thigh Dorsal Glandular warts Shagreen, Shagreen, scattered few glandular warts Glandular warts Glandular warts spinules Horny in males, finely or granular in shagreen female Glandular warts Smooth Smooth Granular Horny Horny spinules in males Granular Glandular warts Granular Glandular warts Glandular warts Granular Lower Lower flank Granular Granular Granular Glandular Granular Granular Granular Glandular folds Horny spinules Horny in males, finely or granular in female shagreen Horny spinules in Horny males Shagreen, few few Shagreen, glandular scattered and horny warts, spinules Glandular warts Shagreen or Shagreen glandular warts Glandular warts Glandular warts Granular Upper flank Glandular warts Granular spinules Horny Glandular Tuberculated, horn-like horn-like Tuberculated, spinules Few glandular warts Few Horny spinules in Horny or males, finely granular in female shagreen Horny spinules in Horny or males, finely granular in female shagreen Shagreen, few scattered scattered few Shagreen, and horny glandular warts spinules (males) Glandular warts Shagreen or with Shagreen glandular warts Glandular warts Smooth Smooth Smooth Posterior dorsum Posterior Glandular warts Smooth, shagreen and Smooth, shagreen males glandular warts, spinules horn like Glandular warts
tuberculated, horn-like horn-like tuberculated, spinules Few glandular warts Few Horny spinules in males, Horny or shagreen finely granular in female Horny spinules in males, Horny or shagreen finely granular in female Shagreen if females, glandular if females, Shagreen spinules ( and horny warts males) Glandular warts Shagreen or with glandular Shagreen warts spinules Horny Glandular warts Smooth, '\ /' shaped ridge Smooth Smooth Anterior dorsum Anterior Glandular warts Smooth, shagreen and Smooth, shagreen males horny glandular warts, spinules Glandular warts spinules Horny Glandular warts Few glandular Few warts Males horny Males horny spinules, females or finely granular shagreen Horny spinules in Horny males Shagreen Glandular warts Shagreen or with Shagreen glandular warts spinules Horny Side of head Glandular warts Shagreen Glandular warts Glandular warts Glandular warts Few glandular warts Few Horny spinules in Horny males, finely granular in female or shagreen Horny spinules in Horny males Shagreen Glandular warts Shagreen or with Shagreen glandular warts spinules Horny Interorbital area Interorbital Glandular warts
Glandular warts Glandular warts Horny spinules in Horny males, finely granular in female or shagreen Horny spinules in males in spinules Horny Smooth or tuberculated Shagreen Glandular warts Glandular warts Glandular warts or with Shagreen glandular warts spinules Horny Snout Glandular warts P. nasutus P. P. nanus P. P. poppiae P. P. mooreorum P. P. oxyrhynchus P. P. malcolmsmithi malcolmsmithi P. microtympanum P. pleurotaenia P. P. maia P. P. pardus P. P. mittermeieri mittermeieri P. P. procax P. P. popularis P. P. nemus P. ocularis P. regius P. Species name papillosus P.
Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3803 Eight new species of Pseudophilautus Wickramasinghe et al. Smooth, few Smooth, few scattered glandular warts Glandular warts Smooth Smooth Smooth Scattered Scattered glandular tubercles Smooth, scattered glandular warts Smooth Dorsal thigh Dorsal Glandular warts Smooth Smooth Smooth Smooth shagreen Weakly Smooth Granular Granular Granular Granular Granular Granular in females Granular Scattered, Scattered, glandular tubercles Granular Granular Lower Lower flank Glandular and warts horny spinules Granular Granular Granular Weakly Weakly areolate Granular Glandular warts Glandular warts, Glandular warts, spinules with horny in males Shagreen with a few with a few Shagreen glandular scattered warts Small tubercules Small tubercules spinules with horny Glandular folds Glandular warts Scattered, glandular Scattered, tubercles Glandular warts and Glandular warts spinules horny horny Shagreen, spinules in males Upper flank Glandular folds, Glandular folds, and glandular warts spinules horny Granular Granular Glandular warts weakly to Shagreen areolate Smooth except for a few a few for Smooth except warts glandular scattered Glandular warts, males Glandular warts, spinules scattered horny Shagreen with a few with a few Shagreen warts, glandular scattered finely tuberculated Small tubercules with Small tubercules spinules horny Glandular warts, males Glandular warts, spinules with horny scattered Smooth Scattered, glandular Scattered, tubercles Smooth and Glandular warts spinules, females horny spinules lack horny spinules horny Shagreen, in males spinules, females Horny spinules lack horny Posterior dorsum Posterior Glandular folds, glandular Glandular folds, spinules and horny warts Shagreen Glandular warts Glandular warts Smooth shagreen Weakly Smooth except for a few a few for Smooth except warts glandular scattered Glandular warts, males horny males horny Glandular warts, spinules scattered Shagreen with a few scattered scattered with a few Shagreen finely glandular warts, tuberculated Small tubercules with horny with horny Small tubercules spinules Glandular warts, males with Glandular warts, spinules scattered horny Glandular warts Scattered, glandular tubercles Scattered, Smooth and horny Glandular warts lack horny spinules, females spinules shagreen Weakly spinules in horny Shagreen, males lack spinules, females Horny spinules horny Anterior dorsum Anterior Glandular folds, glandular Glandular folds, spinules and horny warts Shagreen Glandular warts Glandular warts Glandular warts Smooth except for for Smooth except scattered a few glandular warts Glandular warts Shagreen with Shagreen scattered a few glandular warts Small tubercules Small tubercules spinules with horny Glandular warts Glandular warts Scattered, glandular Scattered, tubercles Side of head Glandular warts Shagreen Glandular warts Glandular warts Glandular warts shagreen Weakly Glandular warts and Glandular warts spinules horny Smooth except for for Smooth except scattered a few glandular warts Glandular warts Horny spinules Horny Shagreen with a few with a few Shagreen glandular scattered warts Small tubercules with Small tubercules spinules horny Glandular warts Glandular warts Scattered, glandular Scattered, tubercles Interorbital area Interorbital Glandular folds, Glandular folds, and glandular warts spinules horny Shagreen shagreen Weakly Glandular warts and Glandular warts spinules horny Smooth except for a for Smooth except glandular scattered few warts Glandular warts Weakly shagreen Weakly spinules Horny Shagreen with a few with a few Shagreen glandular scattered warts Small tubercules with Small tubercules spinules horny Glandular warts Glandular warts Scattered, glandular Scattered, tubercles Snout Glandular warts Shagreen steineri steineri stuarti stuarti P. rus P. P. P. rugatus P. P. semiruber P. P. simba P. P. sordidus P. P. stellatus P. P. P. reticulatus P. P. schneideri P. P. sarasinorum P. silvaticus P. P. silus P. P. singu P. Species name schmarda P. stictomerus P. tanu P.
3804 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Glandular warts Glandular warts Smooth Dorsal thigh Dorsal Shagreen Glandular warts Smooth Granular Granular Granular Lower Lower flank Granular Granular Coarsely Coarsely granular Granular Granular Granular Upper flank spinules Horny (smooth in females and immature males) Glandular warts Glandular warts Weakly tubercular Weakly Smooth Smooth Posterior dorsum Posterior
Horny spinules, Anterior spinules, Anterior Horny with “)(” shaped dorsum of tubercles pattern Glandular warts Horny spinules (smooth in Horny males) and immature females Weakly tubercular Weakly Smooth Smooth Anterior dorsum Anterior Glandular warts Glandular warts Horny spinules Horny (smooth in females and immature males) Shagreen Side of head
Horny spinules Horny Glandular warts Horny spinules Horny (smooth in females males) and immature Shagreen Interorbital area Interorbital
Horny spinules Horny Glandular warts Horny spinules Horny and (smooth in females males) immature Weakly shagreen Weakly Snout P. zorro P. P. zimmeri P. P. viridis P. P. temporalis temporalis P. P. zal P. P. variabilis P. Species name
Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3805 Eight new species of Pseudophilautus Wickramasinghe et al. 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 Figure 6. Pseudophilautus dayawansai sp. nov. holotype (NMSL male SVL 24.5mm (Images 8, 9A,D). Sripada (Peak 2013.02.01 NH), ventral aspect of left hand. Scale bar: 5mm. (© L.J. Wilderness), Ratnapura District, Sabaragamuwa Mendis Wickramasinghe) 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.
Figure 7. Pseudophilautus dayawansai sp. nov. holotype (NMSL 2013.02.01 NH), ventral aspect of left foot. Scale bar: 5mm. (© L.J. Mendis Wickramasinghe) Figure 5. Pseudophilautus dayawansai sp. nov. holotype (NMSL 2013.02.01 NH), head; (A) lateral aspect (B) dorsal aspect (C) ventral aspect. Scale bar: 2mm. (© Dulan Ranga Vidanapathirana)
3806 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. © L.J. Mendis Wickramasinghe
Image 9. Type series of Pseudophilautus dayawansai sp. nov. Left to right, NMSL 2013.02.01 NH, DWC 2013.01.003, DWC 2013.01.004. A-C dorsal aspect and D-F ventral aspect. Scale bar: 5mm. © L.J. Mendis Wickramasinghe © L.J. Mendis Wickramasinghe
Image 8. Pseudophilautus dayawansai sp. nov. holotype in life (NMSL 2013.02.01 NH).
Image 10. Pseudophilautus dayawansai sp. nov. paratype in life Diagnosis: Small sized frog (adult male SVL 24.5–26.2 (DWC 2013.01.003). mm, adult female SVL 30.1mm); head dorsally convex; snout laterally truncate, skin dorsally tuberculated, a single prominent tubercle medially placed; canthus medially; side of head tubercular; anterior dorsum rostralis rounded; loreal region, interorbital space and smooth; posterior dorsum weakly tubercular; median internarial space concave; vomerine teeth absent; lingual dermal ridge absent; upper and lower flank granular; papilla, fringe on fingers and blunt small calcar present; throat shagreen. interorbital area smooth, with a ridge across braking Description of holotype: Small sized frog (SVL
Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3807 Eight new species of Pseudophilautus Wickramasinghe et al. © L.J. Mendis Wickramasinghe © L.J. Mendis Wickramasinghe
Image 12. Pseudophilautus dayawansai sp. nov. unpreserved specimen.
Image 11. Pseudophilautus dayawansai sp. nov. paratype in life prominent, oval, single, all present, III2 and IV2 relatively (DWC 2013.01.004). smaller; inner palmar tubercle single, oval, larger than outer palmer tubercle; outer palmar tubercle single, indistinct; nuptial pads absent; supernumerary tubercles 24.5mm), elongate (SVL/HW 2.3); head large (HL/SVL present on all fingers, and a few indistinct ones on palm; 0.4), about as wide as long (HW/HL 1.0), dorsally convex; femur twice as long as the fourth toe length (FEL/TL-4 snout truncate in lateral aspect, mucronate in dorsal 2.1), foot length longer than thigh (FOL/FEL 1.5), toes aspect, and sub-elliptical in ventral aspect (Fig. 5 A) (ES/ long, thin (Fig. 7); relative length of toes I 3808 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Range 4.4—4.9 7.3—8.5 4.7—5.8 0.9—1.1 4.7—5.1 2.1—2.6 8.0—9.3 5.7—7.0 1.3—1.5 2.0—2.2 2.4—2.7 3.6—4.5 6.2—7.7 4.3—5.1 8.3—9.0 1.5—1.7 0.7—0.9 10.1—12.0 13.6—15.1 10.3—11.5 12.2—13.0 24.5—30.1 SD 0.2 0.6 0.6 0.1 0.2 0.3 1.0 0.7 0.7 0.1 0.7 0.1 0.1 0.4 0.8 0.4 0.3 0.6 0.4 0.1 2.9 0.1 4.6 7.8 5.2 1.0 5.0 2.4 8.7 6.5 1.4 2.1 2.5 4.1 7.1 4.7 8.7 1.6 0.8 11.2 14.4 10.8 12.7 26.9 Mean 4.5 7.7 5.1 0.9 5.0 2.1 9.0 7.0 1.3 2.2 2.6 4.5 7.7 4.3 8.3 1.5 0.7 11.6 14.6 10.3 12.8 26.2 Paratype 2 Paratype 2013.01.004 4.9 8.5 5.8 1.1 4.7 2.6 9.3 6.9 1.5 2.2 2.7 4.1 7.5 5.1 9.0 1.7 0.9 12.0 15.1 11.5 13.0 30.1 Paratype 1 Paratype 2013.01.003 4.4 7.3 4.7 1.1 5.1 2.5 8.0 5.7 2.0 2.4 3.6 1.3 6.2 4.7 8.8 1.5 0.8 10.1 13.6 10.6 12.2 24.5 Holotype 2013.02.01 NH KT SN LAL TBL SVL TAS MN VKL TYH UAL TPD TAD TL-5 TL-4 TL-1 TL-2 TL-3 TND MFE TYW MBE UEW DWC NMSL/ Range 5.2—6.4 1.2—1.4 2.7—3.3 2.8—3.6 2.0—2.4 7.6—9.1 2.8—3.7 5.6—6.8 4.5—5.1 0.5—0.9 5.0—6.5 1.1—1.4 1.1—1.6 4.0—4.8 2.3—3.0 3.8—4.9 9.4—10.8 10.5—12.4 12.1—14.7 12.0—13.4 10.5—12.2 18.1—20.1 SD 0.7 1.0 0.1 0.3 0.4 1.3 0.2 0.7 0.8 0.9 0.5 0.7 0.3 1.1 0.2 0.8 0.9 0.2 0.3 0.5 0.3 0.5 5.6 1.3 3.0 3.2 2.2 8.4 3.2 6.4 4.8 0.7 9.9 5.5 1.3 1.3 4.3 2.6 4.4 11.3 13.5 12.7 11.2 19.3 Mean 5.2 1.2 2.7 3.1 2.3 8.5 3.2 6.7 4.9 0.6 9.6 5.0 1.3 1.1 4.0 2.6 4.4 11.1 13.7 12.8 10.9 20.1 Paratype 2 Paratype 2013.01.004 6.4 1.4 3.3 3.6 2.4 9.1 3.7 6.8 5.1 0.9 6.5 1.4 1.6 4.8 3.0 4.9 12.4 14.7 13.4 12.2 19.7 10.8 Paratype 1 Paratype 2013.01.003 5.2 1.2 2.9 2.8 2.0 7.6 2.8 5.6 4.5 0.5 5.0 1.1 1.2 9.4 4.1 2.3 3.8 10.5 12.1 12.0 10.5 18.1 Holotype 2013.02.01 NH IN IO ES HL DL ED EN GK DB HD FEL DW HW FOL IML DFE DBE FL-1 FL-2 FL-3 FL-4 HNL DWC NMSL/ Table 4. The morphometric measurements (mm) of the type series Pseudophilautus dayawansai sp. nov. measurements 4. The morphometric Table Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3809 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Calcar Present Absent Absent Absent Absent Absent Fringe on Fringe fingers Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Lingual papilla Present Present Present Present Present Present Present Present Present Present Present Present Vomerine teeth Vomerine Absent Flat Flat Flat Flat Flat Flat Flat Flat Flat Internarial space Internarial Concave Flat Flat Flat Convex Flat Flat Convex Flat Flat Flat Convex Flat Flat Convex Flat Convex Convex Convex Convex Interorbital space Interorbital Concave Flat Flat Convex Loreal Loreal region Concave Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Canthus Canthus rostralis Rounded Rounded Oval Rounded Obtusely pointed Obtusely Blunt pointed Obtusely Obtusely pointed Obtusely Rounded Rounded Rounded or Truncate Rounded Rounded Pointed or oval Pointed Rounded Blunt Rounded Rounded Rounded Truncate Obtusely pointed Obtusely Rounded Sharply Pointed Snout lateral Truncate Concave or flat Concave Concave Flat Flat Flat Concave or flat Concave Flat Concave Flat Flat Flat Flat Concave Concave Flat Convex Head dorsally sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners Pseudophilautus dayawansai sp. nov. 32.9–42.9 32.9–42.9 38.4–48.9 21.7–26.4 21.7–26.4 16.3–19.0 33.4–46.8 33.4–46.8 18.9–27.3 18.9–27.3 26.7–29.3 26.7–29.3 17.0–27.5 26.9–44.5 26.9–44.5 23.7–29.4 23.7–29.4 25.4–37.0 25.4–37.0 33.3–41.7 35.3 43.5 23.4–28.8 38.4 33.6–51.1 22.3–32.5 18.3–23.9 18.3–23.9 44.8 16.2–19.2 20.4–24.6 35.8-40.4 17.6–28.9 17.6–28.9 24.5–30.1 17.3–20.2 SVL sp. sp. nov. sp. nov. sp. nov. hallidayi frankenbergi frankenbergi P. P. cavirostris cavirostris P. P. auratus P. caeruleus P. P. fulvus P. P. asankai P. P. P. alto P. P. fergusonianus fergusonianus P. P. folicola P. P. abundus P. adspersus P. P. eximius eximius P. P. extirpo P. P. femoralis femoralis P. P. newtonjayawardanei newtonjayawardanei P. sp. nov. puranappu P. sirilwijesundarai P. sp. nov. P. samarakoon samarakoon P. P. decoris P. dimbullae P. P. karunarathnai karunarathnai P. nov. P. jagathgunawardanai P. sp. nov. P. cuspis P. Species name P. dayawansai P. bambaradeniyai P. sp. nov. Table 5. Characters that differentiate differentiate that 5. Characters Table 3810 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Calcar Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Fringe on Fringe fingers Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Lingual papilla Present Present Present Present Present Present Present Present Present Present Vomerine teeth Vomerine Flat Flat Flat Flat Flat Flat Flat Flat Flat Internarial space Internarial Flat Flat Convex Flat Flat Convex or Convex concave Flat Flat Flat Flat Convex Flat Flat Flat Convex Flat Flat Convex Flat Flat Interorbital space Interorbital Flat Flat Flat Flat Loreal Loreal region Sharp Sharp Sharp Sharp Indistinct Sharp Sharp Sharp Sharp Canthus Canthus rostralis Sharp Obtusely pointed Obtusely Pointed Oval Rounded Rounded Pointed Oval Truncate or oval Truncate Rounded Pointed or oval Pointed Oval Pointed Rounded Rounded Blunt Acuminate Oval Oval pointed Obtusely Oval Rounded Pointed Pointed Blunt Snout lateral Rounded Flat Concave Flat Convex or Convex concave Flat Concave Flat Concave Flat Flat Flat Head dorsally Uneven 20.7 17.4 29.4–35.0 29.4–35.0 34.8 17.5 16.3–18.4 16.3–18.4 23.9–49.0 23.9–49.0 25.1–26.8 18.3–21.7 18.3–21.7 42.5–61.1 17.7–24.7 14.9 25.6–30.3 25.6–30.3 21.3–26.0 21.3–26.0 27.4–42.7 27.4–42.7 19.8 25.7 40.9 46.6 32.1 21.2–23.4 21.2–23.4 22.0–28.6 11.2–22.4 18.6 40 23.1–33.4 18.3–21.9 SVL 27.9 hoffmanni P. nemus P. P. nasutus P. P. mooreorum P. P. nanus P. P. rugatus P. P. mittermeieri mittermeieri P. P. microtympanum P. P. procax P. P. regius P. reticulatus P. P. popularis P. P. malcolmsmithi malcolmsmithi P. P. pleurotaenia pleurotaenia P. P. poppiae P. P. macropus P. P. leucorhinus leucorhinus P. limbus P. lunatus P. P. maia P. P. pardus P. P. hoipolloi P. P. hypomelas P. P. oxyrhynchus P. papillosus P. P. ocularis P. P. hankeni P. Species name P. halyi P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3811 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Calcar Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Fringe on Fringe fingers Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Lingual papilla Absent Present Present Present Present Present Present Present Present Present Vomerine teeth Vomerine Present Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Internarial space Internarial Flat Flat Flat Flat Convex Convex Convex Flat Flat Flat Flat Convex Flat Convex Flat Convex Interorbital space Interorbital Flat Flat Flat Flat Flat Flat Loreal Loreal region Sharp Sharp Sharp Sharp Sharp Sharp Indistinct Sharp Sharp Canthus Canthus rostralis Obtusely pointed Obtusely Obtusely pointed Obtusely Rounded Rounded Blunt Rounded Rounded Pointed Oval Blunt Rounded Oval Oval Blunt or rounded Blunt Pointed Rounded Obtusely pointed Obtusely Snout lateral Oval Rounded Flat Concave Flat Concave Flat Head dorsally Flat Flat 13.5–13.9 13.5–13.9 24.2–32.4 24.2–32.4 39.6–55.3 23.2–36.0 23.2–36.0 30.4–41.6 30.4–41.6 22.2–39.4 22.2–39.4 16.1–16.6 16.1–16.6 12.6–15.6 12.6–15.6 22.9–30.1 22.9–30.1 24.3–31.3 24.3–31.3 31.6 20.0–32.5 35.4–51.8 35.4–51.8 13.4 27.4–36.3 27.4–36.3 19.9–22.8 35.7 28.4–31.0 28.4–31.0 SVL 20.6–24.1 22.6–38.1 17.7–30.0 17.7–30.0 stuarti stuarti steineri steineri P. tanu P. P. P. stellatus P. P. stictomerus P. P. P. sordidus P. P. singu P. P. simba P. P. zorro P. P. silvaticus P. P. zimmeri P. P. zal P. P. silus P. P. semiruber P. P. viridis P. P. schneideri P. P. variabilis P. P. temporalis temporalis P. Species name P. rus P. sarasinorum P. P. schmarda P. 3812 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Shagreen Granular Smooth Throat Granular granular Weakly granular Weakly granular Coarsely granular Weakly Granular Smooth or granular Smooth Weakly granular Weakly Weakly granular Weakly Granular with Granular scattered glandular warts, smooth in females Granular Weakly Weakly granular Smooth or granular Lower flank Lower Weakly Weakly granular Smooth fine, Very horny spinules in Granular males Granular Shagreen Areolate Smooth or with glandular warts Upper flank Weakly Weakly tubercular Tubercular Smooth Smooth Smooth spinules Horny in males spinules Horny Glandular and warts spinules horny Smooth Absent Present Present Present Median dermal ridge Present Present Present Present Horny spinules Horny Weakly tubercular Weakly Horny spinules Horny in females) (Absent Shagreen Shagreen spinules in Horny in males, warty females fine, horny Very spinules Glandular warts spinules and horny lack horny (females spinules) spinules Horny Posterior dorsum Posterior Few horny spinules, horny Few tubercles Prominent Tubercular Shagreen tuberculated, Heavily glandular warts Smooth Spinulate Smooth Prominent tubercles tubercles Prominent spinules and horny Smooth Horny spinules Horny in females) (absent Shagreen with few Shagreen tubercles spinules in Horny in males, warty females fine, horny Very spinules Glandular warts spinules and horny lack horny (females spinules) spinules with a Horny of pattern “)(”shaped tubercles Anterior dorsum Anterior Weakly shagreen Weakly Tubercular Shagreen tuberculated, Heavily glandular warts Smooth Tubercular Smooth Shagreen shagreen Weakly for Smooth except scattered a few glandular warts Smooth fine, horny Very spinules and Glandular warts spinules horny Smooth Side of head Smooth Smooth Smooth Shagreen Glandular warts sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners Pseudophilautus dayawansai sp. nov. Smooth Smooth, with a ridge Smooth, with a ridge breaking across medially Smooth Shagreen, with a Shagreen, ridge prominent tubercle A prominent present for Smooth except scattered a few glandular warts spinules in Horny males fine, horny Very spinules and Glandular warts spinules horny spinules Horny Interorbital area Interorbital Smooth,with two Smooth,with two dermal fringes cross tubercular Weakly Smooth Tubercular Shagreen Glandular warts Smooth Tuberculated, a single Tuberculated, tubercle prominent medially placed Smooth Shagreen with Shagreen, dermal prominent fold for Smooth except scattered a few glandular warts spinules in Horny males fine, horny Very spinules and Glandular warts spinules horny spinules Horny Snout Smooth Shagreen Glandular warts sp. nov. sp. nov. sp. nov. sp. nov. P. bambaradeniyai P. sp. nov. P. puranappu P. P. dayawansai P. Species name abundus P. P. newtonjayawardanei newtonjayawardanei P. sp. nov. samarakoon P. sirilwijesundarai P. sp. nov. adspersus P. alto P. asankai P. caeruleus P. cuspis P. P. karunarathnai karunarathnai P. P. jagathgunawardanai P. sp. nov. auratus P. cavirostris P. Table 6. Characters in the skin structure that differentiate differentiate that in the skin structure 6. Characters Table Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3813 Eight new species of Pseudophilautus Wickramasinghe et al. Glandular, warty, warty, Glandular, or roughly granular Throat Weakly granular Weakly granular Weakly Smooth granular Weakly or Granular smooth Granular Granular Smooth, granular in males Granular Granular but not Granular, rough Smooth Smooth Granular Glandular warts Lower flank Lower Smooth Weakly Weakly granular Glandular Glandular warts Glandular warts Upper flank Smooth Shagreen Smooth Shagreen, males with glandular warts and horny spinules Shagreen Shagreen Glandular with warts, spinules horny in males Glandular bearing warts spinules horny Smooth Smooth or granular Smooth with glandular warts Tip of snout vent to Median dermal ridge Present Shagreen Glandular warts Finely granular glandular warts, Few males with glandular and horny warts spinules horny Shagreen, spinules scattered Shagreen with Glandular warts, spinules in horny males with Shagreen glandular warts Glandular warts bearing horny spinules Posterior dorsum Posterior Smooth with a few Smooth with a few glandular scattered warts Shagreen, with a few with a few Shagreen, glandular warts Smooth Smooth Smooth Smooth or shagreen Smooth Shagreen Glandular warts Finely granular Males Shagreen, and glandular warts spinules horny horny Shagreen, spinules scattered Shagreen with Glandular warts, spinules in horny males with Shagreen glandular warts Glandular warts bearing horny spinules Anterior dorsum Anterior Smooth with a few Smooth with a few glandular scattered warts Shagreen, with a few with a few Shagreen, glandular warts Shagreen Glandular warts Smooth Smooth Finely granular glandular warts Few smooth or Shagreen, scattered with a few glandular warts Glandular warts Shagreen Glandular warts with Shagreen glandular warts Glandular warts bearing horny spinules Smooth Side of head Smooth Shagreen Smooth or shagreen Shagreen Glandular warts Smooth Smooth Finely granular males Shagreen, and glandular warts spinules horny Smooth Shagreen Shagreen with Glandular warts, spinules in horny males with Shagreen glandular warts Glandular warts bearing horny spinules Smooth Interorbital area Interorbital Smooth Shagreen Smooth or shagreen Shagreen Glandular warts Smooth Smooth Finely granular males Shagreen, and glandular warts spinules horny Smooth Shagreen Shagreen with Glandular warts, spinules in horny males with Shagreen glandular warts Glandular warts bearing horny spinules Smooth Snout Smooth Shagreen, with a few with a few Shagreen, glandular warts Smooth or shagreen frankenbergi hallidayi hoffmanni P. dimbullae P. P. decoris P. Species name hypomelas P. P. eximius P. femoralis P. fergusonianus P. folicola P. P. fulvus P. P. halyi P. hankeni P. P. P. leucorhinus P. P. extirpo P. hoipolloi P. 3814 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Throat Granular Smooth Smooth or granular Granular granular Weakly Granular but not Granular rough or with Granular glandular folds Granular granular Weakly Granular Granular Smooth Granular Granular Smooth Lower flank Lower Glandular warts Glandular warts Glandular warts Horny spinules in males Smooth Smooth Upper flank Glandular warts Glandular warts Glandular warts Glandular warts spinules Horny in males Glandular folds Smooth or Shagreen glandular warts Smooth Glandular warts Shagreen, scattered few glandular and warts, spinules horny Tip of snout vent to Median dermal ridge Weakly granular Weakly Shagreen or smooth, Shagreen scattered with a few glandular warts Glandular warts Glandular warts horn Tuberculated, spinules like Glandular warts or with Shagreen glandular warts few Shagreen, glandular scattered and horny warts spinules (males) Posterior dorsum Posterior Smooth Smooth above, and with shagreen glandular warts, males with horn like spinules spinules in Horny males, finely granular in female or shagreen glandular warts Few Glandular warts Smooth Smooth Smooth Weakly granular Weakly Shagreen or smooth, Shagreen scattered with a few glandular warts Glandular warts Glandular warts horn Tuberculated, spinules like Glandular warts or with Shagreen glandular warts with a few Shagreen glandular scattered and horny warts spinules ( males) Anterior dorsum Anterior Smooth, '\ /' shaped ridge Smooth above, and with shagreen glandular warts, males with horn like spinules spinules in Horny males, finely granular in female or shagreen glandular warts Few Glandular warts Weakly granular Weakly Smooth Glandular warts Smooth Glandular warts Glandular warts Glandular warts or with Shagreen glandular warts Smooth Smooth Shagreen Side of head Smooth Shagreen spinules in Horny males glandular warts Few Glandular warts Weakly granular Weakly Smooth Smooth with a few glandular scattered warts Smooth Glandular warts Glandular warts Smooth or with Shagreen glandular warts Smooth Smooth Shagreen Interorbital area Interorbital Smooth Smooth or tuberculated spinules in Horny males glandular warts Few Glandular warts Weakly granular Weakly Smooth Smooth Smooth Glandular warts Glandular warts Glandular warts or with Shagreen glandular warts Smooth Smooth Shagreen Snout Smooth Smooth or tuberculated spinules in Horny males Glandular warts Glandular warts P. limbus P. Species name P. lunatus P. macropus P. maia P. mittermeieri P. nasutus P. nemus P. ocularis P. oxyrhynchus P. pardus P. pleurotaenia P. P. malcolmsmithi P. microtympanum P. mooreorum P. nanus P. papillosus P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3815 Eight new species of Pseudophilautus Wickramasinghe et al. Granular Throat Granular granular Coarsely Smooth or granular granular Weakly Granular Granular Granular in Granular males, smooth in females granular Weakly Smooth Granular Glandular warts Glandular warts Granular Smooth Lower flank Lower Smooth Glandular in Granular females Glandular and warts horny spinules Scattered, glandular tubercles Horny spinules Horny in males, finely or granular in shagreen female Glandular warts Upper flank Shagreen with Shagreen scattered a few glandular warts Smooth Horny spinules Horny Smooth Glandular Glandular folds Glandular glandular folds, and warts spinules horny Small with tubercules spinules horny Glandular warts Glandular warts Scattered, glandular tubercles Median dermal ridge Thin, tip of snout to vent Horny spinules Horny Horny spinules in Horny males, finely granular in female or shagreen Horny spinules Horny in (spinules Absent females) spinules Horny for Smooth except scattered a few glandular warts Glandular warts Glandular warts, males with horny spinules scattered Smooth Smooth glandular Scattered, tubercles Posterior dorsum Posterior Glandular warts Shagreen with a few with a few Shagreen glandular scattered finely warts, tuberculated Glandular folds, and glandular warts spinules horny with Small tubercules spinules horny Smooth Horny spinules Horny Horny spinules in Horny males, finely granular in female or shagreen Horny spinules Horny in (spinules Absent females) spinules Horny for Smooth except scattered a few glandular warts Glandular warts Glandular warts, males with horny spinules scattered Glandular warts Glandular warts glandular Scattered, tubercles Anterior dorsum Anterior Glandular warts Shagreen with a few with a few Shagreen glandular scattered finely warts, tuberculated Glandular folds, and glandular warts spinules horny with Small tubercules spinules horny Glandular warts Horny spinules in Horny males, finely granular in female or shagreen Smooth spinules Horny for Smooth except scattered a few glandular warts Glandular warts Glandular warts Smooth Glandular warts Glandular warts glandular Scattered, tubercles Side of head Glandular warts Shagreen with a few with a few Shagreen glandular scattered warts Glandular warts with Small tubercules spinules horny Smooth Horny spinules in Horny males, finely granular in female or shagreen Smooth spinules Horny for Smooth except scattered a few glandular warts Smooth Glandular warts Smooth Glandular warts Smooth glandular Scattered, tubercles Interorbital area Interorbital Smooth Shagreen with a few with a few Shagreen glandular scattered warts Glandular folds, and glandular warts spinules horny with Small tubercules spinules horny Glandular warts Horny spinules in Horny males, finely granular in female or shagreen Smooth spinules Horny for Smooth except scattered a few glandular warts Smooth Glandular warts Smooth Glandular warts Smooth glandular Scattered, tubercles Snout Smooth Shagreen with a few with a few Shagreen glandular scattered warts Glandular warts with Small tubercules spinules horny P. popularis P. P. poppiae P. Species name P. procax P. regius P. rugatus P. rus P. sarasinorum P. semiruber P. silvaticus P. simba P. singu P. P. silus P. P. reticulates P. schmarda P. schneideri P. 3816 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Granular Throat Smooth Granular granular Weakly Granular Granular Granular Granular Smooth Granular Smooth Lower flank Lower Weakly Weakly areolate Smooth Coarsely granular Glandular and warts spinules horny Glandular warts Upper flank Glandular with warts, spinules horny in males Shagreen weakly to areolate Shagreen, spinules horny in males Smooth Smooth spinules Horny (smooth in and females immature males) Tip of snout posterior to dorsum Tip of snout vent to (narrow) Median dermal ridge Tip of snout to (Very vent narrow) Present Glandular warts Glandular warts, Glandular warts, spinules males horny scattered Shagreen, horny horny Shagreen, spinules in males spinules, Horny lack horny females spinules Glandular warts Glandular warts spinules, and horny lack horny females spinules spinules Horny (smooth in females males) and immature spinules Horny Posterior dorsum Posterior Shagreen Weakly shagreen Weakly Smooth Smooth Glandular warts Glandular warts, Glandular warts, spinules males horny scattered Shagreen, horny horny Shagreen, spinules in males spinules, Horny lack horny females spinules Glandular warts Glandular warts spinules, and horny lack horny females spinules spinules Horny (smooth in females males) and immature tubercular Weakly spinules, Horny with dorsum Anterior “)(”–shaped pattern of tubercles Anterior dorsum Anterior Shagreen Weakly shagreen Weakly Glandular warts Glandular warts Smooth or with glandular warts Smooth Smooth or with glandular warts Smooth spinules Horny (smooth in females males) and immature Shagreen Glandular warts Side of head Shagreen Smooth Weakly shagreen Weakly Glandular warts Glandular warts Smooth spinules Horny Glandular warts and Glandular warts spinules horny Smooth spinules Horny (smooth in females males) and immature Shagreen spinules Horny Interorbital area Interorbital Shagreen Smooth Weakly shagreen Weakly Glandular warts Glandular warts Smooth spinules Horny Glandular warts and Glandular warts spinules horny shagreen Weakly Smooth spinules Horny (smooth in females males) and immature shagreen Weakly spinules Horny Snout Shagreen Smooth stuarti steineri P. zimmeri P. P. sordidus P. Species name stictomerus P. P. P. stellatus P. temporalis P. viridis P. zal P. zorro P. P. tanu P. variabilis P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3817 Eight new species of Pseudophilautus Wickramasinghe et al. and dorsal thigh smooth; dorsally leg weakly tubercular; Range (Manamendra-Arachchi & Pethiyagoda 2005), tarsus weakly tubercular; foot smooth. but the specimen’s locality Dimbula is closer to the Peak Ventral side of body: Throat shagreen; chest weakly Wilderness, hence given the geographical similarity and granular; belly granular; upper arm weakly granular; the unique marking we feel that this specimen could be forearm smooth; thigh weakly granular; leg smooth. Pseudophilautus dayawansai. 