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3789
Pseudophilautus
6
1, 2, 3, 4, 5, 6, 7, 8 Herpetological Foundaon of Sri Lanka, 31/5, Alwis Town, Hendala, Waala, Sri Lanka
1 boiga2000@gmail.com (corresponding author), 2 dulanrangavp@gmail.com, 3 gehanrajeev@gmail.com,
4 harithasihinaya@yahoo.com, 5 aamilachanaka@gmail.com, 6 dharshana.dpl@gmail.com, 7 imeshnu1@gmail.com,
8 nemzy821@gmail.com
hp://dx.doi.org/10.11609/JoTT.o3099.3789-920 | urn:lsid:zoobank.org:pub:8E8AF0BF-38D8-4B31-A67F-5BFD3735F66D
Annemarie Ohler, Muséum naonal d’Histoire naturelle, Paris, France 13 March 2013 (online & print)
Ms # o3099 | Received 14 February 2012 | Final received 11 January 2013 | Finally accepted 19 February 2013
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
© Wickramasinghe et al. 2013. Creave Commons Aribuon 3.0 Unported License. JoT T allows unrestricted use of this arcle in any medium,
reproducon and distribuon by providing adequate credit to the authors and the source of publicaon.
Biodiversity Secretariat of the Ministry of Environment, Nagao Natural Environment Foundaon, and Dilmah Conservaon.
None.
The authors wish to acknowledge the Biodiversity Secretariat of the Ministry of Environment, Sri Lanka for funding and collaborang the
project, Mr. R.H.S.S. Samarathunga (Secretary-Ministry of Environment), Mr. Gamini Gamage (Addional Secretary-Policy Planning), Ms. Padma Abayakoon
(Director), Mr. N.K.G.K. Nannawaththa (Addional Secretary-Natural Resources), Mr. R.A.R. Roopasinghe (Adional Secretary- Administraons), Mr. Ajith de
Silva (Former Director), Mrs. Dakshini Perera, and Mrs. Hasula Wickramasinghe for all the support renderd. To the Nagao Natural Environment Foundaon
for part funding, to the Department of Wildlife Conservaon 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. Sta 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 eld, and late Dr. Amith Munindradasa
fondly remembered for his support and encouragements. The authors wish to acknowledge the contribuons of Mr. Prasanna Samarawickrama, Mr. Chamara
Hearachchi, in preparing the locaon illustraons, to Mr. Charith Pubudu Lakmal and Mr. Lalith Senanayake for preparing images. The authors wish to thank
the Director Naonal Museum of Sri Lanka, Dr. Nanda Wickramasinghe and sta 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 assisng the rst 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 Hege, Mr. Sameera Suranjan Karunarathna, Mr. Gayan
Chathuranga, Mr. Sanjiv de Silva provided valuable literature relevant, and Dr. Neelesh Dahanukar for doing the stascal 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
Foundaon of Sri Lanka (HFS), for various courtesies. Thanks goes to Dr. Sanjay Molur for giving vital suggesons to improve the manuscript, to Ms. Chamila
Weerathunga and Mr. Ranil Nanayakkara for proof reading the nal 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 Conservaon, for funding project
acvies.
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 elevaon in the Sripada World Heritage Site (Peak Wilderness), Central Hills of Sri Lanka.
Detailed descripons 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 disnguishable from one another that could be easily idened in the eld.
The conservaon status of all species described here, have been considered Crically Endangered, except for P. newtonjayawardanei, as all
the new species are recorded from single locaons, and their habitats are under severe threat.
Amphibian, new species,Peak Wilderness, PseudophilautusSri Lanka, Sripada.
ISSN
Online 0974-7907
Print 0974-7893
| www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920
Pseudophilautus
The genus Pseudophilautus consists of 65 known
species currently, all of which are endemic to Sri
Lanka (Manamendra-Arachchi & Pethiyagoda 2005;
Meegaskumbura & Manamendra-Arachchi 2005, 2011;
Meegaskumbura et al. 2007, 2009). These direct
developing rhacophorids (Bossuyt & Dubois 2001;
Meegaskumbura et al. 2002; Bahir et al. 2005; Grosjean
et al. 2008; Li et al. 2008, 2009; Biju & Bossuyt 2009; Yu
et al. 2009) previously belonged to the genus Philautus
which were widely distributed throughout tropical Asia
(Bossuyt & Dubois 2001; Bossuyt et al. 2004; Frost et al.
2006; Yu et al. 2009). But later, considering molecular
phylogenecs, the populaons in Sri Lanka and India
were found to be a separate radiaon belonging to a
separate genus known now as Pseudophilautus (Li et al.
2009; Yu et al. 2010; Meegaskumbura & Manamendra-
Arachchi 2011). The most recent publicaon of this genus
by Meegaskumbura & Manamendra-Arachchi (2011) in
the country described two species which brought the
total number to 65 species. But sadly, a considerable
poron of the total number of idened species, have
already vanished (17 species) (Manamendra-Arachchi &
Pethiyagoda 2005; Meegaskumbura et al. 2007) where
most have not even been sighted aer their inial
descripons, showing an urgency to conserve the few
remaining habitats.
Peak Wilderness as part of the Central Hills was recently
designated as a World Heritage Site. Geographical area
of the sanctuary is about 240sq.km, of which 211.75sq.
km comprises natural or semi natural vegetaon; the
rest is no longer forest and includes tea estates and
village selements. The sanctuary lies in the center
of the western ridge of the Central Highlands, north-
east of Ratnapura and spanning the border of Central
and Sabaragamuwa provinces. Its eastern boundary
is conguous with Horton Plains Naonal Park. Peak
Wilderness occupies the escarpment that rises steeply
from the lowlands to the south and west. Much of the
terrain is very rugged, with altudes ranging from 600m
near Ratnapura to 2,238m at Sripada Peak, Sri Lanka’s
fourth highest peak. It is an important watershed for
three major river systems, the Kelani Ganga, the Kalu
Ganga, and the Walawe Ganga (DWC 2007).
