Description of eight new species of shrub frogs (Ranidae: Rhacophorinae: Philautus) from Sri Lanka

Abstract

Eight new species of Sri Lankan frogs of the genus Philautus are described (P. mooreorum, P. poppiae, P. hoffmanni, P. mittermeieri, P. frankenbergi, P. hallidayi, P. steineri and P. stuarti). The species are diagnosed on the basis of mitochondrial DNA sequence, morphological features and, in two cases, bioacoustics data. Six of the eight species are confined to high elevation cloud forest isolates on the three main mountain ranges of central Sri Lanka. Because of their limited geographic distribution and small extent of remaining habitat, these species are classified as Endangered under the IUCN Red List criteria. These descriptions bring the total number of Sri Lankan Philautus to 61 species, 44 of which are extant.

Full text

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
THE RAFFLES BULLETIN OF ZOOLOGY 2005 Supplement No. 12: 305–338
© National University of Singapore
DESCRIPTION OF EIGHT NEW SPECIES OF SHRUB FROGS
(RANIDAE: RHACOPHORINAE: PHILAUTUS) FROM SRI LANKA
Madhava Meegaskumbura
Department of Biology, Boston University, 5 Cummington Street, Boston, MA, 02215, USA
Wildlife Heritage Trust, 95 Cotta Road, Colombo 8, Sri Lanka
Email: madhava@bu.edu
Kelum Manamendra-Arachchi
Wildlife Heritage Trust, 95 Cotta Road, Colombo 8, Sri Lanka
Email: kelum@slt.lk
ABSTRACT. – Eight new species of Sri Lankan frogs of the genus Philautus are described (P. mooreorum, P.
poppiae, P. hoffmanni, P. mittermeieri, P. frankenbergi, P. hallidayi, P. steineri and P. stuarti). The species are
diagnosed on the basis of mitochondrial DNA sequence, morphological features and, in two cases, bioacoustics
data. Six of the eight species are confined to high elevation cloud forest isolates on the three main mountain
ranges of central Sri Lanka. Because of their limited geographic distribution and small extent of remaining
habitat, these species are classified as Endangered under the IUCN Red List criteria. These descriptions bring
the total number of Sri Lankan Philautus to 61 species, 44 of which are extant.
KEY WORDS. – Rhacophoridae, taxonomy, montane, bioacoustics, evolutionary lineage, conservation
INTRODUCTION
The high endemism and diversity of the direct-developing
rhacophorine frogs of the genus Philautus Gistel, 1848 of Sri
Lanka is now well established (Meegaskumbura et al., 2002;
Bossuyt et. al., 2004; Manamendra-Arachchi & Pethiyagoda,
2005), but many species remain to be described. Molecular
phylogenetic analyses have played an important role in the
recognition and diagnosis of species as well as in the higher-
level systematics of the Sri Lankan radiation. We believe that
cautious description supported by phylogenetic analyses,
utilizing also data on the biology of the species involved,
represents the best approach to the recognition of species.
Otherwise, hypotheses to explain patterns of diversity and
distribution in the Sri Lankan radiation of Philautus risk being
no more than speculative (e.g., Manamendra-Arachchi &
Pethiyagoda, 1998; Meegaskumbura et al., 2002; Dubois,
2004). While we strive for a complete documentation of the
anuran diversity of Sri Lanka, made particularly urgent by the
Globally Threatened status of many of the recently-discovered
species (Stuart et al., 2004), we hope the naming of these
species will facilitate their conservation and further research
on their biology.
Following almost a decade of fieldwork, museum visits to
study type material and molecular analyses (Pethiyagoda &
Manamendra-Arachchi, 1998; Meegaskumbura et al., 2002)
and the taxonomic report of Bossuyt & Dubois (2001),
Manamendra-Arachchi & Pethiyagoda (2005) reviewed the
Sri Lankan Philautus. The latter authors re-described the
name-bearing types, provided descriptions of 27 new species,
recognized 17 species as extinct, and provided a key to all the
valid Sri Lankan species of Philautus. This served to increase
the total number of Philautus in Sri Lanka from six (Kotagama
et al., 1981) to 53 species. The present work furthers this
objective, but it differs from the purely morphological approach
of Manamendra-Arachchi & Pethiyagoda (2005) by doing so
within a phylogenetic framework. Focusing largely on
montane isolates, we provide a hypothesis of phylogeny based
on mitochondrial DNA (with a more complete phylogenetic
analysis of mitochondrial and other data to follow) and
describe eight new species. This brings the total number of
Sri Lankan Philautus to 61.
Based on a “morpho-species” assessment in their original
announcement of this hitherto unsuspected diversity,
Pethiyagoda & Manamendra-Arachchi (1998) suggested that
there could be ~ 200 species of Philautus in Sri Lanka. This
estimate was revised downward when phylogenetic analysis
of mitochondrial DNA (mtDNA) sequences from putative
species suggested that the actual number of distinct lineages
was in fact about half of that originally estimated

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Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
(Meegaskumbura et al., 2002). In addition, the mtDNA
phylogeny provided a phylogenetic framework that facilitated
the identification of apomorphic morphological characters that
were diagnostic of clearly defined mtDNA lineages. Most of
these lineages had already been recognized as potential new
species by Manamendra-Arachchi & Pethiyagoda (2005) using
classical, morphological taxonomy and were not discovered
as a consequence of the molecular analysis alone contra Mace
et al. (2003). While the descriptions and key presented in
Manamendra-Arachchi & Pethiyagoda (2005) do not explicitly
reference the mtDNA tree, both were informed by patterns of
morphological apomorphy implied by the mtDNA tree and
are thus consistent with patterns of morphological character
state distribution in it (a detailed analysis of morphological
evolution in Sri Lankan Philautus will be presented
elsewhere). In addition to informing the diagnosis of species
of Sri Lankan Philautus, the phylogenetic analysis revealed
several distinct mtDNA lineages, which were represented,
however, only by relatively small series of specimens. These
show sufficient morphological and other differentiation to
allow their recognition as species; here we describe eight
such species.
Species Concept. – We consider it important in taxonomic
studies to state explicitly as a testable hypothesis the species
concept used, so as to prevent definition of species in a post-
hoc, arbitrary manner (Sites & Crandall, 1997). More than 25
species “concepts” are recognized in the literature (reviewed
by de Queiroz, 1998; Coyne & Orr, 2004), each with its own
limitations (Hey, 2001). As in our previous treatments of the
Sri Lankan rhacophorine fauna (Meegaskumbura et al., 2002;
Manamendra-Arachchi & Pethiyagoda, 2005), we here adopt
the General Lineage Concept of species (de Queiroz, 1998).
This defines species as independent evolutionary lineages
that are diagnosed by multiple criteria. We find the General
Lineage Concept to be useful and at the same time inclusive
of many of the current species “concepts” in that species are
recognized as segments of evolving lineages which, based
on multiple criteria, appear to be independent from other such
lineages (see de Queiroz, 1998, for a fuller discussion).
Furthermore, the incorporation of multiple criteria for
diagnosing species recognizes the multi-dimensional nature
of species as ecological, morphological and behavioural
entities as well as reproductive and historical entities.
In this analysis, we used mitochondrial DNA (mtDNA)
sequences, external morphology, ecology and, in some
instances, bioacoustics data for diagnosing species. We
consider sets of populations to be evolutionarily independent
species when they are consistently divergent in at least two,
and preferably three or more, of these criteria.
The conservation status of many of the Sri Lankan Philautus
species is precarious, mostly because of habitat loss and the
fragmentation of remaining habitats. Of the 36 extant species
recognized by Manamendra-Arachchi & Pethiyagoda (2005),
seven were classified as Critically Endangered and 17 as
Endangered in terms of the IUCN (1996) criteria for global red
listing. This underlines the urgency to describe and document
the remaining diversity. Six of the eight species described in
this paper are classic montane isolates, restricted to cloud-
forest habitats at high altitude (more than 1,000 m elevation)
in three major mountain ranges: the Central Hills, Rakwana
Hills, and Knuckles Hills (Fig. 1). We have focused on these
especially since the long-term persistence of these montane
isolates is uncertain given that cloud forest habitats, and the
species that rely on them, may be highly susceptible to the
effects of global climate change (Pounds & Puschendorf, 2004)
and also local pressures from habitat loss and degradation.
MATERIALS AND METHODS
Field sampling and measurements were made as described in
Manamendra-Arachchi & Pethiyagoda (2005), except as
mentioned below.
Fig. 1. Map of Sri Lanka depicting the wet zone (WZ), intermediate
zone (IMZ) and the dry zone (DZ), and the three mountain ranges
referred to in this study: Knuckles hills (K), Central hills (C) and
the Rakwana hills (R). The 500 m and 1,000 m contours are shown.
The Central hills have the largest area above 1,000 m and Rakwana
Hills have the smallest. The valley separating Central hills and
Knuckles has a maximum elevation of ~ 550 m and that separating
the Central and Rakwana hills has a maximum elevation of ~ 450 m.
Scale bar: 50 km.
WZ
IMZ
DZ
K
C
R

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
Morphological analysis. – The suit of characters
(continuous and discrete) used by Manamendra-Arachchi &
Pethiyagoda (2005) were analysed for all individuals.
For continuous characters, measurements were made to the
nearest 0.1 mm using dial vernier callipers. All 32 measurements
listed by Manamendra-Arachchi & Pethiyagoda (2005) were
made for the species described herein, but those with high
coefficients of variation or low repeatability were omitted from
the analysis. The following measurements were used for the
current analysis: distance between back of eyes (DBE);
distance between front of eyes (DFE); eye diameter (ED);
eye-to-snout length (ES); femur length (FEL); length of finger
3 (FLIII); pes length (FOL); head length (HL); head width
(HW); inter-narial distance (IN); inter-orbital distance (IO);
lower-arm length (LAL); posterior mandible to eye distance
(MBE); least distance from mandible to anterior eye (MFE);
least distance from mandible to nostril (MN); nostril-to-snout
length (NS); palm length (PAL); snout–vent length (SVL);
tibia length (TBL); and length of toe 4 (TLIV).
Principal components analysis of the character correlation
matrix was used to reduce dimensionality of the continuous
morphological variables and to identify those variables that
best discriminate among species. Various axis rotations were
tested and one selected for optimal interpretability of variation
among the characters. In almost all cases, the first two principal
components explained more than 90% of the variance. Sample
sizes from most populations were small and biased toward
calling males. For consistency, therefore, only mature males
were used in the multivariate morphological analysis. The
single exception was P. hallidayi, for which only a single
male was collected. In this case, mature females were used in
the analysis. In several cases, the small sample sizes likely do
not represent the full range of morphological variance;
nonetheless, the analyses are sufficient to demarcate species
and identify characters that contribute best to species
diagnoses. SYSTAT (Version 11.00.01 for WindowsXP) was
used for statistical analysis.
Molecular analysis. – DNA was extracted from ethanol-
preserved tissues using Qiagen tissue extraction kits and
manufacturer’s protocols. Mitochondrial 12S and 16S
ribosomal RNA gene fragments were amplified using standard
PCR conditions and primers 12Sa and 12Sb (Palumbi, 1996)
which amplified ~ 380bp of the 12S rRNA gene, and 16Sar and
16Sbr (Palumbi, 1996) which amplified ~ 550bp of the 16S
rRNA gene. PCR conditions were as follows: denaturation at
95°C for 40 s, annealing at 45° C for 40 s and extension at 72°
C for 40 s, 35 cycles, with a final extension of 72° C for 5 min.
Products were gel purified and sequenced on an ABI 377 or
ABI 3100 automated sequencer following manufacturer’s
protocols.
Sequences were aligned using Clustal X (Jeanmougin et al.,
1998) and adjusted by eye using Se-Al (ver. 2.0a9; Rambaut,
1996). Positions which were difficult to align and in which we
had low confidence in positional homology were excluded
from subsequent analyses.
Phylogenetic analysis utilized the dataset for the Sri Lankan
Philautus from Meegaskumbura et al. (2002) with 17 additional
sequences. The tree was rooted using two species of Indian
Philautus which form the sister group to the Sri Lankan Philautus
(Meegaskumbura et al., 2002). We removed all other taxa from
the analysis of Meegaskumbura et al. (2002) since supraspecific
taxonomy was not an aim of this study. The data were analyzed
using Bayesian, Maximum Likelihood (ML) and Maximum
Parsimony (MP) criteria. For brevity, we present only the Bayesian
tree, which is identical to the Maximum Likelihood tree and one
of the three equally parsimonious trees. We used Bayesian
inference as implemented in MrBayes (Huelsenbeck & Ronquist,
2001) to generate a phylogenetic hypothesis of relationships
among the taxa and to estimate a general time reversible model of
sequence evolution with gamma-distributed rate variation among
sites and a proportion of invariant sites (GTR+I+G). We ran four
Metropolis-Coupled Markov Chain Monte Carlo (MCMCMC)
chains for 500,000 generations and the summed likelihood of the
four chains converged on a stationary value by 80,000 generations
(the burn-in time). We used the frequency of clades in trees that
were sampled every ten generations from the last 250,000
generations as estimates of the posterior probabilities of those
clades (Huelsenbeck et al., 2001). Uniform priors were used
throughout and branch lengths, topology, and nucleotide
substitution parameters were unconstrained. Maximum
likelihood analysis used a GTR+I+G model of nucleotide
substitution with the parameters estimated from the Bayesian
analysis. A single heuristic search with Tree Bisection and
Reconnection (TBR) branch swapping was conducted using
PAUP*4.0b10 (Swofford, 1998). For tree searches under a
Maximum Parsimony criterion we used 100 heuristic searches
with TBR branch-swapping and random taxon addition as
implemented in PAUP*4.0b10.
Once we identified the divergent mtDNA lineages and their
sister taxa using the 12S and 16S rRNA gene tree, and to
facilitate comparisons with published summaries of
mitochondrial divergence among vertebrate sister species
(Johns & Avise, 1998), we sequenced a 590 bp fragment of
the mitochondrial cytochrome-b gene from the species
described herein and their sister species. For this analysis, a
~ 590 base-pair fragment of the mitochondrial cytochrome-b
gene was amplified using primers CB-J-10933, (5’-
TATGTTCTACCATGAGGACAAATATC-3’) and BSF4 (5’-
CTTCTACTGGTTGTCCTCCGATTCA-3’) (Bossuyt &
Milinkovitch, 2000) under standard PCR conditions:
denaturation at 95° C for 40 s, annealing at 45° C for 40 s and
extension at 72° C for 40 s, 35 cycles, with a final extension of
72° C for 5 min. Products were gel purified and sequenced on
an ABI 377 or ABI 3100 automated sequencer following
manufacturer’s protocols. Sequences were aligned using
translated amino acid sequences using Se-Al (ver. 2.0a9;
Rambaut, 1996).
We used the observed molecular distances (of cytochrome-b
gene fragments) between the new species and their sister
species to estimate molecular divergence times. Since there is
no reliable rate calibration for Philautus (or Rhacophorine
frogs), a crude estimate of the ranges of divergence times for