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 Pseudophilautus jagathgunawardanai sp. nov. anteriorly, its center with a large light brown triangle, (Figs. 8–10; Images 13–20; Tables 7–9) prominent, symmetrically placed on dorsum with the urn:lsid:zoobank.org:act:A5A913CB-9D12-4024-9776-F8E904F3887E base on sacrul hump pointing towards head, its margins blackish-brown and outer area dark brown, three Holotype: NMSL 2013.03.01 NH, 05.ii.2009, adult prominent blackish-brown circular spots, two between female 35.8mm SVL (Image 13, 14A,D). Sripada (Peak eyes, and right in front another circular dark brown spot, Wilderness), Ratnapura District, Sabaragamuwa posterior part from sacrum light brown; lateral body, Province, Sri Lanka (06048’31”N & 80029’10”E; 1642m) and behind eyes dark brown with an olive green tinge; (Fig. 1), coll. L.J.M.W, D.R.V., M.D.G.R., S.C.A., A.W.A.C., limbs dark brown, forelimbs, hind limbs, fingers and toes L.L.D.P. & I.N.B. with maroon cross bands, hind limbs laterally olive green Paratypes: DWC 2013.01.005, adult female, SVL tinge; ventrally light brown with dark brown blotching, 40.4mm (Image 14 B&E); DWC 2013.01.006, adult hands, feet and webbing darker. female, SVL 36.3 mm (Image 14 C&F, 15); data same as Colour in alcohol: Colour pattern remains with a little holotype. fading, maroon to dark brown, venter light brown to off- Diagnosis: Pseudophilautus jagathgunawardanai white with darker marbling, green tinge absent. sp. nov., can be distinguished from known congeners by Paratype and variation:Dorsum colour varies ranging the following combination of characters: Body moderate from a reddish-brown (Image 10) to a lighter brown size (SVL adult female 35.8–40.4 mm); head dorsally, shades (Image 12); in the lighter coloured specimens the loreal region, and interorbital space convex; lateral dorsal colour of foot and hand light yellowish, and limbs snout rounded; canthus rostralis sharp; internarial were cream with brown cross bands. space concave; vomerine teeth, fringe on fingers, Etymology: The species epithet dayawansai is rudimentary webbing on fingers, calcar, and nuptial named after Nihal Dayawansa, Senior Lecturer of the pad absent; lingual papilla present; snout, side of head, Department of Zoology, University of Colombo in anterior dorsum, posterior dorsum, upper flank smooth; recognition of his dedicated service and commitment interorbital area smooth, with two cross dermal fringes; towards amphibian research work in Sri Lanka. The throat and chest weakly granular; underside of thigh species name dayawansai is a noun in the genitive granular; supernumerary tubercles on palm and foot case. present. Comparative details between congeners in Suggested vernacular names: Dayawansage tables 8 and 9. panduru madiya in Sinhala and Dayawansa’s Shrub Frog Description of holotype: Body moderate size (SVL in English. 35.8), elongate (SVL/HW 2.2); head large (HL/SVL 0.5), as Natural History: The species was found in cloud wide as long (HW/HL 1.0), convex above; snout rounded forests of elevation 1550–1900 m above sea level from in lateral and ventral aspects, truncate in dorsal aspect forest floor to about 2m high shrubs. (Fig. 8 A–C) (ES/DFE 0.8, SN/IN 0.6) its length slightly Remarks: Pseudophilautus dayawansai sp. nov., longer than the horizontal diameter of eye (ES/ED 1.1); possesses a unique colour pattern which is not internasal space concave; canthus rostralis sharp, loreal possessed by any of its congeners, hence can easily be region and interorbital space convex, upper eyelid smaller identified. In Dutta & Manamendra-Arachchi 1996: “Fig. than interorbital distance (IO/UEW 1.2), internasal 232. Rhacophorus macropus MCZ 20873 Queenwood, distance equal to upper eyelid (IN/UEW 1.0); distance Dimbula, Sri Lanka” in page 194, closely ressembles between front of eyes almost half the distance between Pseudophilautus dayawansai from its unique mark back of eyes (DBE/DFE 1.9); nostrils oval without flap on head; dark brown marking, forming a clear of skin laterally, closer to tip of snout than to eye (SN/ semicircular blackish margin. Rhacophorus macropus EN 0.6); pupil horizontally elliptical; tympanum distinct, (Pseudophilautus macropus) is restricted to the Knuckles oval (TYH/TYW 1.4), one third the eye diameter (TYH/ 3818 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Figure 8. Pseudophilautus jagathgunawardanai sp. nov. holotype (NMSL 2013.03.01 NH), head; (A) lateral aspect (B) dorsal aspect (C) ventral aspect. Scale bar: 2mm. (© Dulan Ranga Vidanapathirana) Figure 9. Pseudophilautus jagathgunawardanai sp. nov. holotype (NMSL 2013.03.01 NH), ventral aspect of right hand. Scale bar: 5mm (© L.J. Mendis Wickramasinghe) © L.J. Mendis Wickramasinghe Image 13. Pseudophilautus jagathgunawardanai sp. nov. holotype in life (NMSL 2013.03.01 NH). 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 Figure 10. Pseudophilautus jagathgunawardanai sp. nov. holotype than upper arm (LAL/UAL 1.4); fore-arm distinctly (NMSL 2013.03.01 NH), ventral aspect of right foot. Scale bar: 5mm. (© L.J. Mendis Wickramasinghe) enlarged; fingers thin, (Fig. 9), relative length of fingers I Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3819 Eight new species of Pseudophilautus Wickramasinghe et al. © L.J. Mendis Wickramasinghe Image 14. Type series of Pseudophilautus jagathgunawardanai sp. nov. Left to right, NMSL 2013.03.01 NH, DWC 2013.01.005, DWC 2013.01.006. A-B dorsal aspect and C-D ventral aspect. Scale bar: 5mm. © L.J. Mendis Wickramasinghe © L.J. Mendis Wickramasinghe Image 16. Pseudophilautus jagathgunawardanai sp. nov. unpreserved specimen. < II < IV < III (FL-1/FL-3 0.5, FL-2/FL-3 0.6, FL-4/FL-3 0.7); Image 15. Pseudophilautus jagathgunawardanai sp. nov. paratype tips of fingers rounded enlarged, discs present on all in life (DWC 2013.01.006). 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 3820 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. © L.J. Mendis Wickramasinghe © L.J. Mendis Wickramasinghe Image 17. Pseudophilautus jagathgunawardanai sp. nov. unpreserved specimen. Image 18. Pseudophilautus jagathgunawardanai sp. nov. unpreserved specimen. © L.J. Mendis Wickramasinghe © L.J. Mendis Wickramasinghe Image 19. Pseudophilautus jagathgunawardanai sp. nov. unpreserved specimen. Image 20. Pseudophilautus jagathgunawardanai sp. nov. unpreserved specimen. indistinct; inner palmar tubercle single, oval, prominent, larger than outer palmer tubercle; nuptial pads absent; outer palmar tubercle single, oval, indistinct, smaller smaller; supernumerary tubercles concentrated on than the distal subarticular tubercles; supernumerary foot, present on all toes; inner metatarsal tubercle oval tubercles present on all fingers, and palm; prepollex prominent and large, its length 2/3 times the length of absent; femur twice as long as the fourth toe length toe I (IML/TL-1 0.6); outer metatarsal tubercle absent. (FEL/TL-4 2.1); foot length longer than thigh (FOL/FEL Skin on dorsal and lateral part of snout smooth; 1.5); toes thin, (Fig. 10), relative length of toes I < II < V < interorbital space smooth, with two cross dermal fringes III Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3821 Eight new species of Pseudophilautus Wickramasinghe et al. Range 1.7—2.3 1.2—1.6 6.6—7.1 7.2—8.0 3.7—3.8 3.1—3.4 3.5—4.1 5.7—6.5 4.9—6.7 8.9—9.6 6.3—7.5 2.1—2.2 1.2—1.4 9.5—10.1 16.9—18.2 16.9—18.4 12.2—13.3 11.5—12.9 19.5—20.6 12.3—15.9 10.2—11.2 35.8—40.4 SD 0.3 0.2 0.3 0.7 0.3 0.4 0.1 0.8 0.6 0.7 0.5 0.2 0.3 0.4 1.8 0.9 0.4 0.5 0.6 0.1 2.5 0.1 2.0 1.5 6.8 9.8 7.7 3.8 3.2 3.7 6.1 5.7 9.2 7.0 2.1 1.3 17.6 17.5 12.6 12.2 20.1 14.2 10.7 37.5 Mean 2.0 1.6 6.6 9.9 8.0 3.8 3.1 3.6 5.7 6.7 8.9 6.3 2.1 1.4 16.9 17.1 12.2 12.9 19.5 15.9 10.2 36.3 Paratype 2 Paratype 2013.01.006 2.3 1.6 6.7 7.2 3.7 3.4 4.1 6.5 5.5 9.6 7.5 2.2 1.4 18.2 18.4 13.3 10.1 11.5 20.3 12.3 11.2 40.4 Paratype 1 Paratype 2013.01.005 7.1 1.2 7.9 3.8 9.5 3.3 3.5 6.1 4.9 9.2 7.2 2.1 1.7 1.2 17.6 16.9 12.5 12.1 20.6 14.5 10.6 35.8 Holotype 2013.03.01 NH KT SN LAL TBL SVL TAS MN VKL TYH UAL TAD TPD TL-1 TL-2 TL-3 TL-5 TL-4 TND MFE TYW MBE DWC UEW NMSL/ Range 1.2—1.3 1.7—1.9 6.9—7.6 1.8—2.2 3.4—3.9 4.2—4.5 3.1—3.6 1.9—2.3 3.8—4.4 7.4—7.9 4.7—6.2 7.2—8.2 5.1—5.5 3.4—4.5 5.8—6.1 18.9—20.6 15.5—16.3 12.9—14.2 15.9—17.4 27.1—28.8 16.2—17.5 11.1—12.8 SD 0.0 0.9 0.1 0.4 0.7 0.4 0.2 0.2 0.2 0.2 0.2 0.8 0.3 0.3 0.8 0.9 0.5 0.7 0.2 1.0 0.6 0.1 1.2 1.8 1.9 3.7 7.3 4.4 3.3 2.0 4.2 7.6 5.3 7.7 5.3 3.9 5.9 19.5 16.0 13.7 16.6 28.2 16.8 12.2 Mean 1.2 1.7 2.2 3.4 6.9 4.5 3.2 2.0 3.8 7.9 6.2 7.2 5.1 3.9 5.8 18.9 15.5 12.9 16.5 27.1 17.5 11.1 Paratype 2 Paratype 2013.01.006 1.3 1.9 1.9 3.9 7.5 4.2 3.1 2.3 4.4 7.4 4.7 7.7 5.3 4.5 6.1 20.6 16.3 14.2 15.9 28.8 16.6 12.8 Paratype 1 Paratype 2013.01.005 1.2 1.9 1.8 3.7 7.6 4.4 3.6 1.9 4.4 7.5 4.9 8.2 5.5 3.4 5.8 19.2 16.2 14.0 17.4 28.6 16.2 12.7 Holotype 2013.03.01 NH IN IO ES DL HL ED EN GK DB HD FEL DW HW FOL IML DFE DBE FL-1 FL-2 FL-3 FL-4 HNL DWC NMSL/ Table 7. The morphometric measurements (mm) of the type series Pseudophilautus jagathgunawardanai sp. nov. measurements 7. The morphometric Table 3822 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Nuptial pad Nuptial Present Present Present Present Present Absent Calcar Present Present Present Present Present Present Absent Fingers Fingers Rudimentary webbing Present Present Present Present Present Present Present III Fingers and IV Present Present Present Present Present Fingers 1 Fingers and 2 Absent Fringe on Fringe fingers Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Lingual papilla Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Vomerine Vomerine teeth Present Present Present Present Present Present Present Present Concave Internarial Internarial space Flat Flat Flat Flat Flat Flat Flat Flat Convex Concave Interorbital Interorbital space Flat Flat Flat Concave Concave Flat or Concave flat Concave or Concave flat Flat Concave Flat Flat Concave Convex Concave Loreal Loreal region Concave Flat Concave Concave Concave Concave Concave Flat Concave Concave Concave Concave Concave Concave or Concave flat Concave Concave or Concave flat Concave Concave Concave Sharp Rounded Rounded or sharp Canthus Canthus rostralis Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Obtusely Obtusely pointed Snout lateral Pointed or Pointed oval Truncate Blunt Obtusely pointed Oval Blunt Truncate Truncate Truncate Truncate Obtusely pointed Obtusely pointed Sharply Pointed Truncate Convex Flat Head dorsally Concave Concave Flat Flat Flat Concave Flat Flat Flat Concave or flat Concave sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners Pseudophilautus jagathgunawardanai sp. nov. 17.3–20.2 35.8-40.4 18.9–27.3 18.3–23.9 SVL 38.4 20.4–24.6 22.3–32.5 25.4–37.0 33.3–41.7 17.0–27.5 21.7–26.4 16.3–19.0 38.4–48.9 17.6–28.9 44.8 35.3 43.5 23.4–28.8 26.9–44.5 24.5–30.1 16.2–19.2 33.6–51.1 sp. nov. sp. nov. sp. nov. sp. nov. P. bambaradeniyai P. sp. nov. P. jagathgunawardanai P. sp. nov. P. asankai P. decoris P. P. newtonjayawardanei newtonjayawardanei P. sp. nov. samarakoon P. Species name dayawansai P. karunarathnai P. puranappu P. sirilwijesundarai P. sp. nov. abundus P. adspersus P. alto P. auratus P. caeruleus P. cavirostris P. cuspis P. dimbullae P. eximius P. extirpo P. femoralis P. fergusonianus P. Table 8. Characters that differentiate differentiate that 8. Characters Table Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3823 Eight new species of Pseudophilautus Wickramasinghe et al. Nuptial pad Nuptial Present Present Present Present Present Present Present Present Present Calcar Present Present Present Present. Present Present Present Present Fingers Fingers Rudimentary webbing Present Present Present Present Present Present Present Fringe on Fringe fingers Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Lingual papilla Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Vomerine Vomerine teeth Present Present Present Present Present Present Present Present Present Present Present Internarial Internarial space Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Interorbital Interorbital space Concave Concave Flat Flat Flat Flat Concave Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Concave Concave Flat Flat Concave Concave Loreal Loreal region Concave Concave Concave Concave Flat Concave Concave Concave Concave Concave Concave Concave Concave Flat Concave Flat Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Flat Concave Canthus Canthus rostralis Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Indistinct Rounded Rounded Snout lateral Oval Obtusely pointed Oval Pointed Truncate Blunt Pointed Blunt Acuminate Oval Pointed Oval Pointed Oval Pointed Obtusely pointed Truncate Truncate Flat Head dorsally Flat Flat Flat Concave Flat Flat Flat Uneven Concave Flat 23.7–29.4 32.1 SVL 26.7–29.3 33.4–46.8 21.2–23.4 22.0–28.6 11.2–22.4 19.8 25.7 40.9 27.4–42.7 14.9 23.9–49.0 29.4–35.0 34.8 20.7 18.6 40 25.6–30.3 21.3–26.0 17.7–24.7 27.9 18.3–21.9 46.6 16.3–18.4 17.4 23.1–33.4 32.9–42.9 P. folicola P. P. pardus P. Species name frankenbergi P. fulvus P. hoffmanni P. hoipolloi P. hypomelas P. leucorhinus P. limbus P. lunatus P. macropus P. malcolmsmithi P. microtympanum P. mooreorum P. nanus P. nemus P. oxyrhynchus P. papillosus P. pleurotaenia P. poppiae P. popularis P. P. halyi P. hankeni P. maia P. mittermeieri P. nasutus P. ocularis P. P. hallidayi P. 3824 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Nuptial pad Nuptial Present Present Present Present Present Present Present Present Calcar Present Present Present Present Fingers Fingers Rudimentary webbing Present Present Present Present Present Fringe on Fringe fingers Present Present Present Present Present Present Present Present Present Present Present Present Lingual papilla Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Vomerine Vomerine teeth Present Present Present Present Present Present Present Present Present Present Present Present Internarial Internarial space Flat Flat Flat Flat Concave or Convex concave Flat Flat Flat Concave Flat Flat Flat Flat Flat Interorbital Interorbital space Concave Flat Flat Concave or Concave flat Flat Flat Flat Concave Concave Flat Flat Flat Flat Flat Flat Concave Loreal Loreal region Concave Concave Concave Flat Flat Concave Concave Flat Concave Concave Concave Concave Concave Concave Concave Concave Flat Concave Concave Concave Concave Concave Concave Concave Rounded Canthus Canthus rostralis Rounded Rounded Rounded Rounded Rounded Rounded Rounded Rounded Indistinct Rounded Rounded Rounded Rounded Pointed or Pointed oval Snout lateral Pointed Oval Oval Truncate Truncate or Truncate oval Obtusely pointed Pointed Oval Oval Oval Truncate Blunt Obtusely pointed Obtusely pointed or Blunt rounded Blunt Concave Head dorsally Flat Concave Flat Flat Flat Flat Flat Concave 25.1–26.8 17.5 SVL 42.5–61.1 24.3–31.3 39.6–55.3 24.2–32.4 31.6 22.9–30.1 18.3–21.7 20.6–24.1 22.6–38.1 17.7–30.0 19.9–22.8 13.4 35.4–51.8 12.6–15.6 16.1–16.6 22.2–39.4 30.4–41.6 23.2–36.0 13.5–13.9 28.4–31.0 35.7 27.4–36.3 20.0–32.5 P. procax P. P. rugatus P. P. reticulatus P. silvaticus P. stellatus P. stuarti P. zimmeri P. zorro P. Species name regius P. rus P. sarasinorum P. schmarda P. schneideri P. semiruber P. silus P. simba P. singu P. sordidus P. steineri P. stictomerus P. tanu P. temporalis P. variabilis P. viridis P. zal P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3825 Eight new species of Pseudophilautus Wickramasinghe et al. Present Present Supernumerary Supernumerary on tubercles foot Indistinct Absent Absent Absent Present Supernumerary Supernumerary on tubercles palm Absent Absent Granular Underside Underside of thigh Smooth Weakly Weakly granular Chest Weakly Weakly granular Shagreen Throat Shagreen Smooth Shagreen or Granular smooth Areolate Horny spinules Upper flank Granular Granular Weakly granular Weakly Weakly tubercular Tubercular Horny spinules in males Smooth Shagreen Shagreen Shagreen fine, horny Very spinules Posterior Posterior dorsum Weakly Weakly tubercular Spinulate Horny spinules Horny horny Few spinules, Prominent tubercles Tubercular spinules Horny in (Absent females) spinules Horny in males, warty in females Smooth Smooth a for except scattered few glandular warts Shagreen Shagreen Shagreen with Shagreen tubercles few fine, horny Very spinules Anterior Anterior dorsum Prominent Prominent and tubercles spinules horny Weakly shagreen Tubercular spinules Horny in (absent females) spinules Horny in males, warty in females Smooth Smooth except a few for scattered glandular warts Shagreen Shagreen Tubercular Weakly tubercular Weakly shagreen fine, Very horny spinules Side of head sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners Pseudophilautus jagathgunawardanai sp. nov. Smooth, with dermal cross two fringes Smooth Smooth Smooth a for except scattered few glandular warts Shagreen, with a Shagreen, ridge prominent Shagreen Smooth, with across a ridge medially braking Weakly tubercular Smooth A prominent present tubercle spinules in Horny males fine, horny Very spinules Interorbital area Interorbital Tubercular Smooth Smooth except for for Smooth except scattered a few glandular warts Shagreen Shagreen Tuberculated, with Tuberculated, a single prominent medially tubercle placed Smooth, a blunt prominent present tubercle Smooth, with a tubercle blunt with Shagreen, dermal prominent fold spinules in Horny males fine, horny Very spinules Snout Prominent tubercle tubercle Prominent on centre present of snout sp. nov. sp. nov. sp. nov. P. bambaradeniyai P. sp. nov. P. jagathgunawardanai P. sp. nov. P. samarakoon samarakoon P. adspersus P. P. newtonjayawardanei newtonjayawardanei P. sp. nov. auratus P. Species name dayawansai P. karunarathnai P. sp. nov. puranappu P. sirilwijesundarai P. sp. nov. abundus P. alto P. asankai P. Table 9. Characters in the skin structure that differentiate differentiate that in the skin structure 9. Characters Table 3826 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Supernumerary Supernumerary on tubercles foot Absent Absent Absent Absent Absent Supernumerary Supernumerary on tubercles palm Absent Underside Underside of thigh Chest Smooth Smooth Throat Smooth Smooth Smooth Glandular and warts horny spinules Glandular warts Shagreen Shagreen Shagreen, Shagreen, males with glandular and warts horny spinules Shagreen Granular Finely granular Glandular warts, with horny spinules in males Upper flank Granular Glandular and warts spinules horny lack (females spinules) horny Glandular warts Shagreen, spinules horny scattered Shagreen Few glandular Few males warts, with glandular and warts spinules horny Heavily Heavily tuberculated, glandular warts Shagreen with Shagreen, glandular a few warts Finely granular Glandular with warts, spinules horny in males Posterior Posterior dorsum Horny spinules Horny Glandular and warts spinules horny lack (females spinules) horny Glandular warts Shagreen, spinules horny scattered Shagreen Shagreen, Shagreen, Males glandular and warts spinules horny Heavily Heavily tuberculated, glandular warts Shagreen with Shagreen, glandular a few warts Finely granular Glandular with warts, spinules horny in males Anterior Anterior dorsum Horny Horny spinules with a “)(”shaped of pattern tubercles Glandular and warts horny spinules Shagreen, Shagreen, smooth or with a few scattered glandular warts Glandular warts Glandular warts Shagreen Few Few glandular warts Glandular warts Shagreen Shagreen Finely granular Glandular warts Side of head Glandular warts Glandular warts and horny spinules Smooth Smooth Glandular warts Shagreen Shagreen Shagreen, males Shagreen, glandular warts and horny spinules Glandular warts spinules Horny Shagreen Shagreen Finely granular Glandular warts, with horny spinules in males Interorbital area Interorbital Glandular warts and warts Glandular spinules horny Glandular warts Shagreen Shagreen Shagreen, males Shagreen, and glandular warts spinules horny Glandular warts spinules Horny Shagreen with a Shagreen, glandular warts few Finely granular Glandular warts, spinules with horny in males Snout P. caeruleus P. P. decoris P. frankenbergi P. fulvus P. P. fergusonianus fergusonianus P. Species name cavirostris P. cuspis P. dimbullae P. eximius P. extirpo P. femoralis P. folicola P. hallidayi P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3827 Eight new species of Pseudophilautus Wickramasinghe et al. Supernumerary Supernumerary on tubercles foot Absent Absent Absent Absent Absent Supernumerary Supernumerary on tubercles palm Absent Absent Absent Underside Underside of thigh Smooth Smooth Smooth Smooth Chest Smooth Smooth Shagreen Smooth Throat Smooth Smooth Shagreen Smooth Glandular warts Granular Granular with a few scattered glandular warts Granular Glandular bearing warts horny spinules Granular Glandular warts Horny spinules in males Upper flank Glandular warts Glandular warts Smooth with glandular warts Glandular warts Shagreen with Shagreen glandular warts Shagreen or Shagreen smooth, with a scattered few glandular warts Smooth above, and shagreen with glandular males warts, with horn like spinules Glandular warts Glandular warts bearing horny spinules granular Weakly Glandular warts spinules Horny in males, finely or granular in shagreen female Posterior Posterior dorsum Smooth with a scattered few glandular warts Glandular warts Shagreen with Shagreen glandular warts Shagreen or Shagreen smooth, with a scattered few glandular warts Smooth above, and shagreen with glandular males warts, with horn like spinules Glandular warts Glandular warts bearing horny spinules Weakly granular Glandular warts spinules Horny in males, finely or granular in shagreen female Anterior Anterior dorsum Smooth with a scattered few glandular warts Glandular warts Smooth, '\ /' shaped ridge Shagreen Shagreen with glandular warts Glandular warts Shagreen Glandular warts bearing horny spinules Weakly granular Glandular warts Horny spinules in males Side of head Shagreen with Shagreen glandular warts Smooth Smooth or shagreen Smooth Smooth Smooth with a scattered few glandular warts Smooth Smooth Smooth or tuberculated Glandular warts Glandular warts bearing horny spinules granular Weakly Glandular warts spinules in Horny males Interorbital area Interorbital Shagreen with Shagreen glandular warts Glandular warts Glandular warts bearing horny spinules granular Weakly Glandular warts spinules in Horny males Snout P. halyi P. P. leucorhinus leucorhinus P. P. hypomelas hypomelas P. maia P. malcolmsmithi P. Species name hankeni P. hoffmanni P. hoipolloi P. limbus P. lunatus P. macropus P. microtympanum P. mittermeieri P. mooreorum P. 3828 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Supernumerary Supernumerary on tubercles foot Absent Absent Absent Supernumerary Supernumerary on tubercles palm Absent Underside Underside of thigh Smooth Smooth Smooth Chest Smooth Throat Smooth Glandular folds Horny Horny spinules Shagreen or Shagreen glandular warts Glandular Glandular warts Shagreen, scattered few glandular warts, and horny spinules Horny spinules in males, finely or granular in shagreen female Shagreen with a few scattered glandular warts Upper flank Granular Few glandular Few warts Horny spinules Horny Shagreen or Shagreen with glandular warts spinules Horny Tuberculated, Tuberculated, horn like spinules Glandular warts Glandular warts Shagreen, scattered few glandular warts and horny spinules (males) spinules Horny in males, finely or granular in shagreen female with Shagreen scattered a few glandular finely warts, tuberculated Posterior Posterior dorsum Horny spinules Horny (spinules in Absent females) Few glandular Few warts Horny spinules Horny Shagreen or Shagreen with glandular warts spinules Horny Tuberculated, Tuberculated, horn like spinules Glandular warts Glandular warts with Shagreen scattered a few glandular warts and horny spinules ( males) spinules Horny in males, finely or granular in shagreen female with Shagreen scattered a few glandular finely warts, tuberculated Anterior Anterior dorsum spinules Horny (spinules in Absent females) Few Few glandular warts Glandular warts Shagreen Shagreen or with glandular warts Horny spinules Glandular warts Glandular warts Glandular warts Shagreen Horny spinules in males, finely or granular in shagreen female Shagreen with a few scattered glandular warts Side of head Few glandular Few warts Smooth Smooth Smooth Smooth Shagreen or with Shagreen glandular warts spinules Horny Glandular warts Smooth Glandular warts Shagreen spinules Horny in males, finely or granular in shagreen female with Shagreen scattered a few glandular warts Interorbital area Interorbital Glandular warts Glandular warts Shagreen or with Shagreen glandular warts spinules Horny Glandular warts Glandular warts Glandular warts Shagreen spinules Horny in males, finely or granular in female shagreen with Shagreen scattered a few glandular warts Snout P. nanus P. P. popularis P. P. ocularis P. regius P. Species name nasutus P. nemus P. oxyrhynchus P. papillosus P. pardus P. pleurotaenia P. poppiae P. procax P. reticulatus P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3829 Eight new species of Pseudophilautus Wickramasinghe et al. Supernumerary Supernumerary on tubercles foot Supernumerary Supernumerary on tubercles palm Absent Smooth Underside Underside of thigh Smooth Smooth Smooth Smooth Smooth Chest Smooth Granular in males, smooth in females Glandular warts Glandular warts Smooth Throat Smooth Glandular warts Granular Glandular warts Glandular folds Glandular folds, glandular and warts horny spinules Small tubercules with horny spinules Scattered, glandular tubercles Glandular warts, with horny spinules in males Upper flank Glandular warts Glandular and warts horny spinules Smooth a for except scattered few glandular warts Glandular warts Glandular warts Glandular males warts, with horny spinules scattered Glandular folds, glandular warts and horny spinules Small with tubercules spinules horny Scattered, glandular tubercles Glandular males warts, spinules horny scattered Posterior Posterior dorsum Glandular and warts spinules, horny lack females spinules horny Smooth a for except scattered few glandular warts Glandular warts Glandular warts Glandular warts Glandular males warts, with horny spinules scattered Glandular folds, glandular warts and horny spinules Small with tubercules spinules horny Glandular warts Scattered, glandular tubercles Glandular males warts, spinules horny scattered Anterior Anterior dorsum Glandular and warts spinules, horny lack females spinules horny Smooth except a few for scattered glandular warts Glandular warts Glandular warts Glandular warts Glandular warts Glandular warts Small tubercules with horny spinules Glandular warts Scattered, glandular tubercles Glandular warts Side of head Smooth a for except scattered few glandular warts Smooth Smooth Smooth Smooth Glandular warts Glandular folds, glandular warts and horny spinules Small tubercules with horny spinules Glandular warts Scattered, glandular tubercles Glandular warts Glandular warts and horny spinules Interorbital area Interorbital Smooth except for for Smooth except scattered a few glandular warts Glandular warts Glandular warts Small tubercules spinules with horny Glandular warts glandular Scattered, tubercles Glandular warts and warts Glandular spinules horny Snout P. rugatus P. Species name rus P. sarasinorum P. schmarda P. schneideri P. semiruber P. silus P. silvaticus P. simba P. singu P. sordidus P. steineri P. 3830 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Supernumerary Supernumerary on tubercles foot Supernumerary Supernumerary on tubercles palm Smooth Underside Underside of thigh Mid chest Mid chest smooth Chest Smooth Smooth or sha- green Smooth Throat Smooth Smooth or sha- green Granular Granular Granular Glandular warts Upper flank Shagreen Shagreen weakly to areolate Granular Shagreen, Shagreen, horny spinules in males Shagreen spinules Horny (smooth in and females immature males) Weakly tubercular Glandular warts spinules Horny Posterior Posterior dorsum Weakly Weakly shagreen Shagreen, Shagreen, spinules horny in males spinules, Horny lack females spinules horny Shagreen spinules Horny (smooth in and females immature males) Weakly tubercular Glandular warts spinules, Horny Anterior with dorsum “)(”–shaped of pattern tubercles Anterior Anterior dorsum Weakly Weakly shagreen Shagreen, spinules horny in males spinules, Horny lack females spinules horny Shagreen Horny spinules (smooth in and females immature males) Shagreen Glandular warts Glandular warts Weakly Weakly shagreen Side of head Weakly Weakly shagreen Smooth Smooth Smooth Horny spinules Horny Shagreen spinules Horny (smooth in and females males) immature Shagreen Glandular warts spinules Horny Interorbital area Interorbital Weakly shagreen Weakly Horny spinules Horny Shagreen spinules Horny (smooth in females and immature males) shagreen Weakly Glandular warts spinules Horny Snout P. stellatus P. Species name stictomerus P. stuarti P. tanu P. temporalis P. variabilis P. viridis P. zal P. zimmeri P. zorro P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3831 Eight new species of Pseudophilautus Wickramasinghe et al. arm smooth; forearm weakly tubercular; few tubercles Pseudophilautus karunarathnai sp. nov. present on hand; inner and outer thigh smooth, dorsally (Figs. 11–13; Images 21–25; Tables 10–12) weakly tubercular; leg dorsally with tubercles, a small urn:lsid:zoobank.org:act:0B077FB6-CC9D-48AA-8AC8-D5822D4BC0CA calcar like tubercle at heel; tarsus bearing indistinct tubercles; foot weakly tubercular. Holotype: NMSL 2013.04.01 NH, 03.iv.2010, adult Ventral side of body: Throat and chest weakly male 16.2mm SVL (Images 21, 22A,D). Sripada (Peak granular; belly granular; forearm and upper arm weakly Wilderness), Ratnapura District, Sabaragamuwa granular; thigh granular; leg smooth; tarsus with Province, Sri Lanka (06048’31”N & 80028’14”E; 1334m) prominent tubercles. (Fig. 1), coll. L.J.M.W, D.R.V., M.D.G.R., S.C.A. & A.W.A.C.. Colour in life: Dorsum light brown with a green tinge, Paratypes: DWC 2013.01.007, adult female, 19.2mm a dark brown circle on snout and a broad dark brown SVL (Images 22 B,E, 23); DWC 2013.01.008, adult male, band connecting upper eyelids, dark brown triangular 16.3mm (Images 22 C,F, 24); the same date, locality and patch pointed towards head on anterior dorsum, collectors. another broad dark brown cross band on sacrum, side Diagnosis: Pseudophilautus karunarathnai sp. olive green; limbs light brown with a green tinge, fore nov. can be distinguished from known congeners by limbs, hind limbs, fingers and toes with dark brown cross the following combination of characters: body rather bands, finger tips off white; ventral side off white, throat small (males SVL 16.2–16.3 mm, females 19.2mm); and chest marbled with brown blotches, belly, hands, dorsal head and Interorbital space convex; lateral feet and webbing darker (Image 13). snout, canthus rostralis rounded; loreal region and Colour in alcohol: Colour darkens, pattern remains, internarial space concave; lingual papilla and calcar light brown with green tinge changes to dark brown and absent; vomerine teeth and nuptial pad present; snout dark brown patches to maroon, and ventrally off white smooth, a prominent blunt tubercle present; interorbital colour changes to a light brown (Image 14 A&D). area weakly tubercular; anterior dorsum and dorsal Paratype and variation: The species showed quite a thigh smooth; posterior dorsum spinulate; upper flank variation of dorsal colour depending on its surrounding weakly granular; lower flank, throat and chest granular; environment, ranging from maroonish tinges to light supernumerary tubercles on palm and foot absent. brown to cream colour, where they lacked the prominent Comparative characters that distinguish the species from greenish tinge found on holotype. Markings and patterns its congeners are provided in tables 11 and 12. remained the same even with change in colour (Image Description of holotype: Body rather small (SVL 16.2), 16–20). moderately elongate (SVL/HW 2.1); head large (HL/SVL 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 jagathgunawardanai is 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. Figure 11. Pseudophilautus karunarathnai sp. nov. holotype (NMSL 2013.04.01 NH), head; (A) lateral aspect (B) dorsal aspect (C) ventral aspect. Scale bar: 2mm. (© Dulan Ranga Vidanapathirana) 3832 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Figure 12. Pseudophilautus karunarathnai sp. nov. holotype (NMSL 2013.04.01 NH), ventral aspect of left hand. Scale bar: 5mm. ©( L.J. Mendis Wickramasinghe) Figure 13. Pseudophilautus karunarathnai sp. nov. holotype (NMSL 2013.04.01 NH), ventral aspect of left foot. Scale bar: 5mm. (© L.J. Mendis Wickramasinghe) © L.J. Mendis Wickramasinghe Image 21. Pseudophilautus karunarathnai sp. nov. holotype in life (NMSL 2013.04.01 NH). 