Two new species of shrub frogs (Rhacophoridae: Pseudopilautus) from Sri Lanka.

Abstract

Two new species of Sri Lankan shrub frogs of the genus Pseudophilautus are described. These species are diagnosed from their congeners on the basis of morphology, morphometrics and mitochondrial DNA sequence data. Pseudophilautus schneideri, new species, is distinguished from all Sri Lankan Pseudophilautus by its small size (< 22.8 mm SVL), distinct tympanum and supratympanic fold, sharp canthal edges, granular throat, chest and belly, and absence or presence of a vomerine ridge. Pseudophilautus hankeni, new species, is distinguished by its diminutive size (< 21.9 mm SVL), distinct tympanum, rounded canthal edges, tuberculated outer edge of lower arm, tuberculated dermal fold on outer edge of foot, granular throat, chest and belly, and the absence of a vomerine ridge. Pseudophilautus schneideri inhabits shrubs in open areas of the low to mid-elevations of the island’s south-western ‘wet zone’ (rainfall > 2,000 mm•yr-1), including anthropogenic habitats, while P. hankeni is found on shrubs in the understorey of montane forests of the highest peaks (c. 1,200–1,600 m elevation) of the Knuckles region. These descriptions bring the total number of valid species of Sri Lankan Pseudophilautus to 67, 48 of which are extant.

Full text

Accepted by M. Vences: 14 Dec. 2010; published: 24 Jan. 2011
ZOOTAXA
ISSN 1175-5326 (print edition)
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Copyright © 2011 · Magnolia Press
Zootaxa 2747: 1–18 (2011)
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1
Two new species of shrub frogs (Rhacophoridae: Pseudophilautus)
from Sri Lanka
MADHAVA MEEGASKUMBURA1,2,4 & KELUM MANAMENDRA-ARACHCHI3
1Department of Zoology, Faculty of Science, University of Peradeniya, Sri Lanka
2Museum of Comparative Zoology & Harvard University Center for the Environment, 26 Oxford Street, Cambridge MA 02138, USA
3Postgraduate Institute of Archaeology, University of Kelaniya, 407 Bauddhaloka Mawatha Colombo 07
4Corresponding author. E-mail: madhava_m@mac.com
Abstract
Two new species of Sri Lankan shrub frogs of the genus Pseudophilautus are described. These species are diagnosed from
their congeners on the basis of morphology, morphometrics and mitochondrial DNA sequence data. Pseudophilautus sch-
neideri, new species, is distinguished from all Sri Lankan Pseudophilautus by its small size (< 22.8 mm SVL), distinct
tympanum and supratympanic fold, sharp canthal edges, granular throat, chest and belly, and absence or presence of a
vomerine ridge. Pseudophilautus hankeni, new species, is distinguished by its diminutive size (< 21.9 mm SVL), distinct
tympanum, rounded canthal edges, tuberculated outer edge of lower arm, tuberculated dermal fold on outer edge of foot,
granular throat, chest and belly, and the absence of a vomerine ridge. Pseudophilautus schneideri inhabits shrubs in open
areas of the low to mid-elevations of the island’s south-western ‘wet zone’ (rainfall > 2,000 mm•yr-1), including anthro-
pogenic habitats, while P. hankeni is found on shrubs in the understorey of montane forests of the highest peaks (c. 1,200–
1,600 m elevation) of the Knuckles region. These descriptions bring the total number of valid species of Sri Lankan Pseu-
dophilautus to 67, 48 of which are extant.
Key words: Rhacophorinae, taxonomy, molecular systematics, Knuckles Hills, conservation
Introduction
Following the discovery in Sri Lanka of a large radiation of Oriental tree frogs of the genus Pseudophilautus (Mee-
gaskumbura et al. 2002), 39 new species have been described as part of an on-going effort to document this fauna
(Manamendra-Arachchi & Pethiyagoda 2005; Meegaskumbura & Manamedra-Arachchi 2005; Meegaskumbura et
al. 2007; Meegaskumbura et al. 2009). The review and description of 27 new species by Manamendra-Arachchi &
Pethiyagoda (2005), though informed by a phylogeny, was based purely on morphology (given the unavailability
of molecular data for the older type material). Meegaskumbura & Manamendra-Arachchi (2005), however,
described eight more new species using the General Lineage concept (de Queiroz, 1998), where species are consid-
ered as independent evolutionary lineages based on multiple criteria (in this case molecular divergence, morphol-
ogy, ecology and vocalization). Meegaskumbura et al. (2007) added two additional new but extinct species
discovered in historical museum collections, again adopting a purely morphological approach. More recently Mee-
gaskumbura et al. (2009) described two new species from the lowlands of Sri Lanka using molecular, morphologi-
cal and morphometric data. The island’s inventory of Pseudophilautus now stands at 67 species, of which 48 are
extant. Surveys in Sri Lanka since the early 1990s suggest that 19 species, known only from museum specimens
collected in the 19th and early 20th centuries, have since disappeared (Manamendra-Arachchi & Pethiyagoda
2005; Meegaskumbura et al. 2007); these extinctions, together with the high number of Critically Endangered (11)
and Endangered (36) species, highlights the urgent need to describe and name the remaining new species of shrub
frogs discovered by us in Sri Lanka, so that they can be included in the conservation planning process.