The last vertebrate species discovered from
the studied area were Cnemaspis samanalensis
(Wickramasinghe & Munindradasa 2007); Lankascincus
munindradasai and Lankascincus sripadensis
(Wickramasinghe et al. 2007), depicng its diversity.
Because of relavely high altudes, dicult
trails, heavy rainfall, and steep geographic variaons,
proper eld surveys in this region were lacking. Due
to this reason inspite of the geographical and climac
variaons the region was considered to have a lower
faunal diversity. Hence, we carried out the survey with
the intenon of seng a basis for the actual diversity
of herpetofauna of Peak Wilderness. The novel species
described here from the Peak Wilderness, Central Hills,
of Sri Lanka, as results of the preliminary two phases of
an extensive eld survey of the region which lasted for
a period of two years (end of 2009-end of 2011), were
based on morphological and morphometric characters.
Sripada Peak is the highest point in this mountain range,
and because of its religious importance, and aesthec
beauty, pilgrims and travellers have idened six major
trails leading to the top. For the 1st phase, sampling was
carried out in the trail leading from Palabaddala to the
Sripada peak, and in the 2nd phase we chose Kuruvita to
the Sripada peak. Our work here describes a collecon
of eight new species, with each species having a detailed
descripon based on strong morphological characters,
accompanied by colour photographs.
The approximate total distances of the 1st trail (from
Palabaddala to the Sripada peak) was 10km, and the 2nd
trail (from Kuruvita to the Sripada peak) was 10km. The
eld survey commenced at the end of 2009, and the
Phases I and II were completed in December 2011. The
sampling sites were selected considering the accessibility,
representave habitats and spaal distribuon in the
Naonal Park, through an inial reconnaissance survey.
Fieen eld visits were made, each sampling session
spanning eight connuous days. Sampling was done
mostly nocturnally, and photographs of most species
were done in the wild to avoid any confusion of change
in colour aer capvity and to avoid over collecon of
the same species. Specimens collected in the eld were
rst xed in 90% ethanol for two hours and stored in
70% ethanol.
Sex and maturity were determined by examining
secondary sexual characters, or when absent, by
examining the gonads through a small lateral incision
in the specimen. The following morphological
charactetriscs were common to all eight species hence
they have not been menoned under each species;
nostrils oval, without ap of skin laterally, small, close
to p of snout; latero-dorsal folds, lateral line system,
Fejervarya line, and macroglands were absent. We here
| www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920
Pseudophilautus
have explained the terminology used to explain the skin
textures, specially the variaons 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 Conservaon
(DWC), Naonal Wildlife Research and Training Center,
Girithale, Sri Lanka.
The new species were compared with all types
from Sri Lanka, deposited in the NMSL and specimens
deposited in the Natural History Museum, London
(Appendix 4). The specimens that formerly belonged to
the Wildlife Heritage Trust (WHT) bearing WHT numbers
are currently deposited in the NMSL, catalogued under
the same numbers. For the descripon secon, 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 combinaon of characters that best
disnguishes. In the comparison tables for each species,
for convenience of disnguishing the species, an empty
cell in the table denotes either the same character as
that of the species considered, or for few exnct species
when there was no data available. Hence in order to
minimise this error we included all characteriscs and
all species in one table provided in Appendix 1 and 2,
by selecng the best combinaon of characters where
Appendix 1, provides morphological characters that
dierenate Pseudophilautus species and Appendix 2
provides characters in the skin structure that dierenate
Pseudophilautus species. We paid special aenon to
exnct 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 alphabecally herein: Breadth of disk
of the third nger (DB), taken from anterior to posterior
edge of circum-marginal groove; distance between back
of eyes (DBE), measured between posterior edge of
eyes; distance between front of eyes (DFE), measured
between anterior edges of eyes; length of disk of the
third nger (DL), taken from anterior edge of circum-
marginal groove to posterior edge of disk; width of disk
of the third nger (DW), measured across the inner and
outer edges of circum-marginal groove; eye diameter
(ED), horizontal diameter of eye; eye to nostril distance
(EN), measured between anterior most point of eye and
middle of nostril; eye to snout distance (ES), measured
between anterior most point of eye and p of snout;
thigh (femur) length (FEL), distance between vent and
Pseudophilautus
jagathgunawardanai
Pseudophilautus newtonjayawardanei
© L.J. Mendis Wickramasinghe
© L.J. Mendis Wickramasinghe
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Pseudophilautus
knee with both thigh and shank exed; rst nger
length (FL-1), measured between posterior margin of
the most proximal subarcular tubercle or crease of
arculaon and the p of rst nger; second nger
length (FL-2), measured between posterior margin of
the most proximal subarcular tubercle or crease of
arculaon and the p of second nger; third nger
length (FL-3), measured between posterior margin of
the most proximal subarcular tubercle or crease of
arculaon and the p of third nger; fourth nger
length (FL-4), measured between posterior margin of
the most proximal subarcular tubercle or crease of
arculaon and the p of fourth nger; foot length
(FOL), distance between heel and p of fourth toe with
both foot and shank exed; groin to knee distance (GK),
measured between groin and the p of knee; hand
length (HNL), taken from posterior-most margin of inner