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 © L.J. Mendis Wickramasinghe rounded, loreal region concave; interorbital space Image 23. Pseudophilautus karunarathnai sp. nov. paratype in life convex, larger than upper eyelid (IO/UEW 1.1), equal (DWC 2013.01.007). to internasal distance (IN/UEW 1.1); distance between front of eyes 3/5th the distance between back of eyes Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3833 Eight new species of Pseudophilautus Wickramasinghe et al. © L.J. Mendis Wickramasinghe Image 22. Type series of Pseudophilautus karunarathnai sp. nov. Left to right, NMSL 2013.04.01 NH, DWC 2013.01.007, DWC 2013.01.008. A-C dorsal aspect and D-F ventral aspect. Scale bar: 5mm. © L.J. Mendis Wickramasinghe © L.J. Mendis Wickramasinghe Image 24. Pseudophilautus karunarathnai sp. nov. paratype in life (DWC 2013.01.008). Image 25. Pseudophilautus karunarathnai sp. nov. unpreserved (DBE/DFE 1.7); nostrils oval, without flap of skin laterally, specimen. 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), axis; tongue large, lanceolate; lingual papilla absent, tympanum-eye distance 2/3rd tympanum width (TAD/ depression with conical tubercles present on tongue. TYW 0.7); pineal ocellus absent; vomerine teeth present, Arm short, thin (LAL/FEL 0.49, UAL/FEL 0.3); forearm small, few, odontophores oblique and widely separated, shorter than hand length (LAL/HNL 0.9); forearm longer between choanae with an angle of 60° relative to body than upper arm (LAL/UAL 1.5); fingers thin, (Fig. 12), 3834 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Range 1.5—1.6 1.6—1.9 2.5—3.2 5.4—6.5 3.6—5.1 2.4—3.2 4.5—5.8 3.0—3.9 1.0—1.3 0.9—1.0 2.4—3.4 1.4—1.9 7.6—8.8 0.3—0.6 7.9—9.4 3.6—5.0 2.5—3.8 4.7—6.9 6.2—7.9 1.1—1.2 8.8—10.8 16.2—19.2 SD 0.1 0.2 0.4 0.6 1.1 0.8 0.4 0.7 0.5 0.1 0.1 0.5 0.3 0.7 1.7 0.2 0.8 0.7 0.7 1.1 0.9 0.1 1.6 1.8 2.9 6.0 4.4 2.9 5.1 3.4 1.2 0.9 3.0 1.7 8.4 0.5 8.8 4.5 3.2 5.8 7.1 1.2 10.1 17.2 Mean 8.8 1.5 1.6 2.5 5.4 3.6 2.4 4.5 3.0 1.0 0.9 2.4 1.4 7.6 0.3 7.9 3.6 2.5 4.7 6.2 1.1 16.3 Paratype 2 Paratype 2013.01.008 1.6 1.9 3.2 6.5 5.1 3.2 5.8 3.9 1.3 1.0 3.4 1.9 8.8 0.6 9.4 5.0 3.8 6.9 7.9 1.2 10.8 19.2 Paratype 1 Paratype 2013.01.007 1.6 1.9 2.8 1.2 6.1 4.3 3.0 5.2 3.4 0.9 3.3 1.8 8.7 0.6 9.1 4.9 3.2 6.0 7.3 1.2 10.6 16.2 Holotype 2013.04.01 NH KT SN LAL TBL SVL TAS MN VKL TYH UAL TAD TPD TL-1 TL-2 TL-3 TL-4 TL-5 TND MFE TYW MBE UEW DWC NMSL/ Range 1.8—2.0 2.7—3.3 2.0—2.8 4.2—5.8 1.3—1.6 6.9—9.2 4.0—4.5 6.7—8.7 7.0—8.2 0.7—0.8 1.7—1.9 1.9—2.1 1.5—1.9 2.6—3.3 0.4—0.5 5.5—7.4 3.3—4.2 0.7—0.8 0.8—0.9 2.2—3.0 8.2—10.3 11.1—12.9 SD 1.2 0.1 0.3 0.5 0.9 0.8 0.2 1.2 0.3 1.1 0.6 0.1 0.1 0.1 0.2 0.4 0.1 0.9 0.5 0.1 0.1 0.5 sp. nov. of Pseudophilautus karunarathnai 9.5 1.9 3.1 2.5 5.0 1.4 8.3 4.3 7.9 7.6 1.8 0.8 2.0 1.8 3.0 0.5 6.6 3.8 2.8 0.8 0.9 12.0 Mean 8.2 1.8 2.7 2.0 4.2 1.3 6.9 4.0 6.7 7.0 1.7 0.7 1.9 1.5 2.6 0.4 5.5 3.3 2.2 0.7 0.8 11.1 Paratype 2 Paratype 2013.01.008 2.0 3.2 2.8 5.1 1.5 9.2 4.5 8.7 1.9 8.2 0.8 2.1 3.3 1.9 0.5 7.4 4.2 3.0 0.8 0.9 10.3 12.9 Paratype 1 Paratype 2013.01.007 2.0 3.3 2.8 5.8 9.9 1.6 8.8 4.4 8.4 1.9 7.6 0.8 2.0 3.2 1.8 0.5 6.8 4.0 3.0 0.8 0.9 12.0 Holotype 2013.04.01 NH IN IO ES HL DL ED EN GK DB HD FEL DW HW FOL IML DFE DBE FL-2 FL-3 FL-4 FL-1 HNL DWC NMSL/ Table 10. The morphometric measurements (mm) of the type series measurements 10. The morphometric Table Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3835 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Nuptial Nuptial pad Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Present Present Calcar Present Present Present Present Present Present Present Present Absent Present Present Present Present Present Lingual papilla Present Present Present Present Present Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Vomerine Vomerine teeth Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Internarial Internarial space Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Concave Interorbital Interorbital space Concave Concave Flat Flat or flat Concave Flat Concave Concave Flat Flat Concave Flat or flat Concave Flat Flat Flat Flat Flat Concave Concave Concave Flat Concave Flat Flat Flat Flat Convex Loreal region Loreal Convex Flat Flat Flat Flat Concave Sharp Sharp Sharp Sharp Sharp Sharp Sharp Canthus Canthus rostralis Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Rounded Oval Pointed or oval Pointed Truncate Blunt pointed Obtusely Oval Oval Obtusely pointed Obtusely Blunt Truncate pointed Obtusely Truncate Truncate Snout lateral Truncate Blunt Acuminate Obtusely pointed Obtusely pointed Obtusely Sharply Pointed Oval Pointed Truncate Rounded Pointed Truncate Flat Concave Concave Flat Flat Flat Concave Flat Flat Flat Flat Flat Flat Flat or flat Concave Flat Uneven Concave Concave Head dorsally Convex Concave 44.8 35.8-40.4 16.2–19.2 22.3–32.5 40.9 38.4 25.4–37.0 33.3–41.7 21.7–26.4 16.3–19.0 38.4–48.9 26.7–29.3 33.4–46.8 21.2–23.4 22.0–28.6 11.2–24.4 19.8 24.5–30.1 17.0–27.5 18.9–27.3 17.6–28.9 18.3–23.9 35.3 43.5 23.4–28.8 26.9–44.5 23.7–29.4 32.9–42.9 27.9 18.3–21.9 25.7 27.4–42.7 46.6 14.9 17.3–20.2 SVL 33.6–51.1 20.4–24.6 sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. P. jagathgunawardanai P. dimbullae P. P. karunarathnai karunarathnai P. P. sirilwijesundarai P. P. lunatus P. P. dayawansai P. newtonjayawardanei P. puranappu P. abundus P. adspersus P. auratus P. caeruleus P. cavirostris P. frankenbergi P. fulvus P. hoffmanni P. hoipolloi P. hypomelas P. leucorhinus P. Species name samarakoon P. alto P. asankai P. cuspis P. decoris P. eximius P. extirpo P. femoralis P. fergusonianus P. folicola P. hallidayi P. halyi P. hankeni P. limbus P. macropus P. maia P. malcolmsmithi P. P. bambaradeniyai P. sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners sp. nov. Pseudophilautus karunarathnai differentiate that 11. Characters Table 3836 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Nuptial Nuptial pad Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Calcar Present. Present Present Present Present Present Present Present Present Present Lingual papilla Present Present Present Present Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Vomerine Vomerine teeth Absent Absent Absent Absent Absent Absent Absent Internarial Internarial space Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Concave Interorbital Interorbital space Flat Concave Flat Flat Flat Flat Flat Concave Flat Flat Flat Concave Flat Flat Flat Flat Flat Concave or flat Concave Flat Concave Flat Flat Flat Flat Flat Concave Flat Flat Loreal region Loreal Flat Flat Flat Flat Flat Sharp Sharp Canthus Canthus rostralis Sharp Indistinct Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Indistinct Sharp Sharp Pointed Oval Truncate Blunt or oval Pointed Oval Pointed Snout lateral Oval Obtusely pointed Obtusely Pointed Oval Truncate pointed Obtusely Oval Oval Truncate Blunt pointed Obtusely Pointed Truncate or oval Truncate Oval pointed Obtusely or rounded Blunt Blunt Pointed Flat Flat Flat Flat Flat Flat Flat Concave Flat Flat Concave Head dorsally Concave 23.9–49.0 29.4–35.0 20.7 19.9–22.8 34.8 18.6 25.6–30.3 21.3–26.0 17.7–24.7 18.3–21.7 22.6–38.1 17.7–30.0 13.4 35.4–51.8 12.6–15.6 16.1–16.6 22.2–39.4 30.4–41.6 13.5–13.9 28.4–31.0 35.7 20.0–32.5 16.3–18.4 17.4 23.1–33.4 40 32.1 25.1–26.8 42.5–61.1 17.5 20.6–24.1 24.3–31.3 39.6–55.3 23.2–36.0 24.2–32.4 27.4–36.3 31.6 22.9–30.1 SVL P. microtympanum P. P. mooreorum P. nemus P. schneideri P. P. nanus P. oxyrhynchus P. pleurotaenia P. poppiae P. popularis P. regius P. sarasinorum P. schmarda P. semiruber P. silus P. simba P. singu P. sordidus P. steineri P. tanu P. temporalis P. variabilis P. zal P. Species name mittermeieri P. nasutus P. ocularis P. papillosus P. pardus P. procax P. reticulatus P. rugatus P. rus P. silvaticus P. stellatus P. stictomerus P. stuarti P. viridis P. zimmeri P. zorro P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3837 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Super- numerary tubercles on foot Present Present Present Present Present Present Present Present Present Present Absent Super- rnumerary tubercles on palm Present Present Present Present Present Present Present Present Present Present Present Present Granular Chest Granular Throat Shagreen Shagreen Smooth Very fine, Very horny spinules Dorsal thigh Dorsal Weakly tubercular Shagreen Weakly shagreen Tuberculated Tuberculated Granular Very fine, Very horny spinules Lower Lower flank Smooth Weakly Weakly granular Shagreen Areolate spinules Horny Upper flank Tubercular Smooth Smooth Smooth spinules Horny in males Glandular warts and horny spinules Spinulate Shagreen Shagreen fine, horny Very spinules Posterior Posterior dorsum Shagreen Tubercular Weakly tubercular Smooth Smooth Smooth Shagreen with Shagreen tubercles few spinules in Horny in males, warty females fine, horny Very spinules Glandular warts and horny spinules (females lack horny spinules) Prominent Prominent and tubercles spinules horny shagreen Weakly Anterior dorsum Anterior Shagreen Tubercular Weakly tubercular Weakly Smooth, with across a ridge medially braking with a Shagreen, ridge prominent A prominent present tubercle spinules in Horny males fine, horny Very spinules Glandular warts spinules and horny Smooth Smooth,with dermal cross two fringes Smooth Smooth for Smooth except scattered a few glandular warts Interorbital area Interorbital Shagreen Smooth, a blunt prominent present tubercle Tuberculated, with Tuberculated, a single prominent medially tubercle placed Shagreen with Shagreen, dermal prominent fold spinules in Horny males fine, horny Very spinules Glandular warts spinules and horny Snout Prominent tubercle tubercle Prominent on centre present of snout Shagreen sp. nov. sp. nov. sp. nov. P. bambaradeniyai P. sp. nov. P. karunarathnai karunarathnai P. sp. nov. P. dayawansai P. jagathgunawardanai P. sp. nov. newtonjayawardanei P. sp. nov. puranappu P. sirilwijesundarai P. sp. nov. abundus P. alto P. asankai P. caeruleus P. Species name samarakoon P. adspersus P. auratus P. sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners sp. nov. Pseudophilautus karunarathnai differentiate that in the skin structure 12. Characters Table 3838 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Super- numerary tubercles on foot Present Present Present Present Present Present Super- rnumerary tubercles on palm Present Present Present Present Present Present Present Present Present Present Present Present Present Chest Smooth Shagreen Smooth Throat Smooth Smooth Smooth Smooth Shagreen Smooth Dorsal thigh Dorsal Glandular warts Shagreen, glandular few warts Shagreen Glandular warts Glandular warts Glandular with warts horny spinules Lower Lower flank Glandular warts Weakly Weakly granular Glandular Smooth or granular Smooth Upper flank Glandular warts Shagreen Shagreen Glandular warts Smooth Smooth Smooth with glandular warts Smooth or with glandular warts Shagreen, males with glandular warts and horny spinules Shagreen Glandular with warts, spinules horny in males Glandular warts bearing horny spinules Heavily Heavily tuberculated, glandular warts Shagreen, with Shagreen, glandular a few warts Posterior Posterior dorsum Glandular warts Shagreen Finely granular Shagreen with Shagreen glandular warts Smooth Smooth Smooth Smooth or shagreen Smooth Smooth with a scattered few glandular warts Heavily Heavily tuberculated, glandular warts Horny spinules Horny with a “)(”shaped of pattern tubercles Shagreen, with Shagreen, glandular a few warts Glandular warts, with horny spinules in males Glandular warts bearing horny spinules Anterior dorsum Anterior Glandular warts Shagreen Finely granular Males Shagreen, glandular warts and horny spinules horny Shagreen, spinules scattered Shagreen with Shagreen glandular warts Glandular warts Horny spinules Horny Shagreen Glandular warts, with horny spinules in males Glandular warts bearing horny spinules Smooth Smooth Smooth Smooth or shagreen Smooth Interorbital area Interorbital Glandular warts Shagreen Finely granular males Shagreen, glandular warts spinules and horny Shagreen Shagreen with Shagreen glandular warts Glandular warts Horny spinules Horny Shagreen, with Shagreen, glandular a few warts Glandular warts, spinules with horny in males Glandular warts bearing horny spinules Snout Glandular warts Shagreen Finely granular males Shagreen, glandular warts spinules and horny Shagreen Shagreen with Shagreen glandular warts P. cavirostris cavirostris P. P. cuspis P. P. eximius eximius P. extirpo P. hallidayi P. hankeni P. hoffmanni P. Species name decoris P. dimbullae P. femoralis P. fergusonianus P. folicola P. frankenbergi P. fulvus P. halyi P. hoipolloi P. hypomelas P. leucorhinus P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3839 Eight new species of Pseudophilautus Wickramasinghe et al. Super- numerary tubercles on foot Present Present Present Present Present Present Super- rnumerary tubercles on palm Present Present Present Present Present Present Present Present Present Present Present Chest Smooth Smooth Throat Smooth Smooth Smooth Dorsal thigh Dorsal Granular, not Granular, rough Shagreen, scattered few glandular warts Glandular warts Glandular warts Glandular warts Shagreen Glandular warts Glandular warts Glandular warts Lower Lower flank Smooth Horny Horny spinules in males Glandular warts Smooth Smooth Glandular warts Glandular warts Upper flank Glandular folds or Shagreen glandular warts Glandular warts Shagreen, scattered few glandular and warts, spinules horny Glandular warts Smooth spinules Horny in males Glandular warts Tuberculated, horn like spinules Glandular warts Glandular warts Smooth Posterior Posterior dorsum Few glandular Few warts or Shagreen with glandular warts Glandular warts Shagreen, scattered few glandular warts and horny spinules (males) Weakly granular Weakly Smooth or Shagreen smooth, with a scattered few glandular warts Smooth Smooth Glandular warts tuberculated, spinules horn like Glandular warts Glandular warts Smooth, '\ /' shaped ridge Anterior dorsum Anterior Horny spinules Horny in males, finely or granular in shagreen female glandular Few warts or with Shagreen glandular warts Glandular warts with Shagreen scattered a few glandular warts and horny spinules ( males) Weakly granular Weakly Smooth above, and shagreen with glandular males with warts, spinules horn like Glandular warts Glandular warts Smooth Smooth Smooth with a scattered few glandular warts Smooth Smooth Smooth Smooth Interorbital area Interorbital Horny spinules in Horny males glandular Few warts or with Shagreen glandular warts Glandular warts Shagreen Weakly granular Weakly Glandular warts Glandular warts Glandular warts Snout Horny spinules in Horny males Glandular warts or with Shagreen glandular warts Glandular warts Shagreen P. limbus P. P. macropus P. microtympanum P. P. mittermeieri mittermeieri P. nasutus P. nemus P. Species name maia P. malcolmsmithi P. mooreorum P. nanus P. ocularis P. oxyrhynchus P. papillosus P. pardus P. pleurotaenia P. P. lunatus P. 3840 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Super- numerary tubercles on foot Present Present Present Present Present Present Present Present Super- rnumerary tubercles on palm Present Present Present Present Present Present Present Present Present Present Present Present Present Chest Smooth Smooth Throat Smooth Dorsal thigh Dorsal Scattered Scattered glandular tubercles Glandular warts Horny Horny spinules in males, finely or granular in shagreen female Glandular warts Glandular warts Lower Lower flank Scattered, Scattered, glandular tubercles Smooth Smooth Glandular Glandular and warts horny spinules Shagreen with Shagreen scattered a few glandular warts Glandular warts Glandular warts Upper flank Scattered, Scattered, glandular tubercles Glandular warts Smooth Horny spinules Horny in males, finely or granular in shagreen female spinules Horny Smooth Glandular Glandular folds Glandular folds, glandular warts and horny spinules Small with tubercules spinules horny Shagreen with Shagreen scattered a few glandular finely warts, tuberculated Glandular warts Glandular warts Smooth Smooth Posterior Posterior dorsum Scattered, Scattered, glandular tubercles Smooth a for except scattered few glandular warts Smooth Horny spinules Horny in males, finely or granular in shagreen female Horny spinules Horny (spinules Absent in females) with Shagreen scattered a few glandular finely warts, tuberculated Glandular warts Small tubercules with horny spinules Glandular warts Glandular warts Glandular warts Anterior dorsum Anterior Horny spinules Horny spinules Horny Glandular warts, males with horny spinules scattered Glandular folds, glandular warts and horny spinules Scattered, glandular tubercles Horny spinules Horny in males, finely or granular in female shagreen Smooth with Shagreen scattered a few glandular warts for Smooth except scattered a few glandular warts Smooth Small tubercules with horny spinules Smooth Smooth Glandular warts Smooth Interorbital area Interorbital Smooth spinules Horny Glandular warts Glandular folds, glandular warts spinules and horny Scattered, tubercles glandular Horny spinules Horny in males, finely or granular in female shagreen Shagreen with Shagreen scattered a few glandular warts Small tubercules spinules with horny Glandular warts Snout Glandular warts spinules Horny Glandular warts Glandular warts Scattered, glandular tubercles P. poppiae P. P. silus P. P. procax P. reticulatus P. rus P. schneideri P. silvaticus P. Species name popularis P. regius P. rugatus P. sarasinorum P. schmarda P. simba P. singu P. P. semiruber P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3841 Eight new species of Pseudophilautus Wickramasinghe et al. Super- numerary tubercles on foot Present present Present Present Present Present Present Super- rnumerary tubercles on palm Present Present Present Present Present Present Present Present Present Present Present Chest Smooth Smooth Throat Smooth or shagreen Smooth Smooth Dorsal thigh Dorsal Glandular warts Weakly shagreen Shagreen Weakly tubercular Glandular warts Lower Lower flank Weakly areolate Smooth Upper flank Smooth Glandular with warts, spinules horny in males Glandular warts and horny spinules to Shagreen areolate weakly Shagreen, spinules horny in males Smooth spinules Horny (smooth in and females immature males) Shagreen Glandular warts Posterior Posterior dorsum Weakly Weakly tubercular Weakly shagreen Smooth Smooth Glandular warts, Glandular warts, males horny spinules scattered Shagreen Glandular warts Shagreen, horny horny Shagreen, spinules in males Anterior dorsum Anterior Glandular warts Glandular warts and horny spinules, females lack horny spinules spinules, Horny lack females spinules horny Weakly tubercular Weakly shagreen Weakly Glandular warts Shagreen spinules Horny females in (smooth and immature males) Glandular warts Weakly shagreen Weakly Smooth Smooth Smooth Interorbital area Interorbital Glandular warts Glandular warts spinules and horny spinules Horny Shagreen Glandular warts Shagreen spinules Horny (smooth in females and immature males) Glandular warts Snout shagreen Weakly Glandular warts Glandular warts spinules and horny spinules Horny shagreen Weakly P. sordidus P. P. variabilis P. P. stellatus P. tanu P. temporalis P. viridis P. zimmeri P. Species name steineri P. stictomerus P. stuarti P. zal P. 3842 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. relative length of fingers I < II < IV < III (FL-1/FL-3 0.5, FL-2/ Colour in alcohol: Colour pattern remains with a little FL-3 0.6, FL-4/FL-3 0.9) (Table 10); tips of fingers rounded fading. All dark brown patches more prominent in off enlarged, discs present on all fingers, with distinct basal white, ventral side off-white all over (Image 22 A & D). and circum marginal grooves; dermal fringe present on Paratype and variation: Apart from the slightly dark fingers; webbing formula I2—2 1/3 — II2 3 2/3III3+—2 brownish dorsum colouration all specimens observed ½IV; subarticular tubercles prominent, rounded, single, had similar colour not much variation seen (Image 23– all present, IV2 and III2 relatively smaller; inner palmar 25). tubercle indistinct, single, oval, larger than outer palmer Etymology: The species name karunarathnai is named tubercle; nuptial pads present; outer palmar tubercle after Y.G.P. Karunarathna (retired Assistant Director, indistinct, single, oval, smaller than the distal subarticular Department of Wildlife Conservation, Sri Lanka), in great tubercles; supernumerary tubercles absent on fingers appreciation for his commendable services to wildlife indistinct on palm; prepollex absent; femur 1 1/3 times management which has resulted in the minimizing of longer than fourth toe length (FEL/TL-4 2.3); foot length the human-wildlife conflict on the island. The species longer than thigh (FOL/FEL 1.2); toes thin, (Fig. 13), name is a noun in the genitive case. relative length of toes I < II < III < V < IV (TL-1/TL-4 0.4, Suggested vernacular names: Karunarathnage TL-2/TL-4 0.5, TL-3/TL-4 0.7, TL-5/TL-4 0.7); tips of toes panduru madiya and Karunarathna’s Shrub Frog in rounded, enlarged, discs present on all toes with distinct Sinhala and English, respectively. basal and circum marginal grooves; webbing formula Natural history: The species is found in elevations of I2—2-II2+—3+III2+—3-IV2 ½—2+V; dermal fringe present 750–1400 m, from the forest floor to about 1.5m high on inside of toes III, IV and V, thin fringe along exterior shrubs in lowland rain forests and lower montane rain edge of toe V, and absent on metatarsus and tarsal fold; forests. Apart from forests they are also observed in subarticular tubercles prominent, rounded, single, all home gardens without much canopy cover. present, V2 and IV3 relatively smaller; supernumerary Remarks: Pseudophilautus karunarathnai sp. nov. tubercles absent; inner metatarsal tubercle oval possesses a unique colour pattern which is not possessed prominent and large, its length half the length of toe by any of its congeners, hence can easily be identified. I (IML/TL-1 0.5); outer metatarsal tubercle present, smaller than penultimate subarticular tubercles. Pseudophilautus newtonjayawardanei sp. nov. Skin of snout dorsally smooth, a large, prominent, (Figs. 14–16; Images 26–28; Tables 13–15) blunt tubercle present, snout laterally weakly tubercular; urn:lsid:zoobank.org:act:54DAD829-148A-47C4-BCD4-92B994D8E581 interorbital space and upper eyelid weakly tubercular; head laterally weakly tuberculated, a faint median Holotype: NMSL 2013.05.01 NH, 05.ii.2010, adult dermal ridge intermittent with tubercles extends from tip female 38.4mm SVL (Images 26–27). Sripada (Peak 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, Figure 14. Pseudophilautus newtonjayawardanei sp. nov. holotype belly, and hands lighter, feet and webbing dark brown (NMSL 2013.05.01 NH), head; (A) lateral aspect (B) dorsal aspect (C) (Image 21). ventral aspect. Scale bar: 2mm. (© Dulan Ranga Vidanapathirana) Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3843 Eight new species of Pseudophilautus Wickramasinghe et al. Figure 15. Pseudophilautus newtonjayawardanei sp. nov. holotype (NMSL 2013.05.01 NH), ventral aspect of left hand. Scale bar: 5mm. Figure 16. Pseudophilautus newtonjayawardanei sp. nov. holotype (© L.J. Mendis Wickramasinghe) (NMSL 2013.05.01 NH), ventral aspect of left foot. Scale bar: 5mm. (© L.J. Mendis Wickramasinghe) © L.J. Mendis Wickramasinghe Image 26. Pseudophilautus newtonjayawardanei sp. nov. holotype in life (NMSL 2013.05.01 NH). Gehan Rajeev © M.D. Image 28. Pseudophilautus newtonjayawardanei sp. nov. unpreserved specimen. 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. 38.4); dorsal head, interorbital space and internarial Diagnosis: Pseudophilautus newtonjayawardanei sp. space concave; lateral snout, canthus rostralis and nov. can be distinguished from known congeners by the tips of toes rounded; vomerine teeth, lingual papilla, following combination of characters: moderate size (SVL calcar, nuptial pad, and tympanum absent; fringe on 3844 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. © L.J. Mendis Wickramasinghe Image 27. Holotype of Pseudophilautus newtonjayawardanei sp. nov. NMSL 2013.05.01 NH. A dorsal aspect and B ventral aspect. Scale bar: 5mm. Table 13. The morphometric measurements (mm) of the holotype of Pseudophilautus newtonjayawardanei sp. nov. Pseudophilautus newtonjayawardanei sp. nov. NMSL 2013.05.01 NH DB 1.65 FEL 19.03 HL 16.55 MBE 7.41 TL-1 3.3 TYW 0 DBE 14.57 FL-1 3.66 HNL 11.96 MFE 12.17 TL-2 3.8 UAL 7.64 DFE 7.48 FL-2 4.35 HW 16.68 MN 15.03 TL-3 5.61 UEW 3.59 DL 2.06 FL-3 7.28 IML 1.96 SN 1.93 TL-4 8.43 VKL 14.94 DW 2.14 FL-4 6.27 IN 3.32 SVL 38.4 TL-5 6.96 ED 5.76 FOL 28.19 IO 4.53 TAD 0 TND 0 EN 3.97 GK 17.81 KT 17.85 TAS 13.14 TPD 0 ES 5.54 HD 8.85 LAL 10.82 TBL 19.92 TYH 0 fingers present; tarsal tubercle present; snout, side 38.4), elongate (SVL/HW 2.3); head large (HL/SVL 0.4), of head, anterior and posterior dorsum, upper flank about as wide as long (HW/HL 1.0), concave above; and dorsal thigh shagreen; interorbital area shagreen, snout rounded in lateral, dorsal and ventral aspects with a prominent ridge; lower flank weakly granular; (Fig. 14 A–C) (ES/DFE 0.7, SN/IN 0.6), its length equal throat and underside of thigh granular; supernumerary to horizontal diameter of eye (ES/ED 1.0); internasal tubercles present on palm. Comparative characters that space concave; canthus rostralis rounded, loreal distinguish the species from its congeners are provided region concave; interorbital space concave, upper in tables 14 and 15. eyelid smaller than interorbital distance (IO/UEW 1.3); Description of holotype: Body moderate size (SVL internasal distance less than upper eye width (IN/UEW Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3845 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Nuptial Nuptial pad Present Present Present Present Present Present Present Absent Calcar Present Present Present Present Present Present Present Present Tarsal Tarsal tubercle Absent Rounded Tips of Tips toes Triangular Present Fringe on Fringe fingers Absent Absent Absent Absent Absent Absent Lingual papilla Present Present Present Present Present Present Present Present Absent Present Present Vomerine Vomerine teeth Present Present Present Present Present Present Present Present Present Concave Flat Flat Flat Flat Flat Flat Flat Flat Internarial Internarial space Flat Concave Convex Flat Flat Flat Flat Flat Convex Flat Convex Convex Flat Convex Convex Flat Convex Convex Flat Flat Flat Interorbital Interorbital space sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners sp. nov. Rounded Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Canthus Canthus rostralis Truncate Rounded Truncate Truncate pointed Obtusely Blunt pointed Obtusely Sharply Pointed Truncate Blunt Truncate Obtusely pointed Obtusely pointed Obtusely or oval Pointed Oval Snout lateral Truncate Convex Concave Flat Convex Convex Convex Flat Convex Flat Flat Convex Convex Flat Flat Flat Flat Convex Convex Flat or Convex Flat Head dorsally Convex 17.3–20.2 38.4 23.7–29.4 24.5–30.1 35.8-40.4 16.2–19.2 22.3–32.5 25.4–37.0 33.3–41.7 17.0–27.5 21.7–26.4 16.3–19.0 38.4–48.9 17.6–28.9 44.8 35.3 43.5 23.4–28.8 26.9–44.5 26.7–29.3 33.4–46.8 18.9–27.3 18.3–23.9 32.9–42.9 SVL 33.6–51.1 20.4–24.6 sp. nov. sp. nov. sp. nov. P. bambaradeniyai P. sp. nov. P. newtonjayawardanei newtonjayawardanei P. sp. nov. P. puranappu P. samarakoon P. folicola P. Species name dayawansai P. jagathgunawardanai P. sp. nov. karunarathnai P. sp. nov. sirilwijesundarai P. sp. nov. abundus P. adspersus P. alto P. auratus P. caeruleus P. cavirostris P. cuspis P. dimbullae P. eximius P. extirpo P. femoralis P. fergusonianus P. frankenbergi P. fulvus P. P. asankai P. decoris P. hallidayi P. Pseudophilautus newtonjayawardanei Pseudophilautus newtonjayawardanei differentiate that 14. Characters Table 3846 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Nuptial Nuptial pad Present Present Present Present Present Present Present Present Present Calcar Present Present Present Present. Present Present Present Present Tarsal Tarsal tubercle Absent Absent Absent Absent Tips of Tips toes Fringe on Fringe fingers Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Lingual papilla Present Present Present Present Present Present Present Vomerine Vomerine teeth Present Present Present Present Present Present Present Present Present Flat Flat Flat Flat Flat Flat Flat Flat Internarial Internarial space Flat Flat Flat Flat Convex Flat Convex Flat Flat Convex Flat Flat Flat Flat Flat Flat Convex Flat Flat Flat Flat Flat Interorbital Interorbital space Sharp Sharp Indistinct Sharp Sharp Sharp Sharp Sharp Sharp Sharp Canthus Canthus rostralis Pointed Blunt or oval Truncate Blunt Acuminate Oval Oval Pointed Oval Oval pointed Obtusely Pointed Truncate Truncate Oval Oval Pointed pointed Obtusely Oval Truncate Pointed or oval Pointed Snout lateral Uneven Convex Flat or Convex flat Flat Convex Convex Flat Flat Convex Convex Flat Convex Convex Flat Convex Convex Convex Convex Flat Flat Convex Convex Head dorsally 27.9 18.3–21.9 46.6 16.3–18.4 17.4 23.1–33.4 25.1–26.8 42.5–61.1 21.2–23.4 22.0–28.6 11.2–22.4 19.8 25.7 40.9 27.4–42.7 14.9 23.9–49.0 29.4–35.0 34.8 20.7 18.6 40 25.6–30.3 21.3–26.0 17.7–24.7 18.3–21.7 20.6–24.1 32.1 17.5 SVL P. halyi P. P. hankeni P. maia P. mittermeieri P. nasutus P. ocularis P. procax P. reticulatus P. Species name hoffmanni P. hoipolloi P. hypomelas P. leucorhinus P. limbus P. lunatus P. macropus P. malcolmsmithi P. microtympanum P. mooreorum P. nanus P. nemus P. oxyrhynchus P. papillosus P. pleurotaenia P. poppiae P. popularis P. regius P. rus P. P. pardus P. rugatus P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3847 Eight new species of Pseudophilautus Wickramasinghe et al. Nuptial Nuptial pad Present Present Present Present Present Present Calcar Present Present Present Tarsal Tarsal tubercle Absent Absent Absent Absent Absent Tips of Tips toes Absent Absent Fringe on Fringe fingers Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Present Lingual papilla Present Present Present Present Present Present Present Vomerine Vomerine teeth Present Present Present Present Flat Flat Flat Flat Flat Flat Flat Flat Flat Internarial Internarial space Flat Flat Flat Flat Flat Convex Convex Flat Flat Flat Convex Convex Convex Flat Flat Convex Flat Interorbital Interorbital space Sharp Sharp Sharp Sharp Sharp Sharp Sharp Indistinct Sharp Sharp Canthus Canthus rostralis Oval Blunts Obtusely pointed Obtusely Oval Oval Truncate Blunt pointed Obtusely pointed Obtusely Truncate or rounded Blunt Pointed Pointed Snout lateral Flat Convex Flat Convex Convex Convex Convex above Convex Convex Convex Flat or Flat convex Convex Convex Flat Convex Convex Head dorsally 22.6–38.1 24.3–31.3 39.6–55.3 24.2–32.4 31.6 22.9–30.1 17.7–30.0 19.9–22.8 13.4 35.4–51.8 12.6–15.6 16.1–16.6 22.2–39.4 30.4–41.6 23.2–36.0 13.5–13.9 28.4–31.0 35.7 27.4–36.3 20.0–35.5 SVL P. sarasinorum P. P. silvaticus P. stellatus P. stuarti P. zimmeri P. zorro P. Species name schmarda P. schneideri P. semiruber P. silus P. simba P. singu P. sordidus P. steineri P. stictomerus P. tanu P. temporalis P. variabilis P. viridis P. zal P. 3848 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Super- numerary tubercles on palm Present Smooth Underside Underside of thigh Granular Shagreen Shagreen Throat Granular Very fine, Very horny spinules Smooth Weakly tubercular Smooth Weakly shagreen Tubercular Smooth Smooth Smooth Smooth Dorsal thigh Dorsal Shagreen Tuberculated Very fine, Very horny spinules Lower Lower flank Weakly Weakly granular Smooth Areolate Weakly Weakly tubercular Granular Smooth Weakly granular Smooth Granular spinules Horny in males spinules Horny Upper flank Tubercular Shagreen Smooth Few horny horny Few spinules, Prominent tubercles Weakly Weakly tubercular Smooth Spinulate spinules Horny in (Absent females) Smooth Horny spinules in males, warty in females fine, Very spinules horny Posterior Posterior dorsum Tubercular Shagreen Horny spinules Horny sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners sp. nov. Smooth Smooth Smooth Horny spinules in (absent females) Smooth a for except scattered few glandular warts Horny spinules in males, warty in females fine, Very horny spinules Anterior Anterior dorsum Tubercular Shagreen Prominent Prominent tubercles and horny spinules Tubercular Smooth Weakly tubercular Smooth Smooth except a few for scattered glandular warts Smooth fine, Very horny spinules Side of head Smooth Smooth Shagreen Smooth Smooth, with a ridge Smooth, with a ridge breaking across medially Smooth,with two dermal fringes cross tubercular Weakly A prominent present tubercle Smooth for Smooth except scattered a few glandular warts spinules in Horny males fine, horny Very spinules Interorbital area Interorbital Smooth Tubercular Shagreen, with a Shagreen, ridge prominent Smooth Tuberculated, with Tuberculated, a single prominent medially tubercle placed Smooth Smooth, a prominent present tubercle blunt with Shagreen, dermal prominent fold Smooth for Smooth except scattered a few glandular warts spinules in Horny males fine, horny Very spinules Snout Smooth, with a blunt Smooth, with a blunt tubercle Prominent tubercle tubercle Prominent of on centre present snout Shagreen Smooth sp. nov. sp. nov. sp. nov. sp. nov. Species