Here we continue to document the new species discovered in Sri Lanka as a result of our explorations on the
island up to 2005. The species descriptions are based on morphological, morphometric and molecular data, in the
context of the General Lineage concept of species.
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Material and methods
Field sampling and measurements were made as described in Manamendra-Arachchi & Pethiyagoda (2005), except
as mentioned below.
Morphological analysis. The suite of characters and character states used by Manamendra-Arachchi & Pethi-
yagoda (2005) was analyzed for all individuals. Measurements were made to the nearest 0.1 mm using dial vernier
calipers. Measurements with high coefficients of variation or low repeatability were omitted from the analysis, for
which the following were used: distance between back of eyes (DBE); distance between front of eyes (DFE);
length of disk (DL); width of disk (DW); eye diameter (ED); eye-to-nostril distance (EN); eye-to-snout length
(ES); femur length (FEL); length of finger 1 (FLI); length of finger 2 (FLII); length of finger 3 (FLIII); length of
finger 4 (FLIV); pes length (FOL); head length (HL); head width (HW); length of inner metatarsal tubercle (IML);
internarial distance (IN); interorbital 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); length of toe 1 (TLI);
length of toe 2 (TLII); length of toe 3 (TLIII); length of toe 4 (TLIV); length of toe 5 (TLV); diameter of tympanum
(TYD); distance from tympanum to front of eye (TYE); length of upper arm (UAW); and width of upper eyelid
(UEW). Illustration of the webbing pattern follows Manamendra-Arachchi & Pethiyagoda (2005). Measurements
with high coefficients of variation or low repeatability were omitted from the PCA analysis, for which only the fol-
lowing were used: DBE, DFE, DL, DW, ED, EN, ES, FEL, FLI, FLII, FLIII, FLIV, FOL, HL, HW, IN, IO, LAL,
MBE, MFE, MN, NS, PAL, SVL, TL1, TLII, TLIII, TLIV, TLV IML, TYE and TBL.
Principal components analysis (PCA) of the character correlation matrix was used to reduce dimensionality of
the continuous morphological variables and to identify those variables that best discriminate between morphologi-
cally similar species (P. schneideri vs. P. folicola and P. hankeni vs. P. schmarda). Various axis rotations were
tested and one selected for optimal interpretability of variation among the characters. For consistency, only mature
males were used in this analysis. In several cases, the small sample sizes likely do not represent the full range of
morphological variation; nonetheless, the analyses are sufficient to demarcate species and identify characters that
contribute best to their diagnoses. SYSTAT (Version 11.00.01 for Windows XP) was used for statistical analysis.
Molecular analysis. A species, representing P. hankeni (WHT 6302) and two more undescribed species P. cf.
simba WHT 5831 and P. simba WHT 6004 (WHT 3221 was called P. simba in Meegaskumbura et al. 2009, which
is corrected here as P. cf. simba) were added to the data that were analyzed in Meegaskumbura et al. (2009; see
Table 1); P. schneideri (‘Pseudophilautus sp.’ WHT 2667), was already included in that data set (Table 1). Only
12s rRNA and 16s rRNA partial sequences were used to construct the phylogenetic tree, as was done in Meegas-
kumbura et al. (2002), Meegaskumbura & Manamendra-Arachchi (2005) and Meegaskumbura et al. (2009). Cyto-
chrome-b data was used, in addition, to determine the percentage divergences among sister taxa, and PCR
amplification and alignment of sequences was done as explained in Meegaskumbura & Manamendra-Arachchi
(2005).
The data was analyzed using Bayesian, Maximum Likelihood (ML) and Maximum Parsimony (MP) criteria.
For brevity, we present only the Maximum Likelihood tree, which is identical to the Bayesian tree, together with
one of the two 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 100,000 gen-
erations (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 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 2002). For tree searches under a Maximum Parsi-
mony criterion we used 100 heuristic searches with TBR branch-swapping and random taxon addition as imple-
mented in PAUP*4.0b10. Two equally parsimonious trees with tree scores of 1133 were recorded. A bootstrap
analysis (1000 replicates, random stepwise addition with 100 reps.) to determine node support was also carried out
within a maximum-parsimony framework.