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Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
the sister species pairs was derived using published low and
a high estimates of divergence rates. Since some of the
hypervariable regions of the ribosomal genes were removed
from this analysis, only the cytochrome-b fragment was used.
The two rates are 0.69% divergence rate per million years for
all substitutions, which was estimated using frogs (Martin &
Palumbi, 1993) and 1.0% divergence rate per million years for
all substitutions, which was estimated using newts (Tan &
Wake, 1995). A more complete divergence time analysis will
be made with a more complete phylogeny of the Sri Lankan
Philautus.
Bioacoustics analysis. – Bioacoustics data were collected in
the field using a Sony analogue Walkman (WM-D6C) with a
Sennheiser K6 shotgun microphone or a Sony digital Walkman
(TCD-D100) with a Sony ECM-717 microphone. Ambient
relative humidity and temperature were noted during
recordings. Recorded calls were transferred to an Apple G4
iBook using a Griffin iMic USB audio input device. Calls were
analyzed using Raven (ver. 1.4 for Mac OS X).
Since the call terminology is confusing due to the diversity of
signal structures it is necessary explicitly to define the terms
used (Gerhardt & Huber, 2002). Here we use call length (time
in seconds from beginning to end of a single call), pulse rate
((total number of pulses -1)/time, for each call), dominant
frequency (frequency in call that contains the greatest
energy), fundamental frequency (the harmonic with the lowest
frequency), pulse length (time from beginning to end of one
pulse from mid-call), call type (the absence or presence of
frequency modulation within a call), pulse type (absence or
presence of frequency modulation within pulse) and the
number of pulses per call, as described by Cocroft & Ryan
(1995). These characters are potentially phylogenetically
informative and also thought to be important in mate
recognition and sexual selection (Cocroft & Ryan, 1995;
McCraken & Sheldon, 1997; Sullivan, 1992). Some of these
call characters have also been used to distinguish among
anuran species in morphologically cryptic species (eg. Vences
et al., 2000).
Etymological note. – For nearly two decades, the enigmatic
and steady worldwide decline of amphibian populations has
been watched with concern and studied by scientists across
the world. During this time, dozens of species have
disappeared, and hundreds of others are on the brink.
Solutions to this crisis have been scarce, with each result of
ongoing research revealing fresh, hitherto unsuspected
complexity in this problem. Two initiatives have sought to
turn this dismal tide, not just at the level of nations or regions,
but on a global scale: the Declining Amphibian Populations
Task Force (DAPTF) and the Global Amphibian Assessment
(GAA). For more than a decade, the DAPTF has served to
network scientists and conservation managers across the
world, helping to build capacity where it is needed, and to
share information and experience. Over a shorter time frame,
the GAA, through worldwide collaboration perhaps without
precedent in conservation science, succeeded in assessing
the status of every amphibian species (Stuart et al., 2004).
There can be no doubt that both these initiatives have
succeeded only because of the extraordinary zeal, dedication
and resolve of the small number of conservation scientists
who have given of their time, and the handful of private
donors who have given of their wealth, to help save these
wonderful animals.
The results of the GAA focus attention on the fact that with
17 species extinct and a further 44 threatened with extinction,
Sri Lanka’s amphibians are in dire straits. Indeed, we cannot
escape the concern that even the species we describe here
may soon disappear forever. However, the DAPTF and GAA
have given fresh hope that amphibian conservation
programmes on a truly global scale are not just possible, but
realistic. The flow of resources for conservation has
necessarily been from developed countries to those tropical,
developing nations in which the vast majority of amphibian
species dwells, with so few opportunities for workers in the
latter to acknowledge those who make these resources
available. We are grateful therefore, to Rohan Pethiyagoda
(pers. comm.) for his proposal that we recognize eight of the
people who have made the DAPTF and the GAA possible by
naming these new species in their honour. We would like our
action to be viewed as one of the ways in which herpetologists
in developing countries can acknowledge those who
disinterestedly have found the will and the commitment to
support amphibian conservation world-wide.
RESULTS
Molecular phylogenetics. – The final dataset contained
mitochondrial 12S and 16S rRNA gene sequences from 53
individuals (36 from the dataset analyzed by Meegaskumbura
et al. (2002) plus seventeen in addition). Fifty individuals
represent Sri Lankan Philautus; three represent Indian species
(one, P. wynaadensis, is nested within the Sri Lankan clade,
whereas the other two represent the sister group to the Sri
Lankan Philautus: Fig. 2: Meegaskumbura et al., 2002). As
most of the divergent genera that were used by
Meegaskumbura et al. (2002) in their earlier analysis were
removed, alignment of the Indian and Sri Lankan Philautus
sequences was less problematic, resulting in the inclusion of
some sequence regions that were excluded in the previous
analysis. Hence, of the 939 nucleotide positions sequenced,
895 were clearly allignable and were included in this analysis,
whereas only 802 positions were included in the earlier
analysis.
The tree with the highest likelihood resulting from the
Bayesian analysis is shown in Fig. 2. The tree is rooted with
two Indian taxa (Philautus charius and P. signatus) that
represent the sister group to the Sri Lankan Philautus
radiation (Meegaskumbura et al., 2002). We ran 500,000
generations of the MCMCMC algorithm and the summed
likelihood of the four chains reached stationarity by 80,000
generations. The posterior probabilities of clades shown in
Fig. 2 represent the frequency of those clades in the 25,000
trees sampled from the last 250,000 generations and clades
with posterior probability of fifty percent or less were
collapsed. The parameters of the nucleotide substitution

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
model for the most likely tree were as follows. Rate matrix:
R(G-T) = 1, R(C-T) = 78.3408, R(C-G) = 1.2399, R(A-T) = 5.5206,
R(A-G) = 34.0613, R(A-C) = 6.1740. Nucleotide frequency: A =
0.3162, C = 0. 0.2502, G = 0.1877, T =0.2457. Rate variation:
shape parameter for gamma distributed rate variation among
sites (alpha) = 0.745; proportion of invariant sites = 0.382. As
expected, the maximum likelihood tree found via a Tree
Bisection and Reconnection branch-swapping heuristic
search using the above nucleotide substitution parameters
in PAUP*v.4.0b10 had the same topology as the Bayesian
tree, but had slight branch-length differences (tree not
shown). A heuristic search using the Parsimony criterion, TBR
branch swapping with 100 replicates with random taxon
addition, and all characters unordered and weighted equally
gave three equally parsimonious trees (trees not shown), one
of which matched the topology of the Bayesian and ML tree.
The other two MP trees differed only slightly from the
Bayesian and ML trees.
The 22 sequences representing previously described
species are all well separated by relatively long branch
lengths on the tree (see branch lengths for Philautus
Fig. 2. a, The Bayesian tree from 895 base pairs of 12s and 16s rRNA genes of the Sri Lankan and Indian Philautus species. Each included
Sri Lankan sister group is shown in blue and green (new species in blue, their sister species in green). The Indian species, including the
outgroup species, are shown in red. The numbers on the branches depict the Bayesian posterior probabilities (nodes having a posterior
probability of less than 50 have been collapsed). b, Lines indicate distribution of species that are isolated on mountain tops over 1000 m
altitude (red: P. stuarti and P. viridis; light green: P. femoralis, P. mooreorum and P. poppiae; yellow: P. auratus and P. frankenbergi; light blue:
P. steineri and P. microtympanum; purple: P. hoffmani and P. asankai). The three major mountain ranges are Kunckles hills (K), Central hills
(C) and Rakwana hills (R).
a b

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Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
sarasinorum, P. procax, P. schmarda, P. cavirostris, P. alto,
P. viridis, P. femoralis, P. auratus, P. sordidus, P.
microtympanum, P. caeruleus, P. decoris, P. hoipolloi, P.
asankai, P. ocularis, P. zorro, P. pleurotaenia, P. popularis,
P. simba, P. lunatus, P. papillosus and P. limbus: Fig. 2).
However, there are also several unnamed, yet distinct
mitochondrial lineages on the tree that need further
investigation (WHT 2774; WHT 2489; WHT 2484; WHT
2658; WHT 2792; WHT 2797; WHT 2667; WHT 2525; and
WHT 2669: Fig. 2). We selected eight such lineages
(labelled with their new species epithets, P. mooreorum, P.
poppiae, P. hoffmanni, P. frankenbergi, P. hallidayi, P.
steineri, P. mittermieiri and P. stuarti, shown in blue in
Fig. 2) that had adequate sample sizes of voucher
specimens for further investigation. Each of the mtDNA
lineages that we selected for further scrutiny are robustly
placed (posterior probability of 1.00) as closely related
sister lineages to one of the previously recognized species
(P. femoralis, P. asankai, P. auratus, P. cavirostris, P.
microtympanum, P. decoris and P. viridis) (Fig. 2, Table 1),
except for a single instance in which the sister relationship
of P. steineri is not well resolved, although it is placed in a
clade together with P. microtympanum and P. sordidus,
which was moderately supported with a posterior
probability of 0.64. The uncorrected molecular distance
between the new species and their sister species, for the
combined 12S and 16S gene fragment, ranged between 1%
and 8% (Table 1).
We further tested the robustness of the mitochondrial lineages
that were selected as potential species by sequencing a 590
bp fragment of the mitochondrial cytochrome-b gene, which
has been widely used in systematics and taxonomy across
vertebrate taxa (Johns & Avise, 1998). We generated 21
sequences representing the eight new species described
herein and their immediate sister taxa (the same individuals
sequenced for the 12S and 16S fragments were sequenced for
the cytochrome-b fragment). An aim of this analysis was to
compare how the eight putative new Philautus species
compare with trends in molecular divergence among other
vertebrate sister species. Johns & Avise (1998) showed that
90% of putative vertebrate species, including amphibians,
differ by more than 2% genetic distance in the cytochrome-b
gene (uncorrected for unobserved multiple substitutions).
The uncorrected genetic distance between the eight new
species described herein and their respective sister species
ranges from 5.8–15.7% (Table 2), which are moderate to large
mtDNA distances consistent with the lineages representing
different species. This conclusion is supported by analysis
of morphological variation within and among these sister
lineages.
Morphological analysis. – For each of the novel species,
there is a combination of consistent morphological character
states that defines the species and that does not vary within
a given population. These include shagreened, horny, or
smooth body surfaces; snout-angle categories; distinctness
of the tympanum; shape of inter-orbital space; shape of snout
in lateral aspect; presence or absence of glandular warts on
skin; presence or absence of sheath-like undulating
membranes on the posterior margins of limbs; presence or
absence of vomerine ridges; cross-bars on limbs; shapes of
canthal edges; webbing pattern on toes; angle of vomerine
ridge to body axis; presence or absence of calcar on tibio-
tarsal fringe; shape of loreal region; presence or absence of
dermal fringes on fingers; distinctiveness of supratympanic
fold; consistent markings on body; presence or absence of
lingual papilla; and differences in colour. Diagnostic character
state sets and differences among species are discussed in
the species descriptions below.
All the species recognized were effectively differentiated
using principal components analysis (Figs. 5a, 5b, 16, 20, 24,
28, 32). Most of the species pairs differed most strongly on
the first principal component, which in all cases represented
a size axis (snout-vent length). However, in one group (the P.
femoralis, P. mooreorum and P. stuarti clade), P. femoralis
differed from the other species on the second principal
component as well, which represented differences in the
proportional size of hands and feet. In contrast to all others,
Table 1. Percent pairwise uncorrected molecular distance between
the various sister-species groups using a 895 bp fragment of the 12s
and 16s gene, analyzed using PAUP (v. 4b10). Montane isolates
found above 1,000 m altitude are indicated with an asterisk (*).
sister species uncorrected pairwise
molecular distance (%)
P. asankai – P. hoffmanni*1.01
P. auratus – P. frankenbergi*3.00 – 3.13
P. decoris – P. mittermeieri 2.38
P. viridis – P. stuarti*1.45 – 1.56
P. microtympanum – P. steineri*2.34
P. femoralis – P. mooreorum*3.15
P. femoralis – P. poppiae*2.56 – 2.78
P. mooreorum – P. poppiae*2.56 – 2.80
P. cavirostris – P. hallidayi 7.90
Table 2. The percent pairwise uncorrected molecular distance
between the sister species groups using a 590 bp fragment of the
cytochrome-b gene, analyzed in the program PAUP (v. 4b10) and
the time of divergence of the sister species estimated using 0.69 –
1.00% rate of change per million years. Montane isolates found
above 1,000 m are indicated with an asterisk (*).
sister species pairwise molecular
uncorrected divergence
molecular estimate
distance (%) (mya)
P. asankai – P. hoffmanni*6.04 4.16 – 6.04
P. auratus – P. frankenbergi*11.22 –11.97 7.74 –11.97
P. decoris – P. mittermeieri 5.80 4.00 – 5.80
P. viridis – P. stuarti*6.34 – 6.93 4.37 – 6.93
P. microtympanum – P. steineri*11.06 7.63 –11.06
P. femoralis – P. mooreorum*8.0 5.52 – 8.0
P. femoralis – P. poppiae*8.9 6.14 – 8.90
P. mooreorum – P. poppiae*7.7 5.31 – 7.70
P. cavirostri – P. hallidayi 15.7 10.83 –15.70

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P. hallidayi and P. cavirostris differed in head dimensions
(represented on the second principal components axis), as
well as size. The clear discrimination of the new species from
their sister species in multivariate morphological space further
supports our contention that these species represent
independent evolutionary lineages that warrant formal
recognition. Below we detail the comparisons of morphometric
variation among sister species.
Principal components analysis with unrotated axes on the
correlation matrix of continuous characters from Philautus
mooreorum from Knuckles Hills, P. femoralis from the Central
Hills and P. poppiae from the Rakwana Hills showed clear
separation of the three species on two axes (Fig. 5a). Of the
total variance, 79.5% was explained by PC1, which represents
a size axis, with body size (snout–vent length) having the
highest loading and with all other variables having high
positive loadings. Manus and toe dimensions (which load
negatively, but with relatively large absolute values) explain
7.3% of the total variance. All three species separate out well
on the first principal component axis (PC1), with size ascending
in the following order: P. poppiae, P. femoralis, P. mooreorum.
Philautus femoralis separates well from P. poppiae and P.
mooreorum on the second principal component axis (PC2) as
well, with P. femoralis having relatively larger palms, fingers
and toes, for its body size when compared to the other two
species. Philautus poppiae and P. mooreorum almost
completely overlap on PC2 and hence are indistinguishable
on this axis.
The name-bearing type specimen of P. femoralis
(BMNH1947.2.26.89) is a female and is not accompanied by
sufficient information to determine its collection locality within
Sri Lanka. However, based on morphometric data, it appears
to be a member of the Central Hills population (Fig. 5b).
Principal components analysis including the holotype of P.
femoralis with females and males from the three prominent
mountain ranges revealed that the holotype overlaps with
individuals from the Central Hills in size (and, like the Central
Hills specimens, it is distinct from those of the Rakwana and
Knuckles mountains). However, the finger and palm
dimensions are small in BMNH1947.2.26.89, which
distinguishes it from the other three species, including the
Central Hills population of P. femoralis, on PC2. Smaller hands
and feet may be the result of shrinkage in preservative, but
until further evidence is available, we follow Manamendra-
Arachchi & Pethiyagoda (2005) in regarding the Central Hills
population to be P. femoralis sensu stricto.
Unrotated principal components analysis effectively
separates Philautus mittermeieri and P. decoris on a single
axis (PC1; Fig. 16). Of the total variance, 69.35% is explained
by PC1, which is a size axis (the highest factor loading was for
SVL and all other variables had high positive values).
Philautus mittermeieri and P. decoris separate well on PC1
(size), P. decoris being larger than P. mittermeieri. Of the total
variance, 9.1% is explained by PC2, which reflects variance in
eye diameter and distance between eyes. This result, however,
is uninformative as the two species show nearly complete
overlap on this axis.
Philautus frankenbergi and P. auratus, with no rotation (Fig.
20), separate along a single PC axis. Of the total variance,
90.1% was explained by PC1, which again is a size axis. PC2,
which represents variance in inter-narial distance, explains
3.9% of the total variance. Philautus frankenbergi and P.
auratus separate out well on PC1, with P. frankenbergi having
a larger body size relative to P. auratus. There is nearly
complete overlap on PC2, which is uninformative with regard
to species diagnosis.
Philautus hallidayi and P. cavirostris also separate on a single
axis (Fig. 24). With a varimax rotation of axes, 36% of the total
variance is explained by PC1which represents limb dimensions
(length of femur, tibia, foot, lower arm, palm and finger load
positively and most heavily). However, the two species show
nearly complete overlap on PC1 and hence are not diagnosable
by limb dimensions. In contrast, the two species separate
well on PC2, which explains 33% of the variance and represents
variation in head-eye dimensions (distance between front of
eyes, eye-to-snout distance and inter-orbital distance load
positively on PC2). Thus, P. hallidayi can be distinguished
from P. cavirostris by having a smaller eye-to-snout distance,
lesser distance between front of eyes, and lesser inter-orbital
distance.
Philautus steineri and P. microtympanum also separate well
in PC space along a size axis (PC1; Fig. 28), with P. steineri
being larger than P. microtympanum. With unrotated axes,
94.3% of the total variance was explained by the PC1 (size).
Only 2.2% of the total variance was explained by PC2, which
represented palm length (negative loading), finger 3 length
(negative loading) and lower-arm length. Philautus steineri
and P. microtympanum overlapped almost completely on PC2.
Philautus stuarti and P. viridis also show separation by body
size, with P. viridis being larger than P. stuarti (Fig. 32). With
unrotated axes, 74.0% of the total variance is explained by
variance along PC1, which again represents body size.
Variance along PC2 explains 7.0% of the total variance,
representing eye diameter, distance between front of eyes
(negative loading) and inter-orbital distance (negative
loading), but is largely uninformative with regard to species
differences.
Bioacoustics analysis. – Bioacoustic data were limited in
that we were able to obtain good-quality recordings from
multiple males only from the P. viridis (n=4) / P. stuarti (n=3),
and P. hoffmanni (n=3) / P. asankai (n=3) species pairs. For
these pairs, bioacoustics analysis showed clear differences
in call characteristics between the sister species (Table 3).
The calls of both P. hoffmanni and P. asankai comprised a
series of short whistles. These calls differed in call length,
pulse rate, dominant frequency, fundamental frequency, pulse
length, and the number of pulses per call (Fig. 12). Philautus
hoffmanni has a greater call length than P. asankai (0.0638 –
0.692 s vs. 0.385 – 0.409 s)
, greater pulse rate (12.52 – 12.60 s-1,
vs.11.62 – 11.68 s-1), lower fundamental and dominant
frequency (2,386 – 2,399 Hz, vs. 2,859 – 2,870 Hz), shorter
pulse length (0.048 – 0.051 s, vs. 0.069 – 0.071 s) and a greater
number of pulses per call (9 pulses, vs. 3–4 pulses). For these