name dayawansai P. jagathgunawardanai P. sp. nov. karunarathnai P. samarakoon P. sirilwijesundarai P. sp. nov. abundus P. adspersus P. alto P. asankai P. P. puranappu P. P. newtonjayawardanei newtonjayawardanei P. sp. nov. P. bambaradeniyai P. sp. nov. Pseudophilautus newtonjayawardanei Pseudophilautus newtonjayawardanei differentiate that in the skin structure 15. Characters Table Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3849 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Super- numerary tubercles on palm Underside Underside of thigh Smooth Smooth Smooth Smooth Smooth, in granular males Throat Glandular warts Smooth Smooth Smooth Smooth or with glandular warts Smooth Smooth Smooth Smooth with glandular warts, Smooth Smooth Glandular warts Glandular warts Dorsal thigh Dorsal Glandular warts Lower Lower flank Granular or Granular smooth Granular Glandular warts Granular Glandular and warts spinules horny Smooth Smooth or with glandular warts Glandular warts Smooth Finely granular Glandular with warts, spinules horny in males Upper flank Smooth glandular Few males warts, with glandular and warts spinules horny Glandular and warts spinules horny (females lack horny spinules) Heavily tuberculated, glandular warts spinules Horny Glandular warts Smooth granular Finely Glandular with warts, spinules horny in males Posterior Posterior dorsum Smooth or granular Glandular and warts horny spinules (females lack horny spinules) Heavily tuberculated, glandular warts Horny spinules with a “)(”shaped of pattern tubercles Glandular warts Smooth Finely granular Glandular warts, with horny spinules in males Anterior Anterior dorsum Glandular and warts horny spinules Glandular warts Smooth Glandular warts Smooth Finely granular Glandular warts Side of head Few Few glandular warts Glandular warts Glandular warts and Glandular warts spinules horny Glandular warts spinules Horny Glandular warts Smooth Finely granular Smooth Glandular warts, spinules with horny in males Interorbital area Interorbital Shagreen Shagreen Shagreen males Shagreen, and glandular warts spinules horny Shagreen Shagreen with Shagreen glandular warts Glandular warts and Glandular warts spinules horny Glandular warts spinules Horny Glandular warts Smooth Finely granular Smooth with Glandular warts, spinules in horny males Snout Species name caeruleus P. cavirostris P. cuspis P. decoris P. dimbullae P. eximius P. extirpo P. femoralis P. folicola P. frankenbergi P. fulvus P. hallidayi P. halyi P. P. fergusonianus fergusonianus P. P. auratus P. 3850 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Absent Super- numerary tubercles on palm Smooth Smooth Smooth Smooth Underside Underside of thigh Smooth Smooth Shagreen Smooth Throat Smooth Smooth Smooth Smooth, few scattered glandular warts Smooth Glandular with warts horny spinules Smooth Smooth not Granular, rough Smooth Smooth with a few scattered glandular warts Smooth Glandular warts Smooth Dorsal thigh Dorsal Horny Horny spinules in males Smooth Granular with glandular warts Glandular warts Glandular warts Glandular warts Lower Lower flank Granular with Granular scattered few a glandular warts Granular spinules Horny in males Smooth or granular Smooth Smooth Smooth with glandular warts Granular Glandular warts Glandular warts Glandular warts Smooth Smooth Upper flank Glandular warts folds Glandular Glandular bearing warts spinules horny Smooth or shagreen Smooth Smooth Smooth with a scattered few glandular warts Weakly granular Glandular warts Smooth Glandular warts Tuberculated, horn like spinules Glandular warts Posterior Posterior dorsum Smooth glandular Few warts Glandular bearing warts spinules horny Smooth Smooth Smooth with a few scattered glandular warts Weakly granular Glandular warts Smooth, '\ /' shaped ridg Glandular warts Tuberculated, horn like spinules Glandular warts Anterior Anterior dorsum Smooth glandular Few warts Glandular bearing warts horny spinules Smooth Smooth Smooth Weakly granular Glandular warts Smooth Smooth Glandular warts Horny spinules in males Glandular warts Glandular warts Side of head Smooth Few glandular warts Glandular warts bearing horny spinules Smooth Smooth Smooth granular Weakly Smooth with a few glandular scattered warts Smooth Smooth Smooth or tuberculated Glandular warts spinules in Horny males Glandular warts Smooth Interorbital area Interorbital Smooth glandular warts Few Smooth or shagreen Glandular warts Glandular warts bearing horny spinules Smooth Smooth Smooth granular Weakly Smooth Smooth Smooth Smooth or tuberculated Glandular warts spinules in Horny males Glandular warts Glandular warts Snout Smooth Glandular warts Glandular warts Glandular warts bearing horny spinules Species name hoffmanni P. hoipolloi P. hypomelas P. leucorhinus P. limbus P. macropus P. maia P. malcolmsmithi P. microtympanum P. mittermeieri P. mooreorum P. nasutus P. nemus P. P. lunatus P. nanus P. P. hankeni P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3851 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Super- numerary tubercles on palm Smooth Smooth Smooth Underside Underside of thigh Smooth Glandular warts Throat Horny Horny spinules in males, finely or granular in shagreen female Smooth Glandular warts Glandular warts Glandular warts Smooth Smooth Glandular warts Smooth Smooth Smooth, few scattered glandular warts Smooth Smooth Dorsal thigh Dorsal Smooth Smooth Glandular Lower Lower flank Smooth Glandular warts Granular Smooth Glandular Glandular warts Granular folds Glandular Upper flank Horny spinules Horny Smooth Glandular warts Smooth spinules Horny (spinules in Absent females) spinules Horny Smooth a for except scattered few glandular warts Glandular warts Glandular males warts, with horny spinules scattered Posterior Posterior dorsum Horny spinules Horny Smooth Glandular warts Smooth Horny spinules (spinules in Absent females) Horny spinules Smooth a for except scattered few glandular warts Glandular warts Glandular males warts, with horny spinules scattered Anterior Anterior dorsum Horny Horny spinules Smooth Glandular warts Smooth Smooth Horny spinules Smooth except a few for scattered glandular warts Glandular warts Glandular warts Side of head Glandular warts Smooth Glandular warts Smooth Smooth spinules Horny for Smooth except scattered a few glandular warts Smooth Glandular warts Interorbital area Interorbital Shagreen or with Shagreen glandular warts Shagreen spinules Horny in males, finely or shagreen granular in female with a few Shagreen glandular scattered warts Smooth Smooth Glandular warts Smooth Smooth spinules Horny for Smooth except scattered a few glandular warts Smooth Glandular warts Snout Glandular warts Species name oxyrhynchus P. papillosus P. pardus P. pleurotaenia P. poppiae P. procax P. regius P. reticulatus P. rugatus P. rus P. sarasinorum P. P. popularis P. P. ocularis P. 3852 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Super- numerary tubercles on palm Smooth Smooth Underside Underside of thigh Smooth Smooth Smooth or shagreen Throat Smooth Glandular warts Smooth Smooth Smooth Smooth Smooth Glandular warts Smooth, scattered glandular warts Smooth Dorsal thigh Dorsal Smooth Scattered glandular tubercles Glandular warts Weakly areolate Smooth Lower Lower flank Smooth Scattered, glandular tubercles Glandular and warts horny spinules Small tubercules with horny spinules Glandular warts Glandular warts Glandular with warts, spinules horny in males Glandular and warts spinules horny Smooth Upper flank Smooth Scattered, glandular tubercles Glandular folds, glandular and warts spinules horny Small tubercules with horny spinules Glandular warts Smooth Smooth Glandular males warts, spinules horny scattered Glandular and warts horny spinules, lack females spinules horny Horny spinules, lack females spinules horny Posterior Posterior dorsum Smooth Scattered, glandular tubercles Glandular folds, glandular and warts spinules horny Small tubercules with horny spinules Glandular warts Glandular warts Glandular warts Glandular warts, males horny spinules scattered Glandular and warts horny spinules, females lack horny spinules Horny spinules, females lack horny spinules Anterior Anterior dorsum Smooth Scattered, glandular tubercles Glandular folds, glandular and warts horny spinules Small tubercules with horny spinules Glandular warts Glandular warts Glandular warts Glandular warts Smooth or with glandular warts Smooth or with glandular warts Smooth Side of head Smooth Scattered, glandular tubercles Glandular warts Small tubercules Small tubercules spinules with horny Smooth Glandular warts Smooth Glandular warts and Glandular warts spinules horny Smooth spinules Horny Interorbital area Interorbital Scattered, glandular Scattered, tubercles shagreen Weakly Smooth Glandular folds, Glandular folds, and glandular warts spinules horny Small tubercules with Small tubercules spinules horny Smooth Glandular warts Smooth Glandular warts and Glandular warts spinules horny Smooth spinules Horny Snout Smooth Glandular warts Species name schneideri P. silus P. silvaticus P. simba P. singu P. sordidus P. steineri P. stellatus P. stictomerus P. stuarti P. P. semiruber P. P. schmarda P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3853 Eight new species of Pseudophilautus Wickramasinghe et al. Super- numerary tubercles on palm Smooth Underside Underside of thigh Smooth Smooth Throat Smooth Smooth Smooth Weakly tubercular Glandular warts Glandular warts Dorsal thigh Dorsal Lower Lower flank Granular Granular Smooth Granular spinules Horny (smooth in and females immature males) Granular Glandular warts Upper flank Weakly tubercular Smooth Smooth spinules Horny (smooth in and females immature males) Glandular warts spinules Horny Posterior Posterior dorsum Weakly tubercular Smooth Smooth Horny spinules (smooth in and females immature males) Glandular warts Horny spinules, Anterior with dorsum “)(”–shaped of pattern tubercles Anterior Anterior dorsum Smooth Smooth Horny spinules (smooth in and females immature males) Glandular warts Glandular warts Side of head Smooth Smooth spinules Horny (smooth in females and immature males) Glandular warts spinules Horny Interorbital area Interorbital Shagreen Smooth Smooth spinules Horny (smooth in females males) and immature Glandular warts spinules Horny Snout Species name temporalis P. variabilis P. viridis P. zal P. zimmeri P. zorro P. P. tanu P. 3854 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. 0.9); distance between front of eyes half the distance prominent tubercles at heel; tarsal fold absent but tarsus between back of eyes (DBE/DFE 2.0); nostrils oval shagreen with prominent tubercles; foot shagreen. without flap of skin laterally, closer to tip of snout; pupil Ventral side of body: Throat, chest and belly granular; horizontally elliptical; tympanum absent; pineal ocellus upper arm granular; forearm weakly granular; thigh and vomerine teeth absent; tongue large, lanceolate, granular, leg smooth; tarsus weakly granular. lingual papilla absent, but conical tubercles present on Colour in life: Dorsum bronze with dark brown tongue. patches, vertebral stripe off white, a pair of broad dark Arm short, robust (LAL/FEL 0.6, UAL/FEL 0.4); forearm brown longitudinal dorsal bands extends from the back shorter than hand length (LAL/HNL 0.9), longer than of the eye to the groin, dark brown cross band between upper arm (LAL/UAL 1.4); fore arm distinctly enlarged.; eyes, small bronze blotch on snout, dark brown band fingers robust, (Fig. 15), relative length of fingers I < II < on canthal edge, below eye bronze patch, lateral body IV < III (FL-1/FL-3 0.5, FL-2/FL-3 0.6, FL-4/FL-3 0.9) (Table lighter with a green tinge dark brown spots marbled 13); tips of finger disks semi circular, enlarged, discs with off white spots; limbs dorsally bronze, fore limbs, present on all fingers, with distinct basal and circum hind limbs, fingers and toes with brown cross bands, marginal grooves; dermal fringe present on inside of all finger tips off white; ventral side off white all over, hands fingers, but indistinct dermal fringe present on exterior feet and webbing darker (Images 26 & 28). edge of finger IV and hand, absent on ulnar fold; webbing Colour in alcohol: All bronze in life turns to dark formula I2-—2II2-—3+III3—3IV; subarticular tubercles brown, and the colour pattern remains with a little prominent, rounded, single, all present, IV2 and III2 fading (Image 27). relatively smaller; inner palmar tubercle, large, single, Etymology: The species epithet newtonjayawardanei oval, prominent; outer palmar tubercle single, oval, named after Newton Jayawardane, a leading surgeon distinct, larger than the distal subarticular tubercles; by profession and is an ardent naturalist, in great supernumerary tubercles prominent and widespread on appreciation of his dedicated service to the country, palm and on all fingers; prepollex absent; femur 2 1/4 both as a surgeon and a wildlife conservationist. times longer than fourth toe length (FEL/TL-4 2.3); foot Suggested vernacular names: Newton Jayawardanege length longer than thigh (FOL/FEL 1.5); toes thin, (Fig. panduru madiya and Newton Jayawardane’s Shrub Frog 16), relative length of toes I < II < III Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3855 Eight new species of Pseudophilautus Wickramasinghe et al. Figure 17. Pseudophilautus puranappu sp. nov. holotype (NMSL 2013.06.01 NH), head; (A) lateral aspect (B) dorsal aspect (C) ventral aspect. Scale bar: 2mm. (© Dulan Ranga Vidanapathirana) Figure 18. Pseudophilautus puranappu sp. nov. holotype (NMSL 2013.06.01 NH), ventral aspect of left hand. Scale bars: 5mm. (© L.J. Mendis Wickramasinghe) © L.J. Mendis Wickramasinghe Image 29. Pseudophilautus puranappu sp. nov. holotype in life (NMSL 2013.06.01 NH). (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 Figure 19. Pseudophilautus puranappu sp. nov. holotype (NMSL smooth, with a blunt tubercle; interorbital area and 2013.06.01 NH), ventral aspect of left foot. Scale bar: 5mm. (© L.J. Mendis Wickramasinghe) side of head smooth; anterior dorsum and dorsal thigh 3856 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. © L.J. Mendis Wickramasinghe Image 30. Type series of Pseudophilautus puranappu sp. nov. Left to right, NMSL 2013.06.01 NH, DWC 2013.01.009, DWC 2013.01.010. A-C dorsal aspect and D-F ventral aspect. Scale bar: 5mm. weakly shagreen; posterior dorsum few horny spinules, horizontally elliptical; tympanum prominent, oval prominent tubercles; upper flank weakly tubercular; (TYH/TYW 1.2), vertical, smaller than the eye (TYH/ED lower flank granular; throat, chest and underside of 0.3); tympanum-eye distance 4/5th tympanum width thigh weakly granular; supernumerary tubercles on foot (TAD/TYW 0.8); pineal ocellus absent; vomerine teeth indistinct. Comparative characters that distinguish the present, (N=7) prominent, odontophores oblique and species from its congeners are provided in tables 17 and widely separated, between choanae with an angle of 18. 65° relative to body axis; tongue large, tip bifurcate, lanceolate, lingual papilla and conical tubercles present Description of holotype: moderate size (SVL 34.2), on tongue. body moderately elongate (SVL/HW 2.6); head large Arm short (HNL/LAL 1.1); lower-arm longer than (HL/SVL 0.4), concave above; snout rounded in lateral, upper arm (UAL/LAL 0.7); fore-arm distinctly enlarged; truncate in dorsal and nearly rounded in ventral aspects fingers long and thin (Fig. 18), relative length of fingers (Fig. 17 A–C) (ES/DFE 0.7, SN/IN 0.7), its length shorter I < II Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3857 Eight new species of Pseudophilautus Wickramasinghe et al. © L.J. Mendis Wickramasinghe © L.J. Mendis Wickramasinghe Image 31. Pseudophilautus puranappu sp. nov. paratype in life Image 32. Pseudophilautus puranappu sp. nov. paratype in life (DWC 2013.01.009). (DWC 2013.01.010). © L.J. Mendis Wickramasinghe © L.J. Mendis Wickramasinghe Image 33. Pseudophilautus puranappu sp. nov. unpreserved Image 34. Pseudophilautus puranappu sp. nov. unpreserved specimen. specimen. rounded, single, all present, IV2 & III2 relatively smaller; (IML/TL-1 0.50), outer metatarsal tubercle absent. inner palmar tubercle single, oval, prominent, equal to Dorsal snout smooth with a blunt tubercle, lateral outer palmer tubercle; nuptial pads present; outer palmar snout smooth; between eyes smooth; upper eyelid tubercle single, oval, and prominent; supernumerary tubercular; head laterally smooth with prominent tubercle present on all fingers; prepollex absent; femur tubercles present near the gape of mouth; supratympanic about twice as long as the fourth toe length (FEL/TL-4 fold prominent; a median dermal ridge extends from 2.2); foot length longer than thigh (FOL/FEL 1.5); toes tip of snout to back of the head, with tubercles; back long (Fig. 19), thin; relative length of toes I 3858 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Range 3.0—4.7 0.9—2.0 6.1—8.9 5.0—7.6 6.2—8.4 6.3—8.7 3.5—4.8 1.4—2.2 2.3—3.4 1.6—2.4 5.3—8.1 2.9—4.2 8.3—12.1 8.5—12.9 7.7—13.1 9.9—13.6 14.5—21.7 33.6—51.1 15.4—23.2 12.0—17.1 18.6—27.9 12.1—16.9 SD 0.9 3.7 0.6 2.1 2.3 1.6 1.4 1.2 1.3 9.9 0.7 0.4 0.6 0.4 1.6 0.7 2.7 4.1 2.1 2.9 5.1 2.8 3.9 1.4 9.6 7.1 6.0 7.0 7.3 4.0 1.7 2.7 2.0 6.3 3.5 17.6 10.3 39.6 10.1 18.7 11.2 13.7 22.1 13.7 Mean 4.7 2.0 8.9 7.6 8.4 8.7 4.8 2.2 3.4 2.4 8.1 4.2 21.7 12.1 12.9 51.1 13.1 23.2 13.6 17.1 27.9 16.9 Paratype 2 Paratype 2013.01.010 8.5 8.5 3.0 0.9 6.1 5.0 6.4 6.7 3.5 1.4 2.3 1.6 5.5 7.7 9.9 2.9 14.5 33.6 15.4 12.0 18.6 12.1 Paratype 1 Paratype 2013.01.009 1.9 8.3 9.3 4.0 1.3 6.3 5.6 6.2 6.3 3.7 1.6 2.4 5.3 9.6 3.3 16.5 34.2 10.2 17.4 12.1 19.8 12.2 Holotype 2013.06.01 NH KT SN LAL SVL TBL TAS MN VKL TYH TAD UAL TPD TL-4 TL-3 TL-5 TL-2 TL-1 TND MFE TYW MBE UEW DWC NMSL/ sp. nov. Range 3.6—5.5 3.2—4.7 4.9—7.1 3.1—4.3 6.2—8.7 1.7—2.8 1.6—1.9 5.2—6.6 6.3—9.5 1.7—2.1 3.7—6.2 2.7—4.6 0.9—1.4 6.9—10.1 7.2—10.9 10.9—15.6 26.4—37.8 13.6—19.1 15.0—22.9 13.0—20.3 11.9—17.3 17.8—25.8 SD 1.0 2.5 6.4 1.9 0.8 1.2 0.7 1.4 3.2 4.4 0.6 4.0 0.2 0.7 1.9 1.6 3.0 0.2 1.3 1.0 4.6 0.3 of Pseudophilautus puranappu 4.3 8.0 3.8 5.8 3.5 7.1 2.1 1.7 5.8 8.7 7.7 1.9 4.9 3.5 1.1 12.7 30.4 15.5 17.9 15.6 13.8 20.5 Mean 5.5 4.7 7.1 4.3 8.7 2.8 1.9 6.6 9.5 2.1 6.2 4.6 1.4 15.6 37.8 10.1 19.1 22.9 20.3 10.9 17.3 25.8 Paratype 2 Paratype 2013.01.010 3.6 6.9 3.4 4.9 3.1 6.2 2.0 1.7 5.2 8.1 6.3 1.7 3.7 3.1 1.0 11.6 26.4 13.8 15.0 13.6 11.9 17.2 Paratype 1 Paratype 2013.01.009 3.7 6.9 3.2 5.3 3.2 6.5 1.7 1.6 5.7 7.2 7.2 1.9 4.7 2.7 0.9 10.9 27.0 13.6 15.7 13.0 12.2 18.5 Holotype 2013.06.01 NH IN IO ES HL DL ED EN GK DB HD FEL DW HW FOL IML DFE DBE FL-4 FL-3 FL-2 FL-1 HNL DWC NMSL/ Table 16. The morphometric measurements (mm) of the type series measurements 16. The morphometric Table Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3859 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Nuptial pad Nuptial Calcar Absent Present Present Present Present Present Present Present fingers Absent Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Fringe on Fringe papilla Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Lingual Present teeth Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Vomerine Vomerine Flat Flat Flat Flat Flat Flat Flat Flat Flat space Concave Internarial Internarial Flat space Convex Convex Convex Convex Convex Convex Convex Convex Concave Concave Concave Concave Concave Interorbital Interorbital Flat Flat Loreal Loreal region Convex Concave Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp rostralis Canthus Canthus Rounded sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners sp. nov. Oval Blunt Blunt Truncate Truncate Truncate Truncate Truncate Truncate Rounded Snout lateral Pointed or oval Pointed Sharply Pointed Obtusely pointed Obtusely Obtusely pointed Obtusely Obtusely pointed Obtusely Obtusely pointed Obtusely Flat Flat Flat Flat Flat Flat Flat Flat Flat Convex Convex Convex Convex Convex Convex Convex Convex Convex Convex Concave Head dorsally Pseudophilautus puranappu SVL 43.5 35.3 38.4 44.8 35.8-40.4 16.2–19.2 24.5–30.1 23.7–29.4 26.9–44.5 23.4–28.8 18.3–23.9 17.6–28.9 18.9–27.3 17.3–20.2 17.0–27.5 33.4–46.8 26.7–29.3 38.4–48.9 16.3–19.0 21.7–26.4 33.3–41.7 25.4–37.0 20.4–24.6 22.3–32.5 33.6–51.1 sp. sp. nov. sp. nov. sp. nov. sp. nov. P. folicola P. P. fergusonianus fergusonianus P. P. femoralis femoralis P. P. extirpo P. P. eximius eximius P. P. decoris P. P. cuspis P. P. asankai P. nov. P. bambaradeniyai P. alto P. P. newtonjayawardanei newtonjayawardanei P. sp. nov. P. fulvus P. P. frankenbergi frankenbergi P. Species name P. dimbullae P. P. cavirostris cavirostris P. P. caeruleus P. P. auratus P. P. adspersus P. P. abundus P. P. samarakoon samarakoon P. P. sirilwijesundarai P. sp. nov. P. karunarathnai karunarathnai P. P. dayawansai P. jagathgunawardanai P. sp. nov. P. puranappu P. Table 17. Characters that differentiate differentiate that 17. Characters Table 3860 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Nuptial pad Nuptial Calcar Present Present Present Present Present Present Present Present. fingers Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Present Fringe on Fringe papilla Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Lingual Absent teeth Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Vomerine Vomerine Flat Flat Flat Flat Flat Flat Flat Flat space Convex concave Convex or Convex Internarial Internarial space Convex Convex Convex concave Concave Concave Concave Concave Concave Concave Concave Concave Convex or Convex Interorbital Interorbital Flat Flat Flat Flat Loreal Loreal region Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp rostralis Canthus Canthus Indistinct Oval Oval Oval Oval Oval Oval Blunt Blunt Pointed Pointed Pointed Pointed Pointed Truncate Truncate Truncate Acuminate Snout lateral Pointed or oval Pointed Truncate or oval Truncate Obtusely pointed Obtusely Obtusely pointed Obtusely Flat Flat Flat Flat Flat Flat Flat Convex Convex Convex Convex Convex Convex Convex Convex Convex Convex Convex Convex Uneven Convex or flat Convex Head dorsally Convex or Flat Convex 40 SVL 32.1 17.4 14.9 46.6 25.7 19.8 27.9 18.6 34.8 20.7 40.9 42.5–61.1 25.1–26.8 23.1–33.4 16.3–18.4 23.9–49.0 27.4–42.7 18.3–21.7 17.7–24.7 21.3–26.0 18.3–21.9 25.6–30.3 29.4–35.0 11.2–22.4 22.0–28.6 21.2–23.4 32.9–42.9 P. reticulatus P. P. procax P. P. pardus P. P. papillosus P. P. ocularis P. P. nasutus P. P. mittermeieri mittermeieri P. P. microtympanum P. P. malcolmsmithi malcolmsmithi P. P. maia P. P. macropus P. P. limbus P. P. leucorhinus leucorhinus P. P. regius P. P. popularis P. P. poppiae P. P. hankeni P. P. halyi P. pleurotaenia P. Species name P. oxyrhynchus P. P. nanus P. P. mooreorum P. P. hypomelas hypomelas P. P. hoipolloi P. P. hoffmanni P. P. nemus P. P. lunatus P. P. hallidayi P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3861 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Nuptial pad Nuptial Calcar Present Present Present fingers Present Present Present Present Present Present Present Present Present Fringe on Fringe papilla Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Lingual teeth Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Vomerine Vomerine Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat Flat space Internarial Internarial space Convex Convex Convex Convex Convex Convex Convex Concave Concave Concave Interorbital Interorbital Flat Flat Flat Flat Loreal Loreal region Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp rostralis Canthus Canthus Indistinct Oval Oval Oval Oval Blunt Blunt Pointed Pointed Truncate Truncate Snout lateral Blunt or rounded Blunt Obtusely pointed Obtusely Obtusely pointed Obtusely Obtusely pointed Obtusely Flat Flat Flat Flat Flat Convex Convex Convex Convex Convex Convex Convex Convex Convex Convex Convex Convex Convex above above Convex Flat or convex Flat Head dorsally SVL 31.6 35.7 13.4 17.5 22.9–30.1 27.4–36.3 24.2–32.4 23.2–36.0 39.6–55.3 24.3–31.3 20.6–24.1 20.0–32.5 28.4–31.0 13.5–13.9 30.4–41.6 22.2–39.4 16.1–16.6 12.6–15.6 35.4–51.8 17.7–30.0 22.6–38.1 19.9–22.8 P. zorro P. P. zimmeri P. P. viridis P. P. stuarti stuarti P. P. stictomerus P. P. stellatus P. P. silvaticus P. P. rus P. Species name P. zal P. P. variabilis P. P. temporalis temporalis P. P. tanu P. P. steineri steineri P. P. sordidus P. P. singu P. P. simba P. P. silus P. P. semiruber P. P. schmarda P. P. sarasinorum P. P. schneideri P. P. rugatus P. 3862 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Indistinct Super- numerary tubercles on foot Present Present Present Present Present Present Present Present Present Present Weakly Weakly granular Underside Underside of thigh Smooth Weakly Weakly granular Chest Weakly Weakly granular Throat Shagreen Shagreen Weakly Weakly shagreen Dorsal thigh Dorsal Smooth Weakly tubercular Smooth Tuberculated Smooth Smooth Smooth Smooth Tuberculated Granular Lower Lower flank Smooth Granular Weakly Weakly tubercular Upper flank Smooth Granular Shagreen Areolate Smooth Weakly granular Smooth Horny spinules in males Weakly Weakly tubercular Few horny horny Few spinules, Prominent tubercles Posterior Posterior dorsum Smooth Smooth Shagreen Shagreen sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners sp. nov. Smooth Weakly Weakly shagreen Anterior Anterior dorsum Smooth Prominent Prominent tubercles and horny spinules Smooth Tubercular Horny spinules in (absent females) Smooth a for except scattered few glandular warts Horny spinules in males, warty in females Pseudophilautus puranappu Tubercular Smooth Side of head Weakly Weakly tubercular Weakly Weakly shagreen Shagreen Smooth, with a ridge across breaking medially Smooth Interorbital Interorbital area Smooth,with cross two dermal fringes Weakly tubercular Tubercular A prominent A prominent tubercle present Horny spinules in males Shagreen, Shagreen, with a prominent ridge Tuberculated, Tuberculated, with a single prominent tubercle medially placed Smooth, with a blunt tubercle Snout Shagreen, Shagreen, with prominent dermal fold spinules Horny in males Shagreen sp. nov. sp. nov. sp. nov. sp. nov. P. bambaradeniyai P. sp. nov. P. puranappu P. P. dayawansai P. P. jagathgunawardanai P. sp. nov. karunarathnai P. samarakoon P. sirilwijesundarai P. sp. nov. abundus P. alto P. Species name newtonjayawardanei P. sp. nov. adspersus P. Table 18. Characters in the skin structure that differentiate differentiate that in the skin structure 18. Characters Table Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3863 Eight new species of Pseudophilautus Wickramasinghe et al. Super- numerary tubercles on foot Present Present Present Present Underside Underside of thigh Chest Smooth Throat Smooth Smooth Smooth Smooth Dorsal thigh Dorsal Glandular warts Smooth Smooth Smooth Smooth or with glandular warts Smooth Smooth Smooth Smooth with glandular warts, Smooth Very fine, Very horny spinules Lower Lower flank Glandular warts Very fine, Very horny spinules Upper flank Granular Smooth Glandular warts Finely granular Granular or Granular smooth Shagreen Granular Horny Horny spinules Glandular and warts horny spinules Smooth Smooth or glandular with warts Shagreen, males with glandular and warts horny spinules Posterior Posterior dorsum Smooth Smooth Glandular warts Finely granular Shagreen Shagreen Shagreen, with a few glandular warts Very fine, Very horny spinules Anterior Anterior dorsum Smooth or granular Horny spinules with a “)(”shaped of pattern tubercles Smooth Glandular and warts horny spinules (females lack horny spinules) Heavily Heavily tuberculated, glandular warts Glandular warts Finely granular Very fine, Very horny spinules Side of head Shagreen, smooth or with a few scattered glandular warts Glandular and warts horny spinules Shagreen Shagreen Glandular warts Glandular warts Shagreen Finely granular Few glandular warts Very fine, Very horny spinules Interorbital Interorbital area Glandular and warts horny spinules Horny spinules Shagreen Shagreen Glandular warts Glandular warts Shagreen Finely granular Shagreen, males glandular and warts horny spinules Very fine, Very spinules horny Snout Glandular and warts spinules horny spinules Horny Shagreen, with a few glandular warts Shagreen Glandular warts Glandular warts Shagreen Finely granular Shagreen, males glandular and warts spinules horny P. asankai P. P. caeruleus P. cuspis P. eximius P. extirpo P. Species name auratus P. cavirostris P. decoris P. dimbullae P. femoralis P. fergusonianus P. folicola P. 3864 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Super- numerary tubercles on foot Present Present Present Present Absent Present Present Present Underside Underside of thigh Smooth Smooth Smooth Chest Smooth Shagreen Smooth Smooth Throat Smooth Shagreen Smooth Smooth Dorsal thigh Dorsal Smooth Smooth Smooth, few scattered glandular warts Smooth Glandular warts Glandular warts Glandular with warts horny spinules Smooth Smooth Smooth not Granular, rough Smooth Smooth with a few scattered glandular warts Lower Lower flank Glandular Glandular warts Glandular warts Smooth Upper flank Smooth Smooth with glandular warts Glandular warts Glandular warts Glandular warts Granular Shagreen Glandular warts Smooth or granular Granular with a few scattered glandular warts Shagreen Glandular warts, with horny spinules in males Glandular bearing warts horny spinules Smooth Granular Posterior Posterior dorsum Smooth Smooth Smooth with a few scattered glandular warts Smooth Glandular warts Smooth Weakly Weakly granular Shagreen Shagreen with glandular warts Smooth or shagreen Shagreen or smooth, with a few scattered glandular warts Anterior Anterior dorsum Glandular warts, with horny spinules in males Glandular bearing warts horny spinules Smooth Smooth Smooth with a few scattered glandular warts Smooth Glandular warts Smooth, '\ /' shaped ridge Weakly Weakly granular Glandular warts Side of head Glandular warts Shagreen Glandular warts Glandular warts bearing horny spinules Shagreen Shagreen with glandular warts Weakly granular Shagreen Interorbital Interorbital area Glandular warts, with horny spinules in males Glandular warts bearing horny spinules Shagreen Shagreen with glandular warts Weakly granular Shagreen Snout Glandular with warts, spinules horny in males Glandular bearing warts spinules horny Shagreen with Shagreen glandular warts Weakly granular P. frankenbergi frankenbergi P. P. macropus P. P. hoffmanni P. P. hallidayi P. hankeni P. lunatus P. microtympanum P. Species name fulvus P. halyi P. hoipolloi P. hypomelas P. leucorhinus P. limbus P. maia P. malcolmsmithi P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3865 Eight new species of Pseudophilautus Wickramasinghe et al. Super- numerary tubercles on foot Present Present Present Present Underside Underside of thigh Smooth Smooth Smooth Chest Smooth Throat Smooth Smooth Dorsal thigh Dorsal Smooth Smooth Glandular warts Smooth Glandular warts Glandular warts Smooth Smooth Glandular warts Smooth Horny Horny spinules in males, finely or granular in shagreen female Glandular warts Glandular warts Lower Lower flank Smooth Horny Horny spinules in males Smooth Glandular Glandular warts Upper flank Smooth Smooth Granular Glandular folds Glandular warts Shagreen or Shagreen glandular warts Horny Horny spinules in males Smooth Shagreen, scattered few glandular warts, and horny spinules Horny spinules in males, finely or granular in shagreen female Horny spinules Smooth Glandular Glandular warts Posterior Posterior dorsum Glandular warts Smooth Smooth Few glandular Few warts Glandular warts Shagreen or with glandular warts Glandular warts tuberculated, tuberculated, horn like spinules Anterior Anterior dorsum Glandular warts Horny spinules Smooth Smooth Horny spinules (spinules in Absent females) Few glandular Few warts Glandular warts Horny spinules Glandular warts Glandular warts Side of head Glandular warts Glandular warts Horny Horny spinules in males Few glandular warts Shagreen or with glandular warts Glandular warts Shagreen Horny spinules in males, finely or granular in shagreen female Horny spinules Glandular warts Glandular warts Interorbital Interorbital area Horny Horny spinules in males Few glandular warts Shagreen or with glandular warts Glandular warts Shagreen Horny spinules in males, finely or granular in shagreen female Horny spinules Glandular warts Glandular warts Snout Glandular warts Horny spinules Horny in males Glandular warts or Shagreen with glandular warts Glandular warts Shagreen spinules Horny in males, finely granular in or shagreen female Glandular warts spinules Horny P. mittermeieri mittermeieri P. P. nasutus P. P. nemus P. procax P. Species name mooreorum P. nanus P. ocularis P. oxyrhynchus P. papillosus P. pardus P. pleurotaenia P. poppiae P. popularis P. regius P. 3866 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Super- numerary tubercles on foot Present Present Present Present Present Present Underside Underside of thigh Smooth Smooth Smooth Smooth Chest Smooth Smooth Smooth Throat Smooth Glandular warts Smooth Glandular warts Dorsal thigh Dorsal Smooth Scattered Scattered glandular tubercles Smooth Smooth, few scattered glandular warts Smooth Smooth Smooth Smooth Smooth Smooth Glandular warts Glandular warts Lower Lower flank Smooth Scattered, Scattered, glandular