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TWO NEW SRI LANKAN SHRUB FROGS
TABLE 1. Reference numbers, and the Genbank accession numbers for the species used in the phylogenetic analysis.
Species Reference number Genbank Accession numbers
12s 16s
P. alto WHT2723 AY141781 AY141827
P. asankai WHT5107 FJ788141 FJ788160
P. auratus WHT2792 AY141789 AY141835
P. caeruleus WHT2511 AY141764 AY141810
P. cavirostris WHT3299 FJ788137 FJ788156
P. cf. sarasinorum WHT2484 AY141762 AY141808
P. cf. sarasinorum WHT2489 AY141763 AY141809
P. cf. simba WHT5831 GU593345 GU593347
P. cf. simba WHT3221 FJ788148 FJ788167
P. cf. sordidus WHT_H12 AY141791 AY141837
P. cf. sordidus WHT_H15 AY141792 AY141838
P. charius FB AY141840 AY141794
P. decoris WHT3271 FJ788144 FJ788163
P. femoralis WHT2566 AY141771 AY141817
P. femoralis WHT2772 AY141785 AY141831
P. frankenbergi WHT2552 AY141768 AY141814
P. frankenbergi WHT2555 AY141769 AY141815
P. hallidayi WHT_H11 AY141793 AY141839
P. hankeni WHT6302 GU593346 GU593348
P. hoffmanni WHT3223 FJ788142 FJ788161
P. hoipolloi WHT2675 AY141776 AY141822
P. limbus WHT2690 AY141777 AY141823
P. limbus WHT2700 AY141779 AY141825
P. lunatus WHT3283 FJ788150 FJ788169
P. microtympanum WHT2558 AY141770 AY141816
P. mittermeieri KAN2 FJ788143 FJ788162
P. mooreorum WHT3209 FJ788134 FJ788153
P. ocularis WHT2887 FJ788145 FJ788164
P. pappilosus WHT3284 FJ788151 FJ788170
P. pleurotaenia WHT3176 FJ788146 FJ788165
P. poppiae WHT5026 FJ788135 FJ788154
P. poppiae WHT2779 FJ788136 FJ788155
P. popularis WHT3191 FJ788149 FJ788168
P. procax WHT2786 AY141788 AY141834
P. sarasinorum WHT2481 AY141761 AY141807
P. schmarda WHT2715 AY141780 AY141826
P. signatus FB AY141795 AY141841
P. simba WHT6004 GQ204740 GQ204679
P. singu WHT2658 AY141773 AY141819
P. sordidus WHT2699 AY141778 AY141824
P. sp. WHT2515 AY141765 AY141811
continued next page
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As described in Meegaskumbura & Manamendara-Arachchi (2005), 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’- CTTCTACTGGTTGTCCTCCGAT-
TCA-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. 2.0a11; Rambaut 1996).
Results
Morphometric analysis. Pseudophilautus schneideri and P. folicola (the species it morphologically resembles
most closely) separate distinctly from each other in morphological space (Fig. 1, Table 2). Principal components
analysis shows that the two species are distinguished by a combination of body dimensions and nostril to snout dis-
tance. The PC(1) axis, which explains 81 % of the variance, is a size axis (most of the variables load heavily and all
variables have high, positive loadings on this axis; NS does not load heavily, but has a positive value of 0.275). The
PC(2) axis represents 6 % of the variance, with NS (-0.802) and FLIII (-0.588) contributing most heavily. How-
ever, the two species only separate on the body size (i.e. the PC(1)) axis, but completely overlap on the PC(2) axis;
Pseudophilautus schneideri is the smaller of the two species.
TABLE 1. (continued)
Species Reference number Genbank Accession numbers
12s 16s
P. sp. WHT2540 AY141767 AY141813
P. sp. WHT2797 AY141790 AY141836
P. schneideri WHT2667 AY141774 AY141820
P. sp. WHT2669 AY141775 AY141821
P. sp. WHT2525 AY141766 AY141812
P. sp. WHT2774 AY141786 AY141832
P. sp. WHT2729 AY141782 AY141828
P. sp. WHT2731 AY141783 AY141829
P. steineri WHT3210 FJ788138 FJ788157
P. stuarti WHT3207 FJ788139 FJ788158
P. stuarti WHT3208 FJ788140 FJ788159
P. tanu WHT6343 FJ788152 FJ788171
P. viridis WHT2627 AY141772 AY141818
P. viridis WHT2766 AY141784 AY141830
P. wynaadensis FB AY141796 AY141842
P. zorro WHT3175 FJ788147 FJ788166
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FIGURE 1. PC1 vs. PC2 factor scores of the principal components analysis of Pseudophilautus schneideri, n. sp. and P. foli-
cola, show these species to separate well from each other in PC space. Most of the variation is explained by the PC1 axis, which
relates mainly to body size (P. schneideri being smaller). The PC2 axis is mostly explained by nostril-to-snout length and length
of finger 3; here, P. schneideri overlaps completely with P. folicola.