palmar tubercle to p of disk of third nger; head depth
(HD), distance between the apex of eye and the boom
of jaw; head length (HL), distance between angle of
jaws and snout p; head width (HW), measured across
angle of jaws; inner metatarsal tubercle length (IML),
distance between the anterior and posterior edges
of the inner metatarsal tubercle; internarial distance
(IN), least distance between the inner margin of nares;
interorbital width (IO), least distance between the upper
margins of orbits; knee-angle length (KT), distance from
knee-angle to bio-tarsal arculaon; forearm length
(LAL), taken from elbow to posterior-most margin of
inner palmar tubercle; mandible-back of eye distance
(MBE), distance between angle of jaws and posterior-
most point of eye; mandible-front of eye distance (MFE),
taken as distance between angle of jaws and anterior-
most point of eye; mandible-nostril distance (MN), taken
as distance between angle of jaws and middle of nostril;
snout-nostril distance (SN), taken as distances between
middle of nostril and p of snout; snout-vent length
(SVL), measured from p 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 bio-tarsal arculaon and
anterior edge of inner metatarsal tubercle; bia length
(TBL), distance between knee and heel with both
shank and foot exed; rst toe length (TL-1), measured
between posterior margin of the most proximal
subarcular tubercle or crease of arculaon and the p
of rst toe; second toe length (TL-2), measured between
posterior margin of the most proximal subarcular
tubercle or crease of arculaon and the p of second
toe; third toe length (TL-3), measured between posterior
margin of the most proximal subarcular tubercle or
crease of arculaon and the p of third toe; fourth
toe length (TL-4), measured between posterior margin
of the most proximal subarcular tubercle or crease of
arculaon and the p of fourth toe; h toe length
(TL-5), measured between posterior margin of the most
proximal subarcular tubercle or crease of arculaon
and the p of h toe; tympanum-nostril distance
(TND), distance between anterior-most point of inner
margin of inner rim of tympanum and middle point of
nostril; tympanum-front of eye distance (TPD), taken as
distance between anterior-most point of inner margin of
inner rim of tympanum and anterior most point of eye;
tympanum height (TYH), vercal diameter of the inner
rim; tympanum width (TYW), horizontal diameter of the
inner rim; upper arm length (UAW), distance between
axilla and elbow; upper eyelid width (UEW), measured
from bony edge of supraorbital to outer edge of upper
eyelid; vent to knee-angle length (VKL), distance
measured from knee-angle to vent.
Morphometric data of both males and females was
considered for stascal analysis to understand the
morphometric dierence among the individuals of eight
new species of Pseudophilautus. To account for the
size dierence among individuals, all the morphometric
Pseudophilautus
sirilwijesundarai
© L.J. Mendis Wickramasinghe
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Pseudophilautus
3793
characters were taken as %SVL. Principle Component
Analysis (PCA) was performed on the correlaon matrix
between 25 variables using the soware PAST (Hammer
et al. 2001). Dendrogram showing the relaonship
between dierent species was ploed using mean values
of morphometric data for each species using Euclidian
distances and Ward’s method for clustering in PAST.
Geographical coordinates were determined from
GPS readings (Gamin eTrex Gista) with WGS84 (World
Geodec System) datum at the locality. Addional data
for comparison were collected from Pethiyagoda &
Manamendra-Arachchi (2005) and WGS 84 world grid
system.
Collectors are indicated by their inials as follows:
L.J. Mendis Wickramasinghe (L.J.M.W), Dulan Ranga
Vidanapathirana (D.R.V),
M.D. Gehan Rajeev (M.D.G.R),
S. Chathuranga Ariyarathne (S.C.A.), A. W. Amila Chanaka
(A.W.A.C.), L.L. Dharshana Priyantha (L.L.D.P.), Imesh
Nuwan Bandara (I.N.B.).
Pseudophilautus bambaradeniyai
(Figs. 1–4; Images 4–7; Tables 1–3)
urn:lsid:zoobank.org:act:FB630D8A-7354-4307-A343-5B5343921C73
NMSL 2013.01.01 NH, 05.iv.2010, adult
male SVL 17.3mm (Images 4, 5A,D), Sripada (Peak
Wilderness), Ratnapura District, Sabaragamuwa
Province, Sri Lanka (06048’31”N & 8008’14”E; 1334m)
(Fig. 1), coll. L.J.M.W, D.R.V., S.C.A. & A.W.A.C.
DWC 2013.01.001, adult male, SVL
19.9mm (Images 5B,E, 6); DWC 2013.01.002, adult male,
SVL 20.2mm (Images 5C,F, 7); same data as holotype.
Pseudophilautus bambaradeniyai sp.
nov., can be disnguished from known congeners by the
following combinaon of characters: small size frog (SVL
17.3–20.2 mm); head dorsally convex; canthus rostralis
rounded; interorbital space convex; internarial space at;
vomerine teeth, lingual papilla, fringe on ngers, calcar
and nupal pad absent; snout, interorbital area, side of
head, upper and lower ank smooth; anterior dorsum
prominent tubercles and horny spinules, posterior
dorsumhorny spinules. Disnguishing characters
provided in comparison tables 2 and 3.
Small sized frog (SVL
17.3mm), elongate (SVL/HW 2.4); head large (HL/SVL
0.5), convex above; snout truncate in lateral aspect,
mucronate in dorsal aspect, and sub-ellipcal in ventral
aspects (Fig. 2 A–C) (ES/DFE 0.8, SN/IN 0.7), larger
than horizontal diameter of eye (ES/ED 1.1); internasal
space at; canthus rostralis rounded, loreal region
concave; interorbital space convex, larger than upper
eyelid (IO/UEW 1.1), and equal to internasal distance
(IO/IN 1.0); distance between front of eyes 2/3rd the
distance between back of eyes (DBE/DFE 1.6); nostrils
oval, without ap of skin laterally, closer to p of snout
than to eye (SN/EN 0.8); pupil horizontally ellipcal;
tympanum disnct, oval (TYH/TYW 0.8), smaller than
the eye diameter (TYH/ED 0.4), tympanum-eye distance
about 2/5th of tympanum width (TAD/TYW 0.4); pineal
ocellus, vomerine teeth and lingual papilla absent, but
a depression with a conical tubercle present on tongue;
tongue small, lanceolate.