tubercles Glandular and warts horny spinules Upper flank Granular Glandular warts Glandular warts Glandular warts Smooth Scattered, Scattered, glandular tubercles Shagreen Shagreen with a few scattered glandular warts Glandular folds Glandular folds, glandular and warts horny spinules Small tubercules with horny spinules Glandular warts, with horny spinules in males Posterior Posterior dorsum Glandular warts Glandular warts Smooth Smooth Smooth a for except scattered few glandular warts Smooth Scattered, Scattered, glandular tubercles Shagreen Shagreen with a few scattered glandular finely warts, tuberculated Glandular warts, males horny spinules scattered Anterior Anterior dorsum Glandular warts Glandular warts Glandular warts Small tubercules with horny spinules Glandular warts Smooth a for except scattered few glandular warts Smooth Glandular males warts, with horny spinules scattered Glandular folds, glandular and warts horny spinules Scattered, glandular tubercles Shagreen Shagreen with a few scattered glandular warts Glandular warts Side of head Glandular warts Glandular warts Glandular warts Small tubercules with horny spinules Glandular warts Glandular warts Glandular warts Scattered, glandular tubercles Shagreen Shagreen with a few scattered glandular warts Glandular warts Interorbital Interorbital area Smooth except a few for scattered glandular warts Small tubercules with horny spinules Glandular warts Glandular warts Glandular folds, glandular and warts horny spinules Scattered, glandular tubercles Shagreen Shagreen with a few scattered glandular warts Glandular warts Snout Small tubercules with horny spinules Glandular warts Glandular warts Glandular warts Scattered, glandular tubercles P. reticulatus P. P. silus P. P. semiruber P. sordidus P. P. rus P. schneideri P. silvaticus P. Species name rugatus P. sarasinorum P. schmarda P. simba P. singu P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3867 Eight new species of Pseudophilautus Wickramasinghe et al. Super- numerary tubercles on foot Present Present Present Present on Present and toes on foot Present Present Present Underside Underside of thigh Smooth Chest Smooth or sha- green Smooth Mid chest smooth Throat Smooth or shagreen Smooth Smooth Dorsal thigh Dorsal Smooth, scattered glandular warts Smooth Smooth Smooth Smooth Weakly tubercular Glandular warts Glandular warts Smooth Lower Lower flank Weakly Weakly areolate Smooth Upper flank Smooth Granular Granular Granular Granular Glandular and warts horny spinules Shagreen weakly to areolate Shagreen, horny spinules in males Smooth Horny spinules (smooth in and females immature males) Glandular warts Posterior Posterior dorsum Smooth Smooth Weakly tubercular Glandular warts Weakly shagreen Shagreen Anterior Anterior dorsum Glandular and warts horny spinules, females lack horny spinules Horny spinules, females lack horny spinules Smooth Smooth Horny spinules (smooth in and females immature males) Weakly tubercular Glandular warts Horny spinules, Anterior with dorsum “)(”–shaped of pattern tubercles Side of head Weakly shagreen Shagreen Shagreen Glandular warts Glandular warts Glandular and warts horny spinules Interorbital Interorbital area Weakly Weakly shagreen Shagreen Horny Horny spinules Shagreen Glandular warts Horny spinules Glandular and warts spinules horny Snout Weakly shagreen Shagreen spinules Horny (smooth in and females immature males) Horny spinules Horny Weakly shagreen Glandular warts spinules Horny P. steineri steineri P. P. stellatus P. tanu P. temporalis P. variabilis P. viridis P. Species name stictomerus P. stuarti P. zal P. zimmeri P. zorro P. 3868 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Ventral side of body: Throat and chest weakly Pseudophilautus samarakoon sp. nov. granular, belly granular; upper arm and forearm weakly (Figs. 20–22; Images 35–39; Tables 19–21) granular; thigh weakly granular, leg smooth, tarsus urn:lsid:zoobank.org:act:DE23DB51-4DD4-47B0-A974-1F8D70F90E8A weakly granular. Colour in life: Dorsum uniform light brown with Holotype: NMSL 2013.07.01 NH, 03.iv.2010, adult a maroon longitudinal band covering snout to vent female 24.6mm SVL (Images 35, 36 A&D). Sripada (hourglass-shaped), mid dorsally colour discontinues, (Peak Wilderness), Ratnapura District, Sabaragamuwa and gets darker down to vent, a faint strip between eyes Province, Sri Lanka (06048’31”N & 80028’14”E; 1335m) demarcating a fairly lighter triangular region on snout, (Fig. 1), coll. L.J.M.W., D.R.V., S.C.A. & A.W.A.C. canthal edge and tip of snout light brown, laterally lighter Paratypes: DWC 2013.01.011, adult male, 20.4mm with a green tinge; limbs dark brown with a green tinge, SVL (Images 36B,E, 37); DWC 2013.01.012, adult female, forelimbs, hindlimbs, fingers and toes with blackish 22.3mm (Images 36C,F, 38); the same date, locality and maroon bars; ventral side off-white marbled with dark collectors. brown, hands, feet and webbing lighter (Image 29). Diagnosis: Pseudophilautus samarakoon sp. nov. Colour in alcohol: All maroon in life changed to a can be distinguished from known congeners by the blackish tinge, the colour pattern remains with fading following combination of characters: Rather small size (Image 30 A & D). (male SVL 22.3mm; females SVL 22.4–24.6); dorsal head Paratype and variation: Specimens showed dorsal and interorbital space convex; snout laterally truncate; colour variations. Mainly the hour glass maroonish broad canthus rostralis rounded; loreal region concave; patch remains the same in all and only in the holotype internarial space flat; vomerine teeth, lingual papilla the patch discontinued, but the colouration outside this and nuptial pad absent; fringe on fingers and calcar patch on either side of body changes ranging from pink, present; snout prominent tubercle present on centre of and some instances to a yellowish tinge (Image 31–34). snout; interorbital area tubercular; side of head smooth; Etymology: The species epithet puranappu is named anterior and posterior dorsum, upper flank and dorsal after Veera Puran Appu (Veerahennadige Francisco thigh tubercular; lower flank granular; throat and chest Fernando, 1812–1848) Puran Appu was a leader who weakly granular; supernumerary tubercles present on stood up against the might of the British Empire, and foot. died a courageous death, named for his sacrifice for Description of holotype: Body rather small size (SVL freedom of Sri Lanka. The specific name is an invariable 24.6), moderately robust (SVL/HW 2.9); head large size noun in apposition to the generic name. (HL/SVL 0.5), about as wide as long (HW/HL 1.0), convex Suggested vernacular names: Puran Appuge above; snout truncate in lateral, mucronate in dorsal and 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. Figure 20. Pseudophilautus samarakoon sp. nov. holotype (NMSL 2013.07.01 NH), head; (A) lateral aspect (B) dorsal aspect (C) ventral aspect. Scale bar: 2mm. (© Dulan Ranga Vidanapathirana) Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3869 Eight new species of Pseudophilautus Wickramasinghe et al. Figure 21. Pseudophilautus samarakoon sp. nov. holotype (NMSL 2013.07.01 NH), ventral aspect of left hand. Scale bar: 5mm. ©( L.J. Mendis Wickramasinghe) Figure 22. Pseudophilautus samarakoon sp. nov. holotype (NMSL 2013.07.01 NH), ventral aspect of left foot. Scale bar: 5mm. (© L.J. Mendis Wickramasinghe) © L.J. Mendis Wickramasinghe © L.J. Mendis Wickramasinghe Image 35. Pseudophilautus samarakoon sp. nov., holotype in life (NMSL 2013.07.01 NH). Image 37. Pseudophilautus samarakoon sp. nov. paratype in life (DWC 2013.01.011). 3870 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. © L.J. Mendis Wickramasinghe Image 36. Type series of Pseudophilautus samarakoon sp. nov. Left to right, NMSL 2013.07.01 NH, DWC 2013.01.011, DWC 2013.01.012. A-C dorsal aspect and D-F ventral aspect. Scale bar: 5mm. © L.J. Mendis Wickramasinghe © L.J. Mendis Wickramasinghe Image 39. Pseudophilautus samarakoon sp. nov. unpreserved specimen. Image 38. Pseudophilautus samarakoon sp. nov. paratype in life (DWC 2013.01.012). rounded in ventral and aspects (Fig. 20 A–C), its length rounded, loreal region concave; interorbital space mid (ES/DFE 0.8, SN/IN 0.6) longer than horizontal diameter dorsally convex, larger than upper eyelid (IO/UEW 1.6), of eye (ES/ED 1.2); internasal space flat; canthus rostralis larger than internasal distance (IO/IN 1.1); distance Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3871 Eight new species of Pseudophilautus Wickramasinghe et al. between front of eyes 3/5 time the distance between smooth; forearm tubercular; hand smooth; inner and back of eyes (DBE/DFE 1.7); nostrils oval, closer to tip outer thigh smooth, dorsally tubercular, macroglands of snout than to eye (SN/EN 0.6); pupil horizontally absent; leg dorsally tubercular, blunt calcar and small elliptical; tympanum distinct, elliptical (TYH/TYW 1.2), tubercles present at heel; tarsus and foot with a few smaller than the eye (TYH/ED 0.40), tympanum-eye tubercles. Comparative characters that distinguish the distance 4/5th of tympanum width (TAD/TYW 0.8); pineal species from its congeners are provided in tables 20 and ocellus and vomerine teeth absent; tongue lanceolate; 21. lingual papilla absent, with few conical tubercles on Ventral side of body: Throat and chest weakly tongue. granular; belly granular; thigh, leg and tarsus weakly Arm short (LAL/FEL 0.5, UAL/FEL 0.3) thin, lower- granular; vent dorsally shagreen, either side prominent arm shorter than hand length (LAL/HNL 0.8), lower- tubercles present. arm longer than upper arm (LAL/UAL 1.4); fingers thin, Colour in life: Dorsum uniform creamy light brown (Fig. 21), relative length of fingers I < II < IV < III (FL-1/ with dark brown patches, a dark brown band between FL-3 0.5, FL-2/FL-3 0.6, FL-4/FL-3 0.8) (Table 19), tips of eyes and another in front, anteriorly a large dark brown fingers rounded, enlarged, discs present on all fingers, ‘M’ shaped patch with a creamy light brown oval center, with distinct basal and circum marginal grooves; dermal posteriorly two symmetric dark brown diagonal lines; fringe on inside of all fingers, and fringe present on laterally cream colour; limbs dorsally creamy light brown, postaxial edge of finger IV, metacarpal fold weakly forelimbs, hind limbs, fingers and toes with brown present; webbing formula I2 ½–2 ½II2—3 2/3III3+–2 ½; cross bands; ventral side cream, throat brown, feet and subarticular tubercles prominent, rounded, single, all webbing light brown (Image 35). present, IV2 and III2 relatively smaller; inner palmar Colour in alcohol: Colour pattern remains with a little tubercle indistinct, single, oval, larger than outer palmer fading. The colours change from dark brown to brown tubercle; nuptial pads absent; outer palmar tubercle and the cream colour to off white (Images 36 A,D). indistinct, single, oval, smaller than the diatal subarticular Paratype and variation:Dorsum colour varies ranging tubercles; supernumerary tubercles present, indistinct; from a reddish brown to a lighter brown shade, to a prepollex absent; femur twice as long as the fourth toe cream colour. The markings and patterns are the same length (FEL/TL-4 2.2), foot length longer than thigh (FOL/ for all (Images 37–39). The ventral side of thigh granular FEL 1.5), toes thin, (Fig. 22), relative length of toes I < II< III in DWC 2013.01.012, adult female, SVL 22.3mm (Images < V < IV (TL-1/TL-4 0.4, TL-2/TL-4 0.4, TL-3/TL-4 0.7, TL-5/ 36 C,F, 38). TL-4 0.7), tips of toes rounded enlarged, discs present on Etymology: The species name samarakoon is all toes, with distinct basal and circum marginal grooves, named after Ananda Vijith Samarakoon (Assistant webbing formula I2—2-II2+—3III2+—3-IV2 ½—2+V; dermal Director, Department of Wildlife), in great appreciation fringe present on all toes, thin undulating fringe along of his efforts in educating the wildlife officers for the exterior edge of toe V, and metatarsal fold; prominent, conservation of the country’s biodiversity at large. The small tubercles present on outer edge of tarsal fold; species name samarakoon is used as a noun in apposition subarticular tubercles prominent, rounded, single, all to the generic name. present, V2 and IV3, relatively smaller; supernumerary Suggested vernacular names: Samarakoonge tubercles present, indistinct; inner metatarsal tubercle panduru madiya and Samrakoon’s Shrub Frog in Sinhala oval prominent and large, its length 3/4th in length of toe and English, respectively. I (IML/TL-1 0.5); outer metatarsal tubercle indistinct. Natural history: The species was found in elevations Skin between eyes tubercular, with a prominent of 1000–1400 m from the forest floor to about 2m tubercle in the center; a median dermal ridge from tip high shrubs and in bamboo of the lower montane rain of snout to vent (more prominent till back of head) forests. Sympatric with P. bambaradeniyai sp. nov. and P. and a prominent tubercle present on centre of snout; karunarathnei sp. nov.. Most abundantly found near the lateral snout with few tubercles; above nostrils a pair of starting point of Kaluganga River. They were commonly prominent tubercles present; on upper eyelid prominent found perched on bamboo plants close to the stream. 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 3872 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Range 1.2—1.4 1.2—1.4 6.8—9.3 4.8—6.3 3.7—4.7 1.5—2.2 1.8—2.6 3.2—4.1 4.1—6.3 3.8—4.3 3.9—4.6 1.9—2.2 5.9—7.1 3.9—4.6 6.7—7.9 1.0—1.8 0.5—0.9 9.7—11.9 8.2—10.0 10.1—13.4 11.3—15.1 20.4—24.6 SD 1.7 0.1 0.2 1.3 1.9 0.8 0.5 0.4 0.4 0.5 1.2 0.2 0.3 0.2 0.7 0.4 0.7 1.1 1.0 0.4 2.1 0.2 1.4 1.3 8.2 5.6 4.3 1.9 2.3 3.8 5.4 4.0 4.2 2.0 6.7 4.3 7.5 9.3 1.5 0.7 11.7 13.2 10.8 22.4 Mean 1.4 1.2 8.4 5.8 4.7 2.1 2.5 4.0 5.8 4.0 4.2 1.9 7.1 4.6 7.8 9.8 1.6 0.8 11.7 13.2 10.8 22.3 Paratype 2 Paratype 2013.01.012 1.2 1.4 6.8 4.8 3.7 1.5 1.8 3.2 4.1 3.8 3.9 2.2 5.9 3.9 6.7 9.7 8.2 1.0 0.5 10.1 11.3 20.4 Paratype 1 Paratype 2013.01.011 1.2 9.3 6.3 4.5 2.2 2.6 4.1 6.3 4.3 4.6 1.9 7.1 4.4 7.9 1.4 1.8 0.9 13.4 15.1 11.9 10.0 24.6 Holotype 2013.07.01 NH KT SN LAL TBL SVL TAS MN VKL TYH UAL TPD TAD TL-1 TL-2 TL-3 TL-4 TL-5 TND MFE TYW MBE DWC UEW NMSL/ sp. nov. Range 0.3—0.7 7.5—9.2 1.7—2.3 0.9—1.3 2.1—2.8 4.1—5.0 3.2—4.0 4.3—5.5 2.1—2.8 4.5—5.5 5.3—7.9 0.9—1.4 0.8—1.2 2.4—3.2 3.1—3.6 2.2—2.8 3.5—4.4 8.5—11.3 9.4—12.3 9.1—11.2 10.7—13.5 14.3—19.9 SD 0.2 1.6 0.9 1.4 0.3 2.9 0.2 0.4 0.5 0.4 1.4 0.6 0.3 0.5 1.0 1.4 0.3 0.2 0.4 0.3 0.3 0.4 0.5 8.4 9.8 2.1 1.1 2.5 4.6 3.7 4.9 2.4 5.1 6.9 1.1 1.0 2.9 3.4 2.5 4.0 11.7 17.4 10.9 10.3 of Pseudophilautus samarakoon Mean 0.5 8.4 9.7 2.3 1.3 2.7 4.6 3.9 4.9 2.4 5.4 7.5 0.9 1.1 3.0 3.1 2.6 4.1 10.7 18.0 10.9 10.6 Paratype 2 Paratype 2013.01.012 0.3 7.5 8.5 1.7 0.9 2.1 4.1 3.2 9.4 4.3 2.1 4.5 9.1 5.3 0.9 0.8 2.4 3.4 2.2 3.5 10.7 14.3 Paratype 1 Paratype 2013.01.011 0.7 9.2 2.3 1.1 2.8 5.0 4.0 5.5 2.8 5.5 7.9 1.4 1.2 3.2 3.6 2.8 4.4 13.5 11.3 19.9 12.3 11.2 Holotype 2013.07.01 NH IN IO ES DL HL ED EN GK DB HD FEL DW HW FOL IML DFE DBE FL-1 FL-2 FL-3 FL-4 HNL DWC NMSL/ Table 19. The morphometric measurements (mm) of the type series measurements 19. The morphometric Table Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3873 Eight new species of Pseudophilautus Wickramasinghe et al. Present Nuptial Nuptial pad Absent Present Present Present Present Present Present Present Absent Calcar Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Fringe on Fringe fingers Present Absent Absent Absent Absent Absent Absent Present Present Lingual papilla Absent Present Present Present Present Present Present Present Present Present Present Present Present Vomerine Vomerine teeth Absent Present Present Present Present Present Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Internarial Internarial space Flat Concave Concave Concave Concave Concave Concave Concave Concave Flat Flat or flat Concave Flat Concave Concave Flat Flat Concave Flat or flat Concave Flat Flat Flat Flat Flat Interorbital Interorbital space Convex Flat Flat Flat Convex Loreal Loreal region Concave Flat Flat Flat Flat Sharp Sharp Sharp Sharp Sharp Sharp Canthus Canthus rostralis Rounded Sharp Sharp Sharp Sharp Sharp Sharp sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners sp. nov. Rounded Rounded Rounded pointed Obtusely pointed Obtusely Sharply Pointed Rounded Rounded Pointed Blunt Acuminate Snout lateral Truncate Rounded Rounded Pointed or oval Pointed Blunt pointed Obtusely Oval Rounded Blunt Rounded Rounded Rounded Obtusely pointed Obtusely Rounded Head dorsally Convex Concave Flat Concave Concave Flat Flat Flat Concave Flat Flat Flat or Concave flat Flat Flat Flat Uneven Pseudophilautus samarakoon Pseudophilautus samarakoon 16.2–19.2 24.5–30.1 22.3–32.5 SVL 17.3–20.2 20.4–24.6 38.4 23.7–29.4 32.9–42.9 18.3–21.9 33.6–51.1 25.4–37.0 33.3–41.7 17.0–27.5 21.7–26.4 16.3–19.0 38.4–48.9 17.6–28.9 44.8 35.3 43.5 23.4–28.8 26.9–44.5 26.7–29.3 33.4–46.8 21.2–23.4 22.0–28.6 11.2–22.4 35.8-40.4 18.9–27.3 18.3–23.9 27.9 sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. P. sirilwijesundarai P. P. samarakoon samarakoon P. bambaradeniyai P. P. newtonjayawardanei newtonjayawardanei P. folicola P. hallidayi P. hankeni P. Species name dayawansai P. jagathgunawardanai P. karunarathnai P. puranappu P. abundus P. adspersus P. alto P. auratus P. caeruleus P. cavirostris P. cuspis P. dimbullae P. eximius P. extirpo P. femoralis P. fergusonianus P. frankenbergi P. fulvus P. hoffmanni P. hoipolloi P. hypomelas P. P. asankai P. decoris P. halyi P. Table 20. Characters that differentiate differentiate that 20. Characters Table 3874 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Nuptial Nuptial pad Present Present Present Present Present Present Present Present Present Calcar Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Fringe on Fringe fingers Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Present Present Lingual papilla Present Present Present Present Present Present Present Present Present Present Present Vomerine Vomerine teeth Present Present Present Present Present Present Present Present Present Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Internarial Internarial space Concave or Convex concave Concave Concave Concave Concave Flat Concave Concave Concave Flat Concave Flat Flat Concave Flat Flat Flat Flat Flat Concave Flat Flat Flat Concave Flat Flat Flat Flat Interorbital Interorbital space Concave Concave Flat Flat Loreal Loreal region Flat Flat Flat Sharp Sharp Indistinct Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Canthus Canthus rostralis Sharp Sharp Oval Pointed Oval Pointed Oval Pointed pointed Obtusely Rounded Rounded Rounded Rounded pointed Obtusely Pointed Oval Oval Oval Rounded Snout lateral Oval Blunt or oval Pointed Obtusely pointed Obtusely Oval Rounded Pointed Oval Oval Rounded Head dorsally Flat Flat Concave Concave Concave Flat Flat Flat Flat Flat Flat SVL 19.8 17.4 23.1–33.4 25.1–26.8 46.6 42.5–61.1 24.3–31.3 25.7 40.9 27.4–42.7 14.9 23.9–49.0 29.4–35.0 34.8 20.7 18.6 40 25.6–30.3 21.3–26.0 17.7–24.7 18.3–21.7 20.6–24.1 22.6–38.1 17.7–30.0 19.9–22.8 13.4 35.4–51.8 12.6–15.6 16.1–16.6 16.3–18.4 32.1 17.5 P. leucorhinus leucorhinus P. P. nasutus P. ocularis P. procax P. P. maia P. reticulatus P. silvaticus P. Species name limbus P. lunatus P. macropus P. malcolmsmithi P. microtympanum P. mooreorum P. nanus P. nemus P. oxyrhynchus P. papillosus P. pleurotaenia P. poppiae P. popularis P. regius P. rus P. sarasinorum P. schmarda P. schneideri P. semiruber P. silus P. simba P. singu P. P. mittermeieri mittermeieri P. pardus P. rugatus P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3875 Eight new species of Pseudophilautus Wickramasinghe et al. Nuptial Nuptial pad Present Present Present Present Present Calcar Absent Absent Absent Absent Absent Absent Absent Fringe on Fringe fingers Absent Absent Absent Absent Absent Absent Lingual papilla Present Present Present Vomerine Vomerine teeth Present Present Present Present Concave Concave Concave Concave Internarial Internarial space Concave Concave Flat Concave or flat Concave Flat Concave Flat Flat Flat Flat Interorbital Interorbital space Flat Loreal Loreal region Flat Indistinct Sharp Rounded Canthus Canthus rostralis Sharp Sharp Sharp Blunt Rounded pointed Obtusely pointed Obtusely Rounded or rounded Blunt Rounded Blunt Snout lateral Rounded Rounded Pointed Concave Head dorsally Flat Flat Flat Concave SVL 22.2–39.4 24.2–32.4 39.6–55.3 31.6 30.4–41.6 23.2–36.0 13.5–13.9 28.4–31.0 35.7 27.4–36.3 20.0–32.5 22.9–30.1 P. sordidus P. P. stuarti stuarti P. P. stellatus P. zimmeri P. Species name steineri P. stictomerus P. tanu P. temporalis P. variabilis P. viridis P. zal P. P. zorro P. 3876 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Absent Super- numerary on tubercles foot Present Present Chest Weakly Weakly granular Shagreen Shagreen Throat Weakly Weakly granular Smooth Smooth Shagreen Weakly shagreen Smooth Smooth Smooth Smooth Smooth Dorsal thigh Dorsal Very fine, Very horny spinules Tubercular Smooth Lower Lower flank Very fine, Very horny spinules Granular Granular Smooth Smooth Smooth Granular spinules Horny in males Weakly granular Weakly Areolate Upper flank Shagreen or Granular smooth Horny spinules Horny Tubercular Horny spinules Horny Smooth Smooth spinules in Horny in males, warty females Spinulate Shagreen Posterior dorsum Posterior Shagreen Shagreen Very fine, horny fine, horny Very spinules Tubercular sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners sp. nov. Prominent Prominent and tubercles spinules horny Smooth shagreen Weakly for Smooth except scattered a few glandular warts spinules in Horny in males, warty females Smooth Smooth with few Shagreen tubercles Anterior dorsum Anterior Shagreen Shagreen Very fine, horny fine, horny Very spinules Tubercular Pseudophilautus samarakoon Pseudophilautus samarakoon Tubercular Weakly tubercular Weakly shagreen Side of head Shagreen Shagreen Very fine, Very spinules horny Smooth Smooth, with across a ridge breaking medially Smooth Interorbital Interorbital area Shagreen, with Shagreen, a prominent ridge Smooth a for except scattered few glandular warts Shagreen Smooth,with cross two dermal fringes Smooth spinules Horny in males fine, horny Very spinules Tubercular Smooth Snout Shagreen Smooth except a few for scattered glandular warts Shagreen Smooth Shagreen, with prominent dermal fold Smooth Horny spinules in males fine, Very horny spinules Prominent Prominent tubercle on present of centre snout Smooth sp. nov. sp. nov. sp. nov. P. newtonjayawardanei newtonjayawardanei P. sp. nov. adspersus P. auratus P. Species name dayawansai P. jagathgunawardanai P. sp. nov. karunarathnai P. sp. nov. puranappu P. sirilwijesundarai P. sp. nov. abundus P. alto P. asankai P. P. samarakoon samarakoon P. P. bambaradeniyai P. sp. nov. Table 21. Characters in the skin structure that differentiate differentiate that in the skin structure 21. Characters Table Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3877 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Absent Absent Super- numerary on tubercles foot Smooth Chest Smooth Smooth Smooth Smooth Throat Glandular, Glandular, or warty, roughly granular Smooth Smooth Smooth or with glandular warts Smooth Smooth Shagreen, few glandular warts Smooth with Smooth glandular warts, Smooth Smooth Shagreen Glandular warts Dorsal thigh Dorsal Glandular warts Lower Lower flank Glandular warts Smooth Smooth or with glandular warts Smooth Upper flank Glandular warts males Shagreen, with glandular and horny warts spinules Shagreen Shagreen Shagreen Granular Finely granular Granular Glandular warts Glandular warts Glandular warts and horny spinules Horny spinules Horny Smooth Posterior dorsum Posterior Glandular warts glandular Few males warts, with glandular and horny warts spinules horny Shagreen, scattered spinules Shagreen Shagreen with Shagreen, glandular a few warts Finely granular Smooth with Shagreen glandular warts Glandular warts Glandular warts and horny spinules (females lack horny spinules) spinules Horny with a “)(”shaped of pattern tubercles Smooth Anterior dorsum Anterior Glandular warts Males Shagreen, glandular warts spinules and horny horny Shagreen, spinules scattered Shagreen Shagreen with Shagreen, glandular a few warts Finely granular Smooth or granular with Shagreen glandular warts Glandular warts Glandular warts spinules and horny lack (females spinules) horny Side of head Glandular warts glandular Few warts Glandular warts Shagreen Glandular warts Shagreen Shagreen Finely granular Shagreen, smooth or with scattered a few warts glandular with Shagreen warts glandular Glandular and warts spinules horny Interorbital Interorbital area Glandular warts Shagreen, males glandular and warts spinules horny Shagreen Shagreen Glandular warts spinules Horny Shagreen Smooth Shagreen Finely granular Smooth with Shagreen glandular warts Glandular warts Glandular warts and horny spinules Snout Glandular warts Shagreen, males glandular and warts horny spinules Shagreen Shagreen Glandular warts Horny spinules Shagreen Smooth Shagreen, with a few glandular warts Finely granular Smooth Shagreen with glandular warts Glandular and warts horny spinules P. decoris P. fergusonianus P. frankenbergi P. fulvus P. Species name cavirostris P. cuspis P. dimbullae P. eximius P. extirpo P. femoralis P. folicola P. halyi P. P. caeruleus P. 3878 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Absent Absent Absent Super- numerary on tubercles foot Shagreen Smooth Smooth Smooth Chest Shagreen Smooth Smooth Smooth Throat Glandular warts Glandular with warts horny spinules Smooth Smooth Smooth Granular, not rough Smooth Smooth with a few scattered glandular warts Shagreen, few scattered glandular warts Smooth Smooth Smooth, few scattered glandular warts Dorsal thigh Dorsal Smooth Smooth Glandular Glandular warts Glandular warts Lower Lower flank Glandular warts Smooth Smooth Smooth or granular Smooth with glandular warts Glandular warts Glandular warts Glandular warts Granular Upper flank Glandular warts Glandular warts bearing horny spinules Granular with Granular scattered a few glandular warts Glandular warts Glandular warts, Glandular warts, with horny spinules in males Smooth Smooth or shagreen Smooth Smooth with a scattered few glandular warts Smooth Glandular warts Smooth Smooth above, Smooth above, and shagreen with glandular males with warts, spinules horn like Posterior dorsum Posterior Glandular warts Glandular warts bearing horny spinules Weakly granular Weakly or Shagreen smooth, with a scattered few glandular warts Glandular warts Glandular warts, Glandular warts, with horny spinules in males Smooth Smooth or shagreen Smooth Smooth with a scattered few glandular warts Smooth Glandular warts Smooth, '\ /' shaped ridge Smooth above, Smooth above, and with shagreen glandular warts, males with horn spinules like Anterior dorsum Anterior Glandular warts Glandular warts bearing horny spinules Weakly granular Weakly or Shagreen smooth, with a scattered few glandular warts Glandular warts Glandular warts, Glandular warts, with horny spinules in males Shagreen Side of head Glandular bearing warts spinules horny Weakly Weakly granular Glandular warts Glandular warts Glandular warts Smooth Smooth or tuberculated Smooth Interorbital Interorbital area Glandular warts Glandular warts bearing horny spinules Smooth Smooth Smooth Smooth or shagreen Weakly granular Smooth Smooth with a scattered few glandular warts Glandular warts Glandular with warts, spinules horny in males Smooth Snout Smooth Glandular warts bearing horny spinules Smooth Smooth Smooth Smooth or shagreen Weakly granular Smooth Smooth Glandular warts Glandular warts, with horny spinules in males P. hypomelas hypomelas P. P. leucorhinus leucorhinus P. P. hankeni P. maia P. malcolmsmithi P. Species name hoffmanni P. hoipolloi P. limbus P. lunatus P. macropus P. microtympanum P. mittermeieri P. P. hallidayi P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3879 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Absent Absent Absent Super- numerary on tubercles foot Smooth Chest Smooth Throat Smooth Glandular warts Smooth Glandular warts Glandular warts Smooth Smooth Glandular warts Smooth Smooth Glandular warts Smooth Horny Horny spinules in males, finely or granular in shagreen female Dorsal thigh Dorsal Shagreen Glandular Smooth Smooth Lower Lower flank Horny Horny spinules in males Glandular folds or Shagreen glandular warts spinules Horny Glandular Smooth Granular Smooth Smooth Smooth Glandular warts Shagreen, scattered few glandular warts, and horny spinules spinules Horny in males, finely or granular in shagreen female with Shagreen scattered a few glandular warts Upper flank Horny spinules Horny in males Few glandular Few warts or with Shagreen glandular warts spinules Horny spinules Horny Smooth Smooth spinules Horny (spinules Absent in females) Tuberculated, Tuberculated, spinules horn like Glandular warts Glandular warts Shagreen, scattered few glandular warts and horny spinules (males) spinules Horny in males, finely or granular in shagreen female with Shagreen scattered a few glandular finely warts, tuberculated Posterior dorsum Posterior Horny spinules Horny in males, finely or granular in shagreen female Few glandular Few warts or with Shagreen glandular warts spinules Horny spinules Horny Smooth Smooth spinules Horny in (spinules Absent females) tuberculated, horn tuberculated, spinules like Glandular warts Glandular warts with Shagreen scattered a few glandular warts spinules and horny ( males) spinules Horny in males, finely or granular in shagreen female with Shagreen scattered a few glandular warts, finely tuberculated Anterior dorsum Anterior Horny spinules Horny in males, finely or granular in shagreen female Few glandular Few warts or Shagreen with glandular warts Glandular warts spinules Horny Glandular warts Glandular warts Glandular warts Shagreen spinules Horny in males, finely or granular in shagreen female with Shagreen scattered a few warts glandular Side of head Horny spinules Horny in males Few glandular Few warts or Shagreen with glandular warts Smooth spinules Horny Glandular warts Smooth Smooth Glandular warts Smooth Shagreen spinules Horny in males, finely or granular in shagreen female Smooth with Shagreen scattered a few glandular warts Interorbital Interorbital area Horny spinules Horny in males Glandular warts Shagreen or with glandular warts Glandular warts Horny spinules Glandular warts Glandular warts Smooth Glandular warts Smooth Shagreen Horny spinules in males, finely or granular in shagreen female Smooth Shagreen with a few scattered glandular warts Snout Horny Horny spinules in males P. nanus P. ocularis P. popularis P. regius P. Species name nasutus P. nemus P. oxyrhynchus P. papillosus P. pardus P. pleurotaenia P. poppiae P. procax P. reticulatus P. P. mooreorum P. 3880 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Super- numerary on tubercles foot Smooth Smooth Smooth Chest Glandular warts Smooth Smooth Glandular warts Throat Smooth, few Smooth, few scattered glandular warts Smooth Smooth Smooth Smooth Smooth Smooth Glandular warts Smooth, scattered glandular warts Weakly shagreen Scattered Scattered glandular tubercles Smooth Glandular warts Dorsal thigh Dorsal Scattered, Scattered, glandular tubercles Weakly areolate Smooth Glandular warts, horny spinules Lower Lower flank Scattered, glandular tubercles to Shagreen areolate weakly Glandular warts Smooth Glandular warts and horny spinules Granular Glandular folds Glandular folds, glandular warts and horny spinules Small tubercules with horny spinules Glandular warts Glandular warts Glandular warts, with horny spinules in males Upper flank shagreen Weakly Smooth a for except scattered few glandular warts Smooth Glandular warts and horny spinules, females lack horny spinules Glandular warts Glandular warts, males with spinules horny scattered Glandular folds, glandular warts and horny spinules Small tubercules with horny spinules Glandular warts Smooth Smooth Glandular warts, males horny scattered spinules Posterior dorsum Posterior Smooth except for for Smooth except scattered a few glandular warts Smooth Glandular warts and horny spinules, females lack horny spinules shagreen Weakly Glandular warts Glandular warts, males with horny spinules scattered Glandular folds, glandular warts spinules and horny Small tubercules with horny spinules Glandular warts Glandular warts Glandular warts Scattered, glandular tubercles Glandular warts, males horny spinules scattered Anterior dorsum Anterior Glandular warts Glandular warts Glandular warts Small tubercules with horny spinules Glandular warts Glandular warts Glandular warts Scattered, glandular tubercles Glandular warts Weakly shagreen Side of head Smooth Glandular warts Glandular folds, glandular warts and horny spinules Small with tubercules spinules horny Smooth Smooth Glandular warts Smooth Scattered, glandular tubercles Glandular warts Glandular warts and horny spinules shagreen Interorbital Interorbital area Weakly Smooth a for except scattered few glandular warts Smooth Glandular warts Glandular warts Small tubercules with horny spinules Smooth Smooth Glandular warts Smooth Scattered, glandular tubercles Glandular warts Glandular and warts horny spinules Snout Weakly shagreen Smooth except a few for scattered glandular warts Species name rus P. sarasinorum P. schmarda P. schneideri P. semiruber P. silus P. silvaticus P. simba P. singu P. sordidus P. steineri P. stellatus P. P. rugatus P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3881 Eight new species of Pseudophilautus Wickramasinghe et al. Super- numerary on tubercles foot Smooth or shagreen Smooth Mid chest smooth Chest Smooth or shagreen Smooth Smooth Throat Smooth Smooth Smooth Smooth Shagreen Glandular warts Glandular warts Smooth Dorsal thigh Dorsal Smooth Lower Lower flank Granular Shagreen, horny horny Shagreen, spinules in males Smooth Smooth Granular Granular spinules Horny (smooth in and females males) immature Granular Glandular warts Upper flank Shagreen, horny horny Shagreen, spinules in males spinules, Horny lack females spinules horny Smooth Smooth Shagreen spinules Horny (smooth in and females males) immature Glandular warts spinules Horny Posterior dorsum Posterior Shagreen, horny horny Shagreen, spinules in males spinules, Horny lack horny females spinules Smooth Smooth Shagreen spinules Horny (smooth in and females males) immature Glandular warts spinules, Horny dorsum Anterior with “)(”–shaped of pattern tubercles Anterior dorsum Anterior Shagreen spinules Horny (smooth in and females immature males) Shagreen Glandular warts Glandular warts Side of head Horny spinules Horny Shagreen Smooth Smooth spinules Horny (smooth in and females immature males) Shagreen Glandular warts spinules Horny Interorbital Interorbital area Smooth Horny Horny spinules Shagreen Smooth Smooth Horny spinules (smooth in and females immature males) Weakly shagreen Glandular warts Horny spinules Snout Smooth Species name stuarti P. tanu P. temporalis P. variabilis P. viridis P. zal P. zimmeri P. zorro P. P. stictomerus P. 3882 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. 