Pseudophilautus hankeni and P. schmarda also, separate from each other in morphological space (Fig. 2, Table
3). Principal components analysis shows that the two species are distinguished from each other by a combination of
body dimensions, NS, MBE, MFE, and ED. The PC(1) axis, which explains 83 % of the variance, is a size axis
(most variables load heavily and positively on this axis). PC(2) represents 7 % of the variance, with NS (0.507),
MBE (-0.477) and MFE (-0.421) contributing most heavily. However, while these two species separate along the
size (PC1) axis, they completely overlap on the PC(2) axis; P. hankeni has the smaller body dimensions.
Molecular phylogenetics. The final dataset contained 12S and 16S rRNA mitochondrial gene sequences from
58 putative haplotypes. Fifty-five of these represent Sri Lankan Pseudophilautus, while three represent Indian spe-
cies (one, P. wynaadensis, is nested within the Sri Lankan clade, whereas the other two represent the sister group to
the Sri Lankan and nested Indian Pseudophilautus: Meegaskumbura et al. 2002; Bossuyt et al. 2004). Hence, of the
939 nucleotide positions sequenced, 867 were clearly alignable and were included in this analysis.
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TABLE 2. Component loadings for axes 1 and 2 of the principal component analysis, variance explained and percentage of
total variance explained for Pseudophilautus schneideri and P. folicola.
The Maximum Likelihood tree (from the Maximum Likelihood analysis) is shown in Fig. 3. The tree is rooted
with two Indian taxa (Pseudophilautus charius and P. signatus) that represent the sister group to the Sri Lankan
Pseudophilautus radiation (Meegaskumbura et al. 2002). For the Bayesian analysis, we ran 500,000 generations of
the MCMCMC algorithm and the summed likelihood of the four chains reached stationarity by 85,000 generations.
The posterior probabilities of clades shown at nodes in Fig. 3 represent the frequency of those clades in the 25,000
trees sampled from the last 250,000 generations, and clades with posterior probability of 50% or less were col-
lapsed. The parameters of the nucleotide substitution model for the most likely tree were as follows. Rate matrix:
R(G-T) = 0.0021, R(C-T) = 0.6815, R(C-G) = 0.0114, R(A-T) = 0.0308, R(A-G) = 0.2254, R(A-C) = 0.0486.
Nucleotide frequency: A = 0.3243, C = 0. 0.2247, G = 0.1975, T = 0.2534. Rate variation: shape parameter for
gamma distributed rate variation among sites (alpha) = 0.6458; proportion of invariant sites = 0.4020. The maxi-
mum likelihood tree found via a Tree Bisection and Reconnection branch-swapping heuristic search using the
Axis 1 Axis 2
EN 0.989 0.090
HL 0.987 0.081
DBE 0.985 -0.049
SVL 0.978 0.038
FOL 0.973 -0.001
MFE 0.973 0.143
DFE 0.972 0.143
HW 0.971 0.113
ES 0.969 0.051
LAL 0.969 0.122
MN 0.961 0.107
IO 0.958 0.177
MBE 0.957 0.034
TBL 0.956 0.208
FEL 0.952 -0.109
TLV 0.934 0.088
TLI 0.928 0.068
FLIV 0.923 -0.003
IML 0.922 -0.070
TLII 0.919 -0.288
FLI 0.908 -0.185
ED 0.892 0.289
IN 0.891 0.118
FLII 0.839 -0.118
TLIII 0.837 -0.290
PAL 0.806 -0.446
TYE 0.794 0.215
TLIV 0.725 -0.103
FLIII 0.501 -0.588
NS 0.275 -0.802
Variance Explained by Components 24.344 1.762
Percent of Total Variance Explained 81.147 5.872
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above nucleotide substitution parameters in PAUP*v.4.0b10 had the same topology as the Bayesian tree, but had
slight branch-length differences. 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 two equally parsi-
monious trees with the maximum parsimony bootstrap values at nodes (bootstrap values less than 50 % collapsed,
shown below nodes on ML tree; Fig. 3). Bootstrap values towards the base of the Sri Lankan radiation were low,
which results in a basal polytomy. However, as expected, the values closer to the OTUs showed higher bootstrap
values, and relationships of taxa within these better-supported clades were identical to those of the maximum like-
lihood analysis. The relationships of taxa of the clades from which the two new species are described were also
identical to the relationships from the maximum likelihood analysis.