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Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
two species, there was no frequency modulation within the
call or a given pulse and the fundamental frequency is also
the dominant frequency (Fig. 12). Temperature and relative
humidity at which the calls were recorded were 16° C and 85%
for P. asankai and 19° C and 87% for P. hoffmanni.
The calls of both P. stuarti and P. viridis comprised a series of
terse ticks. These calls differed in call length, pulse rate,
dominant frequency, fundamental frequency, pulse length,
and the number of pulses per call (Fig. 33). Philautus stuarti
has a greater call length than P. viridis (1.713 – 1.805 s vs.
1.018 – 1.508 s), greater pulse rate (3.00 – 3.09 s-1, vs. 2.09 –
2.12 s-1), greater fundamental and dominant frequency (2,520
– 2,539 Hz, vs. 2,315 – 2,328 Hz), shorter pulse length (0.051 –
0.059 s, vs. 0.077 – 0.080 se and a greater number of pulses per
call (6 pulses vs. 3–4 pulses). For these two species, there
was no frequency modulation within the call or a given pulse
and the fundamental frequency is also the dominant frequency
(Fig. 33). Temperature and relative humidity at which the calls
were recorded were 16° C and 90% for P. viridis and 18° C,
87% for P. stuarti.
Call characteristics are affected by temperature and body size
(Ryan, 1985; Duellman & Trueb, 1986; Sullivan, 1992).
However, Sullivan (1992) has shown that the characteristic
frequency of a call and call rate do not change significantly
over a 10° C temperature range. The calls analyzed here were
recorded at temperatures within 3° C of each other and at
similar levels of relative humidity. We therefore discount the
possibility that the distinctive vocalizations of these species
are the result of differences in temperature.
TAXONOMY
Philautus mooreorum, new species
(Figs. 3, 4)
Material examined. – Holotype - male, 30.3 mm SVL, WHT 5862,
Hunnasgiriya (Knuckles), elevation 1,100 m (07º23’ N, 80º41’ E),
coll. 17 Oct.2003.
Paratypes - females, 35.0 mm SVL, WHT 2477, Corbett’s Gap
(Knuckles), 1,245 m (07º 22’ N, 80º 51’ E) coll. 6 Jun.1999; 33.8
mm SVL, WHT 6124, Corbett’s Gap (Knuckles), 1,245 m (07º 22’
N, 80º 51’ E) coll. 16 Jun.2004; males, 30.3 mm SVL, WHT 5868,
Hunnasgiriya, same data as holotype, coll. 16 Oct.2003; 31.3 mm
SVL, WHT 3209, Corbett’s Gap (Knuckles), coll. 29 Jun.2001;
29.4 mm SVL, WHT 5869, same data as holotype.
Diagnosis. – (Figs. 4, 5). Philautus mooreorum is distinguished
from all other Sri Lankan congeners by a combination of the
following characters: mature males 29.4–31.3 mm SVL; tympanic
membrane absent; snout-angle category 7–9; canthal edges
rounded; supratympanic fold, lingual papilla, vomerine teeth,
supernumerary tubercles and calcar absent; limbs dorsally
shagreened; toes medially webbed; male without nuptial pads;
dorsally ‘luminous green’ in life.
Description. – (Figs. 3–5). Mature males 29.4–31.3 mm SVL;
mature female 33.8–35.0 mm SVL. Body stout. Head dorsally
flat. Snout-angle category 7–9 (angle of snout 108º–115º);
snout rounded in lateral aspect. Canthal edges rounded. Loreal
region concave. Interorbital space flat. Internasal space
concave. Tympanic membrane absent, tympanic rim absent.
Supratympanic fold indistinct or absent. Pineal ocellus,
vomerine ridge, cephalic ridges, calcar, lingual papilla and co-
ossified skin on skull absent. A lateral dermal fringe present
on fingers. Toes webbed. Tarsal folds absent. Snout,
interorbital space, side of head and dorsum, and upper part of
flank with horny spinules in males. Lower flank smooth. Dorsal
part of forelimb, thigh, shank and pes shagreened. Throat,
chest and belly granular; underside of thigh granular. Inner
vocal slits present in males, nuptial pad absent (but yellow
subdermal glands present on finger 1). Dorsum finely granular
or shagreened in female.
Table 3. Bioacoustics characters for two pairs of sister taxa: P. stuarti – P. viridis and P. asankai – P. hoffmanni.
character P. stuarti P. viridis P. asankai P. hoffmanni
(n=3) (n=4) (n=2) (n=3)
Call length (s) 1.713 – 1.805 1.018 – 1.508 0.385 – 0.409 0.638 – 0.692
Pulse rate (s-1) 3.00 – 3.09 2.09 – 2.12 11.62 – 11.68 12.52 – 12.60
Dominant frequency (Hz) 2,520 – 2,539 2,315 – 2,328 2,859 – 2,870 2,386 – 2,399
Fundamental frequency (Hz) 2,520 – 2,539 2,315 – 2,328 2,859 – 2,870 2,386 – 2,399
Pulse length (s) 0.051 – 0.059 0.077 – 0.080 0.069 – 0.071 0.048 – 0.051
Frequency modulation within call No No No No
Frequency modulation within pulse No No No No
Number of pulses per call 6 3 – 4 3 – 4 9
Fig. 3. Philautus mooreorum, n. sp., WHT 5862, holotype male,
30.3 mm SVL.

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Fig. 4. Philautus mooreorum, n. sp.: a–c, lateral, dorsal and ventral aspects respectively, of head; d, ventral aspect of left hand; e, ventral
aspect of left foot; and f, semi-diagrammatic representation of the left-foot webbing-pattern of holotype, male, WHT5862, 30.3mm SVL.
Scale bar: 1 mm.
a
bc
de f

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Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
Colour in life. – (Fig. 3). Head dorsally and laterally bright
‘luminous’ green. Upper flank yellow and white, lower flank
white. Margins of both lips white. Limbs dorsally green. Outer
edge of lower arm and pes white. Dorsal area of upper arm
white. Fingers and toes dorsally white or pale green. Disks
white. Venter white.
Colour in alcohol. – Dorsally and laterally brownish dark
pink, with or without white pigments. Lower flank, venter,
upper lip and inguinal zone yellow.
Measurements of holotype. – (WHT 5862, in mm) DBE, 11.4;
DFE, 6.4; DL, 1.2; DW, 1.5 ED, 4.2; EN, 3.5; ES, 5.1; FEL, 16.0;
FL I, 2.4; FL II, 3.1; FL III, 5.0; FL IV, 4.1; FOL, 20.4; HL, 12.3;
HW, 13.6; IML, 1.1; IN, 2.6; IO, 3.8; LAL, 6.7; MBE, 4.5; MFE,
8.4; MN, 11.2; NS, 1.5; PAL, 9.1; SVL, 30.3; TBL, 15.5; TL I,
2.4; TL II, 2.6; TL III, 4.7; TL IV, 6.7, TL V, 5.3; TYD, – (tympanum
indistinct); TYE, – (tympanum indistinct); UAW, 6.4; UEW,
2.8.
Distribution. – (Fig. 6) Philautus mooreorum is currently
known from the type locality, Corbett’s Gap, and from
Hunnasgiriya, both in the Knuckles hills.
Etymology. – The species epithet is an eponym in the Latin
genitive plural honouring the benefactors of the Moore
Foundation, Dr. Gordon and Betty Moore (b. California, 1929
and 1928, respectively), in appreciation of their support of
the Global Amphibian Assessment and decades of
philanthropic work in science and conservation.
Remarks. – The P. femoralis group comprises three species:
P. femoralis (Günther, 1864), P. mooreorum and P. poppiae
(Figs. 2, 5), which are separated from each other by a 12S and
16S sequence divergence of 2.56–3.15% and cytochrome-b
sequence divergence of 7.7–8.9% (Tables 1, 2).
Philautus mooreorum keys out as P. femoralis according to
the morphological key of Manamendra-Arachchi &
Pethiyagoda (2005). It may be distinguished from P. femoralis
however, by having the limbs dorsally shagreened (vs. limbs
dorsally smooth in P. femoralis), and its large size (adult male
snout-vent length to 31.3 mm, vs. 27.5 mm in P. femoralis).
Philautus mooreorum is further distinguished from P.
poppiae, by having the head dorsally flat (vs. head dorsally
convex in P. poppiae); limbs dorsally shagreened (vs. limbs
dorsally horny and spinulated); no black dots on dorsum (vs.
black dots present on dorsum); and snout-angle category of
7–9 (vs. snout-angle category of 6 or 7). We note that
unusually for Sri Lankan Philautus, P. mooreorum shows
significant variation in snout-angle, ranging from 108°–115°.
Principal components analysis of metric data (Fig. 5a) shows
clear separation of the three species in body size (snout–
vent length). Philautus mooreorum may further be
Fig. 5. A, PC1 vs. PC2 factor scores of the principal components analysis of male Philautus mooreorum (Knuckles hills, K), P. femoralis
(Central ills, C) and P. poppiae (Rakwana hills, R). Most of the variation is explained by body size, manus and toe dimensions. All three
species separate well on the PC1 axis in relation to body size (P. mooreorum is the largest and P. poppiae is the smallest). Philautus
mooreorum separates out well on the PC2 axis in having palms, fingers and toes relatively larger in comparison to the other two species.
Philautus poppiae and P. mooreorum almost completely overlap on the PC2 axis. B, PC1 vs. PC2 factor scores of the principal components
analysis of female Philautus mooreorum (Knuckles hills, K), P. femoralis (Central hills, C) and P. poppiae (Rakwana hills, R) and the
holotype of P. femoralis BMNH1947.2.26.89 (BM). Most of the total variation is explained by the PC1 axis, which relates mostly body size
(P. mooreorum is the largest and P. poppiae the smallest; P. femoralis and the BMNH holotype overlap in size). Philautus femoralis and its
holotype separate out from the other two species on the PC2 axis, which relates mostly to manus dimensions; the type and the central hills
population do not, however, overlap in this respect. The males are represented by circles, the females by squares.
AB

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distinguished from P. femoralis by having relatively smaller
palm, fingers and toes.
Philautus mooreorum is the largest of the species in the P.
femoralis group, with a male snout–vent length range of 29.4
–31.3 mm; and P. poppiae the smallest (male snout–vent
length 21.3–24.7 mm). Philautus femoralis is intermediate in
size, with a male snout–vent length range of 26.3–28.8 mm.
The holotype of the P. femoralis (BMNH 1947.2.26.89) is a
female. In principal components analysis this specimen clearly
separates from the other members of the group (Fig. 5b).
Though it may be distinguished from the other species by its
relatively smaller manus dimensions, it overlaps with P.
femoralis in SVL. Until further evidence is available, we follow
Manamendra-Arachchi & Pethiyagoda (2005) in considering
the Central Hills population as conspecific with P. femoralis
sensu stricto.
Though P. mooreorum has been recorded from elevations of
1,100–1,245 m, we expect it to be present on the higher peaks
of the Knuckles mountain range where suitable cloud forest
habitat persists. Philautus femoralis occurs only in the higher
parts (above 1,000 m) of the Central Hills, whereas P. poppiae
is restricted to the Rakwana Hills, at elevations between 1,060
and 1,270 m. These species are isolated from one other by the
deep valleys that separate these mountain ranges.
Specialised to the forest sub-canopy and shrubs in the
understorey of closed-canopy cloud forest (though occurring
also in areas under-planted with cardamom), the three species
of the P. femoralis group occupy similar microhabitats. Males
vocalize 1–3 m above ground, perched on leaves (the species
is strictly arboreal). We noted that the density of all three
species is greater in somewhat marshy habitats, possibly
because these frogs depend for reproduction on environments
with high relative humidity (Bahir et al., 2005).
Conservation status. – (Fig. 5). The limited Extent of
Occurrence (~10 km2, in the Corbett’s Gap and Hunnasgiriya
region of the Knuckles mountain range), suggests that the
species should be considered Endangered (criteria B1 a, b(iii)).
Restriction to a single forest site, low abundance and strictly
arboreal behaviour renders these frogs susceptible to habitat
modification (forest clearing) and stress during periods of
drought, with chemicals used in cardamom agriculture also
posing a potential threat. We advocate periodic monitoring
to assess population trends and address possible negative
impacts.
Philautus poppiae, new species
(Figs. 7, 8)
Material examined. – Holotype - male, 24.7 mm SVL, WHT 3285,
Handapan Ella Plains (near Suriyakanda), elevation 1,270 m
(06º26’42"N, 80º36’35"E), coll. 5 Jul.2001.
Paratypes - males, 22.7 mm SVL, WHT 2030, coll. 5 Aug.1997;
21.3 mm SVL, WHT 2029, coll. 5 Aug.1997; 23.9 mm SVL, WHT
2475, coll. 29 May.1999; 24.0 mm SVL, WHT 2778, coll. 24
Jul.1999; 23.0 mm SVL, WHT 2781, coll. 24 Jul.1999; 22.5 mm
SVL, WHT 3533; 22.5 mm SVL, WHT 3534; 24.1 mm SVL, WHT
3535; 24.3 mm SVL, WHT 3536, coll. 14 Jan.1999, Morningside
(near Rakwana), elevation 1060 m (06º24’ N, 80º38’ E).
Others - female, 26.0 mm SVL, WHT 3543, Morningside (near
Rakwana), elevation 1060 m (06º24’ N, 80º38’ E), coll. 22 Feb.1996.
Juvenile, 18.6 mm SVL, WHT 3274, Morningside (near Rakwana),
elevation 1060 m (06º24’ N, 80º38’ E), coll. 14 Jul.2001.
Diagnosis. – (Fig. 8). Philautus poppiae is distinguished from
all other Sri Lankan congeners by a combination of the
Fig. 6. Distribution of Philautus mooreorum, n. sp., in Sri Lanka. Fig. 7. Philautus poppiae, n. sp., WHT 3285, holotype male, 24.7
mm SVL.