Arm short, thin (LAL/FEL 0.5, UAL/FEL 0.3); forearm
shorter than hand length (LAL/HNL 0.9), longer than
upper arm (LAL/UAL 1.4); ngers thin, (Fig. 3), relave
length of ngers I < II < IV < III (FL-1/FL-3 0.5, FL-2/
FL-3 0.7, FL-4/FL-3 0.9) (Table 1), and webbing formula
I2½—2 2/3II2-— 3½III2½—2-IV; ps of ngers rounded
enlarged, discs present on all ngers, with disnct basal
and circum marginal grooves; dermal fringes absent;
rudimentary webbing present on all ngers; subarcular
tubercles prominent, oval, single, all present, III2 and
IV2 relavely smaller, supernumerary tubercles present
on nger III and IV; inner palmar tubercle indisnct,
single, oval, larger than outer palmar tubercle; outer
palmar tubercle indisnct, single, oval, smaller than the
subarcular tubercles; nupal pads absent; prepollex
absent; femur 2 2/5 mes longer than fourth toe length
(FEL/TL-4 2.4); foot length longer than thigh (FOL/FEL
1.3); toes thin, (Fig. 4), relave length of toes I < II < III <
V < IV (TL-1/TL-4 0.3, TL-2/TL-4 0.4, TL-3/TL-4 0.7, TL-5/
TL-4 0.7); webbing formula I2+—2-II2+—3-III2+—3-IV3—
2V; ps of toes rounded, enlarged, discs present on all
toes with disnct basal and circum marginal grooves;
thin dermal fringe present on all toes, thin fringe along
postaxial edge of toe V, small tubercles present on outer
edge of tarsal fold; subarcular tubercles prominent,
oval, single, all present, IV2, IV3, and V2 relavely
smaller; supernumerary tubercles present, indisnct, on
toe III and IV; inner metatarsal tubercle oval prominent
and large, its length 3/4 mes in length of toe I (IML/TL-1
0.6); outer metatarsal tubercle indisnct.
Skin of snout dorsally, laterally and interorbital space
smooth; upper eyelid tubercular; head laterally smooth;
dorsum anterior with prominent tubercles and horny
spinules, posterior with prominent horny spinules;
upper and lower part of ank smooth; supratympanic
fold disnct; upper arm, forearm, and hand smooth;
thigh inner and outer smooth, dorsally with tubercles,
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Pseudophilautus
● Pseudophilautus bambaradeniyai P karunarathnai
Psamarakoon ” P dayawansai P sirilwijesundarai ♦P jagathgunawardanai” P
newtonjayawardanei■P puranappu
▲
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Pseudophilautus
macroglands absent; leg smooth with few tubercles;
calcar absent, small tubercles present at heel; vent
dorsally shagreen, either side shagreen.
Ventral side of body: Throat and chest weakly
granular; belly granular; upper arm weakly granular to
Pseudophilautus bambaradeniyai
Pseudophilautus bambaradeniyai
Pseudophilautus bambaradeniyai
Pseudophilautus bambaradeniyai
© L.J. Mendis Wickramasinghe
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Pseudophilautus
3796
smooth; forearm smooth; thigh, leg and tarsus smooth.
Dorsum dark brown with blackish
blotches, large dark brown marking covering much of the
anterior, blackish cross band between eyes, a prominent
o-white vertebral stripe from the p of snout to anus,
and connuing down the hind limbs symmetrically,
lateral body lighter; limbs dark brown, fore limbs, hind
limbs, ngers and toes with blackish-brown cross bands;
ventral side o white with light brown blotches, hands,
feet and webbing lighter (Image 4).
Colour paern remains with a
lile fading, o-white vertebral stripe prominent on dark
brown, ventral side o-white all over (Images 5 A,D).
Dorsum colour varies
ranging from orange to a lighter brownish nge and the
thickness of the vertebral line varies; in some this line
disconnues towards limbs (Images 5 B,E, 6).