FIGURE 2. PC1 vs. PC2 factor scores of the principal components analysis of Pseudophilautus hankeni, n. sp. and P.
schmarda, show these species to separate fairly well from each other in PC space. Most of the variation is explained by the PC1
axis, which relates mainly to body size (P. hankeni being smaller). The PC2 axis is mostly explained by NS, MBE and MFE
(NS loads positively, and the other two negatively); here, P. hankeni overlaps completely with P. schmarda.
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FIGURE 3. Maximum likelihood tree of 12s and 16s rRNA gene fragments, with posterior probabilities from the Bayesian
analysis shown above nodes and Maximum Parsimony Bootstrap values shown below nodes. Pseudophilautus schneideri and
P. hankeni are indicated by asterisks.
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TABLE 3. Component loadings for axes 1 and 2 of the principal component analysis, variance explained and percentage of
total variance explained for Pseudophilautus hankeni and P. schmarda.
Pseudophilautus schneideri, new species
(Figs. 4–7)
Material examined. Holotype: mature male, 22.8 mm SVL, WHT6355, Kudawa, Sinharaja World Heritage Site,
alt. 381 m (6°26’N, 80°25’E), coll. 9 June 1999; K.M.-A. & M.M.
Paratypes: 7, mature male, 19.9 mm SVL, WHT6354; mature male, 21.0 mm SVL, WHT6353; mature male, 21.0
mm SVL, WHT6357; mature male, 20.2 mm SVL, WHT6356; collection data same as holotype. Two mature
males, 20.4 mm SVL, WHT6349; 20.7 mm SVL, WHT6350, Kanneliya Forest Reserve, alt. 41 m (6°15’N,
80°20’E), coll. 05 VI 1999, M.M. Mature female, 20.7 mm SVL, WHT2667, Elpitiya forest reserve, alt. 31 m
(6°11’N, 80°11’E), coll. 14 September 1999, K.M.-A. & M.M.
Diagnosis. Pseudophilautus schneideri is assigned to the genus Pseudophilautus as they are well nested within
the Sri Lankan monophyletic group (Figs. 1 & 2) of frogs (Meegaskumbura et al. 2002; Bossuyt et al. 2004; Yu et
al. 2010) and are characterized by terrestrial direct development (Bossuyt and Dubois 2000). Pseudophilautus sch-
neideri is distinguished from all other Sri Lankan congeners by a combination of the following characters: size
small, mature individuals 19.9–22.8 (female 20.7 mm SVL) mm SVL; tympanum distinct; supratympanic fold dis-
tinct; canthal edges sharp; vomerine ridge absent or present; throat, chest and belly granular.
Description. (Based on Holotype WHT6355) Body elongate. Head laterally convex. Snout obtusely pointed
in dorsal view, pointed in lateral view. Canthal edges sharp. Loreal region flat. Interorbital space convex. Internasal
space flat. Nostrils oval. Pupil oval, horizontal. Tympanum distinct, oval, vertical. Pineal ocellus absent. Vomerine
ridge present only on right side, bearing small teeth, between, anterior to and proximal to choanae, angled at about
45° to body axis. Tongue moderate, emarginate, not bearing a lingual papilla. Supratympanic fold distinct.
Cephalic ridges absent. Co-ossified skin on head absent. Both upper and lower arms short. Fingers thin. Relative
length of fingers, 1 < 2 < 4 < 3. Tips of fingers with discs, with circum-marginal grooves. Fingers lack a lateral der-
mal fringe. Webbing on fingers absent. Subarticular tubercles on fingers distinct, oval, single, some absent IV 2
(penultimate subarticular tubercle). Prepollex oval, distinct. Two palmar tubercles, oval, distinct. Supernumerary
tubercles absent. Thigh slender. Shank slender. Toes thin. Relative length of toes, 1 < 2 < 3 < 5 < 4. Tips of toes
with discs, with circum-marginal groves. Webbing present on toes. Subarticular tubercles on toes distinct, oval, sin-
Axis 1 Axis 2
HL 0.979 -0.111
DBE 0.968 -0.085
MN 0.965 -0.230
ES 0.958 0.156
TBL 0.955 0.117
EN 0.942 0.210
FOL 0.937 0.250
HW 0.931 0.065
DFE 0.922 0.203
IN 0.917 0.137
SVL 0.915 0.004
ED 0.898 -0.308
MFE 0.874 -0.421
MBE 0.776 -0.477
NS 0.700 0.507
Variance Explained by Components 12.495 1.038
Percent of Total Variance Explained 83.195 6.922
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gle, all present. Inner metatarsal tubercle distinct, oval. Outer metatarsal tubercle absent. Tarsal fold absent. Super-
numerary tubercles present on toes and foot. Tarsal tubercle absent. Small tubercles with horny spinules on dorsal
and lateral parts of head and body and dorsal side of flank. Lower flank granular. Dorsolateral fold absent. Dorsal
and lateral parts of upper arm, lower arm, thigh, shank and foot smooth. Throat, chest and belly granular, underside
of thigh smooth. Nuptial pad present on inner edge of 1st finger and on prepollex, creamy yellow, oval. Vocal sacs
and internal vocal slits present.