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Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
Fig. 8. Philautus poppiae, n. sp.: a–c, lateral, dorsal and ventral aspects respectively, of head; d, ventral aspect of left manus; e, ventral aspect
of left pes; and f, semi-diagrammatic representation of the left-pes webbing-pattern of holotype, male, WHT3285, 24.7 mm SVL. Scale bar:
1 mm.
a
bc
de f

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following characters: mature males 21.3–24.7 mm SVL;
tympanum membrane absent; lower tympanic rim distinct;
snout-angle category 6 or 7; dorsal surface of body with horn-
like spinules in males; canthal edges rounded; supratympanic
fold present; lingual papilla, vomerine teeth, supernumerary
tubercles and calcar absent; toes medially webbed; male
without nuptial pads; belly granular; dorsally ‘luminous green’
and dorsum with black dots in life.
Description. – (Figs. 6, 7). Mature males 21.3–24.7 mm SVL;
mature female 26.0 mm SVL. Body stout. Head dorsally convex.
Snout-angle category 6 or 7 (angle of snout 100º–108º); snout
rounded in lateral aspect. Canthal edges rounded. Loreal
region concave. Interorbital space flat. Internasal space
concave. Tympanic membrane absent, lower tympanic rim
distinct. Supratympanic fold present. Pineal ocellus, vomerine
ridge, cephalic ridges, calcar, lingual papilla and co-ossified
skin on skull absent. A lateral dermal fringe present on fingers.
Toes webbed. Tarsal folds absent. Snout, interorbital space,
side of head and dorsum, upper part of flank, dorsal area of
forelimb, thigh, shank and pes with horny spinules in males.
Lower flank smooth. Throat, chest and belly granular;
underside of thigh smooth. Inner vocal slits present in males;
nuptial pad absent, but yellow subdermal glands present on
finger 1. Dorsum finely granular or shagreened in female.
Colour in life. – (Fig. 7). dorsal and lateral parts of head and
dorsum bright ‘luminous’ green; some specimens with yellow
or red spots on dorsum. Dorsum spotted with black. Flank
yellow. Inguinal zone and anterior thigh brownish-yellow.
Edges of both lower and upper lips yellow or white. Upper
arm yellow or greenish-yellow dorsally. Outer edge of lower
arm with a longitudinal white band. Inner side of both upper
and lower arms yellow. Outer edges of shank, pes and toe 5
white. Pes ventrally white. Venter pale yellow.
Colour in alcohol. – Dorsum and flanks ashy pink with
whitish yellow (rarely with red) patches and scattered black
dots. Lower flank yellow. Upper lip yellow. Limbs dorsally
ashy pink with scattered black dots. Posterior thigh yellow.
Venter pale yellow.
Measurements of holotype. – (WHT 3285, in mm) DBE, 9.2;
DFE, 5.0; DL, 1.1; DW, 1.5 ED, 3.6; EN, 3.0; ES, 4.2; FEL, 10.9;
FL I, 2.1; FL II, 2.5; FL III, 4.5; FL IV, 4.3; FOL, 16.6; HL, 10.3;
HW, 10.1; IML, 1.1; IN, 2.1; IO, 3.1; LAL, 5.4; MBE, 3.7; MFE,
6.5; MN, 9.2; NS, 1.2; PAL, 7.7; SVL, 24.7; TBL, 12.2; TL I, 2.2;
TL II, 2.6; TL III, 3.9; TL IV, 5.7, TL V, 4.0; TYD, – (tympanum
indistinct); TYE, – (tympanum indistinct); UAW, 4.4; UEW,
2.2.
Distribution. – (Fig. 9). Philautus poppiae shows a very
restricted distribution, being known only from the type locality
(Handapan Ella, elevation 1,270 m; 06º26’42"N, 80º36’35"E)
and Morningside Forest, near Rakwana (elevation 1,060 m;
06º24’ N, 80º38’ E, about 10 km from the type locality).
Etymology. – The species epithet is an eponym, Latinized in
the feminine genitive singular, for Poppy Valentina Meyer (b.
27 Nov.2003), in honor of her parents, George A. Meyer (b.
Pennsylvania, 1956) and Maria Semple (b. California, 1964),
for their support of the Global Amphibian Assessment and
ongoing commitment to amphibian conservation around the
world.
Remarks. – According to the key of Manamendra-
Arachchi & Pethiyagoda (2005), P. poppiae keys out as P.
femoralis (Günther, 1864). Philautus poppiae may be
distinguished from P. femoralis by having dorsal surface
of head flat (vs. dorsal surface of head convex in P.
femoralis); limbs dorsally horny, spinulated (vs. limbs
dorsally smooth); black dots present on dorsum (vs. black
dots absent on dorsum); and snout-angle category of 6 or
7 (vs. snout-angle category 8). It may be distinguished
from P. mooreorum by having the head dorsally convex
(vs. head dorsally flat in P. mooreorum); limbs dorsally
horny and spinulated (vs. limbs dorsally shagreened);
black dots present on dorsum (vs. black dots absent on
dorsum); snout angle category 6 or 7 (vs. snout angle
category of 7–9).
Philautus poppiae is the smallest of the three species in the
P. femoralis group, with a male snout–vent length up to 24.7
mm (see also Remarks in account of P. mooreorum). While
the manus and pes dimensions of P. poppiae are similar to
those of P. mooreorum, they are distinctly less than those of
P. femoralis (Fig. 5).
Philautus poppiae occurs only in the montane cloud forests
in the Rakwana mountains, above 1,060 m elevation, while P.
mooreorum and P. femoralis are restricted to the cloud forests
of the Knuckles and Central Hills, respectively. Only a very
small part of the Rakwana Hills exceed an elevation of 1,000
m, from which we conclude that the area of occupancy of P.
poppiae is extremely small, even if it were to occur on peaks
as yet unexplored.
Fig. 9. Distribution of Philautus poppiae, n. sp., in Sri Lanka.

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Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
Conservation status. – (Fig. 9). Extent of Occurrence, c. 10
km2: Morningside and Handapan Ella region of the Rakwana
Hills. Outcome: Endangered (criteria B1 a, b(iii)). Restriction
to a single forest site, low abundance and strictly arboreal
behaviour renders these frogs susceptible to habitat
modification (forest clearing) and stress during periods of
drought, with chemicals used in cardamom agriculture also
posing a potential threat. We advocate periodic monitoring
of this population to assess trends and address negative
impacts.
Philautus hoffmanni, new species
(Figs. 10, 11)
Material examined. – Holotype - male, 23.4 mm SVL, WHT 6120,
Corbett’s Gap (Knuckles Hills), elevation 1,245 m (07º22’ N, 80º51’
E), coll. 15 Jan.2004.
Paratypes - males, 21.2 mm SVL, WHT 3542; 21.7 mm SVL, WHT
3223; 23.1 mm SVL, WHT 3222, coll. 29 Jun.2001, all from type
locality.
Diagnosis. – (Fig. 11). Philautus hoffmanni is distinguished
from all other Sri Lankan congeners by a combination of the
following characters: mature males 21.2–23.4 mm SVL;
tympanum present, distinct; snout-angle category 6 or 7;
snout rounded in lateral aspect; interorbital space flat; fine,
horny spinules absent on lower flank; canthal edges rounded;
vomerine ridge and lingual papilla absent; nuptial pads absent
in males.
Description. – (Figs. 10, 11). Mature males 21.2–23.4 mm SVL.
Body stout. Head dorsally flat. Snout-angle category 6 or 7
(angle of snout 103º–107º); snout rounded in lateral aspect.
Canthal edges rounded. Loreal region flat. Interorbital space
flat. Internasal space flat. Tympanum distinct. Supratympanic
fold indistinct. Pineal ocellus, vomerine ridge, cephalic ridges,
lingual papilla and co-ossified skin on skull absent. A lateral
dermal fringe present on fingers. Toes webbed. Tarsal folds
and calcar absent. Snout, interorbital space, side of head,
dorsum and upper flank smooth; lower flank glandular. Dorsal
part of forelimb, thigh, shank and pes smooth. Throat, chest
and belly granular; under thigh smooth. Inner vocal slits
present in males and nuptial pad absent.
Colour in life. – (Fig. 10) (based on WHT 3222). Dorsally ash
and brown. Loreal region brown. Both upper and lower lips
ash. Limbs dorsally ashy brown. Digits dorsally bright yellow
and ash. Flank ash. Inguinal zone, posterior and anterior thigh
ash with golden-yellow dots. Ventral aspect of digits, limbs
and body golden yellow.
Colour in alcohol. – Dorsal parts of head and body greyish-
brown with brown dots. Upper flank brown and lower flank
yellow. Inguinal zone brown. Loreal region greyish-brown,
tympanic region and tympanum brown. Limbs dorsally greyish
brown. Venter yellow.
Measurements of holotype. – (WHT 6120, in mm), DBE, 8.5;
DFE, 4.8; DL, 1.1; DW, 1.5; ED, 3.0; EN, 2.5; ES, 4.0; FEL, 10.5;
FL I, 1.8; FL II, 2.4; FL III, 3.8; FL IV, 3.1; FOL, 14.7; HL, 9.5;
HW, 9.7; IML, 1.1; IN, 2.2; IO, 2.8; LAL, 4.4; MBE, 3.1; MFE,
5.7; MN, 8.2; NS, 1.6; PAL, 7.2; SVL, 23.4; TBL, 11.2; TL I, 1.9;
TL II, 2.1; TL III, 3.5; TL IV, 5.4, TL V, 3.9; TYD, – outer rim of
tympanum indistinct; TYE, –outer rim of tympanum indistinct;
UAW, 3.6; UEW, 2.1.
Distribution. – (Fig. 13). The species was recorded only from
Corbett’s Gap (Knuckles hills), elevation 1,245 m (07º22’ N,
80º51’ E).
Etymology. – The species name is an eponym in the Latin
genitive singular honouring Luc Hoffmann (b. Switzerland,
1923), Honorary President of World Wildlife Fund – France
and Director Emeritus of Wetlands International who, as
benefactor to the MAVA Foundation, generously supported
the Global Amphibian Assessment.
Remarks. –Philautus hoffmanni is the sister species of P.
asankai Manamendra-Arachchi & Pethiyagoda, 2005 (Fig.
2). These two species are separated from each other by a 12S
and 16S sequence divergence of 1.01% and a cytochrome-b
divergence of 6.04% (Tables 1, 2). Philautus hoffmanni keys
out as P. asankai according to the key of Manamendra-
Arachchi & Pethiyagoda, 2005, but can be distinguished from
that species by having the interorbital space flat (vs.
Fig. 10. Philautus hoffmanni, n. sp., a, WHT 6120, holotype male,
23.4 mm SVL; b, WHT 3542, paratype male, 21.2 mm SVL; WHT
3222, paratype male, 23.1 mm SVL.
a
b

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Fig. 11. Philautus hoffmanni, n. sp.: a–c: lateral, dorsal and ventral aspects respectively, of head; d, ventral aspect of left manus; e, ventral
aspect of left pes; and f, semi-diagrammatic representation of the left-pes webbing-pattern of holotype, male, WHT6120, 23.4 mm SVL.
Scale bar: 1 mm.
a
bc
de
f

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Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
interorbital space convex); tympanum distinct (vs. tympanum
indistinct); fine, horn-like spinules absent on lower flank (vs.
fine, horn-like spinules present on lower flank).
The vocalizations (Table 3; Fig 12) of Philautus hoffmanni
and P. asankai consist of a series of short whistles. The
advertisement call of P. asankai may be distinguished from
that of P. hoffmanni by the shorter call length (0.385 – 0.409 s,
vs. 0.638 – 0.692 s), lower pulse rate (11.62 – 11.68 s-1, vs.
12.52 – 12.6.60 s-1), higher dominant and fundamental
frequencies (2,859 – 2,870 Hz, vs. 2,396 – 2,399 Hz), greater
pulse length (0.069 – 0.071 s, vs. 0.048 – 0.051 s) and the
smaller number of pulses per call (3 or 4, vs. 9).
Both species are restricted to montane regions: P. asankai
occurs only in the highest peaks (1,300–1,900 m elevation) of
the Central Hills, while P. hoffmanni is restricted to the type
locality, Corbett’s Gap, elevation 1,245 m (07º22’ N, 80º51’ E).
Future sampling may show that P. hoffmanni occurs also in
other cloud-forest areas of the Knuckles Hills.
Philautus hoffmanni and P. asankai utilize similar
microhabitats, inhabiting shrubs in gaps within closed-canopy
cloud forests, and cardamom plantations within these. The
ability to live in gaps has enabled P. asankai to persist in
anthropogenic habitats. Philautus hoffmanni, however, was
observed only in close proximity to relatively undisturbed
montane forests. Males of both species vocalize while perched
on leaves ~0.3–1 m above ground. The diurnal resting habitat
of these frogs is under a leaf, or on a leaf axil, often in well-
illuminated habitats.
Conservation status. – (Fig. 13). The limited Extent of
Occurrence (~10 km2 in the Corbett’s Gap region of the
Knuckles mountain range) suggests that the species should
be considered Endangered (criteria B1 a, b(iii)). Its restriction
to a single forest site, low abundance, and dependence on a
closed canopy forest, makes this frog susceptible to habitat
modification (clearing) and stress during periods of drought.
Continuous population monitoring is recommended.
Philautus mittermeieri, new species
(Figs. 14–15)
Material examined. – Holotype - male, 18.4 mm SVL, WHT 3522,
Kottawa (Galle), elevation 60 m (06º’06’ N, 80º20’ E), coll. 22 May
2004.
Paratypes - males, 17.8 mm SVL, WHT 2668, Beraliya forest
(Elpitiya), elevation 150 m (06º16’ N, 80º11’ E), coll. 14 Sep.1999;
16.3 mm SVL, WHT 3523; 17.7 mm SVL, WHT 3524; 17.7 mm
SVL, WHT 3525; 17.2 mm SVL, WHT 3526, all from type locality.
Diagnosis. – (Figs. 14, 15). Philautus mittermeieri is
distinguished from all other Sri Lankan congeners by a
a
b
Fig. 13. Distribution of Philautus hoffmanni, n. sp., in Sri Lanka.
Fig. 12. Uncalibrated waveform envelope and spectrogram of the
whistle-like advertisement calls of a, Philautus asankai and b, P.
hoffmanni. The advertisement call of P. asankai may be distinguished
from that of P. hoffmanni by the shorter call length (0.385–0.409 s,
vs. 0.638–0.692 s), lower pulse rate (11.62–11.68 s-1, vs. 12.52 –
12.60 s-1), higher dominant and fundamental frequencies (2,859–
2,870 Hz, vs. 2,396–2,399 Hz), greater pulse length (0.069–0.071
s, vs. 0.048–0.051 s) and smaller number of pulses per call (3–4, vs.
9). There is no frequency modulation within a call or within a
pulse.