The species epithet bambaradeniyai
Pseudophilautusbambaradeniyai
Pseudophilautusbambaradeniyai
Pseudophilautusbambaradeniyai
© L.J. Mendis Wickramasinghe
© L.J. Mendis Wickramasinghe
© L.J. Mendis Wickramasinghe
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Pseudophilautus
3797
0.4 0.5 0.5 0.5 0.0 0.4—0.5 9.4 9.6 9.6 9.5 0.1 9.4—9.6
6.8 7.6 7.3 7.2 0.4 6.8—7.6 4.6 4.8 4.8 4.7 0.1 4.6—4.8
4.3 4.3 4.3 4.3 0.0 4.3—4.3 3.2 3.0 3.5 3.2 0.2 3.0—3.5
0.7 0.7 0.8 0.7 0.0 0.7—0.8 6.1 6.3 6.0 6.2 0.1 6.0—6.3
0.8 0.8 0.9 0.9 0.0 0.8—0.9 7.5 8.0 7.8 7.8 0.3 7.5—8.0
3.2 3.4 3.2 3.3 0.1 3.2—3.4 1.5 1.5 1.3 1.5 0.1 1.3—1.5
1.9 2.1 2.1 2.0 0.1 1.9—2.1 17.3 19.6 20.2 19.0 1.5 17.3—20.2
3.4 3.3 3.2 3.3 0.1 3.2—3.4 0.6 0.6 0.6 0.6 0.0 0.6—0.6
10.1 9.6 10.2 10.0 0.4 9.6—10.2 6.2 6.3 6.8 6.4 0.3 6.2—6.8
1.5 1.7 1.7 1.6 0.1 1.5—1.7 10.5 10.8 10.9 10.7 0.2 10.5—10.9
2.1 2.3 2.1 2.2 0.1 2.1—2.3 1.4 1.5 1.4 1.4 0.1 1.4—1.5
3.2 3.7 3.5 3.5 0.3 3.2—3.7 1.5 1.7 1.7 1.7 0.1 1.5—1.7
2.8 2.9 3.0 2.9 0.1 2.8—3.0 2.9 2.9 3.0 2.9 0.0 2.9—3.0
13.3 14.1 14.2 13.9 0.5 13.3—
14.2 4.2 4.7 4.6 4.5 0.2 4.2—4.7
9.2 8.7 9.2 9.0 0.3 8.7—9.2 3.1 3.3 3.2 3.2 0.1 3.1—3.3
4.2 4.4 4.6 4.4 0.2 4.2—4.6 5.3 5.5 5.6 5.5 0.2 5.3—5.6
8.4 8.6 8.5 8.5 0.1 8.4—8.6 3.6 3.7 3.7 3.7 0.1 3.6—3.7
5.3 6.4 6.1 5.9 0.6 5.3—6.4 1.3 1.5 1.5 1.4 0.1 1.3—1.5
7.4 8.3 8.2 8.0 0.5 7.4—8.3 1.5 1.5 1.5 1.5 0.0 1.5—1.5
0.9 0.8 0.9 0.9 0.1 0.8—0.9 3.4 3.5 3.5 3.5 0.1 3.4—3.5
2.1 2.4 2.0 2.2 0.2 2.0—2.4 1.9 2.2 2.1 2.1 0.2 1.9—2.2
2.2 2.7 2.6 2.5 0.3 2.2—2.7 9.1 8.9 8.7 8.9 0.2 8.7—9.1
Pseudophilautus bambaradeniyai
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Pseudophilautus
3798
Species name
P. bambaradeniyai
sp. nov. 17.3–20.2 Convex Truncate Rounded Convex Flat Absent Absent Absent Absent Absent