FIGURE 4. Pseudophilautus schneideri (WHT 6127), in life, Sinharaja World Heritage Site.
Coloration in life. Head dorsally light brown, laterally dark brown. Supratympanic area, upper tympanum and
interorbital area black. Mid-dorsum dark brown with tiny black and dark brown spots. Both upper and lower areas
of flank pale brown with dark-brown spots. Inguinal area yellowish brown with dark-brown spots. Dorsal and lat-
eral parts of limbs pale brown with dark-brown spots. Lower arm with 1, thigh with 3 and shank with 3 dark-brown
crossbars. Fingers and toes pale yellowish brown with dark-brown pigments. Outer edge of ventral side of foot
dark brown. Ventral side of foot pale yellowish brown with dark-brown patches. Throat, margins of throat, chest,
belly, underside of thigh and webbing pale yellowish with dark-brown pigments.
Coloration in alcohol (description based on holotype, WHT6355). Snout dorsally pale brown, laterally dark
brown. Supratympanic area, middle of tympanum and interorbital area dark brown. Mid-dorsum dark brown with
tiny dark spots. Both upper and lower areas of flank pale brown with dark-brown spots. Inguinal zone yellowish
pale brown. Both upper and lower lips dark brown. Dorsal and lateral parts of limbs pale brown with dark-brown
spots. Lower arm with 1, thigh with 1 and shank with 2 dark-brown cross-bars. Fingers and toes pale yellowish
brown with dark-brown pigments. Outer edge of ventral side of foot dark brown. Ventral side of foot pale yellowish
brown with dark-brown patches. Throat, margins of throat, chest, belly, underside of thigh and webbing pale yel-
lowish with dark-brown pigments. Paratype, WHT6354: A thin yellow line on mid dorsum from tip of snout to
vent, on mid-thigh, on mid-flank and rear edge of foot.
Measurements of holotype (WHT6355, in mm): DBE, 8.2; DFE, 5.0; DL, 1.0; DW, 1.2; ED, 3.4; EN, 2.6; ES,
4.1; FEL, 10.7; FL I, 1.8; FL II, 2.3; FL III, 3.4; FL IV, 3.1; FOL, 14.4; HL, 9.7; HW, 8.9; IML, 1.0; IN, 2.3; IO,
2.3; LAL, 4.5; MBE, 2.9; MFE, 6.0; MN, 9.2; NS, 1.6; PAL, 6.6; SVL, 22.8; TBL, 11.4; TL I, 1.6; TL II, 2.0; TL
III, 3.2; TL IV, 4.3; TL V, 3.4; TYD, 0.5; TYE, 1.5; UAW, 3.6; UEW, 2.3.
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TWO NEW SRI LANKAN SHRUB FROGS
Etymology. The species name is a patronym in the genitive singular in honour of the evolutionary biologist
and herpetologist Professor Christopher J. Schneider (Department of Biology, Boston University, USA).
Remarks. Morphologically, P. schneideri resembles P. folicola. It is distinguished from the latter, however, by
possessing an obtusely pointed snout in lateral aspect (vs. snout rounded or truncate in lateral aspect); having the
loreal region flat (vs. concave); lacking (vs. possessing) a lateral dermal fringe on fingers; lacking (vs. possessing)
supernumerary tubercles on the palm; and having the underside of the thigh smooth (vs. granular).
FIGURE 5. Pseudophilautus schneideri n. sp.: a, lateral; b, dorsal; and c, ventral aspects, respectively, of head of holotype,
male, WHT 6355, 22.8 mm SVL. Scale bar: 1 mm.
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FIGURE 6. Pseudophilautus schneideri n. sp.: a, ventral aspect of left manus; b, ventral aspect of left pes; and c, semi-dia-
grammatic representation of the left-pes webbing pattern of the holotype, male,WHT 6355, 22.8 mm SVL. Scale bar: 1 mm.
Distribution. We observed males of P. schneideri perched on leaves of shrubs, 0.5–1.0 m above ground in
open habitats (the species has hitherto not been observed in closed-canopy rainforest). Individuals were observed
also in forest edges and in anthropogenic habitats such as home gardens and tea plantations. Most individuals were
observed in the undergrowth of neglected tea plantations with extensive weedy undergrowth adjacent to rainforest
(the species is apparently absent in plantations from which the undergrowth has been cleared. Considering its cur-
rent distribution, however, the extent of available habitat for P. schneideri is relatively high: it may indeed have a
much wider distribution than recorded here. Given that this species uses tea plantations as a surrogate for scru-
bland, intensive agro-chemical usage in this habitat may affect these populations.