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 Figure 23. Pseudophilautus sirilwijesundarai sp. nov. holotype the following combination of characters: Moderate size (NMSL 2013.08.01 NH), head; (A) lateral aspect (B) dorsal aspect (C) ventral aspect. Scale bar: 5mm. (© L.J. Mendis Wickramasinghe) (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, Figure 24. Pseudophilautus sirilwijesundarai sp. nov. holotype closer to tip of snout than to eye (SN/EN 0.5); tympanum (NMSL 2013.08.01 NH), ventral aspect of left hand. Scale bar: 5mm. indistinct, oval (TYH/TYW 1.2), vertical, smaller than the (© L.J. Mendis Wickramasinghe) 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 than upper arm (LAL/UAL 1.4); fingers long and thin, (Fig. oblique and widely separated, between choanae with 24), relative length of fingers I < II < IV < III (FL-1/FL-3 0.4, an angle of 700 relative to body axis; tongue large, tip FL-2/FL-3 0.6, FL-4/FL-3 0.6) (Tables 22), tips of fingers bifurcate, lanceolate, lingual papilla absent, but conical rounded, enlarged, discs present on fingers I – II – III – IV, tubercles present on tongue. with distinct basal and circum marginal grooves; dermal Arm short (LAL/FEL 0.5, UAL/FEL 0.4) thin, lower-arm fringe on inside of all fingers, webbing formula I2 2/3 – 2 shorter than hand length (LAL/HNL 0.9), lower-arm longer ½II2 – 3-III2 2/3 – 2 2/3IV; prepollex absent; subarticular Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3883 Eight new species of Pseudophilautus Wickramasinghe et al. 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 2/3rd 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 Figure 25. Pseudophilautus sirilwijesundarai sp. nov. holotype back of the head; anterior dorsum shagreen with few (NMSL 2013.08.01 NH), ventral aspect of left foot. Scale bar: 5mm. tubercles, posterior dorsum shagreen; upper part of (© L.J. Mendis Wickramasinghe) 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 © L.J. Mendis Wickramasinghe 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). Image 40. Pseudophilautus sirilwijesundarai sp. nov. holotype in life (NMSL 2013.08.01 NH). 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 tubercles prominent, rounded, single, all present, IV2 named after Siril Wijesundara (Director, Royal Botanical and III2 relatively smaller; inner palmar tubercle single, gardens, Peradeniya, Sri Lanka), in great appreciation of oval, indistinct, larger than outer palmer tubercle; his service to the country. The species name is a noun in 3884 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. © L.J. Mendis Wickramasinghe Image 41. Type series of Pseudophilautus sirilwijesundarai sp. nov. Left to right, NMSL 2013.08.01 NH, DWC 2013.01.013. A-B dorsal aspect and C-D ventral aspect. Scale bar: 5mm. 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 © L.J. Mendis Wickramasinghe on trees along side mosses, from about 10m tall tree branches. Image 42. Pseudophilautus sirilwijesundarai sp. nov. paratype in life (DWC 2013.01.013). Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3885 Eight new species of Pseudophilautus Wickramasinghe et al. Range 1.1—1.4 2.1—4.4 1.5—1.8 4.9—6.3 1.2—1.6 1.0—1.4 4.6—6.7 7.4—9.9 3.8—5.5 5.9—8.9 4.4—6.2 4.7—7.4 2.9—3.6 2.2—3.0 6.0—9.4 7.6—12.1 9.3—13.1 7.2—11.0 22.3—32.5 10.6—18.2 10.6—15.8 13.0—21.1 SD 2.9 0.1 1.6 3.6 0.8 0.1 0.5 0.2 0.1 0.7 0.9 1.3 1.3 0.6 1.1 0.6 0.9 0.2 2.7 0.3 1.2 1.9 1.3 9.9 3.3 1.6 5.6 1.4 1.2 5.7 8.6 9.1 4.7 7.4 5.3 6.0 3.2 2.6 7.7 17.0 27.4 11.2 14.4 13.2 Mean 1.1 7.6 2.1 1.5 4.9 1.2 1.0 4.6 7.4 9.3 7.2 3.8 5.9 4.4 4.7 2.9 2.2 6.0 13.0 22.3 10.6 10.6 Paratype 1 Paratype 2013.01.13 1.4 4.4 1.8 6.3 1.6 1.4 6.7 9.9 5.5 8.9 6.2 7.4 3.6 3.0 9.4 21.1 12.1 32.5 13.1 11.0 18.2 15.8 Holotype 2013.08.01 NH KT SN LAL TBL SVL TAS MN VKL TAD TYH UAL TPD TL-3 TL-4 TL-5 TL-2 TL-1 TND MFE TYW MBE UEW NMSL/ DWC Range 2.8—3.5 3.5—5.3 2.5—3.4 2.5—4.9 0.8—1.8 4.0—5.2 1.2—1.7 5.0—7.4 1.0—1.7 4.8—5.9 5.9—6.8 4.7—6.8 0.6—1.0 3.3—4.3 2.7—4.4 8.1—10.9 9.8—13.1 8.9—12.4 12.3—18.1 10.1—12.7 17.2—27.3 10.8—18.5 SD 0.2 0.7 2.1 0.3 0.9 1.0 1.2 0.4 0.4 0.9 0.2 0.9 0.2 0.4 3.6 2.7 0.3 0.7 0.2 0.4 1.2 0.6 of Pseudophilautus sirilwejasundarai sp. nov. 3.1 4.4 3.0 3.7 9.5 1.3 4.6 1.4 6.2 1.3 5.3 6.4 5.8 0.8 3.8 3.5 15.2 11.5 11.4 22.2 14.6 10.7 Mean 2.8 3.5 2.5 9.8 2.5 8.1 0.8 4.0 1.2 5.0 1.0 4.8 5.9 8.9 4.7 0.6 3.3 2.7 12.3 10.1 17.2 10.8 Paratype 1 Paratype 2013.01.13 3.5 5.3 3.4 4.9 1.8 5.2 1.7 7.4 1.7 5.9 6.8 6.8 1.0 4.3 4.4 18.1 13.1 10.9 12.7 27.3 18.5 12.4 Holotype 2013.08.01 NH IN IO ES HL DL ED EN GK DB HD FEL DW HW FOL IML DFE DBE FL-1 FL-4 FL-3 FL-2 HNL NMSL/ DWC Table 22. The morphometric measurements (mm) of the type series measurements 22. The morphometric Table 3886 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Present Nuptial pad Nuptial Absent Present Present Present Present Present Present Present Present Present Calcar Present Absent Present Present Present Present Absent Fringe on Fringe fingers Present Absent Absent Absent Absent Lingual papilla Present Present Absent Present Present Present Present Vomerine Vomerine teeth Absent Absent Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Internarial Internarial space Flat Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Interorbital Interorbital space Concave Flat Flat Convex Concave Concave Concave Flat Flat or Concave flat Flat Flat Flat or Concave flat Flat Concave Flat Flat Flat Flat Loreal Loreal region Concave Flat Flat Canthus Canthus rostralis Sharp Sharp Rounded Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Snout lateral Obtusely pointed Obtusely Truncate Rounded Blunt Truncate Acuminate Pointed or oval Pointed Truncate pointed Obtusely Truncate Truncate Oval Sharply Pointed Obtusely pointed Obtusely Obtusely pointed Obtusely Blunt Head dorsally Flat Flat Convex Flat Concave Concave Concave Flat Flat Flat or flat Concave Flat Concave Flat Flat Uneven sp. nov. among congeners of the genus Pseudophilautus of the genus among congeners Pseudophilautus sirilwijesundarai sp. nov. SVL 18.3–23.9 44.8 35.3 20.0–32.5 22.3–32.5 43.5 17.3–20.2 17–22.4 16.2–19.2 38.4 33.6–51.1 25.4–37.0 33.3–41.7 17.0–27.5 20.4–24.6 33.4–46.8 26.7–29.3 23.4–28.8 26.9–44.5 23.7–29.4 38.4–48.9 17.6–28.9 16.3–19.0 18.9–27.3 21.7–26.4 27.9 32.9–42.9 sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. Species name P. decoris P. dimbullae P. eximius P. P. bambaradeniyai P. P. extirpo P. P. dayawansai P. jagathgunawardanai P. P. karunarathnai karunarathnai P. newtonjayawardanei P. P. puranappu P. samarakoon P. sirilwijesundarai P. abundus P. adspersus P. alto P. P. fulvus P. P. frankenbergi P. P. femoralis P. fergusonianus P. folicola P. P. cavirostris P. cuspis P. P. caeruleus P. P. asankai P. auratus P. P. halyi P. P. hallidayi P. Table 23. Characters that differentiate differentiate that 23. Characters Table Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3887 Eight new species of Pseudophilautus Wickramasinghe et al. Present Present Present Present Present Nuptial pad Nuptial present Present Present Present Present Present Present Present Calcar Present Present Present Absent Absent Absent Absent Absent Fringe on Fringe fingers Absent Absent Absent Absent Absent Absent Absent Present Present Present Present Lingual papilla Present Present Absent Absent Absent Absent Absent Absent Absent Absent Vomerine Vomerine teeth Absent Absent Absent Absent Absent Absent Absent Absent Absent Concave Concave Concave Concave Concave Concave Concave Concave Concave Internarial Internarial space Concave Concave Concave Concave Concave Concave Concave Convex or Convex concave Concave Flat Concave Flat Flat Concave Flat Flat Flat Concave Concave Flat Flat Flat Concave Interorbital Interorbital space Flat Flat Flat Concave Flat Flat Flat Concave Concave Flat Flat Flat Loreal Loreal region Flat Flat Sharp Sharp Sharp Sharp Indistinct Sharp Canthus Canthus rostralis Sharp Sharp Sharp Oval Truncate Pointed Oval Pointed Oval Blunt Pointed Truncate Truncate Oval Pointed pointed Obtusely Snout lateral Pointed or oval Pointed Oval Pointed Blunt Acuminate Oval pointed Obtusely Truncate or oval Truncate Oval Concave Flat Flat Flat Head dorsally Concave Flat Flat Flat Concave Flat Flat 27.4–42.7 46.6 14.9 23.9–49.0 16.3–18.4 29.4–35.0 34.8 18.6 40 32.1 25.6–30.3 21.3–26.0 17.7–24.7 40.9 17.4 20.7 SVL 25.1–26.8 18.3–21.7 18.3–21.9 21.2–23.4 22.0–28.6 11.2–22.4 19.8 25.7 23.1–33.4 42.5–61.1 17.5 20.6–24.1 P. macropus P. maia P. malcolmsmithi P. microtympanum P. mittermeieri P. mooreorum P. nanus P. P. oxyrhynchus P. papillosus P. P. pardus P. pleurotaenia P. poppiae P. popularis P. P. lunatus P. P. nasutus P. nemus P. Species name P. procax P. regius P. P. hankeni P. hoffmanni P. hoipolloi P. hypomelas P. leucorhinus P. limbus P. P. ocularis P. P. reticulatus P. rugatus P. rus P. 3888 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Present Nuptial pad Nuptial Present Present Present Present Present Calcar Present Present Present Fringe on Fringe fingers Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Lingual papilla Present Present Present Present Vomerine Vomerine teeth Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Internarial Internarial space Concave Concave Concave Concave Concave Concave Concave Concave Concave Interorbital Interorbital space Flat Flat Concave Flat Concave or Concave flat Flat Flat Flat Flat Flat Concave Flat Flat Loreal Loreal region Flat Flat Flat Flat Canthus Canthus rostralis Sharp Sharp Sharp Sharp Sharp Sharp Sharp Sharp Indistinct Snout lateral Blunt Blunt Obtusely pointed Obtusely Pointed Oval Oval Truncate pointed Obtusely Pointed Oval pointed Obtusely Truncate or rounded Blunt Head dorsally Flat Concave Flat Concave Flat Flat SVL 31.6 22.2–39.4 30.4–41.6 17.7–30.0 19.9–22.8 13.4 35.4–51.8 12.6–15.6 16.1–16.6 39.6–55.3 23.2–36.0 22.6–38.1 22.9–30.1 24.3–31.3 24.2–32.4 13.5–13.9 28.4–31.0 35.7 27.4–36.3 20.0–32.5 Species name P. sordidus P. steineri P. P. schmarda P. schneideri P. semiruber P. silus P. simba P. singu P. stellatus P. stictomerus P. P. sarasinorum P. P. zorro P. P. silvaticus P. stuarti P. tanu P. temporalis P. variabilis P. viridis P. zal P. P. zimmeri P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3889 Eight new species of Pseudophilautus Wickramasinghe et al. Weakly granular Weakly Weakly granular Weakly Throat Shagreen granular Weakly Granular Smooth or granular glandular Granular, smooth in warts, female Smooth Smooth or warty, Glandular, granular roughly granular Weakly Granular granular Weakly granular Coarsely granular Weakly Granular Granular Smooth Weakly tubercular Weakly Upper flank Areolate Smooth Granular Shagreen or smooth Granular and Glandular warts spinules horny Smooth Smooth or with glandular warts Glandular warts Shagreen Weakly granular Weakly Tubercular Smooth Granular spinules in Horny males spinules Horny Smooth spinules, horny Few tubercles Prominent Posterior dorsum Posterior Shagreen spinules Horny tubercular Weakly Glandular warts and horny and horny Glandular warts lack spinules (females spinules) horny tuberculated, Heavily glandular warts spinules Horny Glandular warts Spinulate Tubercular in spinules (Absent Horny females) Smooth spinules in males, Horny in females warty spinules fine, horny Very Smooth Anterior dorsum Anterior with few Shagreen tubercles tubercles Prominent spinules and horny Smooth Glandular warts Glandular warts spinules and horny lack horny (females spinules) tuberculated, Heavily glandular warts spinules with a Horny of pattern “)(”shaped tubercles Glandular warts Smooth Tubercular spinules Horny in females) (absent for Smooth except scattered a few glandular warts spinules in Horny in males, warty females fine, horny Very spinules Smooth Side of head shagreen Weakly Smooth Tubercular Glandular warts and Glandular warts spinules horny Glandular warts Smooth Glandular warts Smooth a few for Smooth except warts glandular scattered Smooth spinules fine, horny Very Weakly tubercular Weakly Smooth Smooth Smooth,with two cross cross Smooth,with two dermal fringes Interorbital area Interorbital tubercle A prominent present Smooth across Smooth, with a ridge medially braking Glandular warts and horny and horny Glandular warts spinules Glandular warts spinules Horny Glandular warts Shagreen Shagreen, with a prominent with a prominent Shagreen, ridge Smooth a few for Smooth except warts glandular scattered spinules in males Horny spinules fine, horny Very Shagreen Smooth Smooth, a prominent blunt blunt Smooth, a prominent present tubercle Smooth Dorsal snout Dorsal Shagreen, with prominent with prominent Shagreen, dermal fold Smooth a single Tuberculated, medially tubercle prominent placed Glandular warts and horny and horny Glandular warts spinules Glandular warts spinules Horny Glandular warts Shagreen Shagreen Smooth a few for Smooth except warts glandular scattered spinules in males Horny spinules fine, horny Very Shagreen Prominent tubercle present present tubercle Prominent of snout on centre Smooth, with a blunt tubercle Smooth, with a blunt sp. nov. sp. nov. sp. nov. P. karunarathnai karunarathnai P. sp. nov. Species name jagathgunawardanai P. sp. nov. P. samarakoon samarakoon P. P. sirilwijesundarai P. sp. nov. bambaradeniyai P. sp. nov. dayawansai P. P. puranappu P. caeruleus P. cavirostris P. cuspis P. decoris P. dimbullae P. P. newtonjayawardanei newtonjayawardanei P. sp. nov. abundus P. adspersus P. alto P. asankai P. auratus P. Pseudophilautus of the genus among congeners Pseudophilautus sirilwijesundarai sp. nov. differentiate that in the skin structure 24. Characters Table 3890 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Weakly granular Weakly Smooth but not Granular, rough Granular Smooth Throat Smooth Granular or smooth Granular Weakly granular Weakly Smooth Granular Granular Smooth, granular in males Granular Granular Granular Smooth or granular Granular Smooth Shagreen, males with Shagreen, and glandular warts spinules horny Smooth with glandular warts Granular Glandular warts Upper flank Smooth Granular Granular Finely granular Shagreen Shagreen Glandular warts, spinules with horny in males Glandular warts Glandular warts bearing horny spinules Smooth Smooth or granular Smooth with a few Granular glandular scattered warts Glandular warts Glandular warts male glandular warts, Few and with glandular warts spinules horny Smooth with a few Smooth with a few warts glandular scattered granular Weakly Smooth Posterior dorsum Posterior Smooth Smooth Smooth Finely granular with Glandular warts, spinules in males horny bearing Glandular warts spinules horny Smooth Glandular warts Smooth Smooth with a few Smooth with a few glandular scattered warts granular Weakly Smooth Anterior dorsum Anterior Smooth Smooth or granular Smooth Finely granular with Glandular warts, spinules in horny males Glandular warts bearing horny spinules Smooth Glandular warts Smooth, '\ /' shaped ridge Smooth granular Weakly Smooth Glandular warts Smooth Side of head Glandular warts Finely granular glandular warts Few Glandular warts bearing Glandular warts spinules horny Smooth Smooth Smooth Smooth Smooth Smooth granular Weakly Smooth scattered Smooth with a few glandular warts Smooth Shagreen Interorbital area Interorbital Shagreen Smooth Finely granular males glandular Shagreen, spinules and horny warts Shagreen with horny Glandular warts, spinules in males with glandular Shagreen warts bearing Glandular warts spinules horny Smooth Smooth or shagreen Smooth Smooth Smooth Smooth granular Weakly Smooth Smooth Smooth with a few Shagreen, glandular warts Dorsal snout Dorsal Smooth Shagreen Finely granular males glandular Shagreen, spinules and horny warts Shagreen with horny Glandular warts, spinules in males with glandular Shagreen warts bearing Glandular warts spinules horny Smooth Smooth or shagreen Smooth Smooth Species name hypomelas P. P. folicola P. P. leucorhinus leucorhinus P. limbus P. lunatus P. macropus P. P. eximius eximius P. extirpo P. P. frankenbergi frankenbergi P. P. femoralis femoralis P. fergusonianus P. fulvus P. hallidayi P. halyi P. hankeni P. hoffmanni P. hoipolloi P. maia P. malcolmsmithi P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3891 Eight new species of Pseudophilautus Wickramasinghe et al. Throat Granular Granular Granular granular Weakly Granular but not Granular rough or with Granular glandular folds Smooth Granular granular Weakly Granular Granular Granular granular Coarsely Granular Smooth or granular granular Weakly Granular Glandular warts Upper flank Granular Glandular warts spinules in Horny males Glandular folds Smooth Smooth or Shagreen glandular warts Smooth Glandular warts Granular few Shagreen, glandular scattered and horny warts, spinules spinules Horny (male), finely or shagreen granular (female) spinules Horny Smooth Glandular with a few Shagreen glandular scattered warts Glandular warts Granular Glandular folds Posterior dorsum Posterior Glandular warts glandular warts Few horn like Tuberculated, spinules Glandular warts Smooth Glandular warts Smooth spinules Horny spinules (spinules Horny in females) Absent spinules Horny a few for Smooth except warts glandular scattered Glandular warts males Glandular warts, spinules with horny scattered Anterior dorsum Anterior Smooth above, Smooth above, and with shagreen glandular warts, males with horn like spinules Glandular warts glandular warts Few horn tuberculated, spinules like Glandular warts Smooth Glandular warts Smooth spinules Horny spinules Horny in (spinules Absent females) spinules Horny for Smooth except scattered a few glandular warts Glandular warts Glandular warts, males with horny spinules scattered Side of head Glandular warts spinules in males Horny glandular warts Few Glandular warts Glandular warts Smooth Glandular warts Smooth Glandular warts Smooth spinules Horny a few for Smooth except warts glandular scattered Glandular warts Glandular warts Interorbital area Interorbital Glandular warts spinules in males Horny glandular warts Few Glandular warts Smooth or with glandular Shagreen warts Smooth Glandular warts Smooth Shagreen spinules (male), finely Horny (femal) or shagreen granular Smooth Smooth spinules Horny with a few Shagreen warts glandular scattered a few for Smooth except warts glandular scattered Smooth Glandular warts Dorsal snout Dorsal Smooth or tuberculated Glandular warts spinules in males Horny Glandular warts Glandular warts Glandular warts or with glandular Shagreen warts Smooth Glandular warts Smooth Shagreen spinules (male), finely Horny (female) or shagreen granular Glandular warts Smooth spinules Horny with a few Shagreen warts glandular scattered a few for Smooth except warts glandular scattered Smooth Glandular warts Species name P. microtympanum P. P. mittermeieri mittermeieri P. mooreorum P. nanus P. nasutus P. nemus P. ocularis P. oxyrhynchus P. papillosus P. pardus P. pleurotaenia P. poppiae P. popularis P. procax P. regius P. reticulatus P. rugatus P. rus P. sarasinorum P. 3892 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Granular Weakly granular Weakly granular Weakly Throat Glandular warts Granular Smooth Granular in males, Granular smooth in females Granular Smooth Granular Granular Granular Granular Smooth Granular Smooth Granular Smooth Upper flank Glandular folds, and glandular warts spinules horny with Small tubercules spinules horny Smooth Glandular warts Glandular warts horny Shagreen, spinules in males Granular Smooth Granular spinules Horny (smooth in females males) and immature Granular Granular Glandular warts Scattered, glandular Scattered, tubercles Glandular warts, spinules with horny in males and Glandular warts spinules horny Horny spinules, females spinules, females Horny spinules lack horny tubercular Weakly Posterior dorsum Posterior glandular Glandular folds, spinules and horny warts with Small tubercules spinules horny Smooth Glandular warts Smooth Smooth Smooth Smooth spinules (smooth Horny and immature in females males) Glandular warts spinules Horny Scattered, glandular Scattered, tubercles males Glandular warts, spinules scattered horny and horny Glandular warts lack spinules, females spinules horny Shagreen, horny horny Shagreen, spinules in males spinules, Horny lack horny females spinules Anterior dorsum Anterior Glandular folds, and glandular warts spinules horny with Small tubercules spinules horny Smooth Glandular warts Glandular warts Glandular warts Shagreen with Shagreen glandular warts Smooth Smooth spinules Horny (smooth in females males) and immature Glandular warts spinules, Horny with dorsum Anterior “)(”–shaped pattern of tubercles Scattered, glandular Scattered, tubercles Glandular warts, spinules males horny scattered Glandular warts spinules, and horny lack horny females spinules Smooth or with glandular warts Smooth Side of head Glandular warts with Small tubercules spinules horny Smooth Glandular warts Glandular warts Glandular warts Smooth Smooth spinules (smooth Horny and immature in female male) Glandular warts Glandular warts Scattered, glandular Scattered, tubercles Glandular warts Smooth or with glandular warts Shagreen Smooth spinules Horny Interorbital area Interorbital glandular Glandular folds, spinules and horny warts with horny Small tubercules spinules Smooth Smooth Glandular warts Smooth shagreen Weakly Smooth Smooth spinules (smooth Horny and immature in females males) Shagreen Glandular warts spinules Horny Glandular warts and horny Glandular warts spinules Shagreen Smooth spinules Horny Scattered, glandular tubercles Scattered, Dorsal snout Dorsal shagreen Weakly Glandular warts with horny Small tubercules spinules Smooth Smooth Glandular warts Smooth Smooth Smooth spinules (smooth in Horny males) and immature females shagreen Weakly Glandular warts spinules Horny Glandular warts and horny Glandular warts spinules P. tanu P. Species name stictomerus P. stuarti P. P. singu P. P. schmarda P. schneideri P. semiruber P. silus P. silvaticus P. simba P. P. temporalis temporalis P. variabilis P. viridis P. zal P. zimmeri P. zorro P. P. sordidus P. steineri P. stellatus P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3893 Eight new species of Pseudophilautus Wickramasinghe et al. DISCUSSION Peak, is a place of worship of all religions in the country and a place of aesthetic beauty. Millions visit this site PCA extracted 10 factors with eigenvalue more every year during the pilgrim season which lasts for a than unity. First three PCA axes explained total 61.29% period of six months, which is the time where a large of the total variation in the data with 34.92% being amount of garbage gets collected, and the natural forest explained by the first component, 16.58% by the second gets over exploited. Illegal gem mining on either sides and 9.78% by the third. PCA ordination on the first two of the river bank within the forest is another potential axes (Fig. 26a) could virtually separate Pseudophilautus threat to the amphibian diversity. Tea cultivation in the newtonjayawardanei, P. sirilwijesundarai, P. surrounding areas are slowly expanding. Illegal tree bambaradeniyai and P. puranappu, however the felling to cultivate tea, has become a major threat in the remaining species showed overlaps within the convex area. hulls. PCA ordination on the first three axes (Fig. 26b) could Amongst the most important findings in this region separate all the eight species. Correlation values for the is the forest die back possibly due to pollution and/ variables along the first three PCA axes are provided in Fig. or climate change (Image 43). It has never been 27. These characters define the separation of different documented in this region before and a large area has species from related species. For instance, variables with now been effected with the natural forest cover slowly negative loadings on first and third principle axis and declining. positive on the second principle component define the Most importantly, although these species have separation ofP. newtonjayawardanei from other species been described from a single locality, it must be as this species have high values for such variables. emphasised that all these species possess quite unique Dendrogram depicting the similarity and differences morphological characters and are well distinguishable among the species (Fig. 28) suggested that the eight from one another. P. newtonjayawardanei sp. nov. as species are divided into three morphometrical clusters. mentioned under its natural history notes is a very rare Pseudophilautus newtonjayawardanei was distinctly species and nothing is known about its behaviour as different from all other species and its separation only two specimens have been observed by us till date. was supported by high bootstrap value of 100. Out The high altitude, low temperatures, misty conditions, of the remaining species, the cluster comprising of tall canopy dwelling habit and small size make it very Pseudophilautus bambaradeniyai, P. karunarathnai, P. difficult to locate the species. Hence, we preserved samarakoon and P. sirilwijesundarai was separated from only a single specimen of the two. We have critically the cluster of P. dayawansai, P. jagathgunawardanai and compared the novel species with the 17 extinct species P. puranappu with high value of bootstrap of 86. of this genus, as several species morphologically similar Statistical analysis presented here has limited have been recorded during these surveys, where further application as the number of individuals in each species studies are ongoing to clarify their identity. The quite is less. However, the analysis does reveal that proposed large number of species descriptions in this area in the species are morphometrically distinct from each other. recent past is due to the increased number of systematic Most importantly, P. newtonjayawardanei, which is surveys being carried out in the region, and many more represented by a single individual, is distinctly different species await their discoveries. from all other species described in the current study. Although Peak Wilderness is part of the recently Further morphometric data of different species could declared Central Hills World Heritage Site, and is one of be helpful in determining the morphological species the few remaining virgin forests with a continuous natural boundaries among the proposed species (Appendix 3). forest cover and altitudinal gradient forest influenced by The conservation status of all species described different climatic zones (Image 44–45), the diversity of here, except for P. newtonjayawardanei sp. nov., have this area remains relatively unknown and hence under been considered Critically Endangered, according to the represented. The new species described here is just a IUCN Red List criteria, because for all species the extent glimpse of what remains to be uncovered. At the current of occurrence is <100km2, are recorded from single rate of destruction of the natural habitats in the region the locations, and their habitats are under severe threat. amphibians are at a higher risk of extinction, where they P. newtonjayawardanei sp. nov., will have to be treated may be extripated without ever being known to science. as Data Deficient as it is described based on a single The novel descriptions presented here hopefully will set specimen. a basis for future conservation efforts in the region. The highest point of this mountain range the Sripada 3894 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Figure 26. Principle Component Analysis (PCA) of morphometric data of eight new species of Pseudophilautus along the first two (a) and three (b) principle axes. Percentages in parenthesis are percent variation explained by each principle axis. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3895 Eight new species of Pseudophilautus Wickramasinghe et al. 1 a 0.8 0.6 0.4 0.2463 0.2338 0.2204 0.208 0.2063 0.1944 0.1831 0.1796 0.1781 0.1787 0.1734 0.1555 0.1485 0.1307 0.2 0.1202 0.1127 0.09973 0.08294 0.02828 0.02005 0 Loading -0.2 -0.0442 -0.01055 -0.01322 -0.01445 -0.116 -0.0675 -0.02565 -0.03941 -0.0901 -0.006387 -0.05856 -0.06704 -0.07168 -0.1242 -0.08775 -0.09065 -0.1591 -0.1834 -0.1789 -0.4 -0.1931 -0.1964 -0.3051 -0.6 -0.3306 -0.8 -1 IN IO DL HL KT ES SN ED EN GK HD DB MN HW IML DW TBL LAL FEL VKL TAD TAS TPD TYH UAL HNL DFE FOL TL-1 TL-2 TL-3 TL-4 TL-5 TND MFE DBE FL-1 FL-2 FL-3 FL-4 MBE TYW UEW 1 b 0.8 0.6 0.4 0.2365 0.2224 0.2174 0.2076 0.2024 0.1927 0.1954 0.1914 0.1877 0.1828 0.1886 0.1813 0.184 0.1736 0.1701 0.161 0.1619 0.1551 0.1527 0.1525 0.1492 0.1499 0.1457 0.1404 0.1481 0.1375 0.1438 0.1423 0.1356 0.128 0.1261 0.1183 0.2 0.1198 0.1097 0.1043 0.09881 0.08902 0.07052 0.05077 0.04469 0.03464 0.01595 0 Loading -0.2 -0.02188 -0.4 -0.6 -0.8 -1 IN IO DL HL KT ES SN ED EN GK HD DB MN HW IML DW TBL LAL FEL VKL TAD TAS TPD TYH UAL HNL DFE FOL TL-1 TL-2 TL-3 TL-4 TL-5 TND MFE DBE FL-1 FL-2 FL-3 FL-4 MBE TYW UEW 1 c 0.8 0.6 0.4 0.3254 0.3093 0.2876 0.24 0.2329 0.1964 0.1582 0.1438 0.2 0.1311 0.1124 0.105 0.1105 0.105 0.08642 0.04795 0.05013 0.05307 0.03884 0.04535 0.0373 0.03458 0.01674 0 Loading -0.2 -0.01077 -0.15 -0.02042 -0.03885 -0.04277 -0.007484 -0.005289 -0.1112 -0.05207 -0.1025 -0.1092 -0.1404 -0.09766 -0.1464 -0.1482 -0.4 -0.1835 -0.1921 -0.2147 -0.2313 -0.2355 -0.2681 -0.6 -0.8 -1 IN IO HL KT DL ES SN GK HD ED EN DB MN HW IML DW TBL LAL FEL VKL UAL TPD TYH TAD TAS HNL FOL DFE TL-1 TL-2 TL-3 TL-4 TL-5 TND MFE FL-1 FL-2 FL-3 FL-4 DBE MBE TYW UEW Figure 27. PCA factor loadings. Correlations for each variable are shown along the first (a), second (b) and third (c) principle component. 3896 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Pseudophilautus sirilwijesundarai 30 Pseudophilautus samarakoon 60 Pseudophilautus karunarathnai 54 Pseudophilautus bambaradeniyai 86 Pseudophilautus jagathgunawardanai 54 Pseudophilautus dayawansai 74 100 Pseudophilautus puranappu Pseudophilautus newtonjayawardanei 5101520253035404550 Distance Figure 28. Dendrogram of morphometric relationship among eight new species of Pseudophilautus. Clustering is performed using Ward’s method on Euclidian distance. Numbers along the branches are bootstrap values in percentage of 10,000 iterations. © L.J. Mendis Wickramasinghe © Dulan Ranga Vidanapathirana © Dulan Ranga Image 43. The forest die back spread over a large area of the Sripada Image 45. Canopy cover of the sub montane forest of Peak Peak, caused due to climate change/pollution. Wilderness. © L.J. Mendis Wickramasinghe Image 44. Panaromic view of the Sripada massif (Peak Wilderness), on a cloudy morning, captured from the Horton Plains National Park. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3897 Eight new species of Pseudophilautus Wickramasinghe et al. REFERENCES Li, J., J. Che, R.W. Murphy, H. Zhao, E. Zhao, D. Rao & Y. Zhang (2009). New insights to the molecular phylogenetics and generic assessment Bahir, M.M., M. Meegaskumbura, K. Manamendra-Arachchi, in the Rhacophoridae (Amphibia: Anura) based on five nuclear and C.J. Schneider & R. Pethiyagoda (2005). Reproduction and three mitochondrial genes, with comments on the evolution of terrestrial direct development in Sri Lankan shrub frogs (Ranidae: reproduction. Molecular Phylogenetics and Evolution 53: 509–522. Rhacophorinae: Philautus). The Raffles Bulletin of Zoology, Manamendra-Arachchi, K. & R. Pethiyagoda (2005). The Sri Supplement No. 12: 339–350. Lankan Shrub-frogs of the genus philautus Gistel, 1848 (Ranidae: Biju, S.D. & F. Bossuyt (2009). Systematics and phylogeny of Philautus Rhacophorinae) with description of 27 new species. The Raffles Gistel, 1848 (Anura, Rhacophoridae) in the Western Ghats of India, Bulletin of Zoology 12: 163–303. with descriptions of 12 new species. Zoological Journal of Linnaean Meegaskumbura, M. & K. Manamendra-Arachchi (2005). Description Society 155: 374–444. of eight new species of Shrub-frogs (Ranidae: Rhacophorinae: Bossuyt, F. & A. Dubois (2001). A review of the frog genus Philautus Philautus) from Sri Lanka. The Raffles Bulletin of Zoology 12: 305– Gistel, 1848 (Amphibia, Anura, Ranidae, Rhacophorinae). Zeylanica 338. 