P. dayawansai sp. nov. 24.5–30.1 Concave Concave Present Present Present
P. jagathgunawardanai
sp. nov. 35.8-40.4 Rounded Sharp Concave Present
P. karunarathnai sp. nov. 16.2–19.2 Rounded Concave Present Present Present
P. newtonjayawardanei
sp. nov. 38.4 Concave Rounded Concave Concave Present
P. puranappu sp. nov. 33.6–51.1 Concave Rounded Flat Concave Present Present Present
P. samarakoon sp. nov. 20.4–24.6 Present Present
P. sirilwijesundarai
sp. nov. 22.3–32.5 Rounded Present Present
P. abundus 25.4–37.0 Concave Pointed or oval Flat Concave Present Present Present
P. adspersus 33.3–41.7 Flat Sharp Present
P. alto 17.0–27.5 Obtusely pointed Concave Present Present Present
P. asankai 18.9–27.3 Obtusely pointed Present
P. auratus 21.7–26.4 Flat Blunt Flat Concave Present Present
P. caeruleus 16.3–19.0 Flat Obtusely pointed Sharp Flat Present
P. cavirostris 38.4–48.9 Concave Oval Sharp Concave Concave Present Present Present
P. cuspis 17.6–28.9 Sharply Pointed Sharp Present Present
P. decoris 18.3–23.9 Flat Obtusely pointed Concave Concave or at Present Present
P. dimbullae 44.8 Sharp Flat Present Present
P. eximius 35.3 Flat Rounded Sharp Flat Present Present
P. exrpo 43.5 Flat Blunt Sharp Concave Concave Present Present
P. femoralis 23.4–28.8 Flat Flat Concave Present
P. fergusonianus 26.9–44.5 Concave or
at Rounded Concave or at Concave Present Present Present
P. folicola 23.7–29.4 Flat Rounded or truncate Sharp Flat Concave Present Present
P. frankenbergi 26.7–29.3 Flat Rounded Sharp Flat Present
P. fulvus 33.4–46.8 Rounded Sharp Flat Concave Present Present Present Present
P. hallidayi 32.9–42.9 Rounded Flat Present Present Present
Pseudophilautus bambaradeniyai Pseudophilautus
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Pseudophilautus
3799
Species name
P. halyi 27.9 Uneven Rounded Sharp Flat Present Present
P. hankeni 18.3–21.9 Pointed Concave
P. homanni 21.2–23.4 Flat Rounded Flat Present
P. hoipolloi 22.0–28.6 Blunt Flat Concave Present
P. hypomelas 11.2–22.4 Acuminate Absent or present
P. leucorhinus 19.8 Flat Oval Sharp Concave Concave
P. limbus 25.7 Concave Obtusely pointed Concave Concave Present Present Present
P. lunatus 40.9 Flat Oval Concave Concave Present Present Present Present
P. macropus 27.4–42.7 Convex or
concave Oval Flat Present Present Present Present
P. maia 46.6 Concave Sharp Concave Present Present
P. malcolmsmithi 14.9 Pointed Sharp Flat Present
P. microtympanum 23.9–49.0 Oval Flat Concave Present Present Present
P. miermeieri 16.3–18.4 Pointed Concave Concave Present Present.
P. mooreorum 29.4–35.0 Flat Rounded Flat Concave Present
P. nanus 34.8 Oval Flat Concave Present Present
P. nasutus 17.4 Pointed Sharp Flat Concave Present
P. nemus 20.7 Obtusely pointed Flat Present
P. ocularis 23.1–33.4 Flat Blunt Flat Concave or at Present Present
P. oxyrhynchus 18.6 Flat Pointed Sharp Concave Concave Present
P. papillosus 40 Sharp Concave Present Present Present Present Present
P. pardus 32.1 Rounded Flat Concave
P. pleurotaenia 25.6–30.3 Concave Present
P. poppiae 21.3–26.0 Rounded Flat Concave Present
P. popularis 17.7–24.7 Rounded Flat
P. procax 25.1–26.8 Concave Pointed or oval Flat Concave Present Present Present
P. regius 18.3–21.7 Flat Oval Concave Concave Present Present
P. reculatus 42.5–61.1 Convex or at Sharp Convex or
concave Present Present Present Present Present
P. rugatus 17.5 Rounded Sharp Concave
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Pseudophilautus
Species name
P. rus 20.6–24.1 Flat Oval Flat Present
P. sarasinorum 22.6–38.1 Flat Rounded Flat Concave Present
P. schmarda 17.7–30.0 Obtusely pointed Sharp Concave Present Present
P. schneideri 19.9–22.8 Pointed Sharp Present
P. semiruber 13.4 Oval Concave Present
P. silus 35.4–51.8 Oval Sharp Flat Present Present
P. silvacus 24.3–31.3 Oval Sharp Flat Present Present
P. simba 12.6–15.6
P. singu 16.1–16.6 Rounded Concave Present
P. sordidus 22.2–39.4 Blunt Present Present
P. steineri 30.4–41.6 Flat Rounded Sharp Flat Concave Present Present Present
P. stellatus 39.6–55.3 Concave Rounded Concave Concave Present Present Present
P. sctomerus 23.2–36.0 Obtusely pointed Present Present
P. stuar 24.2–32.4 Flat Rounded Sharp Flat
P. tanu 13.5–13.9 Obtusely pointed Flat
P. temporalis 28.4–31.0 Rounded Concave Concave Present
P. variabilis 35.7 Flat Flat Concave Present
P. viridis 27.4–36.3 Blunt or rounded Flat Present
P. zal 20.0–32.5 Rounded Concave Present Present
P. zimmeri 31.6 Blunts Sharp Flat Present Present Present
P. zorro 22.9–30.1 Concave Pointed Sharp Flat Concave Present Present
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Pseudophilautus
P. bambaradeniyai
sp. nov. Smooth Smooth Smooth Prominent tubercles and
horny spinules Horny spinules Smooth Smooth Tuberculated
P. dayawansai sp. nov.
Tuberculated, a single
prominent tubercle
medially
Smooth, with a ridge
across breaking
medially
Tubercular Smooth Weakly tubercular Granular Granular Smooth
P. jagathgunawardanai
sp. nov. Smooth,with two
cross dermal fringes Smooth Smooth Weakly
granular
P. karunarathnai sp. nov. Smooth, a prominent
blunt tubercle Weakly tubercular Weakly tubercular Smooth Spinulate Weakly granular Granular Smooth
P. newtonjayawardanei
sp. nov. Shagreen Shagreen, with a
prominent ridge Shagreen Shagreen Shagreen Shagreen Weakly
granular Shagreen
P. puranappu sp. nov. Smooth, with a blunt
tubercle Weakly shagreen Few horny spinules,
Prominent tubercles Weakly tubercular Granular Weakly shagreen
P. samarakoon sp. nov.
Prominent tubercle
present on centre of
snout
Tubercular Tubercular Tubercular Tubercular Granular
P. sirilwijesundarai
sp. nov.
Shagreen, with
prominent dermal fold
A prominent tubercle
present Weakly shagreen Shagreen with few tubercles Shagreen Areolate Granular Smooth
P. abundus Horny spinules (absent in
females) Granular Smooth
P. adspersus
Smooth except for a
few scaered glandular
warts
Smooth except for
a few scaered
glandular warts
Smooth except for
a few scaered
glandular warts
Smooth except for a few
scaered glandular warts Smooth Granular Granular Smooth
P. alto Horny spinules in males Horny spinules in
males Horny spinules in males,
warty in females
Horny spinules in males,
warty in females