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TWO NEW SRI LANKAN SHRUB FROGS
FIGURE 7. Distribution of Pseudophilautus schneideri, n. sp. (circles) and Pseudophilautus hankeni, n. sp., (squares), in Sri
Lanka.
Pseudophilautus hankeni, new species
(Figs. 7–10)
Material examined. Holotype: mature male, 21.9 mm SVL, WHT6304, Bambarella, Knuckles, alt. 1490 m
(7°26’N, 80°46’E), M.M., coll. 23 V 1999.
Paratypes: mature males, 19.0 mm SVL, WHT6310; 19.6 mm SVL, WHT6300; 20.7 mm SVL, WHT6302; 20.0
mm SVL, WHT6298; 21.4 mm SVL, WHT6295; 18.3 mm SVL, WHT6308; Bambarella, Knuckles, alt. 1600 m
(7° 25’N, 80° 47’E), coll. 23 V 1999, M.M..
Diagnosis. Philatus hankeni is distinguished from all Sri Lankan congeners by the combination of the follow-
ing characters: size small, mature individuals 18.3–21.9 mm SVL; tympanum distinct; canthal edges rounded;
vomerine ridge absent; outer edge of lower arm and outer edge of foot with a tuberculated dermal fold; throat, chest
and belly granular. (For diagnosis from its sister species, P. schmarda, see Remarks, below.)
Description. (based on holotype, WHT6304, and six paratypes, WHT6310, WHT6300, WHT6302,
WHT6298, WHT6295, WHT6308). Pseudophilautus hankeni is assigned to the genus Pseudophilautus as they are
well nested (Figs. 1 & 2) within the Sri Lankan monophyletic group of frogs (Meegaskumbura et al. 2002; Bossuyt
et al. 2004) and are characterized by terrestrial direct development (Bossuyt and Dubois 2000). Body stout. Head
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convex in lateral view. Snout rounded in dorsal view, pointed in lateral view. Canthal edges rounded. Loreal region
concave. Interorbital space convex. Internasal space concave. Nostrils oval.
Pupil oval, horizontal. Tympanum distinct, oval, oblique, its outer rim indistinct. Pineal ocellus absent. Vom-
erine ridge absent. Tongue moderate, emarginate, not bearing a lingual papilla. Supratympanic fold distinct.
Cephalic ridges absent. Co-ossified skin on head absent. Upper arm short, lower arm short, strong. Fingers thin.
Relative length of fingers, 1 < 2 < 4 < 3. Tips of fingers with discs bearing circum-marginal grooves. Fingers with-
out lateral dermal fringe. Outer edge of 4th finger and outer edge of lower arm with tuberculated dermal fold. Web-
bing on fingers absent. Subarticular tubercles on fingers prominent, oval, single, some absent IV 2 (penultimate
subarticular tubercle). Prepollex oval, distinct. Two palmar tubercles, oval, distinct. Supernumerary tubercles pres-
ent on palm. Thigh slender. Shank slender. Toes strong. Relative length of toes, 1 < 2 < 3 < 5 < 4. Tips of toes with
discs, with circum marginal groves. Webbing present on toes (absent on toe I). Subarticular tubercles on toes prom-
inent, oval, single, some absent IV 2. Inner metatarsal tubercle distinct, oval. Tarsal fold present. Outer metatarsal
tubercle absent. Outer edge of 5th toe and outer edge of foot with a tuberculated dermal fold. Supernumerary tuber-
cles present on foot. Tarsal tubercle present. Dorsal and lateral parts of head and body and upper part of flank with
glandular warts bearing horny spinules. Lower part of flank granular. Dorsolateral fold absent. Dorsal and lateral
parts of upper arm, lower arm, thigh, shank and foot with glandular warts with horny spinules. Nuptial pad absent,
but pale yellow subdermal glands on inner edge of 1st finger and on prepollex. Throat, chest and belly granular.
Underside of thigh and shank smooth. Vocal sacs and internal vocal slits present.
Coloration in life. Dorsal and lateral parts of head brown with symmetrical darker brown patches. Upper areas
of eyelids and posterior area of interorbital with distinct black patches. Upper flank grayish brown, lower flank pale
yellow-gray. Inguinal zone pale yellow with dark-brown patches. Loreal region brown. Tympanic region and tym-
panum brown with small darker brown patches. Both upper and lower lips pale brown. Dorsal and lateral parts of
limbs brown. Lower arm with 2, thigh with 2, shank with 3 and foot with 4 wide, brown patches. Throat, margins
of throat, chest and belly pale yellow or white. Underside of thigh and webbing pale yellow. Fingers and toes dor-
sally light brown.