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combination of the following characters: mature males 16.3–
18.4 mm SVL; tympanum distinct; snout-angle category 4 or
5; snout pointed in lateral aspect; canthal edges rounded;
vomerine ridge absent’ tarsal folds and calcar present; dorsal
surface of forelimb, thigh, shank and pes smooth; a prominent
sheath-like undulating fringe present on posterior margin of
pes and lower arm; nuptial pads absent in male.
Description. – (Figs 13, 14). Mature males 16.3–18.4 mm SVL.
Body slender. Head dorsally convex. Snout-angle category 4
or 5 (angle of snout 90º–95º); snout pointed in lateral aspect.
Canthal edges rounded. Loreal region concave. Interorbital
space concave. Internasal space concave. Tympanum distinct,
vertically orientated. Supratympanic fold distinct. Pineal
ocellus, vomerine ridge, cephalic ridges, lingual papilla and
co-ossified skin on skull absent. A lateral dermal fringe present
on fingers. Toes webbed. Tarsal folds and calcar present.
Snout, interorbital space, side of head and dorsum, both upper
and lower flank with glandular warts. Dorsal part of forelimb,
thigh, shank and pes smooth. Throat, chest, belly granular
and underside of thigh granular. Inner vocal slits present in
males; nuptial pad absent.
Colour in life. – (Fig. 14). Mid-dorsum dark ashy-olive.
Tubercles on head orange. Posterior part of back with orange
pigments. Side of head ashy brown. Supratympanic fold
orangish-light brown. Lower lip ash. Inguinal area dark brown.
Flank ashy-yellow with dark-brown patches. Anterior thigh
with dark-brown cross markings. Posterior thigh and tibia
dark brown. Throat, chest, and upper abdomen ash; lower
abdomen ashy-yellow. Fingers ventrally ash. Lower arm with
4, thigh with 3 and shank with 3 dark, ashy-brown crossbars.
Ventral area of thigh, shank, pes and toes dark brown. Venter
yellow.
Colour in alcohol. – Dorsal parts of head and body grey
with brown patches. Both upper and lower flanks grey with
brown pigments. Upper lip and loreal region grey. Inguinal
zone dark brown. Tympanic region and tympanum grey with
brown pigments. Limbs dorsally grey, with indistinct
crossbars; posterior part of thigh brown; area around vent
yellow. Venter pale yellow.
Measurements of holotype. – (WHT 3522, in mm) DBE, 7.1;
DFE, 4.6; DL, 0.8; DW, 1.1; ED, 3.3; EN, 2.5; ES, 3.7; FEL, 8.6;
FL I, 1.3; FL II, 1.6; FL III, 2.6; FL IV, 2.1; FOL, 11.7; HL, 8.3;
HW, 7.8; IML, 0.9; IN, 2.0; IO, 2.2; LAL, 3.6; MBE, 2.9; MFE,
5.1; MN, 7.3; NS, 1.3; PAL, 5.1; SVL, 18.4; TBL, 9.5; TL I, 1.1;
TL II, 1.6; TL III, 2.6; TL IV, 4.1, TL V, 2.5; TYD, 0.5; TYE, 1.1;
UAW, 3.4; UEW, 2.1.
Distribution. – (Fig. 17) Philautus mittermeieri is a lowland
(60–150 m) species that has been observed in two widely
spaced localities, Kottawa (60 m elevation, 06º06’ N, 80º20’ E);
and Beraliya Forest, Elpitiya (150 m elevation, 06º16’ N, 80º11’
E).
Etymology. – The species name is a patronym in the Latin
genitive singular, in honour of Russell Mittermeier (b. New
York, 1949), President of Conservation International, for his
special commitment to the Global Amphibian Assessment and
for continuing support of amphibian conservation worldwide.
Dr. Mittermeier’s lifetime focus on protection of threatened
species has resulted in effective conservation programmes in
many parts of the world.
Remarks. – The sister species of P. mittermeieri, is P. decoris
Manamendra-Arachchi & Pethiyagoda, 2005 (Fig. 2). The two
species are separated from each other by a 12S and 16S
divergence of 2.38% and a cytochrome-b sequence divergence
of 5.80% (Tables 1, 2).
Philautus mittermeieri keys out as P. decoris according to
the morphological key of Manamendra-Arachchi &
Pethiyagoda (2005). It may, however, be distinguished from P.
decoris by having the snout obtusely pointed in lateral aspect
(vs. snout sharply pointed); glandular warts present on dorsal
surface of forelimb, thigh, shank and pes (vs. dorsal surface
of forelimb, thigh, shank and pes smooth); and a prominent
sheath-like undulating fringe present on posterior margin of
pes and lower arm (vs. a feebly-defined sheath-like undulating
fringe present on posterior margin of pes and lower arm).
Principal components analysis shows clear separation of the
two species by size, with P. mittermeieri being the smaller
species (up to 18.4 mm SVL in P. mittermeieri, vs. up to 20.7
mm in P. decoris; Fig. 16).
Philautus decoris occurs only on the highest peaks (above
1,000 m elevation) of the Rakwana Hills (Morningside Forest),
P. mittermeieri being a lowland species (60–150 m elevation).
The two species are thus isolated from each other also by
altitudinal constraints.
Philautus mittermeieri is a habitat specialist inhabiting the
shrub understorey of closed-canopy lowland rainforests. At
night, males vocalize in chorus from leaves, up to ~ 1 m above
ground. The calling site of P. mittermeieri is similar to that of
P. decoris, and calling behaviour similar in the two species in
that they are group callers, where several males call in chorus.
Fig. 14. Philautus mittermeieri, n. sp., WHT 3522, holotype male,
18.4 mm SVL.

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Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
Fig. 15. Philautus mittermeieri, n. sp.: a–c, lateral, dorsal and ventral aspects respectively, of head; d, ventral aspect of left manus; e, ventral
aspect of left pes; and f, semi-diagrammatic representation of the left-pes webbing-pattern of holotype, male, WHT3522, 18.4 mm SVL.
Scale bar: 1 mm.
a
bc
de f

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Philautus frankenbergi, new species
(Figs. 18, 19)
Material examined. – Holotype - male, 29.3 mm SVL, WHT 2554,
Namunukula Peak, elevation 1,850–1,980 m (06º56’ N, 81º07’ E),
coll. 2 Sep.1999.
Paratypes - males, 27.4 mm SVL, WHT 2726; 26.7 mm SVL, WHT
2727; 29.2 mm SVL, WHT 2728; Horton Plains National Park,
elevation 2135 m (06º46’ N, 81º07’ E), coll. 20 Sep.1999. Males,
27.4 mm SVL, WHT 2551; 28.7 mm SVL, WHT 2552; 28.7 mm
SVL, WHT 2555; 27.5 mm SVL, WHT 2556 (all from type locality).
Diagnosis. – (Fig. 19). Philautus frankenbergi is
distinguished from all other Sri Lankan congeners by a
combination of the following characters: mature males 26.7–
29.3 mm SVL; snout-angle category 5–7 (97º–106º); vomerine
ridge absent; lateral dermal fringe present on fingers; canthal
edges sharp; horny spinules on dorsum; limbs with distinct
cross-bars; supernumerary tubercles absent on pes; nuptial
pad absent in males.
Description. – (Figs. 18, 19). Mature males 26.7–29.3 mm SVL.
Body stout. Head dorsally flat. Snout-angle category 5–7
(angle of snout 97º–106º), rounded in lateral aspect. Canthal
edges sharp. Loreal region concave. Interorbital and internasal
spaces flat. Tympanum distinct, oval, vertically orientated.
Vomerine ridge absent. Pineal ocellus, lingual papilla, cephalic
ridges, tarsal tubercle, tarsal fold and co-ossified skin on skull
absent. Supratympanic fold distinct. Lateral dermal fringe
present on fingers. Rudimentary webbing present on fingers.
Supernumerary tubercles present or absent on palm and
absent on pes. Toes webbed. Snout, interorbital space,
dorsum and upper flank shagreened; side of head with
glandular warts; lower flank granular. Dorsal part of forelimb
and shank with glandular warts; dorsal parts of thigh and pes
smooth. Horny spinules scattered on dorsum. Throat, chest,
belly and underside of thigh granular. Inner vocal slits present
in males, nuptial pad absent but subdermal nuptial glands
present on inner surface of prepollex.
Colour in life. – (Fig. 18) (based on holotype, WHT 2554).
Dorsum uniform ashy brown. Canthal edges, loreal and
temporal regions dark brown. Lower half of tympanum light
brown. Inguinal zone marbled in black and white. Flank ashy
brown with white patches. Discs dorsally ashy yellow. Limbs
dorsally pale brown with dark-brown crossbars. Some
specimens (e.g., paratype WHT 2551) have dark-brown dorsal
markings, pale reddish-brown dots on dorsum, a dark-brown
interorbital bar, and rarely, white patches on dorsum.
Colour in alcohol. – (based on holotype, WHT 2554). Dorsally
ashy brown. Both upper and lower flanks white with dark-
brown patches. Inguinal zone with white and dark-brown
patches. Loreal region, tympanic region and tympanum ashy
brown. Upper lip ashy brown. Limbs dorsally ashy brown
with indistinct crossbars. Thigh dorsally ashy brown with
white patches and indistinct crossbars. Venter and webbing
pale yellow with brown pigments.
Fig. 16. PC1 vs. PC2 factor scores of the principal components
analysis of Philautus mittermeieri, n. sp., (wet-zone lowlands, L)
and P. decoris (Rakwana hills, R). Most of the variation is explained
by the PC1 axis, which relates mostly to body size (P. decors is the
larger species). The two species overlap on the PC2 axis, which
relates mostly to eye diameter and distance between eyes.
Fig. 17. Distribution of Philautus mittermeieri, n. sp., in Sri Lanka.
Conservation status. – (Fig. 17). Extent of Occurrence: 2,000
km2: Kottawa and Beraliya. Some parts of the range of P.
mittermeieri are subject to rapid land-use change. Outcome:
Vulnerable (criteria B1 a, b(iii)).

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Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
Measurements of holotype. – (WHT 2554, in mm) DBE, 11.2;
DFE, 6.4; DL, 1.4; DW, 1.7, ED, 4.7; EN, 3.1; ES, 5.0; FEL, 12.8;
FL I, 2.3; FL II, 3.3; FL III, 5.1; FL IV, 4.7; FOL, 19.2; HL, 12.4;
HW, 12.4; IML, 1.4; IN, 2.5; IO, 3.3; LAL, 6.0; MBE, 4.3; MFE,
8.0; MN, 10.8; NS, 2.0; PAL, 9.3; SVL, 29.3; TBL, 13.5; TL I,
2.4; TL II, 2.9; TL III, 4.8; TL IV, 7.1, TL V, 5.3; TYD, 0.6; TYE,
1.8; UAW, 6.7; UEW, 3.7.
Distribution. – (Fig. 21). Philautus frankenbergi is restricted
to the highest peaks of the Central Hills: Horton Plains
National Park (06º46’ N, 81º07’ E, elevation 2,135 m) and
Namunukula Peak (06º56’ N, 80º47’ E, alt. 1,850–1,980 m).
Etymology. – The species name honours the late Regina
Bauer Frankenberg (USA, 1908–1991), a resident of New York
City, who was passionately devoted to the welfare of animals.
Through her will, Ms. Frankenberg established a foundation
devoted exclusively to animal welfare, including the protection
of threatened species through conservation and research.
The Regina Bauer Frankenberg Foundation supported the
involvement of NatureServe in carrying out the Global
Amphibian Assessment.
Remarks. – The sister species of P. frankenbergi, is P. auratus
Manamendra-Arachchi & Pethiyagoda, 2005 (Fig. 2). These
two species are separated from each other in 12S and 16S
sequence divergence of 3.01–3.13% and a cytochrome-b
sequence divergence of 11.6% (Tables 1, 2).
Philautus frankenbergi keys out as P. auratus in the key of
Manamendra-Arachchi & Pethiyagoda (2005), from which
species it differs by having the canthal edges sharp (vs. canthal
edges rounded in P. auratus); vomerine ridge absent (vs.
vomerine ridge present); horny spinules present on dorsum
(vs. horny spinules on absent dorsum); and limbs with distinct
crossbars (vs. limbs with indistinct crossbars).
Principal components analysis reveals that the two species
separate well by size, with P. frankenbergi being the larger
species (male snout–vent length up to 29.3 mm, vs. male snout-
vent length up to only 23.3 mm in P. auratus) (Fig. 20).
The ranges of the two species are separated by a 1,450 m-
deep valley. Philautus frankenbergi occurs only on the
highest peaks (above 1,850 m elevation) of the Central Hills
(including the Namunukula peak area), whereas P. auratus
occurs only in Rakwana Hills, at elevations of about 1,000 m.
Philautus frankenbergi and P. auratus occupy similar
microhabitats in the sub-canopy of closed-canopy montane
forests and cardamom plantations within these. Males of both
species call from about 1–3 m above ground, while perched
on branches or large leaves.
Conservation status. – (Fig. 21). The limited Extent of
Occurrence of Philautus frankenbergi (~ 100 km2, in the
highest peaks of the Central Hills and Namunukula mountain)
suggests that the species should be considered Endangered
(criteria B1 a, b(iii)). Dependence on relatively undisturbed
closed-canopy cloud forest renders this frog susceptible to
Fig. 18. Philautus frankenbergi, n. sp., a, WHT 2554, holotype
male, 29.3 mm SVL; b, WHT 2726, paratype male, 27.4 mm SVL;
and c, WHT 2551, paratype male, 27.4 mm SVL.
a
b
c

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Fig. 19. Philautus frankenbergi, n. sp.: a–c, lateral, dorsal and ventral aspects respectively, of head; d, ventral aspect of right manus; e, ventral
aspect of right pes; and f, semi-diagrammatic representation of the left-pes webbing-pattern of holotype, male, WHT2554, 29.3 mm SVL.
Scale bar: 1 mm.
a
bc
de
f

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Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
Philautus hallidayi, new species
(Figs. 22, 23)
Material examined. – Holotype - female, 42.9 mm SVL, WHT
3575, Hanthana range, Kandy, elevation 510–800 m (07º15’ N,
80º34’ E), coll. 24 May 2003.
Paratypes - females, 38.8 mm SVL, WHT 3577; 36.7 mm SVL,
WHT 3576, from type locality. Male, 32.9 mm SVL, WHT 6072,
Tonacombe Estate, Namunukula, elevation 1320 m (06º52’ N, 81º07’
E), coll. 23 Apr.2004; 37.9 mm SVL, WHT 3573, type locality, 6
Sep.1999.
Diagnosis. – (Figs. 22, 23). Philautus hallidayi is
distinguished from all other Sri Lankan congeners by a
combination of the following characters: snout-angle category
5 or 6; vomerine ridge absent; lateral dermal fringe present on
fingers; supernumerary tubercles present on pes; canthal
edges rounded; toes 1–3 not fully webbed; toe 5 fully webbed
only in a single specimen; angle of vomerine ridge to body
axis ~ 70º; tuberculated fringe on posterior margin of lower
arm and pes absent; calcar absent; nuptial pad present in
males.
Description. – (Figs. 22, 23). Mature male 32.9 mm SVL,
females 36.7–42.9 mm SVL. Body stout. Head dorsally convex
or flat. Snout-angle category 5 or 6 (angle of snout 97º–103º);
snout rounded in lateral aspect. Canthal edges rounded. Loreal
region concave. Interorbital space flat. Internasal space flat
or concave. Tympanum rather distinct, rounded. Vomerine
ridge present (left ridge absent in holotype), bearing about 3
or 4 small teeth, angled at about 70º relative to body axis.
Pineal ocellus, lingual papilla, cephalic ridges, tarsal tubercle,
tarsal fold and co-ossified skin on skull absent. Supratympanic
fold distinct. Lateral dermal fringe present on fingers. Discs
on fingers and toes oval. Distinct glandular warts present on
palm (including outer edge of palm) and outer edge of lower
arm. Supernumerary tubercles present on palm, absent on
pes. Toes webbed. Distinct glandular warts present on outer
edge of pes and on tibio-tarsal articulation. Snout, interorbital
space, side of head, dorsum and upper flank with glandular
warts; lower flank granular. Snout, interorbital space, dorsum
and upper flank with horny spinules in males. Dorsal part of
forelimb, thigh, shank and pes with glandular warts. Throat
Fig. 20. PC1 vs. PC2 factor scores of the principal components
analysis of Philautus frankenbergi, n. sp. (Central hills, C) and P.
auratus (Rakwana hills, R). Most of the total variation is explained
by the PC1 axis, which relates mainly to body size (P. frankenbergi
is the larger species). These two species overlap on the PC2 axis,
which relates mostly to inter-narial distance.
-2 -1 0 1
PC1
-2
-1
0
1
2
3
PC2
C
R
Fig. 21. Distribution of Philautus frankenbergi, n. sp., in Sri Lanka.
habitat modification (land-use change) and stress during
periods of drought. Continuous population monitoring is
recommended. Fig. 22. Philautus hallidayi, n. sp., WHT 3575, holotype female,
42.9 mm SVL.