6(1): 1–112. Meegaskumbura, M. & K. Manamendra-Arachchi (2011). Two new Bossuyt F., M. Meegaskumbura, N, Beenaerts, D.J. Gower, R. species of shrub frogs (Rhacophoridae: Pseudophilautus) from Sri Pethiyagoda, K. Roelants, A. Mannaert, M. Wilkinson, M.M. Bahir, Lanka. Zootaxa 2747: 1–18. K. Manamendra-Arachchi, P.K.L. Ng, C.J. Schneider, O.V. Oommen Meegaskumbura, M., F. Bossuyt, R. Pethiyagoda, K. Manamendra- & M.C. Milinkovitch (2004). Local Endemism Within the Western Ararchchi, M. Bahir, M.C. Milinkovitch & C.J. Schneider (2002). Sri Ghats-Sri Lanka Biodiversity Hotspot. Science 306: 479–481. Lanka: an amphibian hotspot. Science 298: 379. Dutta, S.K. & K. Manamendra-Archchi (1996). The Amphibian Fauna Meegaskumbura, M., K. Manamendra-Arachchi & R. Pethiyagoda of Sri Lanka. Wildlife Heritage Trust of Sri Lanka, Colombo, 230pp. (2009). Two new species of shrub frogs (Rhacophoridae: Philautus) DWC (2007). Biodiversity Baseline Survey: Peak Wilderness Sanctuary. from the lowlands of Sri Lanka. Zootaxa 2122: 51–68. Consultancy Services Report prepared by Green, M.J.B. (ed.), De Meegaskumbura, M., K. Manamendra-Arachchi, C.J. Schneider & R. Alwis, S.M.D.A.U., Dayawansa, P.N., How, R., Singhakumara, B.M.P., Pethiyagoda (2007). New species amongst Sri Lanka’s extinct shrub Weerakoon, D. and Wijesinghe, M.R. ARD Inc in association with frogs. Zootaxa 1397: 1–15. Infotech IDEAS and GREENTECH Consultants. Sri Lanka Protected Wickramasinghe, L.J.M., R. Rodrigo, N. Dayawansa & U.L.D. Jayantha Areas Management and Wildlife Conservation Project (PAM&WCP/ (2007). Two new species of Lankascincus (Squmamata: Scincidae) CONSULT/02/BDBS), Department of Wildlife Conservation, Ministry from Sripada Sanctuary (Peak Wilderness), in Sri Lanka. Zootaxa of Environment and Natural Resources, Colombo. 1–44pp. 1612: 1–24. Frost, D.R., T. Grant, J.N. Faivovich, R.H. Bain, A. Haas, C.L.F.B. Wickramasinghe, L.J.M. & D.I.A. Munindradasa (2007). Review of the Haddad, R.O. de Sa, A. Channing, M. Wilkinson, S.C. Donnellan, Genus Cnemaspis Strauch, 1887 (Sauria: Gekkonidae) in Sri Lanka C.J. Raxworthy, J.A. Campbell, B.L. Blotto, P. Moler, R.C. Drewes, with the description of five new species. Zootaxa 1490: 1–63. R.A. Nussbaum, J.D. Lynch, D.M. Green & W.C. Wheeler (2006). Wickramasinghe, L.J.M., D.R. Vidanapathirana, S. Airyarathne, The Amphibian Tree of Life. Bulletin of the American Museum of G. Rajeev, A. Chanaka, J. Pastorini, G. Chathuranga & N. Natural History 297: 1–370. Wickramasinghe (2013). Lost and found: One of the world’s most Grosjean, S., M. Delorme, A. Dubois & A. Ohler (2008). Evolution of elusive amphibians, Pseudophilautus stellatus (Kelaart, 1853) reproduction in the Rhacophoridae (Amphibia, Anura). Journal of rediscovered. Zootaxa 3260(1): 112–128. Zoological Systematics and Evolutionary Research 46(2): 169–176. Yu, G., M. Zhang & J. Yang (2010). Generic allocation of Indian and Sri Hammer, Ø., D.A.T. Harper & P.D. Ryan (2001). PAST: Paleontological Lankan Philautus (Anura: Rhacophoridae) inferred from 12S and 16S Statistics Software Package for Education and Data Analysis. rRNA genes. Biochemical Systematics and Ecology 38: 402–409. Palaeontologia Electronica 4(1): 1-9. Yu, G., D. Rao, M. Zhang & J. Yang (2009). Re-examination of the Li, J., J. Che, R.H. Bain, E. Zhao & Y. Zhang (2008). Molecular phylogeny phylogeny of Rhacophoridae (Anura) based on mitochondrial and of Rhacophoridae (Anura): a framework of taxonomic reassignment nuclear DNA. Molecular Phylogenetics and Evolution 50: 571–579. of species within the genera Aquixalus, Chiromantis, Rhacophorus and Philautus. Molecular Phylogenetics and Evolution 48: 302–312. 3898 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Oval, vertical Oval, Rather distinct, rounded distinct, Rather Half–circle or oval, horizontal or oval, Half–circle Distinct, oval, vertical oval, Distinct, Not discernible or Feebly Not discernible or Feebly and subdermal defined Distinct, oval, vertically vertically oval, Distinct, orientated Oval, vertical Oval, Oval, oblique or vertical Oval, Not discernible Rounded or oval, oblique or oval, Rounded Oval, vertical Oval, Absent (subdermal) Absent Indistinct, oval, vertical oval, Indistinct, Not discernible (subdermal) Oval, vertical Oval, eliptical Distinct, Not discernible, its outer rim Not discernible, its outer Indistinct Indistinct Oblique Oval Distinct, oval, vertical oval, Distinct, Oval Oval, vertical Oval, Oval, rounded Oval, Oval, vertical Oval, Oval Distinct, oval, vertically vertically oval, Distinct, Elongate Tympanum Distinct, oval, oblique oval, Distinct, Flat Concave or flat Concave Concave Concave Flat Flat Concave Concave Flat Concave Concave Concave Flat Concave Concave Flat Flat Concave Flat Concave Concave or flat Concave Concave Flat Concave Concave Flat Concave or flat Concave Internarial space Internarial Concave Flat Flat Flat Flat Convex Flat Convex Flat Concave or flat Concave Concave or flat Concave Flat Flat Convex Concave Flat Convex Flat Concave Flat Convex Concave Convex Convex Concave Concave Convex Flat Interorbital space Interorbital Convex Pseudophilautus. Concave Concave Concave Flat Concave Flat Concave Concave Concave Concave or flat Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Convex Concave or flat Concave Concave Concave Concave Concave Loreal region Loreal Rounded Sharp Rounded Rounded Sharp Rounded Sharp Rounded Sharp Sharp Rounded Rounded Rounded Rounded Sharp Rounded Rounded Sharp Rounded Sharp Rounded Rounded or sharp Rounded Sharp Sharp Rounded Sharp Rounded Sharp Canthus rostralis Canthus Pointed Rounded Rounded Blunt Rounded Obtusely pointed Obtusely Rounded Obtusely pointed Obtusely Rounded or Truncate Rounded Truncate Rounded Pointed or oval Pointed Truncate Rounded Blunt Rounded Truncate Rounded Rounded Truncate Rounded Obtusely pointed Obtusely Rounded Sharply Pointed Truncate Oval Truncate Obtusely pointed Obtusely Snout lateral Uneven Convex or Flat Convex Flat Convex Convex Flat Convex Flat Flat Concave or flat Concave Concave Flat Convex Flat Concave Convex Flat Concave Convex Convex Flat Convex Convex Convex Concave Convex Flat Head dorsally 18.3–21.9 27.9 32.9–42.9 21.7–26.4 33.4–46.8 18.9–27.3 26.7–29.3 17.0–27.5 23.7–29.4 33.3–41.7 26.9–44.5 25.4–37.0 23.4–28.8 22.3–32.5 43.5 33.6–51.1 20.4–24.6 35.3 38.4 44.8 16.2–19.2 18.3–23.9 35.8-40.4 17.6–28.9 24.5–30.1 38.4–48.9 17.3–20.2 16.3–19.0 SVL sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. P. hankeni P. P. halyi P. P. hallidayi P. P. auratus P. P. fulvus P. P. asankai P. P. frankenbergi P. P. alto P. P. folicola P. P. adspersus P. P. fergusonianus P. P. abundus P. P. femoralis P. P. samarakoon samarakoon P. sirilwijesundarai P. P. extirpo P. P. puranappu P. P. eximius P. P. newtonjayawardanei newtonjayawardanei P. P. dimbullae P. P. karunarathnai karunarathnai P. P. decoris P. P. jagathgunawardanai P. P. cuspis P. P. dayawansai P. P. cavirostris P. P. bambaradeniyai P. Species name caeruleus P. Appendix 1. A complete list of morphological characters considered to differentiate new species among congeners of the genus of the congeners new species among differentiate to considered characters of morphological list Appendix 1. A complete Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3899 Eight new species of Pseudophilautus Wickramasinghe et al. Oval Oval, oblique Oval, Oval, oblique Oval, Absent Faintly Discernible Faintly Vertically orientated Vertically Oval (vertical or oblique) (vertical Oval Discernible Oval, oblique Oval, Oval, vertical Oval, Oval, vertical Oval, Oval, oblique Oval, Oval, oblique Oval, Oval, horizontal or vertical horizontal Oval, Lower tympanic rim distinct Lower Oval, oblique Oval, Half circle, horizontal Half circle, Oval, vertical Oval, Oval, vertical Oval, Not discernible Oval, oblique Oval, Rounded Oval, oblique Oval, Distinct, oval or rounded, or oval Distinct, oblique Oval, vertical Oval, Oval or half circle, horizontal or half circle, Oval Distinct Oval, vertical Oval, Oval, vertical Oval, Tympanum Oval, vertical Oval, Oval, vertical Oval, Concave Concave Concave Flat Concave Concave Concave Convex or Convex concave Concave Concave Flat Concave Flat Flat Flat Concave Concave Concave Concave Concave Concave Flat Flat Concave Concave Flat Concave or flat Concave Flat Flat Flat Internarial space Internarial Concave Flat Flat Flat Flat Flat Convex Concave Convex or concave Convex Flat Concave Flat Flat Concave Flat Flat Flat Concave Convex Concave Flat Concave Concave Convex Concave Flat Concave Flat Flat Convex Flat Interorbital space Interorbital Convex Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Concave Flat Concave Concave Concave Flat Flat Flat Concave Flat Flat Concave Loreal region Loreal Sharp Sharp Rounded Rounded Rounded Rounded Sharp Rounded Sharp Rounded Rounded Sharp Rounded Sharp Rounded Rounded Rounded Rounded Rounded Rounded Indistinct Sharp Sharp Rounded Sharp Rounded Rounded Rounded Rounded Sharp Rounded Canthus rostralis Canthus Obtusely pointed Obtusely Pointed Rounded Oval Oval Rounded Rounded Pointed Truncate or oval Truncate Oval Oval Pointed Pointed or oval Pointed Truncate Rounded Oval Rounded Oval Truncate Obtusely pointed Obtusely Rounded Oval Truncate Acuminate Pointed Blunt Oval Blunt Rounded Pointed Obtusely pointed Obtusely Snout lateral Convex Flat Convex Flat Flat Convex Convex Convex or flat Convex Convex Flat Convex Concave Concave Convex Convex or Convex concave Convex Flat Convex Concave Convex Flat Convex Convex Flat Convex Convex Flat Flat Convex Head dorsally 17.7–30.0 17.4 22.6–38.1 34.8 20.6–24.1 29.4–35.0 17.5 16.3–18.4 42.5–61.1 23.9–49.0 18.3–21.7 14.9 25.1–26.8 46.6 17.7–24.7 27.4–42.7 21.3–26.0 40.9 25.6–30.3 25.7 32.1 19.8 40 11.2–22.4 18.6 22.0–28.6 13.4 23.1–33.4 21.2–23.4 19.9–22.8 20.7 SVL P. schmarda P. P. nasutus P. P. sarasinorum P. P. nanus P. P. rus P. P. mooreorum P. P. rugatus P. P. mittermeieri P. P. reticulatus P. P. microtympanum P. P. regius P. P. malcolmsmithi P. P. procax P. P. maia P. P. popularis P. P. macropus P. P. poppiae P. P. lunatus P. P. pleurotaenia P. P. limbus P. P. pardus P. P. leucorhinus P. P. papillosus P. P. hypomelas P. P. oxyrhynchus P. P. hoipolloi P. P. semiruber P. P. ocularis P. P. hoffmanni P. P. schneideri P. P. nemus P. Species name 3900 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Oval or vertical (rarely (rarely or vertical Oval Indistinct) Feebly defined Feebly Oval, vertical Oval, Present Distinct, oval, vertical oval, Distinct, Present Vertically orientated, orientated, Vertically crescentic Present Present Oval, vertical Oval, Present Indistinct Present Oval, oblique Oval, Absent Oval, oblique Oval, Present Oval, oblique, its outer rim oblique, its outer Oval, distinct Present Oval,vertical Fingers Rudimentary webbing Rudimentary Fingers Rounded Oval, vertical Oval, Oval, oblique Oval, III and IV Fingers Oval, vertical Oval, Tympanum Present Oval Flat Present Concave Present Concave Present Flat Present Flat Absent Present Concave or flat Concave Present Concave Present Concave Absent Flat Present Concave Absent Flat Fringe on fingers Fringe Flat Concave Flat Present Flat Internarial space Internarial Present Concave Flat Absent Flat Absent Concave Flat Absent Flat Absent Absent Concave or flat Concave Absent Concave Absent Flat Absent Convex Absent Convex Absent Convex Co–ossified skin Flat Flat Flat Absent Flat Interorbital space Interorbital Absent Convex Concave Concave Absent Concave Absent Concave Flat Absent Concave Absent Absent Concave Absent Concave Absent Concave Absent Concave Absent Concave Absent Flat Cephalic ridges Concave Concave Concave Absent Concave Loreal region Loreal Absent Rounded or sharp Rounded Indistinct Distinct Rounded Prominent Rounded Distinct Rounded Prominent Sharp Prominent Prominent Rounded Prominent Rounded Distinct Sharp Prominent Rounded Prominent Rounded Distinct Rounded Supratympanic fold Supratympanic Sharp Sharp Sharp Distinct Sharp Canthus rostralis Canthus defined Feebly Rounded Blunt or rounded Blunt Absent Truncate Absent Rounded Present Obtusely pointed Obtusely Absent Rounded Present Absent Obtusely pointed Obtusely Absent Rounded Absent Rounded Present Blunt Present Rounded Absent Truncate Lingual papilla Oval Pointed Oval Absent Blunt Snout lateral Absent Convex Convex Absent Flat Absent Convex Present Convex Present Flat Present Absent Flat or convex Flat Absent Concave Present Flat Absent Convex Absent Convex Absent Convex Vomerine teeth Vomerine Convex Concave Convex above Convex Present Convex Head dorsally Absent 20.0–32.5 27.4–36.3 Absent 35.7 Absent 28.4–31.0 Absent 13.5–13.9 Absent 24.2–32.4 Absent Absent 23.2–36.0 Absent 39.6–55.3 Absent 30.4–41.6 Absent 22.2–39.4 Absent 16.1–16.6 Absent 12.6–15.6 Pineal ocellus 24.3–31.3 22.9–30.1 35.4–51.8 Absent 31.6 SVL Absent sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. sp. nov. P. zal P. P. asankai P. P. viridis P. P. alto P. P. variabilis P. P. adspersus P. P. temporalis P. P. abundus P. P. tanu P. P. samarakoon samarakoon P. sirilwijesundarai P. P. stuarti P. P. puranappu P. P. stictomerus P. P. newtonjayawardanei newtonjayawardanei P. P. stellatus P. P. karunarathnai karunarathnai P. P. steineri P. P. jagathgunawardanai P. P. sordidus P. P. dayawansai P. P. singu P. P. bambaradeniyai P. P. simba P. Species name P. silvaticus P. P. zorro P. P. silus P. auratus P. P. zimmeri P. Species name Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3901 Eight new species of Pseudophilautus Wickramasinghe et al. Present Present Present Present Present Fingers 1 and 2 Fingers Present Present Present Present Fingers Rudimentary webbing Rudimentary Fingers Absent Present Absent Present Absent Present Absent Present Present Present Present Present Present Present Present Absent Present Present Present Present Present Absent Present Absent Absent Absent Present Present Absent Present Fringe on fingers Fringe Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Absent Absent Absent Absent Absent Absent Absent Absent Co–ossified skin Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Two fronto– Two ridges parietal Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Two fronto– Two ridges parietal Absent Absent Absent Absent Absent Absent Absent Absent Cephalic ridges Distinct Prominent Distinct Distinct Prominent Distinct Prominent Distinct Indistinct or absent Indistinct Distinct Distinct Prominent Distinct Absent Distinct Prominent Prominent Distinct Distinct Distinct Prominent Prominent Prominent Feebly defined Feebly Distinct Prominent Prominent Not prominent Not prominent Indistinct Supratympanic fold Supratympanic Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent or present Absent Absent Absent Absent Absent Absent Absent Present Present Present Present Absent Present Present Absent Absent Absent Absent Absent or present Absent Present Absent Lingual papilla Absent Absent Absent Present Absent Present Present Absent Absent Present Absent Present Present Absent Present Absent Absent Present Absent Present Present Present Absent Absent Absent Absent Present Absent Absent Absent Vomerine teeth Vomerine Absent Absent Present Absent Absent Absent Absent Absent Absent Absent or Absent present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Pineal ocellus P. hankeni P. P. halyi P. P. nemus P. P. hallidayi P. P. nasutus P. P. fulvus P. P. nanus P. P. frankenbergi P. P. mooreorum P. P. folicola P. P. mittermeieri P. P. fergusonianus P. P. malcolmsmithi P. P. microtympanum P. P. femoralis P. P. maia P. P. extirpo P. P. macropus P. P. eximius P. P. lunatus P. P. dimbullae P. P. limbus P. P. decoris P. P. leucorhinus P. P. cuspis P. P. hypomelas P. P. cavirostris P. P. hoipolloi P. P. caeruleus P. P. hoffmanni P. Species name 3902 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Absent Present Present Present Absent Present Present Absent Present Absent or present Absent Fingers Rudimentary webbing Rudimentary Fingers Present Absent Present Present Present Absent Present Absent Present Absent Absent Present Absent Present Absent Present Absent Absent Present Present Present Present Absent Absent Absent Present Absent Absent Absent Present Present Fringe on fingers Fringe Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Co–ossified skin Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Cephalic ridges Absent Feebly defined Feebly Distinct Prominent Prominent Prominent Prominent Distinct Prominent Distinct Prominent Distinct Distinct or prominent Distinct Absent Distinct Prominent Distinct Distinct Prominent Absent Present Distinct Not prominent Distinct Indistinct Distinct Distinct Distinct Distinct Distinct Distinct or indistinct Distinct Distinct Supratympanic fold Supratympanic Absent Absent Absent Absent Absent Present Absent Absent Absent Absent Absent Present Absent Absent Absent Present Absent Absent Absent Absent Absent Present Absent Present Present Absent Absent Present Absent Absent Lingual papilla Absent Present Present Absent Absent Absent Absent Present Absent Absent Present Present Absent Absent Absent Present Present Absent Present Absent Present Absent Present Absent Absent Present Absent Absent Present Present Present Vomerine teeth Vomerine Absent Absent Present Absent Absent Absent Absent Absent Absent Absent Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent or Absent present Absent Present Absent Absent Absent Absent Absent Absent Absent Absent Pineal ocellus P. semiruber P. P. schneideri P. P. zimmeri P. P. schmarda P. P. zal P. P. sarasinorum P. P. viridis P. P. rus P. P. variabilis P. P. rugatus P. P. temporalis P. P. reticulatus P. P. tanu P. P. regius P. P. stuarti P. P. procax P. P. stictomerus P. P. popularis P. P. stellatus P. P. poppiae P. P. steineri P. P. pleurotaenia P. P. sordidus P. P. pardus P. P. singu P. P. papillosus P. P. simba P. P. oxyrhynchus P. P. silvaticus P. P. ocularis P. P. silus P. Species name Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3903 Eight new species of Pseudophilautus Wickramasinghe et al. Fingers Rudimentary webbing Rudimentary Fingers Absent Present Absent Absent Present Absent Absent Absent Present Absent Absent Absent Absent Present Present Absent Absent Calcar Absent Absent Present Absent Absent Absent Absent Fringe on fingers Fringe Present Present Present Present Present Absent Present Present Present Tarsal tubercle Tarsal Absent Co–ossified skin Absent Present, tuberculated Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Tarsal fold Tarsal Absent Present, Present, undulating Absent Absent Absent Absent Cephalic ridges Absent Present Present Present Present Present Present Present Present Fringe on toes Fringe Prominent On outer edge of edge On outer toe fifth Supratympanic fold Supratympanic Medially webbed 1, 2, 3 and 5 fully webbed Medially webbed Medially webbed Medially webbed Fully webbed Medially webbed Present Present Present Present Present Present Medially webbed Present Webbed Present Medially webbed Webbing on toes Webbing Medially webbed Absent Medially webbed Medially webbed Medially webbed Medially webbed Medially webbed Lingual papilla Medially webbed Oval Oval Oval Oval Oval Oval Oval Oval Rounded Rounded Rounded Rounded Rounded Rounded Oval Rounded Oval Oval Oval Tips of toes Tips Triangular Absent Oval Oval Oval Oval Oval Vomerine teeth Vomerine Present Absent Absent Absent Present Absent Absent Absent Present Absent Present Absent Present Absent Absent but Absent subdermal glands nuptial Present Absent Absent Nuptial pad Nuptial Present Absent Absent Pineal ocellus P. cavirostris P. cuspis P. P. caeruleus P. P. auratus P. P. asankai P. P. adspersus P. alto P. P. abundus P. P. sirilwijesundarai P. P. samarakoon P. P. puranappu P. P. newtonjayawardanei P. P. karunarathnai P. P. jagathgunawardanai P. P. fulvus P. P. dayawansai P. P. frankenbergi P. P. bambaradeniyai P. P. folicola P. Appendix 1. continued Species name P. dimbullae P. eximius P. extirpo P. femoralis P. fergusonianus P. P. zorro P. P. decoris P. Species name 3904 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Fingers Rudimentary webbing Rudimentary Fingers Present Absent Present Present Absent Absent Present. Absent Absent Present Present Present Present Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Fringe on fingers Fringe Absent Present Absent Absent Present Absent Present Co–ossified skin Absent Absent Absent Absent Absent Absent Absent Present. Absent Absent Absent Absent Absent. Absent Absent Absent Absent Absent Absent Present Absent Absent Absent Absent Absent Absent Cephalic ridges Absent Absent Supratympanic fold Supratympanic Medially webbed 2, 3 Fully webbed Medially webbed Medially webbed Medially webbed Medially webbed Webbed Medially webbed Webbed 1, 2, 3 and 5 fully webbed Webbed Medially webbed Medially webbed Webbed 2, 3 and 5 Fully webbed Present Medially webbed toe 4 webbing 4 webbing toe penultimate to subarticular tubercle side on outer Medially webbed Medially webbed Present Webbed Medially webbed Medially webbed Webbed Webbed 2, 3 and 4 Fully webbed Lingual papilla Webbed Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Vomerine teeth Vomerine Oval Absent Absent Present Present Absent Present Absent Present Absent Present Absent Present Absent Present Absent Absent Absent Absent Present Present Absent Present Absent Pineal ocellus P. pardus P. P. ocularis P. papillosus P. P. oxyrhynchus P. P. nemus P. P. nasutus P. P. nanus P. P. mooreorum P. P. maia P. malcolmsmithi P. P. microtympanum P. P. mittermeieri P. P. lunatus P. macropus P. P. leucorhinus P. P. limbus P. P. reticulatus P. P. hypomelas P. P. regius P. P. hoipolloi P. P. procax P. P. hankeni P. P. hoffmanni P. P. popularis P. P. halyi P. P. poppiae P. P. hallidayi P. P. pleurotaenia P. Species name Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3905 Eight new species of Pseudophilautus Wickramasinghe et al. Fingers Rudimentary webbing Rudimentary Fingers Absent Absent Absent Absent Absent Present Absent Absent Absent Absent Absent Present Absent Present Fringe on fingers Fringe Absent Absent Present Absent Absent Absent Absent Absent Absent Present Present Co–ossified skin Absent Absent Absent Absent Absent Absent Absent Absent Absent Present Absent Absent Absent Absent Absent Absent Absent presernt Absent Absent Absent Absent Cephalic ridges Absent Present Absent Present Poorly developed developed Poorly 5 on toe fringe Supratympanic fold Supratympanic Present Present Medially webbed Medially webbed Medially webbed Webbed Medially webbed Webbed Present Present Medially webbed Rudimentary webbing Rudimentary Medially webbed Medially webbed Present Present Medially webbed Medially or fully 1, 2, 3 toes webbed; and 5 fully webbed Medially webbed Medially webbed Medially webbed Lingual papilla Medially webbed Rounded or oval Rounded Oval Oval Oval Oval Oval Oval Oval Rounded Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Oval Vomerine teeth Vomerine Oval Present Absent Absent Absent Absent Present Present Absent Absent Absent Absent Present Absent Absent Absent Absent Present Present Pineal ocellus Absent P. variabilis P. P. zal P. P. tanu P. P. temporalis P. P. viridis P. P. stuarti P. P. stellatus P. P. stictomerus P. P. steineri P. P. singu P. P. sordidus P. P. simba P. P. silvaticus P. P. silus P. P. schneideri P. P. semiruber P. P. schmarda P. P. rugatus P. P. sarasinorum P. P. zimmeri P. P. zorro P. Species name P. rus P. 3906 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Median dermal ridge on Median dermal ridge mid–dorsum Absent Absent tip of from dermal ridge Faint vent snout to back of head Tip of snout to vent Tip of snout to till back of head) (Prominent back of head Tip of snout to with tubercles vent Tip of snout to till back of head) (Prominent back of head Tip of snout to Tip of snout to vent (narrow) vent Tip of snout to tuberculated Heavily Posterior dorsum Posterior spinules Horny tubercular Weakly Smooth Spinulate Shagreen spinules, horny Few tubercles Prominent Tubercular Shagreen in spinules (Absent Horny females) Smooth spinules in males, Horny in females warty spinules fine, horny Very Shagreen and horny Glandular warts lack spinules (females spinules) horny tuberculated, Heavily glandular warts spinules Horny Glandular warts Shagreen Smooth Anterior dorsum Anterior and horny tubercles Prominent spinules Smooth Smooth Smooth Shagreen shagreen Weakly Tubercular tubercles with few Shagreen in spinules (absent Horny females) Smooth except for a few a few for Smooth except warts glandular scattered spinules in males, warty Horny in females spinules fine, horny Very Shagreen and horny Glandular warts lack horny spinules (females spinules) glandular tuberculated, Heavily warts spinules with a “)(”shaped Horny of tubercles pattern Glandular warts Shagreen Smooth species. Side of head Smooth Tubercular Smooth tubercular Weakly Shagreen Smooth Smooth shagreen Weakly Smooth Smooth except for a few a few for Smooth except warts glandular scattered Smooth spinules fine, horny Very Shagreen and horny Glandular warts spinules Glandular warts Smooth Smooth Glandular warts Shagreen Pseudophilautus Interorbital area Interorbital Smooth across Smooth, with a ridge medially braking cross Smooth,with two dermal fringes tubercular Weakly with a prominent Shagreen, ridge Smooth Tubercular tubercle A prominent present Smooth Smooth except for a few a few for Smooth except warts glandular scattered spinules in males Horny spinules fine, horny Very Shagreen and horny Glandular warts spinules Glandular warts spinules Horny Smooth Glandular warts Shagreen Snout Smooth with a single Tuberculated, medially tubercle prominent placed Smooth blunt Smooth, a prominent present tubercle Shagreen Smooth, with a blunt tubercle present tubercle Prominent of snout on centre with prominent Shagreen, dermal fold Smooth Smooth except for a few a few for Smooth except warts glandular scattered spinules in males Horny spinules fine, horny Very Shagreen and horny Glandular warts spinules Glandular warts spinules Horny Smooth Glandular warts Shagreen sp. nov. sp. nov. sp. nov. sp. nov. P. bambaradeniyai P. sp. nov. dayawansai P. jagathgunawardanai P. sp. nov. karunarathnai P. newtonjayawardanei P. sp. nov. puranappu P. samarakoon P. sirilwijesundarai P. sp. nov. abundus P. Species name adspersus P. alto P. asankai P. auratus P. caeruleus P. cavirostris P. cuspis P. eximius P. P. decoris P. dimbullae P. differentiate that in the skin structure of characters list Appendix 2. A complete Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3907 Eight new species of Pseudophilautus Wickramasinghe et al. Median dermal ridge on Median dermal ridge mid–dorsum Tip of snout to vent Tip of snout to Thin median dermal ridge Tip of from on mid dorsum back of head snout to Side of head shagreen, spinules on males with horny dorsum Posterior dorsum Posterior with a few Shagreen, glandular warts Finely granular males glandular warts, Few and with glandular warts spinules horny Smooth spinules horny Shagreen, scattered Shagreen with horny Glandular warts, spinules in males with glandular Shagreen warts bearing Glandular warts spinules horny Smooth Smooth or shagreen Smooth Smooth with a few warts glandular scattered granular Weakly Smooth or smooth, with Shagreen glandular scattered a few warts Glandular warts Smooth shagreen Smooth above, and with glandular warts, males with horn like spinules Shagreen, with a few glandular with a few Shagreen, warts Anterior dorsum Anterior Glandular warts bearing horny bearing horny Glandular warts spinules scattered Smooth with a few glandular warts Finely granular Males glandular warts Shagreen, spinules and horny Smooth or granular spinules horny Shagreen, scattered Shagreen with horny Glandular warts, spinules in males with glandular warts Shagreen Smooth Smooth or shagreen Smooth granular Weakly Smooth or smooth, with a few Shagreen warts glandular scattered Glandular warts Smooth, '\ /' shaped ridge and shagreen Smooth above, males with with glandular warts, spinules horn like Shagreen Side of head Glandular warts bearing horny bearing horny Glandular warts spinules Smooth Finely granular glandular warts Few smooth or with a Shagreen, warts glandular scattered few Glandular warts Shagreen Glandular warts with glandular warts Shagreen Smooth Smooth or shagreen Smooth granular Weakly Smooth Glandular warts Smooth Smooth Shagreen Shagreen Interorbital area Interorbital Glandular warts bearing Glandular warts spinules horny Smooth Finely granular males glandular Shagreen, spinules and horny warts Smooth Shagreen Shagreen with horny Glandular warts, spinules in males with glandular Shagreen warts Smooth Smooth or shagreen Smooth granular Weakly Smooth Smooth with a few warts glandular scattered Smooth Smooth Smooth or tuberculated Shagreen, with a few with a few Shagreen, glandular warts Snout Glandular warts bearing Glandular warts spinules horny Smooth Finely granular males glandular Shagreen, spinules and horny warts Smooth Shagreen Shagreen with horny Glandular warts, spinules in males with glandular Shagreen warts Smooth Smooth or shagreen Smooth granular Weakly Smooth Smooth Smooth Smooth Smooth or tuberculated P. extirpo P. P. hankeni P. leucorhinus P. Species name femoralis P. fergusonianus P. folicola P. frankenbergi P. fulvus P. hallidayi P. halyi P. hoffmanni P. hoipolloi P. hypomelas P. limbus P. lunatus P. macropus P. maia P. malcolmsmithi P. microtympanum P. 3908 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Median dermal ridge on Median dermal ridge mid–dorsum Tip of snout to vent Tip of snout to or finely granular Dorsum in female shagreen Finely tuberculated Posterior dorsum Posterior Glandular warts Horny spinules in males, Horny or shagreen finely granular in female glandular warts Few horn like Tuberculated, spinules Glandular warts or with glandular Shagreen warts Smooth Glandular warts Smooth scattered few Shagreen, and horny glandular warts spinules (males) spinules in males, Horny or shagreen finely granular in female spinules Horny spinules (spinules Horny in females) Absent spinules Horny with a few Shagreen warts, glandular scattered finely tuberculated a few for Smooth except warts glandular scattered Glandular warts males with Glandular warts, spinules scattered horny glandular Glandular folds, spinules and horny warts Anterior dorsum Anterior Glandular warts Glandular warts spinules Horny a few for Smooth except warts glandular scattered Horny spinules in males, finely Horny in female or shagreen granular glandular warts Few spinules horn like tuberculated, or with glandular Shagreen warts Smooth Glandular warts Smooth scattered with a few Shagreen and horny glandular warts spinules ( males) spinules in males, finely Horny in female or shagreen granular spinules (spinules Absent Horny in females) spinules Horny scattered with a few Shagreen finely glandular warts, tuberculated Glandular warts males with Glandular warts, spinules scattered horny glandular warts Glandular folds, spinules and horny Side of head Glandular warts Glandular warts Glandular warts a few for Smooth except warts glandular scattered Horny spinules in males Horny glandular warts Few Glandular warts or with glandular Shagreen warts Smooth Glandular warts Smooth Shagreen spinules in males, finely Horny in female or shagreen granular Smooth spinules Horny scattered with a few Shagreen glandular warts Glandular warts Glandular warts Glandular warts Interorbital area Interorbital Glandular warts Smooth Smooth a few for Smooth except warts glandular scattered Horny spinules in males Horny glandular warts Few Glandular warts or with glandular Shagreen warts Smooth Glandular warts Smooth Shagreen spinules in males, Horny or shagreen finely granular in female Smooth spinules Horny with a few Shagreen warts glandular scattered Smooth Glandular warts glandular Glandular folds, spinules and horny warts Snout Glandular warts Glandular warts Glandular warts a few for Smooth except warts glandular scattered Horny spinules in males Horny Glandular warts Glandular warts or with glandular Shagreen warts Smooth Glandular warts Smooth Shagreen spinules in males, Horny or shagreen finely granular in female Smooth spinules Horny with a few Shagreen warts glandular scattered Smooth Glandular warts Glandular warts P. mittermeieri mittermeieri P. P. nemus P. popularis P. rugatus P. Species name mooreorum P. nanus P. nasutus P. ocularis P. oxyrhynchus P. papillosus P. pardus P. pleurotaenia P. poppiae P. procax P. regius P. reticulatus P. rus P. sarasinorum P. schmarda P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3909 