Horny spinules in
males Granular Smooth
P. asankai Very ne, horny
spinules
Very ne, horny
spinules
Very ne, horny
spinules Very ne, horny spinules Very ne, horny spinules Horny spinules
Very ne,
horny
spinules
Very ne, horny
spinules
P. auratus Shagreen Shagreen Shagreen Shagreen Shagreen Granular or smooth Granular Smooth
P. caeruleus Glandular warts and
horny spinules
Glandular warts and
horny spinules
Glandular warts
and horny spinules
Glandular warts and horny
spinules (females lack horny
spinules)
Glandular warts and
horny spinules (females
lack horny spinules)
Glandular warts and
horny spinules
Granular
in males Smooth
P. cavirostris Glandular warts Glandular warts Glandular warts Heavily tuberculated,
glandular warts
Heavily tuberculated,
glandular warts Granular Smooth
P. cuspis Horny spinules Horny spinules Horny spinules with a “)
(”shaped paern of tubercles Smooth or with
glandular warts
P. decoris Glandular warts Glandular warts Glandular warts Glandular warts Glandular warts Glandular warts Glandular
warts Glandular warts
P. dimbullae Shagreen Shagreen Shagreen Shagreen Shagreen Shagreen Granular Smooth
Pseudophilautus bambaradeniyai Pseudophilautus
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Pseudophilautus
P. eximius Smooth Smooth Weakly
granular Smooth
P. exrpo Shagreen, with a few
glandular warts Shagreen Shagreen Shagreen, with a few
glandular warts
Shagreen, with a few
glandular warts Granular Granular Shagreen, few
glandular warts
P. femoralis Finely granular Finely granular Finely granular Finely granular Finely granular Finely granular Granular Smooth
P. fergusonianus
Shagreen, males
glandular warts and
horny spinules
Shagreen, males
glandular warts and
horny spinules
Few glandular
warts
Shagreen, males glandular
warts and horny spinules
Few glandular warts,
males glandular warts and
horny spinules
Shagreen, males
with glandular warts
and horny spinules
Granular Smooth with
glandular warts,
P. folicola
Shagreen, smooth
or with a few
scaered glandular
warts
Smooth or granular Smooth Granular Granular Smooth
P. frankenbergi Shagreen Shagreen Glandular warts Shagreen, horny spinules
scaered
Shagreen, horny spinules
scaered Shagreen Granular Smooth
P. fulvus Shagreen Shagreen Shagreen Shagreen Shagreen Shagreen Granular Shagreen
P. hallidayi Glandular warts, with
horny spinules in males
Glandular warts,
with horny spinules
in males
Glandular warts Glandular warts, with horny
spinules in males
Glandular warts, with
horny spinules in males
Glandular warts,
with horny spinules
in males
Granular Glandular warts
P. halyi Shagreen with
glandular warts
Shagreen with
glandular warts
Shagreen with
glandular warts
Shagreen with glandular
warts
Shagreen with glandular
warts Glandular warts Granular Glandular warts
P. hankeni Glandular warts
bearing horny spinules
Glandular warts
bearing horny
spinules
Glandular warts
bearing horny
spinules
Glandular warts bearing
horny spinules
Glandular warts bearing
horny spinules
Glandular warts
bearing horny
spinules
Granular
Glandular warts
with horny
spinules
P. homanni Smooth Smooth Glandular Smooth
P. hoipolloi Smooth or shagreen Smooth or shagreen Smooth
P. hypomelas Smooth Smooth
P. leucorhinus Smooth with a few scaered
glandular warts
Smooth with a few
scaered glandular warts
Smooth with
glandular warts
Granular
with
glandular
warts
Smooth
P. limbus Weakly granular Weakly granular Weakly granular Weakly granular Weakly granular Granular Granular Granular, not
rough
P. lunatus Smooth Smooth Glandular warts Granular Smooth
P. macropus
Smooth with a few
scaered glandular
warts
Glandular warts Shagreen or smooth, with a
few scaered glandular warts
Shagreen or smooth, with
a few scaered glandular
warts
Granular with a few
scaered glandular
warts
Granular
Smooth with a
few scaered
glandular warts
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Pseudophilautus
P. maia Glandular warts Glandular warts Glandular warts Glandular
warts Smooth
P. malcolmsmithi Smooth, '\ /' shaped ridge Smooth Glandular warts Glandular
warts
Smooth, few
scaered
glandular warts
P. microtympanum Smooth or
tuberculated Shagreen
Smooth, shagreen and
glandular warts, males horny
spinules
Smooth, shagreen and
glandular warts, males
horn like spinules
Granular Granular
Shagreen,
few scaered
glandular warts
P. miermeieri Glandular warts Glandular warts Glandular warts Glandular warts Glandular warts Glandular warts Glandular
warts Smooth
P. mooreorum Horny spinules in males Horny spinules in
males
Horny spinules in
males
Horny spinules in males,
nely granular or shagreen
in female
Horny spinules in
males, nely granular or
shagreen in female
Horny spinules in
males
Horny
spinules in
males
Shagreen
P. nanus Glandular warts Few glandular warts Few glandular
warts Few glandular warts Few glandular warts Glandular folds Granular Smooth
P. nasutus Glandular warts Glandular warts Glandular warts tuberculated, horn-like
spinules
Tuberculated, horn-like
spinules Granular Glandular warts
P. nemus Glandular warts Glandular warts Glandular warts Glandular warts Granular Smooth
P. ocularis Shagreen or with
glandular warts
Shagreen or with
glandular warts
Shagreen or with
glandular warts
Shagreen or with glandular
warts
Shagreen or with
glandular warts
Shagreen or
glandular warts Granular Glandular warts
P. oxyrhynchus Smooth Smooth
P. papillosus Glandular warts Glandular warts Glandular warts Glandular warts Glandular warts Glandular warts Granular Glandular warts
P. pardus Smooth Smooth Granular Granular Smooth
P. pleurotaenia Shagreen Shagreen Shagreen
Shagreen if females, glandular
warts and horny spinules (
males)
Shagreen, few scaered
glandular warts and horny
spinules (males)
Shagreen, few
scaered glandular
warts, and horny
spinules
Granular Smooth
P. poppiae
Horny spinules in
males, nely granular
or shagreen in female
Horny spinules in
males, nely granular
or shagreen in female
Males horny
spinules, females
nely granular or
shagreen
Horny spinules in males,
nely granular or shagreen
in female
Horny spinules in
males, nely granular or
shagreen in female
Horny spinules
in males, nely
granular or
shagreen in female
Horny spinules
in males, nely
granular or
shagreen in
female