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Fig. 23. Philautus hallidayi, n. sp.: a–c, lateral, dorsal and ventral aspects respectively, of head; d, ventral aspect of left manus (arrow
indicates nuptial pad); e, ventral aspect of left pes; and f, semi-diagrammatic representation of the left-pes webbing-pattern of paratype,
male, WHT 6072, 32.9 mm SVL. Scale bar: 1 mm.
a
bc
de f

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Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
and chest smooth, belly and underside of thigh granular.
Throat and chest granular in males. Inner vocal slits and nuptial
pad present in males.
Colour in life. – (Fig. 22) (based on holotype, WHT 3575).
Dorsum and lateral head brown, with symmetrical, dark-brown
markings and white patches; both upper and lower flanks
brown with dark-brown and white patches. Inguinal zone dark
brown with white markings. Limbs dorsally brown with dark-
brown crossbars and white dots. Posterior thigh dark brown.
Venter pale brown; webbing dark brown.
Colour in alcohol. – (based on holotype, WHT 3575). Dorsally
brown with symmetrical dark-brown markings. Interorbital bar
dark brown. Both upper and lower flanks brown with dark-
brown and white patches. Inguinal zone dark brown with white
markings. Loreal region and tympanic region brown with
dark-brown patches; tympanum yellowish brown, its outer
rim brown; mid-tympanum dark brown. Upper lip brown with
white patches. Limbs dorsally brown with dark-brown
crossbars and white dots. Posterior part of thigh dark brown.
Venter pale yellowish brown; webbing dark brown.
Measurements of holotype. – (WHT 3575, in mm) DBE, 15.3;
DFE, 9.4; DL, 1.5; DW, 2.7, ED, 5.7; EN, 5.0; ES, 7.3; FEL, 24.5;
FL I, 4.4; FL II, 4.9; FL III, 7.5; FL IV, 5.6; FOL, 30.6; HL, 18.5;
HW, 18.4; IML, 1.5; IN, 3.8; IO, 3.9; LAL, 8.7; MBE, 6.3; MFE,
10.9; MN, 16.2; NS, 2.4; PAL, 13.4; SVL, 42.9; TBL, 24.4; TL I,
3.4; TL II, 4.1; TL III, 6.7; TL IV, 9.5, TL V, 7.0; TYD, 0.8; TYE,
2.4; UAW, 7.4; UEW, 3.4.
Distribution. – (Fig. 25). Philautus hallidayi is known only
from two locations in Sri Lanka’s wet-zone: Hanthana Range,
Kandy and Tonacombe Estate, Namunukula.
Etymology. – The species name, in the Latin genitive singular,
is a patronym honouring Timothy Richard Halliday (b.
England, 1945), since 1994 International Director of the IUCN/
SSC Task Force on Declining Amphibian Populations
(DAPTF), recognizing also his three decades of research on
amphibians and his exceptional commitment to advancing our
understanding of the global amphibian decline crisis.
Remarks. – The sister species of P. hallidayi, is P. cavirostris
(Günther, 1869) (Fig. 2). These two species are separated from
each other by a 12S and 16S divergence of 7.9% and a
cytochrome-b sequence divergence of 15.7% (Tables 1, 2).
Philautus hallidayi keys out as P. cavirostris (Günther, 1869)
in the key of Manamendra-Arachchi & Pethiyagoda (2005),
but may be distinguished from that species by having canthal
edges rounded (vs. canthal edges sharp); toes 1, 2, 3 and 5
not fully webbed (vs. toes 1, 2, 3 and 5 fully webbed); angle of
vomerine ridge to body axis ~ 70º (vs. angle of vomerine ridge
to body axis ~ 45º); absence of tuberculated fringe on posterior
margin of lower arm and pes (vs. presence of a tuberculated
fringe on posterior margin of lower arm and pes); and absence
of calcar on tibio-tarsal articulation (vs. presence of calcar on
tibio-tarsal articulation).
Fig. 24. PC1 vs. PC2 factor scores of the principal components
analysis of Philautus hallidayi, n. sp. (Hantana, Kandy, H) and P.
cavirostris (lowlands and mid-elevations, L). About one-third of
the variation is explained by the PC1 axis, which relates to pes and
manus dimensions; there is, however, some overlap of these
characters. Another third of the variation is explained by the PC2
axis, which relates to distance between front of eyes, eye-to-snout
distance and inter-orbital distance (these dimensions are all relatively
smaller in P. hallidayi than in P. cavirostris).
-3 -2 -1 0 1 2
PC1
-2
-1
0
1
2
3
PC2
H
L
Fig. 25. Distribution of Philautus hallidayi, n. sp., in Sri Lanka.
Principal components analysis of Philautus hallidayi and P.
shows that the two species may be distinguished by distance
between front of eyes, eye-to-snout distance and interorbital
distance (Fig. 24). Philautus hallidayi may be distinguished
by having a smaller eye-to-snout distance (up to 7.3 mm, vs.
up to 9.5 mm); smaller distance between the front of eyes (up

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
to 9.4 mm, vs. up to 12.1 mm); and smaller inter-orbital distance
(up to 4.0 mm, vs. up to 5.1 mm), compared to P. cavirostris.
Microhabitat utilization by P. hallidayi differs from that of P.
cavirostris. Philautus hallidayi is a habitat specialist usually
observed perched on large boulders within closed-canopy
habitats, often in proximity to large streams (boulders are
common in the vicinity of streams in the mid-elevations of Sri
Lanka, and this apparent correlation could in fact be a
coincidence). Philautus cavirostris, on the other hand, is a
shrub and sub-canopy species usually found 0.3–2 m above
ground, perched on branches in closed-canopy forests.
Conservation status. – (Fig. 25). Though this species has a
large range, it is only common where optimal habitat
conditions (mentioned above) are present. It may also be at
risk from water-borne pollutants (agricultural effluents).
Outcome: Vulnerable (criteria B1 a, b(iii)).
Philautus steineri, new species
(Figs. 26, 27)
Material examined. – Holotype - male, 31.2 mm SVL, WHT 3210,
Corbett’s Gap (Knuckles Hills), elevation 1245 m (07º22’ N, 80º51’
E), coll. 29 Jun.2001.
Paratypes - (all from type locality), male, 30.5 mm SVL, WHT
3519; male, 30.2 mm SVL, WHT 3521; female, 41.6 mm SVL,
WHT 3520, coll. 5 Jun.1999. Male, 30.6 mm SVL, WHT 6116,
coll. 17 Oct.2003; female, 30.4 mm SVL, WHT 3518, coll. 1998.
Diagnosis. – (Figs. 26, 27). Philautus steineri is distinguished
from all other Sri Lankan congeners by a combination of the
following characters: snout-angle category 5 or 6; vomerine
ridge present; lateral dermal fringe absent on fingers; an angle
of about 85° between vomerine ridge and body axis;
supernumerary tubercles present on both palm and on pes;
posterior margin of thigh pale brown; nuptial pad present in
males.
Description. – (Figs. 26, 27). Mature males 30.2–31.2 mm SVL;
mature female 30.4–41.6 mm SVL. Body stout. Head dorsally
flat. Snout-angle category 5 or 6 (angle of snout 98º–103º);
snout rounded in lateral aspect. Canthal edges sharp. Loreal
region concave. Interorbital space flat. Internasal space
concave. Tympanum distinct, oval, oblique. Vomerine ridge
present, bearing about 5 small teeth, angled at about 85º relative
to body axis, shorter than the distance between them. Pineal
ocellus present or absent. Lingual papilla, cephalic ridges,
calcar, and co-ossified skin on skull absent. Supratympanic
fold distinct. A lateral dermal fringe present on fingers.
Supernumerary tubercles present on both palm and pes. Toes
webbed. Tarsal folds absent. Snout, interorbital space, dorsum
and upper flank with glandular warts and horny spinules
(females lack horny spinules on dorsum); side of head smooth
or with glandular warts; lower flank granular. Dorsal part of
forelimb, thigh, shank and pes smooth with scattered glandular
warts. Throat, chest, belly and underside of thigh granular. A
feebly-defined dermal fringe mid-extends from tip of snout to
posterior dorsum. Inner vocal slits and nuptial pad present in
males.
Colour in life. – (Fig. 26) (based on WHT 3210). Mid-dorsum
brown; dorso-lateral area light green; upper flank green with
black patches, lower flank ashy green. A dark brownish-black
stripe present on upper flank. Interorbital bar dark brown.
Posterior head and mid-back brownish-green. Loreal and
tympanic regions and tympanum dark brown with light-green
stripes. Upper edge of supratympanic fold and canthal edges
light brown; lower lip white with brown bands. Chin, chest,
abdomen and limbs ventrally pale brown with dark-brown
patches. Limbs dorsally brown with dark-brown crossbars.
Digits pale-ashy brown. Posterior thigh green.
Colour in alcohol. – Mid-dorsum dark brown; flank and inguinal
zone ashy brown. Loreal region tympanic region and tympanum
dark brown. Limbs dorsally brown or ashy brown; posterior thigh
pale brown. Venter and webbing yellowish brown.
Measurements of holotype. – (WHT 3210, in mm) DBE, 12.7;
DFE, 6.8; DL, 1.1; DW, 1.9, ED, 4.7; EN, 3.3; ES, 5.4; FEL, 15.6;
FL I, 2.6; FL II, 3.3; FL III, 5.9; FL IV, 4.6; FOL, 22.5; HL, 13.4;
HW, 13.0; IML, 1.4; IN, 3.2; IO, 3.7; LAL, 7.2; MBE, 5.2; MFE,
8.5; MN, 11.7; NS, 2.3; PAL, 9.5; SVL, 31.2; TBL, 16.4; TL I,
2.2; TL II, 3.1; TL III, 5.1; TL IV, 7.7, TL V, 5.2; TYD, 2.0; TYE,
1.6; UAW, 5.0; UEW, 3.7.
Distribution. – (Fig. 29) Philautus steineri has been recorded
only from the type locality, the Corbett’s Gap region in the
Knuckles mountain range (elevation 1,245 m; 07º22’ N, 80º
51’E).
Etymology. – The species epithet, in the Latin genitive
singular, honours Achim Steiner (b. Brazil, 1961), Director
General (2001–) of IUCN, The World Conservation Union, a
champion of the Global Amphibian Assessment.
Remarks. – The sister species of P. steineri, is P.
microtympanum (Günther, 1859) (Fig. 2). These two species
are separated from each other by a 12S and 16S sequence
divergence of 2.34% and cytochrome-b sequence divergence
of 11.6% (Tables 1, 2).
Principal components analysis of morphological variables
reveals that P. steineri and P. microtympanum separate well
Fig. 26. Philautus steineri, n. sp., WHT 3210, holotype male, 31.2
mm SVL.

330
Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
Fig. 27. Philautus steineri, n. sp.: a–c, lateral, dorsal and ventral aspects respectively, of head; d, ventral aspect of left manus (arrow indicates
nuptial pad); e, ventral aspect of left pes; and f, semi-diagrammatic representation of the left-pes webbing-pattern of holotype, male,
WHT3210, 31.2mm SVL. Scale bar: 1 mm.
a
bc
de f

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
by size (Fig. 28). Philautus steineri is the larger species (male
SVL to 31.2 mm), compared to P. microtympanum (male SVL
to 25.6 mm).
According to the key of Manamendra-Arachchi &
Pethiyagoda (2005), P. steineri keys out as P. microtympanum
(Günther, 1869) and can be diagnosed morphologically by
having snout-angle category 5 or 6 (vs. snout-angle category
7); an angle of 45° between vomerine ridge and body axis (vs.
angle of 85° between vomerine ridge and body axis); pale
brown posterior margin of thigh (vs. dark-brown posterior
margin of thigh); and pale-brown blotches absent on posterior
margin of thigh (vs. pale-brown blotches present on posterior
margin of thigh).
Philautus steineri is restricted to the highest peaks (~ 1,245
m) of the Knuckles Hills, while P. microtympanum occurs only
in the Central Hills, above 1,555 m. The two populations are
separated from each other by a distance of ~ 100 km and the
Mahaweli River valley, which descends to about 500 m at
Kandy (Figs. 1, 29).
Philautus steineri is a habitat generalist, inhabiting the leaf
litter, shrubs and sub-canopy trees, both in open and closed-
canopy habitats. Males of both species call from 0.3–3 m
above ground, perched on branches.
Conservation status. – (Fig. 29). The Extent of Occurrence of
P. steineri is approximately 10 km2 in the Corbett’s Gap region
at the southern end of the Knuckles mountain range. The
species is considered Endangered (criteria B1 a, b(iii)). Its
restriction to a single forest site and relatively low abundance
make this frog susceptible to habitat modification (forest
clearing). Periodic population monitoring is recommended.
Philautus stuarti, new species
(Figs. 30, 31)
Material examined. – Holotype - male, 25.3 mm SVL, WHT 3208,
Corbett’s Gap (Knuckles Hills), elevation 1,245 m (07º22’ N, 80º
51’ E), coll. 29 Jun.2001.
Paratypes - (all from type locality), male, 24.2 mm SVL, WHT
3207; male, 25.2 mm SVL, WHT 3206; female, 32.4 mm SVL,
WHT 3218, coll. 29 Jun.2001. male, 25.0 mm SVL, WHT 3527,
coll. 1998; male, 25.1 mm SVL, WHT 3574, coll. 1998.
Diagnosis. – (Fig. 31). Philautus stuarti is distinguished from
all other Sri Lankan congeners by a combination of the
Fig. 28. PC1 vs. PC2 factor scores of the principal components
analysis of Philautus steineri, n. sp., (Knuckles hills, K) and P.
microtympanum (Central hills, C). Most of the variation is explained
by the PC1 axis, which relates mostly to body size (P. steineri is the
larger species). The two species overlap completely on the PC2
axis, which relates mostly to palm length, finger 3 length and lower-
arm length.
Fig. 29. Distribution of Philautus steineri, n. sp., in Sri Lanka.
Fig. 30. Philautus stuarti, n. sp., WHT 3208, holotype male, 25.3
mm SVL.