Eight new species of Pseudophilautus Wickramasinghe et al. Median dermal ridge on Median dermal ridge mid–dorsum Thin median dermal ridge Thin median dermal ridge tip of from on mid dorsum back of head, than snout to vent to posterior Tip of snout to (feebly–defined) dorsum Absent (Very vent Tip of snout to narrow) to tip of snout from Present, back of head ending with a tubercle prominent large (narrow) vent Tip of snout to Posterior dorsum Posterior with horny Small tubercules spinules Smooth Glandular warts Smooth Smooth glandular Scattered, tubercles males Glandular warts, spinules scattered horny and horny Glandular warts lack horny spinules, females spinules shagreen Weakly spinules horny Shagreen, in males spinules, females Horny spinules lack horny Shagreen Smooth Smooth spinules (smooth Horny and immature in females males) tubercular Weakly Glandular warts spinules Horny Anterior dorsum Anterior with horny Small tubercules spinules Smooth Glandular warts Glandular warts Glandular warts glandular tubercles Scattered, males horny Glandular warts, spinules scattered and horny Glandular warts lack horny spinules, females spinules shagreen Weakly spinules in horny Shagreen, males lack spinules, females Horny spinules horny Shagreen Smooth Smooth spinules (smooth in Horny males) and immature females tubercular Weakly Glandular warts dorsum spinules, Anterior Horny of with “)(”–shaped pattern tubercles Side of head with horny Small tubercules spinules Smooth Glandular warts Glandular warts Glandular warts glandular tubercles Scattered, Glandular warts Smooth or with glandular warts shagreen Weakly Smooth or with glandular warts Smooth Shagreen Smooth Smooth spinules (smooth in Horny males) and immature females Shagreen Glandular warts Glandular warts Interorbital area Interorbital with horny Small tubercules spinules Smooth Smooth Glandular warts Smooth glandular Scattered, tubercles Glandular warts and horny Glandular warts spinules shagreen Weakly Smooth spinules Horny Shagreen Smooth Smooth spinules (smooth Horny and immature in females males) Shagreen Glandular warts spinules Horny Snout with horny Small tubercules spinules Smooth Smooth Glandular warts Smooth glandular Scattered, tubercles Glandular warts and horny Glandular warts spinules shagreen Weakly Smooth spinules Horny Shagreen Smooth Smooth spinules (smooth Horny and immature in females males) shagreen Weakly Glandular warts spinules Horny P. schneideri P. Species name semiruber P. silus P. silvaticus P. simba P. singu P. sordidus P. steineri P. stellatus P. stictomerus P. stuarti P. tanu P. temporalis P. variabilis P. viridis P. zal P. zimmeri P. zorro P. 3910 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Glandular warts, posterior posterior Glandular warts, arm with an of lower edge fold dermal undulating Smooth Smooth glandular few Shagreen, warts Foot Smooth Smooth tubercular Weakly Smooth Shagreen tubercles Smooth, with few V on toe tubercles Few Smooth Smooth Granular Smooth spinules fine, horny Very or smooth Shagreen Smooth present fringe A tuberculated margin on posterior Smooth or with glandular warts Glandular warts Smooth Smooth glandular few Shagreen, warts Shank tubercles Smooth with few tubercular Weakly Tubercular Smooth Shagreen Tubercular Tubercular Smooth Smooth Smooth Smooth spinules fine, horny Very Smooth Smooth Smooth scattered Glandular folds, glandular warts Glandular warts Smooth Smooth glandular warts few Shagreen, Dorsal thigh Dorsal Tuberculated Smooth tubercular Weakly Smooth Shagreen shagreen Weakly Tuberculated Smooth Smooth Smooth Smooth spinules fine, horny Very Smooth Smooth Smooth Smooth or with glandular warts Posterior edge of lower arm of lower edge Posterior fold dermal with an undulating Shagreen with small glandular Shagreen warts Smooth glandular with a few Shagreen, warts Dorsal and lateral forelimbs and lateral Dorsal Smooth Shagreen Upper arm smooth, forearm tubercular weakly Smooth Shagreen Upper arm smooth, forearm tubercular weakly Upper arm smooth, forearm tubercular shagreen Upper arm weakly with few shagreen forearm tubercles Smooth Glandular warts Smooth spinules fine, horny Very or smooth Shagreen Smooth dermal fold A tuberculated of lower edge on outer Present arm Smooth or with glandular warts Glandular warts Granular granular Weakly Granular Lower flank Lower Smooth Granular granular Weakly Granular granular Weakly Granular Granular Granular Granular Granular Granular spinules fine, horny Very Granular in males Granular Granular Smooth or granular Glandular warts Shagreen Smooth Granular Upper flank Smooth Granular Smooth granular Weakly Shagreen tubercular Weakly Tubercular Areolate Smooth Granular spinules in males Horny spinules Horny or smooth Granular and horny Glandular warts spinules Smooth Smooth or with glandular warts P. dimbullae P. eximius P. extirpo P. Appendix 2. contiuned Species name P. bambaradeniyai P. dayawansai P. jagathgunawardanai P. karunarathnai P. newtonjayawardanei P. puranappu P. samarakoon P. sirilwijesundarai P. abundus P. adspersus P. alto P. asankai P. auratus P. caeruleus P. cavirostris P. cuspis P. decoris P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3911 Eight new species of Pseudophilautus Wickramasinghe et al. Foot Smooth warts, glandular with Smooth side prominent posterior glandular tubercles Smooth Smooth Smooth Glandular warts Glandular warts with horny Glandular warts spinules Smooth Smooth Smooth Smooth not rough Granular, scattered Smooth, few glandular warts glandular Smooth, scattered Smooth scattered Smooth, few glandular warts scattered few Shagreen, glandular warts Smooth Shagreen. warts Shank spinules Horny Smooth with glandular warts, Smooth Glandular warts Smooth Glandular warts Glandular warts with horny Glandular warts spinules Smooth Smooth Smooth scattered Smooth, few glandular warts not rough Granular, scattered Smooth few glandular warts scattered Smooth few glandular warts Smooth scattered Smooth, few glandular warts scattered few Shagreen, glandular warts Smooth Shagreen Smooth Smooth with glandular warts, Smooth Smooth Shagreen Glandular warts Glandular warts with horny Glandular warts spinules Smooth Smooth Smooth Smooth not rough Granular, Smooth scattered Smooth with a few glandular warts Dorsal thigh Dorsal Smooth, few scattered glandular scattered Smooth, few warts scattered few Shagreen, glandular warts Smooth Shagreen Smooth Smooth Smooth with glandular warts, arm side of lower posterior glandular tubercles prominent Smooth Glandular warts Shagreen Glandular warts Glandular warts with horny Glandular warts spinules Smooth Smooth Smooth scattered Smooth, with a few glandular warts not rough Granular, Smooth scattered with a few Shagreen glandular warts Dorsal and lateral forelimbs and lateral Dorsal Smooth, with a few scattered scattered Smooth, with a few glandular warts scattered with a few Shagreen, glandular warts Smooth Shagreen Smooth Granular Granular Granular Granular Granular Granular Granular Granular Glandular Smooth or granular Smooth with glandular Granular warts Granular Granular Granular Lower flank Lower Glandular warts Granular Glandular warts spinules in males Horny Glandular warts Finely granular males with Shagreen, and horny glandular warts spinules Granular Shagreen Shagreen with horny Glandular warts, spinules in males Glandular warts bearing Glandular warts spinules horny Smooth Smooth or granular Smooth Smooth with glandular warts Granular Glandular warts with a few Granular warts glandular scattered Upper flank Glandular warts Granular Glandular warts spinules in males Horny Glandular warts P. femoralis femoralis P. fergusonianus P. folicola P. frankenbergi P. fulvus P. hallidayi P. halyi P. hankeni P. hoffmanni P. hoipolloi P. hypomelas P. leucorhinus P. limbus P. lunatus P. macropus P. Species name P. malcolmsmithi malcolmsmithi P. microtympanum P. mittermeieri P. mooreorum P. P. maia P. 3912 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Foot Smooth Glandular warts Smooth Glandular warts Smooth Glandular warts Smooth Smooth spinules in males, Horny or shagreen finely granular in female Smooth Smooth Smooth Smooth scattered Smooth, few glandular warts Smooth Glandular warts Glandular warts Smooth Smooth Smooth Shank Smooth Glandular warts Glandular warts Glandular warts Smooth Glandular warts Smooth Smooth spinules in males, Horny or shagreen finely granular in female Glandular warts Smooth Glandular warts Smooth scattered Smooth, few glandular warts Smooth Smooth Glandular warts Smooth Smooth Smooth Dorsal thigh Dorsal Smooth Glandular warts Smooth Glandular warts Smooth Glandular warts Smooth Smooth spinules in males, finely Horny in female or shagreen granular Glandular warts Smooth Glandular warts Smooth glandular scattered Smooth, few warts Smooth Smooth Glandular warts Smooth Smooth Smooth Dorsal and lateral forelimbs and lateral Dorsal of edge outer Glandular warts, arm with a tuberculated lower dermal fold Smooth Glandular warts Glandular warts Glandular warts Smooth Glandular warts Smooth Smooth spinules in males, finely Horny in female or shagreen granular Glandular warts Smooth Glandular warts Shagreen scattered Smooth with a few glandular warts Smooth Smooth Smooth Smooth Smooth Lower flank Lower and horny Glandular warts spinules Granular Granular Granular Granular Smooth Granular Granular Granular Smooth Granular Granular Glandular Granular Granular Granular in females Granular Granular Smooth Granular Upper flank glandular Glandular folds, spinules and horny warts Glandular folds Smooth Smooth or glandular warts Shagreen Smooth Glandular warts Granular scattered few Shagreen, and horny glandular warts, spinules spinules in males, Horny or shagreen finely granular in female spinules Horny Smooth Glandular with a few Shagreen warts glandular scattered Glandular warts Granular Glandular folds with horny Small tubercules spinules Smooth Glandular warts P. nanus P. nasutus P. nemus P. ocularis P. oxyrhynchus P. papillosus P. pardus P. pleurotaenia P. poppiae P. popularis P. procax P. regius P. reticulatus P. rugatus P. rus P. sarasinorum P. schmarda P. schneideri P. semiruber P. silus P. Species name Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3913 Eight new species of Pseudophilautus Wickramasinghe et al. Smooth glandular tubercles Scattered Granular glandular Smooth, scattered warts shagreen Weakly Smooth Smooth Smooth Smooth Smooth Shagreen Smooth Glandular warts Smooth tubercles Supernumerary on foot Present Present Foot Glandular warts Present Present Absent Present Indistinct Present Present Absent Smooth glandular Scattered tubercles Glandular warts Smooth, scattered glandular warts shagreen Weakly Smooth Smooth Smooth Smooth Smooth Shagreen tubercular Weakly Glandular warts Glandular warts tubercles Supernumerary on palm Present Present Shank Glandular warts Present Absent Present Present Present Present Present Smooth glandular tubercles Scattered Glandular warts glandular Smooth, scattered warts shagreen Weakly Smooth Smooth Smooth Smooth Smooth Shagreen tubercular Weakly Glandular warts Glandular warts of thigh Underside Smooth granular Weakly Dorsal thigh Dorsal Smooth Granular Granular Granular granular Weakly granular Weakly Granular Granular Smooth glandular tubercles Scattered Glandular warts Smooth with scattered glandular warts shagreen Weakly Glandular warts Smooth Smooth scattered Smooth with a few glandular warts arm part of lower Dorsal smooth, upper arm granular Shagreen tubercular Weakly Glandular warts Smooth Belly Granular Granular Dorsal and lateral forelimbs and lateral Dorsal Glandular warts Granular Granular Granular Granular Granular granular Coarsely Granular Granular glandular Scattered, tubercles Granular Granular areolate Weakly Granular Smooth Granular granular Coarsely Granular Granular Granular Granular Granular Chest granular Weakly granular Weakly Lower flank Lower Granular Weakly granular Weakly Granular Granular granular Weakly granular Weakly granular Weakly granular Coarsely Scattered, glandular Scattered, tubercles with horny Glandular warts, spinules in males and horny Glandular warts spinules areolate weakly to Shagreen spinules horny Shagreen, in males Smooth Granular Smooth Granular spinules (smooth Horny and immature in females males) Granular Granular Glandular warts Throat granular Weakly Shagreen Upper flank Glandular warts Weakly granular Weakly Granular Granular granular Weakly granular Weakly Shagreen granular Coarsely P. singu P. sordidus P. steineri P. stellatus P. stictomerus P. stuarti P. tanu P. temporalis P. variabilis P. viridis P. zal P. zimmeri P. zorro P. Appendix 2. continued Species name bambaradeniyai P. dayawansai P. P. silvaticus P. simba P. Species name jagathgunawardanai P. P. karunarathnai P. newtonjayawardanei P. puranappu P. samarakoon P. sirilwijesundarai P. abundus P. 3914 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Supernumerary tubercles tubercles Supernumerary on foot Present Present Absent Present Present Absent Present Present Absent Absent Absent Present Absent Present Present Absent Absent Present Present Absent Absent Present Present Supernumerary tubercles tubercles Supernumerary on palm Present Present Absent Present Present Present Present Present Present Present Present Absent Present or present Absent Present Present Present Present Present Absent Present Absent Present Present Present Present Present Underside of thigh Underside granular Weakly Granular Granular Granular Granular Granular Granular Granular granular Coarsely granular Coarsely granular Coarsely granular Coarsely Granular Granular Granular Granular Granular Granular Smooth Smooth Granular granular Weakly granular Coarsely Granular Smooth granular. Smooth, distally Granular Granular Granular Belly Coarsely granular Coarsely Granular Granular Granular in males Granular Granular granular Coarsely granular Coarsely Granular Granular Granular Granular granular Coarsely granular Coarsely granular. Coarsely Granular Granular Granular Granular Granular Granular Granular Granular granular Coarsely Smooth Granular Granular Granular Granular Chest Coarsely granular Coarsely Granular Granular Smooth or granular in males, smooth Granular in females Smooth granular Weakly granular Weakly or smooth Granular Shagreen Granular Smooth or granular granular Coarsely granular Weakly granular Weakly Smooth, granular Granular Granular in males Smooth, granular Granular Granular Granular Smooth but not rough Granular, Smooth Granular Granular Smooth Granular Throat Weakly granular Weakly Granular Granular Smooth or granular with scattered Granular smooth in glandular warts, females Smooth granular Weakly Smooth or smooth Granular Shagreen Granular Smooth or roughly warty, Glandular, granular Smooth granular Weakly granular Weakly Granular Granular in males Smooth, granular Granular Granular Granular Smooth but not rough Granular, Smooth Smooth or granular Granular Smooth Granular P. adspersus P. alto P. asankai P. auratus P. caeruleus P. Species name cavirostris P. dimbullae P. femoralis P. folicola P. hoipolloi P. limbus P. P. cuspis P. decoris P. eximius P. extirpo P. fergusonianus P. frankenbergi P. fulvus P. hallidayi P. halyi P. hankeni P. hoffmanni P. hypomelas P. leucorhinus P. lunatus P. macropus P. maia P. malcolmsmithi P. microtympanum P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3915 Eight new species of Pseudophilautus Wickramasinghe et al. Supernumerary tubercles tubercles Supernumerary on foot Absent Present Present Absent Absent Present Absent Present Present Present Present Present Present Present Present Present Supernumerary tubercles tubercles Supernumerary on palm Absent Present Present Present Present Present Present Absent Present Present Present some mature Present, individuals with a smooth palm, some with a granulated Present Present Present Present Absent Present Present Present Present Present Present Present Present Underside of thigh Underside Granular Granular Smooth Granular Granular or glandular folds Granular Smooth Granular granular Coarsely Granular Smooth Granular Granular Granular Granular granular Coarsely Granular Smooth Granular Smooth Smooth Granular Granular granular Coarsely Smooth Granular Granular Granular Belly Granular Granular Granular Granular granular Coarsely Granular Granular granular Coarsely Granular Granular granular Coarsely Granular Granular Coarsely granular Coarsely Granular granular Coarsely or with glandular Granular folds Granular Granular Granular Granular Granular Granular Smooth Granular in males Granular Granular Granular Chest Granular Granular Smooth Granular granular Coarsely Granular Granular mid– for Smooth except which is granular area, Smooth Granular granular Weakly Granular Granular Weakly granular Weakly Granular granular Coarsely or with glandular Granular folds Granular granular Coarsely Granular Smooth or granular Granular Granular Smooth Granular in males, smooth Granular in females Smooth granular Weakly Throat Granular Granular Smooth Granular granular Weakly Granular Granular granular Weakly Glandular warts Glandular warts granular Weakly Granular Granular Weakly granular Weakly Granular but not rough Granular or with glandular Granular folds Granular granular Coarsely Granular Smooth or granular Granular Granular Smooth Granular in males, smooth Granular in females Smooth granular Weakly P. mittermeieri mittermeieri P. Species name mooreorum P. oxyrhynchus P. papillosus P. pardus P. pleurotaenia P. poppiae P. rugatus P. sarasinorum P. schmarda P. simba P. singu P. steineri P. P. nanus P. nasutus P. nemus P. ocularis P. popularis P. procax P. regius P. reticulatus P. rus P. schneideri P. semiruber P. silus P. silvaticus P. sordidus P. stellatus P. 3916 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. Supernumerary tubercles tubercles Supernumerary on foot Present and on foot on toes Present Present Present Present Present Supernumerary tubercles tubercles Supernumerary on palm Present Present I–III and on palm Fingers Present Present Present Present Present Present Underside of thigh Underside Granular Granular Granular granular Weakly Granular Granular Granular Granular Smooth Belly Granular Granular Granular Granular Granular granular Coarsely Granular Granular Granular Chest Smooth or shagreen Granular Granular smooth Mid chest Granular Smooth Granular Granular Granular Throat Smooth or shagreen Granular Granular Smooth Granular Smooth Granular Granular Granular P. stictomerus P. Species name stuarti P. zal P. zorro P. P. tanu P. temporalis P. variabilis P. viridis P. zimmeri P. Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3917 Eight new species of Pseudophilautus Wickramasinghe et al. Appendix 3. Analysis of morphometric data Appendix 4. Material examined Morphometric data of both males and females was considered Pseudophilautus abundus (Manamendra-Arachchi & Pethiyagoda, for statistical analysis to understand the morphometric difference 2005). Holotype: WHT 3006; Labugama Forest Reserve, Labugama, among the individuals of 8 new species of Pseudophilautus. To account elevation 78m (06051’N & 80010’E); Paratypes: WHT 2302, 2303, for the size difference among individuals, all the morphometric 3457, 3459, 3494, 3496; Dediyagala Forest Reserve, Akuressa, characters were taken as %SVL. Principle Component Analysis (PCA) 150m (06010’N & 80026’E); WHT 1711 3455, 3456, 3495; Kanneliya was performed on the correlation matrix between 25 variables using (Galle), 150m (06015’N & 80020’E). the software PAST (Hammer et al. 2001). Dendrogram showing the Pseudophilautus adspersus (Günther, 1872). Holotype: BMNH relationship between different species was plotted using mean values 1947.2.6.23, Ceylon (Sri Lanka). of morphometric data for each species using Euclidian distances and Pseudophilautus alto (Manamendra-Arachchi & Pethiyagoda, 2005). Ward’s method for clustering in PAST. Holotype: WHT 2721; Paratypes: WHT 2723, 2718, 2719, 2720; PCA extracted 10 factors with eigenvalue more than unity. First 2722, 2724; Horton Plains National Park, 2135m (06046’N & three PCA axes explained total 61.29% of the total variation in the data 80047’E). with 34.92% being explained by the first component, 16.58% by the Pseudophilautus asankai (Manamendra-Arachchi & Pethiyagoda, second and 9.78% by the third. PCA ordination on the first two axes (Fig. 2005). Holotype: WHT 3507; Paratypes: WHT 3504, 3505, 3506, 26a) could virtually separate Pseudophilautus newtonjayawardanei, P. 5425; Agra Arboretum, Torrington Estate, near Agarapatana, sirilwijesundarai, Pseudophilautus bambaradeniyai and P. puranappu, 1665m (06050’36”N & 80040’40”E); WHT 2100, Moray Estate, however the remaining species showed overlaps within the convex Rajamally, 1370m (06048’N & 80031’E); WHT 5472, Dayagama hulls. PCA ordination on the first three axes (Fig. 26b) could separate Estate, 3rd division, Dayagama, 1830m (06050’N & 80040’E). all the eight species. Correlation values for the variables along the Pseudophilautus auratus (Manamendra-Arachchi & Pethiyagoda, first three PCA axes are provided in Fig. 27. These characters define 2005). Holotype: WHT 2356; Paratypes: WHT 2357, 2375, 2433, the separation of different species from related species. For instance, 2782; Morningside Forest Reserve (near Rakwana), 1060m variables with negative loadings on first and third principle axis and (06024’N & 80038’E); WHT 3282, Handapan Ella Plains (near positive on the second principle component define the separation of Rakwana), 1270m (06026’N & 80036’E); WHT 3298, Sinharaja P. newtonjayawardanei from other species as this species have high World Heritage Site (near Kudawa), 513m (06025’N & 080025’ E). values for such variables. Pseudophilautus caeruleus (Manamendra-Arachchi & Pethiyagoda, Dendrogram depicting the similarity and differences among the 2005). Holotype: WHT 2514; Paratypes: WHT 2511, 2512, 2513; species (Fig. 28) suggested that the eight species are divided into Bogawanthalawa–Balangoda road (near Udugama), 810m three morphometrical clusters. Pseudophilautus newtonjayawardanei (06044’N & 80041’E); WHT 2101A; 2101B, Moray Estate, Rajamally, was distinctly different from all other species and its separation was 1370 m (06048’N & 80031’E). supported by high bootstrap value of 100. Out of the remaining Pseudophilautus cavirostris (Günther, 1869). Holotype: BMNH species, the cluster comprising of Pseudophilautus bambaradeniyai, P. 1947.2.7.83, Southern Ceylon (Southern Sri Lanka); WHT 1294, karunarathnai, P. samarakoon and P. sirilwijesundarai was separated Haycock (Hiniduma, Galle), 660m (06020’N & 80018’E); WHT 3299; from the cluster of P. dayawansai, P. jagathgunawardanai and P. 3300, Sinharaja World Heritage Site, Weddagala, 513m (06025’N puranappu with high value of bootstrap of 86. & 80025’E); WHT 3389, Kitulgala, 200m (07000’N & 80024’E); WHT Statistical analysis presented here has limited application as the 2318; Kosmulla near Neluwa, 450m (06023’N & 80023’E); WHT number of individuals in each species is less. However, the analysis 2046; 2425, Pathanegala (Knuckles), 1,000m (07033’N & 80044’E); does reveal that proposed species are morphometrically distinct WHT 2045; Kadugannawa, 450m (07015’N & 80030’E); WHT 3483; from each other. Most importantly, P. newtonjayawardanei, which is Pussellawe, 986m (07000’N & 80054’E). represented by a single individual, is distinctly different from all other Pseudophilautus cuspis (Manamendra-Arachchi & Pethiyagoda, species described in the current study. Further morphometric data of 2005). Holotype: WHT 1177(e); Paratypes: WHT 1177(a), 1177(b), different species could be helpful in determining the morphological 1177(c), 1177(d), 1177(f), 1177(g), 1177(h), 1177(i), 1177(j), species boundaries among the proposed species. Koskulana (near Panapola), 460m (06025’N & 80027’E). Pseudophilautus decoris (Manamendra-Arachchi & Pethiyagoda, 2005). Holotype: mature female, 23.9mm SVL, WHT 2358; Paratypes: WHT 3194, 3257, 3258, 3265, 3266, 3267, 3268, 3269, 3271, 3270, 3272; Morningside Forest Reserve (near Rakwana), 1060m (06024’N & 80038’E). Pseudophilautus femoralis (Günther, 1864). Holotype: 1947.2.26.89, Ceylon (Sri Lanka); WHT2701; 2702, 2703, 2704, 2705, 2706, 2707, 3540, 3537, 3538, 3539, 2478, 2479; 07.vi.1999, Horton Plains National Park, 2135m (06046’N & 80047’E). Pseudophilautus fergusonianus (Ahl, 1927). Holotype: BMNH 1947.2.27.61, Ceylon (Sri Lanka); WHT 731, 2233; Kumaradola, Monaragala, 305m (06053’N & 81022’E); WHT 1012, Kitulhela (near Moragahapitiya), Monaragala, 520m (07020’N & 81028’E); WHT 2400, 2401; Sera Ella (near Pottotawela), Knuckles, 460m (07035’N & 80045’E); WHT 2038, 2039, 2040, Puwakpitiya (near Laggala), Knuckles, 450m (07034’N & 80045’E); WHT 3360, Deniyaya, 460m (06021’N & 80034’E); WHT 3361, 3362, 3363, 3364, 3365; Pitadeniya (near Watugala), 320m (06022’N & 80028’E) WHT 3168, 3177, 3195; Gannoruwa Forest reserve, Kandy, 684m (07017’N & 80035’E); WHT 3178, 3179, 27, 3180, 3181, 3182; Puwakpitiya, Knuckles, 414m (07034’N & 80044’E); WHT 3229, Hantana, Kandy, 600m (07015’N & 80037’E); WHT 3380, WHT 3381; NMSL 2006.64.1, Medapitiya, Wasgamuwa. Pseudophilautus folicola (Manamendra-Arachchi & Pethiyagoda, 2005). Holotype: mature female, 29.4mm SVL, WHT 2645; Paratypes: 3918 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 Eight new species of Pseudophilautus Wickramasinghe et al. WHT 2646, 2647, 2649, 2650, 2651, 2652; Kottawa (Galle), 60m Pseudophilautus nanus (Günther, 1869). Lectotype: BMNH 1947.2.7.78; (06006’N & 80020’E). BMNH 1947.2.7.79, BMNH 1947.2.7.82, BMNH 1947.2.7.86, Pseudophilautus frankenbergi (Meegaskumbura & Manamendra- southern Ceylon (Southern Sri Lanka) Arachchi, 2005). Holotype: WHT 2554; Paratypes: WHT 2551, Pseudophilautus nasutus (Günther, 1869). Holotype: BMNH 2552, 2555, 2556; Namunukula Peak, 1980m (06056’N & 81007’E); 1947.2.6.21, Ceylon (Sri Lanka). WHT 2726, 2727, 2728; Horton Plains National Park, 2135m Pseudophilautus nemus (Manamendra-Arachchi & Pethiyagoda, (06046’N & 81007’E). 2005). Holotype: WHT 1319; Haycock (Hiniduma), 660m (06020’N Pseudophilautus fulvus (Manamendra-Arachchi & Pethiyagoda, 2005). & 80018’E). Holotype: WHT 2949; Paratypes: WHT 2948, Laggala (Knuckles), Pseudophilautus ocularis (Manamendra-Arachchi & Pethiyagoda, 1220m (07033’N & 80044’E); WHT 3112, 3114, 3463, Moussakanda, 2005). Holotype: WHT 2360; Paratypes: WHT 3273, 2376, 2377, Gammaduwa, Knuckles, 915m (07034’N & 80042’E); WHT 3121; 2378; Morningside Forest Reserve (near Rakwana), 1060m Puwakpitiya (near Laggala), Knuckles, 450m (07034’N & 80045’E). (06024’N & 80038’E); WHT 3288, 3289, 3290, 3291, 3292, 3293; Pseudophilautus hallidayi (Meegaskumbura & Manamendra-Arachchi, Handapan Ella Plains (near Suriyakanda), 1270m (06026’N & 2005). Holotype: WHT 3575; Paratypes: WHT 3573, 3576, 3577; 80036’E). Hanthana range, Kandy, 510–800 m (07015’N & 80034’E); WHT Pseudophilautus oxyrhynchus (Günther, 1872). Lectotype of Ixalus 6072, Tonacombe Estate, Namunukula, 1320m (06052’N & oxyrhynchus (here designated), mature female, 18.6mm SVL, 81007’E). BMNH 1947.2.6.40, ‘Ceylon’, coll. K.G.H.K. Thwaites. Pseudophilautus halyi (Boulenger, 1904). Holotype: BMNH 1947.2.6.16, Pseudophilautus papillosus (Manamendra-Arachchi & Pethiyagoda, Pattipola (6051’N & 80050’E, 1,890m), Ceylon (Sri Lanka). 2005). Holotype: WHT 3284; Handapan Ella Plains (near Rakwana), Pseudophilautus hoffmanni (Meegaskumbura & Manamendra- 1270m (06026’N & 080036’E). Arachchi, 2005). Holotype: WHT 6120; Paratypes: WHT 3222, Pseudophilautus pardus (Meegaskumbura, Manamendra-Arachcchi, 3223, 3542; Corbett’s Gap (Knuckles Hills), 1245m (07022’N & Schneider & Pethiyagoda, 2007). Holotype: BMNH 1947.2.7.96. 80051’E). Pseudophilautus pleurotaenia (Boulenger, 1904). BMNH 1947.2.7.64, Pseudophilautus hoipolloi (Manamendra-Arachchi & Pethiyagoda, Kandy, Sri Lanka; WHT 3176, 5824; Gannoruwa Forest Reserve, 2005). Holotype: WHT 2673; Paratypes: WHT 2674, 2675, Haycock Kandy, 684m (07017’N & 80035’E); WHT 5860, 5861; Nawalapitiya, (Hiniduma), 660m (06020’N & 80018’E); WHT 2405, 2406, 2407, 700m (07003’N & 80032’E). 2408, 2409; Kahaduwatta (Galle), 15m (06004’N & 80012’E); WHT Pseudophilautus poppiae (Meegaskumbura & Manamendra- 3196, 3197, 3198, 3199, 3200; Udugama (Galle), 30m (06014’N Arachchi, 2005). Holotype: WHT 3285; Handapan Ella Plains (near & 80020’E); WHT 3201, Kodagoda (Galle), 45m (06002’N & Suriyakanda), 1270m (06026’N & 80036’E); Paratypes: WHT 2030, 080023’E). 2029, 2475, 2778, 2781, 3533, 3534, 3535, 3536; Morningside Pseudophilautus hypomelas (Günther, 1876). Lectotype BMNH (near Rakwana), 1060m (06024’N & 80038’E). 1947.2.27.8, Ceylon (Sri Lanka); Paralectotypes: BMNH Pseudophilautus procax (Manamendra-Arachchi & Pethiyagoda, 1947.2.7.47; BMNH 1947.2.7.48; BMNH 1947.2.7.49; BMNH 2005). Holotype: WHT 3279; Paratypes: WHT 2786, 2787, 3277, 1947.2.7.50; BMNH 1947.2.7.51; BMNH 1947.2.7.52; BMNH 3278, 3280, 3281; Morningside Forest Reserve (near Rakwana), 1947.2.7.53; BMNH 1947.2.7.54; BMNH 1947.2.7.55; BMNH 1060m (06024’N & 080038’E). 1947.2.7.56; BMNH 1947.2.7.57, Ceylon (Sri Lanka). BMNH Pseudophilautus regius (Manamendra-Arachchi & Pethiyagoda, 1947.2.27.9; BMNH 1947.2.27.10, Le Vallon Estate in Nilambe 2005). Holotype: WHT 3122; Paratypes: WHT 5827, 5828, 5829; District, Ceylon (Sri Lanka). Angammadilla, Polonnaruwa, 90m (07051’N & 80055’E). Pseudophilautus limbus (Manamendra-Arachchi & Pethiyagoda, 2005). Pseudophilautus reticulates (Günther, 1864). Holotype: BMNH Holotype: WHT 2700; Haycock, 560m (06020’N & 80018’E). 1947.2.8.60, Ceylon, (Sri Lanka); WHT 2286, 2287, 2290, 2291; Pseudophilautus lunatus (Manamendra-Arachchi & Pethiyagoda, Kosmulla (near Nelluwa), 320m (06024’N & 80023’E); WHT 2288, 2005). Holotype: WHT 3283; Handapan Ella Plains, 1270m 2289; Dediyagala Forest Reserve, Akuressa, 150m (06010’N & (06026’N & 80036’E). 80026’E); WHT 2344, Yagirala, 30m (06022’N & 80010’E); WHT Pseudophilautus macropus (Günther, 1869). Holotype: BMNH 3230, Induruwa, (Ratnapura), 150m (06045’N & 80026’E); NMSL 1947.2.8.78, southern Ceylon (southern Sri Lanka); WHT 3390, 2006.66.1; Gileemale forest, 150m (06045’ N, 080026’ E); WHT 3391, 3392, 3393, 3394, 3395, 3396, 3397, 3398, 3399, 3400, 2520, 2521; Check Poleat Gap (near Norton Bridge), 800m 3401, 3402, 3403, 3404, 3405, 3406, Mousakanda (Gammaduwa), (06056’N & 80030’E); WHT 3366, Haycock (Hiniduma), 660m Knuckles, 760m (07034’N & 80042’E); WHT 3183, Divulgahapathana, (06020’N & 80018’E) Knuckles, 603m (07033’N & 80043’E); WHT 5900, Bambaraella, Pseudophilautus rus (Manamendra-Arachchi & Pethiyagoda, 2005). Knuckles, 1260m (07024’N & 80047’E). Holotype: WHT 3474; Paratypes: WHT 3475, 3476, 3477; Pseudophilautus maia (Meegaskumbura, Manamendra-Arachcchi, Kiribatkumbura (near Peradeniya), 450m (07016’N & 80034’E); Schneider & Pethiyagoda, 2007). Holotype: BMNH 76.3.21.18; WHT 5437, 5439, 5436; WHT 5440, 5438, 5432, 5435, 5434; Paratype: BMNH 76.3.21.19. Both Holotype and Paratype collected Pilimatalawa (near Peradeniya), 658m (07015’N & 80034’E). at “Poojagodde” (Poojagoda) Estate, Ramboda, Sri Lanka. Pseudophilautus sarasinorum (Müller, 1887). WHT 2480, 2481, Pseudophilautus microtympanum (Günther, 1859). Lectotype: BMNH 2482, 2483; Bogawanthalawa-Balangoda road, 1300m (06045’N 1947.2.8.48, Ceylon (Sri Lanka), WHT 2710, 2708, 2712, 2713, & 80042’E); WHT 2426, 2427, 2428, 2429; Corbett’s Gap, 1000m 3436, 3437, 3438, 3439, 3440, 3441, 3442, 3443, 3444, 3446; (07022’N & 80050’E). Horton Plains National Park, 2135m (06046’N & 80047’E); WHT Pseudophilautus schmarda (Kelaart, 1854). Neotype: WHT 3353; 3260, Agra Arboretum, near Agarapatana, 1555m (06051’N & Horton Plains National Park, 2135m (06046’N & 80047’E). 80041’E). Pseudophilautus silus (Manamendra-Arachchi & Pethiyagoda, 2005). Pseudophilautus mittermeieri (Meegaskumbura & Manamendra- Holotype: WHT 3453; Paratypes: WHT 3451, 3452, 3454, 3460; Arachchi, 2005). Holotype: WHT 3522; Paratypes: WHT 3523, Agra Arboretum, near Agarapatana, 1555m (06051’N & 80041’E); 3524, 3525, 3526; Kottawa, Galle, 60m (06006’N & 80020’E); WHT WHT 3412, Tangamalai Sanctuary, near Haputale, 1600m (06046’N 2668, Beraliya forest, Elpitiya, 150m (06016’N & 80011’E). & 80055’E) Pseudophilautus mooreorum (Meegaskumbura & Manamendra- Pseudophilautus silvaticus (Manamendra-Arachchi & Pethiyagoda, Arachchi, 2005). Holotype: WHT 5862; Paratypes: WHT 5868, 2005). Holotype: WHT 3295; Paratypes: WHT 3296, 3378, 3379, 5869; Hunnasgiriya (Knuckles), 1100m (07023’N & 80041’E); WHT 3461, 3462; Handapan Ella Plains (near Suriyakanda), 1270m 2477, 3209, 6124; Corbett’s Gap (Knuckles), 1245m 0 (07 22’N & (06026’N & 80036’E); WHT 3275, 3276, Morningside Forest Reserve 80051’E). (near Rakwana), 1060m (06024’N & 80038’E); WHT 3310, 3316, Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920 3919 Eight new species of Pseudophilautus Wickramasinghe et al. Sinharaja World Heritage Site (near Kudawa), 513m (06025’N & Sinhala Abstract idrdxYh# uOH l÷lr f,dal Wreu rlaIs; l,dmhg wh;a 80025’E). Y%S mdo rlaIs;fhys —WÉp;ajh fjkiaùug wkqj Wr.hska iy WNhcSùkaf.a Pseudophilautus simba (Manamendra-Arachchi & Pethiyagoda, úúO;ajh fjkiajkq ,nk wdldrh˜ ms 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. 3920 Journal of Threatened Taxa | www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920