P. popularis Glandular warts Glandular warts Horny spinules Horny spinules Granular Glandular warts
P. procax Granular Smooth
P. regius Horny spinules Horny spinules Horny spinules Horny spinules Glandular Glandular Glandular warts
| www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920
Pseudophilautus
P. reculatus
Shagreen with a few
scaered glandular
warts
Shagreen with a few
scaered glandular
warts
Shagreen with
a few scaered
glandular warts
Shagreen with a few scaered
glandular warts, nely
tuberculated
Shagreen with a few
scaered glandular warts,
nely tuberculated
Shagreen with a few
scaered glandular
warts
Granular Smooth
P. rugatus
Smooth except for a
few scaered glandular
warts
Smooth except for
a few scaered
glandular warts
Smooth except for
a few scaered
glandular warts
Smooth except for a few
scaered glandular warts
Smooth except for a few
scaered glandular warts Glandular warts Granular
Smooth, few
scaered
glandular warts
P. rus Glandular warts Glandular warts Glandular warts Granular Granular Smooth
P. sarasinorum Glandular warts Glandular warts Glandular warts Glandular warts, males with
horny spinules scaered
Glandular warts, males
with horny spinules
scaered
Glandular folds Granular
in females
P. schmarda Glandular warts
Glandular folds,
glandular warts and
horny spinules
Glandular warts Glandular folds, glandular
warts and horny spinules
Glandular folds, glandular
warts and horny spinules
Glandular folds,
glandular warts and
horny spinules
Glandular
warts and
horny
spinules
Glandular warts
P. schneideri Small tubercules with
horny spinules
Small tubercules with
horny spinules
Small tubercules
with horny spinules
Small tubercules with horny
spinules
Small tubercules with
horny spinules
Small tubercules
with horny spinules Granular Smooth
P. semiruber Smooth Smooth Smooth
P. silus Glandular warts Glandular warts Glandular warts Glandular warts Granular Smooth
P. silvacus Glandular warts Glandular warts Glandular warts Glandular warts Smooth Glandular warts Granular Smooth
P. simba Glandular warts Glandular warts Smooth Granular Smooth
P. singu Scaered, glandular
tubercles
Scaered, glandular
tubercles
Scaered, glandular
tubercles Scaered, glandular tubercles Scaered, glandular
tubercles
Scaered, glandular
tubercles
Scaered,
glandular
tubercles
Scaered
glandular
tubercles
P. sordidus Glandular warts Glandular warts Glandular warts Glandular warts, males horny
spinules scaered
Glandular warts, males
horny spinules scaered
Glandular warts,
with horny spinules
in males
Granular Glandular warts
P. steineri Glandular warts and
horny spinules
Glandular warts and
horny spinules
Glandular warts and horny
spinules, females lack horny
spinules
Glandular warts and
horny spinules, females
lack horny spinules
Glandular warts and
horny spinules Granular
Smooth,
scaered
glandular warts
P. stellatus Weakly shagreen Weakly shagreen Weakly shagreen Weakly shagreen Weakly shagreen Shagreen to weakly
areolate
Weakly
areolate Weakly shagreen
P. sctomerus Shagreen, horny spinules in
males
Shagreen, horny spinules
in males
Shagreen, horny
spinules in males Granular Smooth
P. stuar Horny spinules Horny spinules Horny spinules, females lack
horny spinules
Horny spinules, females
lack horny spinules
P. tanu Shagreen Shagreen Shagreen Shagreen Shagreen Granular Granular Smooth
| www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920
Pseudophilautus
P. temporalis Smooth Smooth Coarsely
granular Smooth
P. variabilis Smooth Smooth Granular Granular Smooth
P. viridis
Horny spinules
(smooth in females and
immature males)
Horny spinules
(smooth in females
and immature males)
Horny spinules
(smooth in females
and immature
males)
Horny spinules (smooth in
females and immature males)
Horny spinules
(smooth in females
and immature
males)
Granular Shagreen
P. zal Weakly shagreen Shagreen Shagreen Weakly tubercular Weakly tubercular Granular Granular Glandular warts
P. zimmeri Glandular warts Glandular warts Glandular warts Glandular warts Glandular warts Granular Granular Glandular warts
P. zorro Horny spinules Horny spinules Glandular warts
Horny spinules, Anterior
dorsum with “)(” shaped
paern of tubercles
Glandular warts Granular Glandular warts
| www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920
Pseudophilautus
is named aer Channa Bambaradeniya, a leading
wetland scienst who has been working immensely to
protect the wetlands and its ecosystems in Sri Lanka, in
great appreciaon of his dedicated eorts towards the
conservaon of the biodiversity at large in the country.
The species name bambaradeniyai is a noun in the
genive case.
Bambaradeniyage
panduru madiya, and Bambaradeniya’s Shrub Frog in
Sinhala and English, respecvely.
The species was found in elevaons
of 750–1400 m from the forest oor to about 1.5m high
shrubs in lowland rain forests to lower montane rain
forests.
Pseudophilautus dayawansai
(Figs. 5–7; Images 8–12; Tables 4–6)
urn:lsid:zoobank.org:act:3D56DDDE-770F-4D0E-BBE3-04C357922CDE
NMSL 2013.02.01 NH, 23.x.2011, adult
male SVL 24.5mm (Images 8, 9A,D). Sripada (Peak
Wilderness), Ratnapura District, Sabaragamuwa
Province, Sri Lanka (06048’30”N & 80029’19”E; altude
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..
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);
informaon same as holotype.
Pseudophilautus dayawansai
Pseudophilautus dayawansai
Pseudophilautus dayawansai
| www.threatenedtaxa.org | 13 March 2013 | 5(4): 3789–3920
Pseudophilautus
Pseudophilautus dayawansai
© L.J. Mendis Wickramasinghe
Small sized frog (adult male SVL 24.5–26.2
mm, adult female SVL 30.1mm); head dorsally convex;
snout laterally truncate, skin dorsally tuberculated, a
single prominent tubercle medially placed; canthus
rostralis rounded; loreal region, interorbital space and
internarial space concave; vomerine teeth absent; lingual
papilla, fringe on ngers and blunt small calcar present;
interorbital area smooth, with a ridge across braking
medially; side of head tubercular; anterior dorsum
smooth; posterior dorsum weakly tubercular; median
dermal ridge absent; upper and lower ankgranular;
throat shagreen.
Small sized frog (SVL
Pseudophilautus dayawansai
© L.J. Mendis Wickramasinghe
Pseudophilautus dayawansai
© L.J. Mendis Wickramasinghe