332
Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
Fig. 31. Philautus stuarti, n. sp.,: a–c, lateral, dorsal and ventral aspects respectively, of head; d, ventral aspect of left manus; e, ventral aspect
of left pes; and f, semi-diagrammatic representation of the left-pes webbing-pattern of holotype, male, WHT3208, 25.3mm SVL. Scale bar:
1 mm.
a
bc
de
f

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
following characters: snout-angle category 7; vomerine ridge
absent; lateral dermal fringe absent on fingers; canthal edges
sharp; supernumerary tubercles present on both palm and
pes; nuptial pad absent in males.
Description. – (Figs. 30, 31). Mature males 24.2–25.3 mm SVL;
mature female 32.4 mm SVL. Body stout. Head dorsally flat.
Snout-angle category 7 (angle of snout 105º–107º); snout
rounded in lateral aspect. Canthal edges sharp. Loreal region
concave. Interorbital and internasal spaces flat. Tympanum
distinct, vertically orientated, crescentic. Vomerine ridge
absent. Pineal ocellus, lingual papilla , cephalic ridges, calcar,
and co-ossified skin on skull absent. Supratympanic fold
prominent. Lateral dermal fringe absent on fingers.
Supernumerary tubercles present on both palm and on pes.
Toes webbed. Tarsal folds absent. Snout, interorbital space,
and both anterior and posterior dorsum with horny spinules;
side of head, both upper and lower flanks smooth. Dorsal
part of forelimb, thigh, shank and pes smooth. Throat, chest,
belly and underside of thigh granular. Females lack horny
spinules on dorsum. Inner vocal slits present, nuptial pad
absent in males.
Colour in life. – (Fig. 30) (based on WHT 3208 and WHT
3206). Dorsally light green with ashy green, ashy yellow or
black dots. Loreal region, tympanic region and tympanum
green. Canthal edges, supratympanic fold and both upper
and lower lips yellow (lips white or brown in some specimens).
Digits dorsally yellow. Mid-flank yellow. Chin and chest
yellow; abdomen ashy flesh colour, granules on abdomen
white; area around vent ash. Limbs and digits ventrally ashy
yellow; anterior and posterior thigh brown.
Colour in alcohol. – (Based on WHT 3208). Dorsal and lateral
parts of head and body ashy light blue; upper flank ashy blue
with black patches; lower flank white, inguinal zone dark brown.
Loreal region, tympanic region and tympanum ashy blue.
Upper lip grey. Dorsal parts of limbs ashy blue; posterior
thigh brown. Throat, margin of throat and vocal sacs yellow
with brown dots; chest and belly yellow; ventral thigh and
web yellow with brown dots.
Measurements of holotype. – (WHT 3208, in mm) DBE, 10.1;
DFE, 6.0; DL, 1.0; DW, 1.4, ED, 3.7; EN, 2.6; ES, 4.6; FEL, 13.1;
FL I, 1.9; FL II, 2.7; FL III, 4.3; FL IV, 3.1; FOL, 17.3; HL, 10.4;
HW, 11.4; IML, 1.1; IN, 2.6; IO, 3.2; LAL, 5.1; MBE, 4.0; MFE,
6.6; MN, 9.0; NS, 1.9; PAL, 7.5; SVL, 25.3; TBL, 13.3; TL I, 1.8;
TL II, 2.3; TL III, 4.1; TL IV, 5.9, TL V, 4.3; TYD, 0.8; TYE, 1.4;
UAW, 4.1; UEW, 2.4.
a
b
Fig. 32. PC1 vs. PC2 factor scores of the principal components
analysis of Philautus stuarti, n. sp. (Knuckles hills, K) and P. viridis
(Central hills, C). Most of the variation is explained by the PC1
axis, which relates mainly to body size (P. viridis is the larger
species). The two species overlap on the PC2 axis, which relates
mainly to inter-orbital distance and distance between front of eyes.
Fig. 33. Uncalibrated waveform envelope and spectrogram of the
advertisement calls of a, Philautus stuarti and b, P. viridis. The call
of P. stuarti may be distinguished from that of P. viridis by the
greater call length (1.731–1.805 s, vs. 1.018–1.508 s), greater pulse
rate (3.00–3.09 s-1, vs. 2.09–2.12 s-1), lower dominant and
fundamental frequency (2,520–2,539 Hz, vs. 2,315–2,328 Hz),
smaller pulse length (0.051–0.059 s, vs. 0.077–0.080 s) and the
greater number of pulses per call (6, vs. 3 or 4). There is no frequency
modulation within a call or within a pulse.

334
Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
The vocalisations of these two species (Table 3; Fig. 33)
consist of a series of terse ticks. The advertisement call of P.
stuarti may be distinguished from that of P. viridis by the
greater call length (1.731 – 1.805 s, vs. 1.018 – 1.508 s), greater
pulse rate (3.00 – 3.09 s-1, vs. 2.09 – 2.12 s-1), lower dominant
and fundamental frequencies (2,520 – 2,539 Hz ,vs. 2,315 –
2,328 Hz), smaller pulse length (0.051 – 0.059 s, vs. 0.077 –
0.080 s) and the greater number of pulses per call (6, vs. 3 or
4).
Philautus viridis occurs only on the highest peaks of the
Central Hills (above 1,000 m), P. stuarti being restricted to the
Knuckles Hills at about 1,245 m. Populations of P. stuarti and
P. viridis are separated from each other by the Mahaweli River
valley, which separates the Central Hills from the Knuckles
Hills (Figs. 1, 34), descending to an altitude of about 500 m at
Kandy.
The microhabitat utilized by P. stuarti is similar to that of P.
viridis: both species are habitat specialists inhabiting the
understorey of closed-canopy montane forests, including
cardamom plantations within these. Males of both species
call from 2–3 m above ground, while perched on leaves.
Conservation status. – (Fig. 34). Extent of Occurrence ~10
km2. The species is known from only a single population. Its
low abundance and the susceptibility of its habitat to change
as a result of agricultural expansion lead to it being considered
Endangered (criteria B1 a, b(iii)). Population monitoring is
recommended.
DISCUSSION
Phylogenetic analysis of mitochondrial 12S and 16S ribosomal
sequences revealed at least 17 distinct lineages that may
represent undescribed species (Fig. 2). Close examination of
eight of these revealed diagnostic morphological characters
which, when combined with the separation of these taxa in
multivariate morphological space, differences in advertisement
calls for some species, and differences in ecology for some,
support their recognition as distinct species at this time.
While we used genetic analyses to help identify distinct
evolutionary lineage, we emphasize that there is no genetic
benchmark to designate species. At a minimum, factors such
as the rate and pattern of sequence variation in the taxon
under investigation and the gene region addressed have to
be considered in interpreting genetic distance data for species
delimitation. This is particularly true for organellar DNA which
may have a history distinct from the organismal phylogeny.
Nonetheless, based on experience or review of published data,
some workers have suggested various amounts of genetic
divergence in mitochondrial DNA (mtDNA) as benchmarks
for species-level differences. For example, Bradley & Baker
(2001) suggested that more than 2% genetic divergence of
the cytochrome-b gene indicates the possibility of species-
level divergence in several groups of mammals. Johns & Avise
(1998) found that 90% of putative sister species across a
wide range of vertebrate taxa showed more than 2% molecular
divergence in the mitochondrial cytochrome-b gene. However,
Fig. 34. Distribution of Philautus stuarti, n. sp., in Sri Lanka.
Distribution. – (Fig. 34). Philautus stuarti is currently known
only from a singe location, Corbett’s Gap, in the southern end
of the Knuckles Hills (07º22’ N, 80º51’ E), 1,245 m elevation.
Etymology. – The species epithet is a patronym in the Latin
genitive singular in honour of Simon Nicolas Stuart (b.
England, 1956). A former Acting Director General of IUCN
(2000–2001), Dr. Stuart has a legacy of establishing
international conservation priorities through global
biodiversity studies and enabling conservation projects for
threatened species worldwide. As Head of program of the
Species Survival Commission (1991–2000), he led the
development of the new IUCN Red List categories. As current
Senior Director of the IUCN SSC/CABS Biodiversity
Assessment, Dr. Stuart continues to enhance the Red List of
threatened species by leading global species assessments.
Remarks. – The sister species of P. stuarti, is P. viridis
Manamendra-Arachchi & Pethiyagoda, 2005 (Fig. 2). These
two species are separated from each other by a mtDNA
sequence divergence of 6.6% and from all other Sri Lankan
Philautus in mtDNA sequence divergence of more than 13.8%
(Table 1, 2).
Principal components analysis of morphological variables
show that P. viridis and P. stuarti separate well in morphological
space by size (Fig. 32). Philautus stuarti is the smaller species
when compared to P. viridis: males reach 25.3 mm SVL (cf.
29.6 mm in P. viridis).
According to the key of Manamendra-Arachchi &
Pethiyagoda (2005), P. stuarti keys out as P. viridis from which
species it may be distinguished, by having snout-angle
category 7 (vs. snout-angle category 8) and canthal edges
sharp (vs. canthal edges indistinct).

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
well-known radiations such as the cichlid fauna of the African
rift lakes have virtually no mtDNA differences between closely
related species that have nevertheless been demonstrated to
be robust ecological and sexual entities (Moran et al., 1994;
Bowers et al., 1994; Reinthal & Meyer, 1997). Here we have
used a portion of the mitochondrial 12S and 16S gene fragment
to integrate the current analysis with previously published
data (Meegaskumbura et al., 2002), because it is extensively
used in molecular systematic studies of frogs (de Sá & Hillis,
1990; Richards & Moore, 1998; Emerson et al., 2000; Vences
et. al., 2004) and because it has been used to differentiate
species that are difficult to diagnose morphologically in a
wide range of frogs (Dawood et al., 2002; Chek et al., 2001;
Donnellan et al., 1999; Mahony et al., 2001; Knowles et al.,
2004). All species described here are distinguished by 1–8%
12S and 16S mitochondrial ribosomal gene sequence
divergence between sister species.
Investigation of cytochrome-b sequence divergence among
the new species described here and their respective sister
species showed that they differ substantially (5.8–15.7%) and
thus fit the pattern of greater than 2% cytochrome-b
divergence that is common among vertebrate species (Johns
& Avise, 1998). Some taxonomists advocate that genetic
distance alone should not be used as a criterion to designate
species (Ferguson, 1998), especially if it is low; Hebert et al.
(2004) strongly recommended coupling analysis of molecular
data with traditional taxonomic tools to distinguish between
species. We agree, and think it important to consider multiple
sources of data in diagnosing independent evolutionary
lineages, especially when the molecular data are derived from
organellar gene sequences.
The new species separate from their sister species in
multivariate morphological space largely by size but also, in
some cases, by combinations of other characters. As has
been shown in many disparate frog taxa, the anuran body
plan is conservative (Richards & Moore, 1996; Liu et al., 2000;
Check et al., 2001). Though superficially more or less similar,
all the frog species described here are diagnosable on the
basis of both morphological and molecular characters, with
each species possessing a suite of diagnostic morphological
characters that distinguished it from its congeners. Many of
these same characters have also been used to diagnose other
species in the genus Philautus (Biju & Bossuyt, 2005; Bossuyt
& Dubois, 2001; Das & Chanda, 1998; Inger & Stuebing, 1996;
Ohler et al., 2002; Stuebing & Wong, 2000).
Some of the characters that distinguish the sister species
discussed in this analysis, such as body size and call
characteristics, have been shown to be important in mate
choice and sexual selection (Ryan, 1985; Gerhardt, 1991;
Giacoma & Castellano, 2001). In many groups of frogs, size-
assorted mating preference is pronounced (Tsuji, 2004; Boll
& Linsenmair, 1998). In addition, body size affects other
characters such as the tone of the call and the call rate (Smith
& Roberts, 2003; Tarano & Herrera, 2003). The significant
body size and call differences that exist between the sister
species described here suggests that these species are
different reproductive entities as well. Thus, by multiple
criteria, the new species described here represent independent
evolutionary lineages that warrant recognition as species.
Conservation must aim to preserve the processes that generate
and sustain biodiversity (Crandall et al., 2000; Cowling &
Pressey, 2001; Pressey et al., 2003; Rodrigues, et al., 2004).
Most of the new species described here, along with their
sister species, show a repeated biogeographic pattern of sister
species being isolated on adjacent mountain ranges at high
altitudes, among the three main mountain ranges in Central
Sri Lanka (Fig. 2). For example, each of the three species of
the P. femoralis group (P. femoralis, P. mooreorum, and P.
poppiae) is isolated in cloud forests of one of the three
prominent mountain ranges: the Knuckles, Central and
Rakwana Hills. This same pattern is seen in all other cloud
forest sister species described here (P. viridis Central Hills
and P. stuarti Knuckles Hills; P. microtympanum Central Hills
and P. sterneri Knuckles Hills; P. auratus Rakwana Hills and
P. frankenbergi Central Hills; P. asankai Central Hills and P.
hoffmanni Knuckles Hills). This is also consistent with the
geographic predicted by the montane isolation hypothesis
proposed by Moreau (1966), which postulates that cool, wet
adapted species were pushed to mountaintops during
Pleistocene interglacials. However, the mtDNA distances
among the sister species treated here suggests divergence
that antedates the Pleistocene (Table 2).
Mitochondrial DNA divergence among sister species isolates
suggests divergence times in the Late Miocene to Early
Pliocene (12 – 4 mya). However, these are crude estimates at
best; calculating divergence times from molecular data is
fraught with difficulty, particularly given the lack of calibration
points with which to estimate nucleotide substitution rates
within the Sri Lankan radiation. Even with good calibration,
stochastic variance in the substitution process and sampling
variance introduced by limited sampling of both taxa and
nucleotides is likely to increase the variance associated with
any estimate.
It is difficult to identify the initial barrier that separated the
montane species but the current factors that separate these
species are low altitude saddles, which are substantially
warmer and drier than the cloud forest habitats in which the
montane species are found. The valleys that separate the
three prominent mountain massifs dip to an elevation of at
most 550 m, and apparently present a significant barrier to
dispersal for species that are adapted to the cool wet cloud
forest habitats found above 1,000 m elevation in the three
mountain ranges. While it is difficult to infer the process of
speciation from geographic distribution alone, the repeated
pattern displayed by the taxa described here strongly suggests
a similar response by several taxa to external forcing factors,
likely past climate change (Schneider et al., 1998; Foster, 2001;
Pounds & Puschendorf, 2004). We do not have sufficient
information on paleoclimate in Sri Lanka to make strong
inference of causality, but the levels of mtDNA divergence
among species on mountain tops suggests that their origin
lies with climate change in the Late Miocene - Pliocene which
resulted in the retreat of cool, wet habitats to higher elevations
in response to climate warming and perhaps drying

336
Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
(Premathilake & Risberg, 2003; Manamendra-Arachchi et al.,
2005). Interestingly, this scenario may be repeated in the
agamid lizards of Sri Lanka (Schulte et al., 2002). The isolation
of populations in cool, wet montane habitats makes them
vulnerable to the effects of global warming and we are
concerned that many of these species will lose their optimal
habitats, and thus face extinction, as a consequence of climate
warming over the next several decades.
ACKNOWLEDGEMENTS
We thank Rohan Pethiyagoda, Christopher J. Schneider and
James Hanken for suggestions to improve this manuscript
and financially supporting our work; Mohomed M. Bahir for
comments on the manuscript and extensive field work
associated with this study; Sudath Nanayakkara for field work
and support at WHT’s Agrapatana field station; Suyama
Meegaskumbura for comments on the manuscript and
assistance with field work; and Sudesh Batuwita for field work.
We are very grateful for the critical comments of an anonymous
reviewer, Robert F. Inger (Field Museum of Natural History,
Chicago), and Don Church (Conservation International), to
improve this paper. We also thank Mark Wilkinson, Barry
Clarke and David Gower (BMNH) for facilities to work in their
institution, hospitality and much valuable assistance. We are
grateful to the Forest Department of Sri Lanka for research
permits to work in their reserves, enthusiastic encouragement
and support, and accommodation during visits; and the
Department of Wildlife Conservation Sri Lanka for collection
and export permits. This study was supported financially by
a grant from the Nagao Environmental Foundation to KM-A;
National Science Foundation (grant No. DEB 0345885);
National Geographic Society (Grant No. 7612-04); a Declining
Amphibian Populations Task Force seed grant to Rohan
Pethiyagoda; the Society of Systematic Biologists and
SigmaXi (graduate student awards to MM); and Boston
University.
LITERATURE CITED
Bahir, M. M., M. Meegaskumbura, K. Manamendra-Arachchi, C. J.
Schneider & R. Pethiyagoda, 2005. Reproduction and terrestrial
direct development in Sri Lankan shrub frogs (Amphibia: Ranidae:
Rhacophorinae: Philautus). In: Yeo, D. C. J., P. K. L. Ng & R.
Pethiyagoda (eds.), Contributions to biodiversity exploration
and research in Sri Lanka. The Raffles Bulletin of Zoology,
Supplement No. 12: 339–350.
Biju. S. D. & F. Bossuyt, 2005. A new species of frog (Ranidae,
Racophorinae, Philautus) from the rainforest canopy of the
Western Ghats, India. Current Science, 88(1): 175–178.
Boll, S. & K. E. Linsenmair, 1998. Size-dependent male reproductive
success and size-assortative mating in the midwife toad Alytes
obstetricans. Amphibia-Reptilia, 19(1): 75–89.
Bossuyt, F. & M. C. Milinkovitch, 2000. Convergent adaptive
radiation in Madagascan and Asian Ranid frogs reveal covariation
between larval and adult traits. Proceedings of the National
Academy of Sciences of the United States of America, 97(12):
6585–6590.
Bossuyt, F. & A. Dubois, 2001. A review of the frog genus Philautus
Gistel, 1848 (Amphibia, Anura, Ranidae, Rhacophorinae).
Zeylanica, 6: 1–112.
Bossuyt, F., M. Meegaskumbura, N. Beenaerts, D. J. Gower, R.
Pethiyagoda, K. Roelants, A. Mannaert, M. Wilkinson, M. M.
Bahir, K. Manamendra-Arachchi, P. K. L. Ng, C. J. Schneider, O.
V. Oommen & M. C. Milinkovitch, 2004. Local endemism within
the Western Ghats-Sri Lanka biodiversity hotspot. Science, 306:
479–481.
Bowers N., J. R. Stauffer & T. D. Kocher, 1994. Intra- and interspecific
mitochondrial DNA sequence variation within two species of
rock dwelling cichlids. Molecular Phylogenetics and Evolution,
3: 7582
Bradley, R. D. & R. J. Baker, 2001. A test of the genetic species
concept: Cytochrome-b sequences and mammals. Journal of
Mammalogy, 82(4): 960–973.
Chek, A. A., S. C. Lougheed, J. P. Bogart & P. T. Boag, 2001.
Perception and history: Molecular phylogeny of a diverse group
of neotropical frogs, the 30-chromosome Hyla (Anura: Hylidae).
Molecular Phylogenetics and Evolution, 18(3): 370–385.
Cocroft, R. B. & M. J. Ryan, 1995. Patterns of advertisement call
evolution in toads and chorus frogs. Animal Behaviour, 49: 283-
303.
Cowling, R. M. & R. L. Pressey, 2001. Rapid plant diversification:
Planning for an evolutionary future. Proceedings of the National
Academy of Sciences of the United States of America, 98(10):
5452–5457.
Coyne, J. A. & H. A. Orr, 2004. Speciation. Sinauer Assocites,
Massachusetts. 545 pp.
Crandall, K. A., R. P. Olaf, Bininda-Emonds, G. M. Mace & K.
Robert, 2000. Considering evolutionary processes in conservation
biology. Trends in Ecology and Evolution, 15(7): 290–295.
Das, I. & S. K. Chanda, 1998. A new species of Philautus (Anura:
Rhacophoridae) from the Eastern Ghats, south-eastern India.
Journal of South Asian Natural History, 3(1) 103–112.
Dawood, A., A. Channing, & J. P. Bogart, 2002. A molecular
phylogeny of the frog genus Tomopterna in southern Africa:
Examining species boundaries with mitochondrial 12S rRNA
sequence data. Molecular Phylogenetics and Evolution, 22(3):
407–413.
de Sá, R. O. & D. M. Hillis, 1990. Phylogenetic relationships of the
pipid frogs Xenopus and Silurana: an integrating of ribosomal
DNA and morphology. Molecular Biology and Evolution, 7: 365–
376.
Donnellan, S. C., K. McGuigan, R. Knowles, M. Mahony & C.
Moritz, 1999. Genetic evidence for species boundaries in frogs
of the Litoria citropa species-group (Anura: Hylidae). Australian
Journal of Zoology, 47(3): 275–293.
Dubois A., 2004. Developmental pathway, speciation and
supraspecific taxonomy in amphibians, 1: Why are there so many
frog-species in Sri Lanka? Alytes, 22(1–2): 19–27.
Duellman W. E. & L. Trueb, 1986. Biology of Amphibians. McGraw-
Hill, New York. 696 pp.
Emerson, S. B., R. F. Inger, & D. Iskandar, 2000. Molecular
systematics of the fanged frogs of Southeast Asia. Molecular
Phylogenetics and Evolution, 16: 131–142.
Foster, P., 2001. The potential negative impacts of global climate
change on tropical montane cloud forests. Earth-Science Reviews,

337
THE RAFFLES BULLETIN OF ZOOLOGY 2005
55(1–2): 73–106.
Gerhardt, H., 1991. Female mate choice in treefrogs. Static and
dynamic acoustic criteria. Animal Behaviour, 42: 615–635.
Gerhardt, C. H. & F. Huber, 2002. Acoustic communication in insects
and anurans. The University of Chicago Press, Chicago. 542
pp.
Giacoma, C. & S. Castellano, 2001. Advertisement call variation and
speciation in the Bufo viridis complex. In: Ryan, M. J. (ed.),
Anuran communication. Smithsonian Institution Press,
Washington & London. pp. 205–219.
Gistel, J., 1848. Naturgeschichte des Thierreichs für höhere Schulen.
Hoffmann, Stuttgart. xi+216+iv pp., 32 pls.
Günther, A., 1859. Catalogue of the Batrachia Salientia in the
collection of the British Museum. Taylor & Francis, London.
xvi+160 pp, 12 pls.
Günther, A., 1864. The reptiles of British India. Ray Society,
London. xxvii+452 pp, 26 pls.
Günther, A., 1869. First account of species of tailless batrachians
added to the collection of the British Museum. Proceedings of
the Zoological Society of London, 1868: 478–490, pls. 37–40.
Hebert, P. D. N., E. H. Penton, J. M. Burns, D. H. Janzen & W.
Hallwachs, 2004. Ten species in one: DNA barcoding reveals
cryptic species in the neotropical skipper butterfly Astraptes
fulgerator. Proceedings of the National Academy of Sciences of
the United States of America, 101(41): 14812–14817.
Hey, J., 2001. Genes, categories and species. Oxford University
Press, Oxford. 217 pp.
Huelsenbeck, J. P. & F. Ronquist, 2001. MRBAYES: Bayesian
inference of phylogenetic trees. Bioinformatics, 17(8): 754–755.
Huelsenbeck, J. P., F. Ronquist, R. Neilsen & J. P. Bollback, 2001.
Bayesian inference of phylogeny and its impact on evolutionary
biology. Science, 294: 2310–2314.
Inger, R. F. & R. B. Stuebing, 1996. Two new species of frogs from
Southern Sarawak. The Raflles Bulletin of Zoology, 44(2) 543–
549.
Jeanmougin, F., J. D. Thompson, M. Gouy, D. G. Higgins & T. J.
Gibson, 1998. Multiple sequence alignment with Clustal x. Trends
in Biochemical Sciences, 23(10): 403–405.
Kotagama, S. W., K. D. Arudpragasam & I. Kotalawala, 1981. A
check list of Amphibia of Sri Lanka. National Science Council,
Colombo. 8 pp.
Liu, W. Z., A. Lathrop, J. Z. Fu, D. T. Yang & R. W. Murphy, 2000.
Phylogeny of east Asian bufonids inferred from mitochondrial
DNA sequences (Anura: Amphibia). Molecular Phylogenetics
and Evolution, 14(3): 423–435.
Mace, G. M., J.L. Gittleman & A. Purvis, 2003. Preserving the Tree
of Life. Science 300(5626): 1707–1709.
Mahony M., R. Knowless, R. Foster & S. Donnellan, 2001.
Systematics of the Litoria citropa (Anura: Hylidae) complex in
Nothern New South Wales and Southern Queensland, Australia,
with the description of a new species. Records of the Australian
Musuem, 53: 37–48.
Manamendra-Arachchi, K. & R. Pethiyagoda, 2005. The Sri Lankan
shrub-frogs of the genus Philautus Gistel, 1848 (Ranidae:
Rhacophorinae), with description of 27 new species. In: Yeo, D.
C. J., P. K. L. Ng & R. Pethiyagoda (eds.), Contributions to
biodiversity exploration and research in Sri Lanka. The Raffles
Bulletin of Zoology, Supplement No. 12: 163–303.
Manamendra-Arachchi, K., R. Pethiyagoda, R. Dissanayake & M.
Meegaskumbura, 2005. An extinct tiger from the Late Quaternary
of Sri Lanka. In: Yeo, D. C. J., P. K. L. Ng & R. Pethiyagoda
(eds.), Contributions to biodiversity exploration and research in
Sri Lanka. The Raffles Bulletin of Zoology, Supplement No. 12:
423–434.
Martin, A. P. & S. R. Palumbi, 1993. Body size, metabolic rate,
generation time, and the molecular clock. Proceedings of the
National Academy of Science USA, 90: 4087 – 4091.
McCraken K. G. & F. H. Sheldon, 1997. Avian vocalizations and
phylogenetic signal. Proceedings of the National Academy of
Sciences of theUSA, 94(8): 3833–3836.
Meegaskumbura, M., F. Bossuyt, R. Pethiyagoda, K. Manamendra-
Ararchchi, M. Bahir, M. C. Milinkovitch & C. J. Schneider,
2002. Sri Lanka: an amphibian hotspot. Science, 298: 379.
Moran P., I. Kornfield & P. Reinthal, 1994. Molecular systematics
and radiation of the haplochromine cichlids (Teleostei:
Perciformes) of Lake Malawi. Copeia, 1994: 274–288.
Moreau, R. E., 1966. The bird faunas of Africa and its islands.
Academic Press, New York. 424 pp.
Palumbi, S. R., 1996. Nucleic Acids II: The polymerase chain reaction.
In: Hillis, D. M., C. Moritz & B. K. Mable (eds.), Molecular
Systematics. Sinauer Associates Inc., Sunderland, Massachusetts.
pp. 205–248.
Ohler, A., S. R. Swan & J. C. Daltry, 2002. A recent survey of the
amphibian fauna of the Cardamom Mountains, Southwest
Cambodia with descriptions of three new species. The Raffles
Bulletin of Zoology, 50(2): 465–481.
Pethiyagoda, R. & K. Manamendra-Arachchi, 1998. Evaluating Sri
Lanka’s amphibian diversity. Occasional Papers of the Wildlife
Heritage Trust, 2: 1–12.
Pounds, J. A. & R. Puschendorf, 2004. Ecology — Clouded futures.
Nature, 427 (6970): 107–109.
Premathilake, R. & J. Risberg, 2003. Late Quaternary climate history
of the Horton Plains, central Sri Lanka. Quaternary Science
Review, 22: 1525–1541.
Pressey, R. L., R. M. Cowling & M. Rouget, 2003. Formulating
conservation targets for biodiversity pattern and process in the
Cape Floristic Region, South Africa. Biological Conservation,
112(1–2): 99–127.
Queiroz, K. de, 1998. The General Lineage Concept of species. In:
Howard, D. J. & S. H. Berlocher (eds.), Endless forms: species
and speciation. Oxford University Press, Oxford. pp. 57–75.
Rambaut, A., 1996. Se-Al: Sequence Alignment Editor. Available at
http://evolve.zoo.ox.ac.uk/.
Reinthal, P. & A. Meyer, 1997. Molecular phylogenetic tests of
speciation models in African cichlid fishes. In Givinish, T. & K.
Sytsma (eds.), Molecular Phylogenetics of Adaptive Radiations.
Cambridge University Press, Cambridge and Boston. pp 375–
390.
Richards, C. M. & W. S. Moore, 1996. A phylogeny for the African
treefrog family Hyperoliidae based on mitochondrial DNA.
Molecular Phylogenetics and Evolution, 5: 522–532.
Richards, C. M. & W. Moore, 1998. A molecular phylogeny of the

338
Meegaskumbura & Manamendra-Arachchi: Eight new species of Sri Lankan Philautus
Old World tree frog family Racophoridae. Journal of Herpetology,
8: 41–46.
Rodrigues, A. S. L., S. J. Andelman, M. I. Bakarr, L. Boitani, T. M.
Brooks, R. M. Cowling, L. D. C. Fishpool, G. A. B. da Fonseca,
K. J. Gaston, M. Hoffmann, J. S. Long, P. A. Marquet, J. D.
Pilgrim, R. L. Pressey, J. Schipper, W. Sechrest, S. N. Stuart, L.
G. Underhill, R. W. Waller, M. E. J. Watts & X. Yan, 2004.
Effectiveness of the global protected area network in representing
species diversity. Nature, 428(6983): 640–643.
Ryan M. J., 1985. The Tungara Frog: a study in sexual selection and
communication. University of Chicago Press, Chicago. 246 pp.
Schneider C.J., M. Cunningham & C. Moritz, 1998. Comparative
phylogeography and the evolution of reptiles and amphibians
endemic to the Wet Tropics rainforests of Australia. Molecular
Ecology, 7: 487–498
Schulte, J. A., J. R. Macey, R. Pethiyagoda & A. Larson, 2002.
Rostral horn evolution among agamid lizards of the genus
Ceratophora endemic to Sri Lanka. Molecular Phylogenetics and
Evolution, 22(1): 111–117.
Sites, J. W. & K. A. Crandall, 1997. Testing species boundaries in
biodiversity studies. Conservation Biology 11(6): 1289–1297.
Smith M. J., & J. D. Roberts, 2003. Call structure may affect male
mating success in the quacking frog Crinia georgiana (Anura:
Myobatrachidae). Behavioral Ecology and Sociobiology, 53(4):
221–226.
Stuart, S., J. S. Chanson, N. A. Cox, B. E. Young, A. S. L. Rodrigues,
D. L. Fischman & R. W. Waller, 2004. Status and trends of
amphibian declines and extinctions worldwide. Science, 306:
1783–1786.
Stuebing, R. B. & A. Wong, 2000. A new species of frog, Philautus
erythrophthalmus (Rhacophoridae) from southwestern Sabah,
Malaysia. The Raffles Bulletin of Zoology, 48(2): 293–296.
Sullivan, B. K., 1992. Sexual selection and calling behavior in the
American toad (Bufo americanus). Copeia, 1992 (1): 1–7.
Swofford, D. L., 2002. PAUP, phylogenetic analysis using parsimony
(and other methods), v. 4b10. Sinauer Associates, Sunderland,
MA.
Tan, A. M. & D. Wake, 1995. MtDNA phylogeography of the
California newt Taricha torosa (Caudata, Salamandridae).
Molecular Phylogenetics and Evolution, 4: 383–394.
Tarano, Z. & E. A. Harrera, 2003. Female preference for call traits
and male mating success in the neotropical frog Physalaemus
enesefae. Ethology, 109(2): 121–134.
Tsuji, H., 2004. Reproductive ecology and mating success of male
Limnonectes kuhlii, a fanged frog from Taiwan. Herpetologica,
60(2): 155–167.
Vences, M., F. Glaw, J. Riccardo & G. Schimmenti, 2000. A new
species of Heterixalus (Amphibia, Hyperoliidae) from western
Madagascar, African Zoology, 35(2) 269–276.
Vences, M., J. Kosuch, S. Lötters, A. Widmer, K. H. Jungfer, J.
Köhler & M. Veith, 2000. Phylogeny and classification of poison
frogs (Amphibia: Dendrobatidae), based on mitochondrial 16S
and 12S ribosomal gene sequences. Molecular Phylogenetics and
Evolution, 15(1): 34–40.