FIGURE 8. Pseudophilautus hankeni (WHT 6295), in life, Bambarella.
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TWO NEW SRI LANKAN SHRUB FROGS
FIGURE 9. Pseudophilautus hankeni n. sp.: a, lateral; b, dorsal; and c, ventral aspects, respectively, of head of holotype, male,
WHT 6304, 21.9 mm SVL. Scale bar: 1 mm.
Coloration in alcohol (based on holotype, WHT6304). Dorsal and lateral parts of head grayish brown with
symmetrical brown patches. Upper areas of eyelids and posterior area of interorbital with distinct black patches.
Upper part of flank grayish brown, lower part pale gray. Inguinal zone white with dark-brown patches. Loreal
region gray. Tympanic region and tympanum grayish brown. Both upper and lower lips pale gray. Dorsal and lat-
eral parts of limbs gray. Lower arm with 2, thigh with 2, shank with 3 and foot with 4 wide brown patches. Throat,
margins of throat, chest and belly pale gray or white. Underside of thigh and webbing pale yellow. Fingers and toes
dorsally pale gray.
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FIGURE 10. Pseudophilautus hankeni n. sp.: a, ventral aspect of left manus; b, ventral aspect of left pes; and c, semi-diagram-
matic representation of the left-pes webbing pattern of the holotype, male, WHT 6304, 21.9 mm SVL. Scale bar: 1 mm.
Measurements of holotype (WHT6304, in mm): DBE, 7.4; DFE, 4.6; DL, 0.8; DW, 1.3; ED, 2.7; EN, 2.2; ES,
3.8; FEL, 9.6; FL I, 1.6; FL II, 2.2; FL III, 4.0; FL IV, 3.0; FOL, 13.1; HL, 8.4; HW, 8.5; IML, 0.9; IN, 2.0; IO, 2.1;
LAL, 3.9; MBE, 2.8; MFE, 5.2; MN, 7.3; NS, 1.2; PAL, 6.4; SVL, 21.9; TBL, 10.1; TL I, 1.6; TL II, 2.0; TL III,
3.2; TL IV, 4.8; TL V, 3.3; TYD, 0.6; TYE, 1.1; UAW, 4.2; UEW, 1.8.
Etymology. The species name honors the developmental biologist, director of the Museum of Comparative
Zoology (Harvard University, USA), and Alexander Agassiz Professor of Zoology, James Hanken. It is Latinized
as a noun in the genitive singular case.
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TWO NEW SRI LANKAN SHRUB FROGS
Remarks. Pseudophilautus hankeni resembles P. schmarda morphologically, but it can be distinguished from
the latter species as follows: canthal edges rounded (vs. sharp in P. schmarda); toes with (vs. without) a lateral der-
mal fringe; rudimentary webbing present (vs. absent) on toes; and presence (vs. absence) of a subdermal nuptial
pad.
Distribution. Pseudophilautus hankeni shows a very restricted distribution, being known hitherto only from its
type locality at Bambarella and Riverston regions in Knuckles. The species seems to be restricted to the highest
elevations of the Knuckles mountains (peaks over 1200 m elevation). It was observed on the leaves of shrubs (0.3–
1.0 m above ground) under the cover of the montane cloud forest canopy. Given its restricted, high-elevation habi-
tat, it may be at risk from climate warming, further encroachments into the forest from lower elevations, and agro-
chemical usage in nearby tea plantations.
Discussion
Given its extremely restricted range and specialized montane-forest habitat, P. hankeni is clearly at a high risk of
extinction from habitat loss and change. Meegaskumbura & Manamendra-Archchi (2005) highlighted and dis-
cussed the conservation implications of six such Pseudophilautus species with similar upper montane distributions.
Pseudophilautus hankeni also faces these same threats, such as further restriction of habitat due to future global
warming, and given that it already occurs on the highest peaks—the unavailability of yet higher and cooler habi-
tats. Pseudophilautus schmarda, the sister species of P. hankeni, lives on high elevation mountains of the island’s
Central Hills. The two species, however, are separated from each other by the deep (and therefore warmer) valley
of the Mahaweli River, which apparently forms a barrier either is unlikely to cross. The available area of forest for
the Endangered P. schmarda, however, is much greater than that for P. hankeni, making the latter more vulnerable
to extinction.
Pseudophilautus schneideri is a forest-edge species, so far found only from the undergrowth of tea estates
adjacent to secondary forests and also at night perched on shrubs along open trails. Because it has not been
observed more than 100m away from the forests, it probably depends on the natural forest for increased humidity,
lower temperature and other biological resources such as food species. However, given this frog’s tolerance of sec-
ondary habitats, given suitable conditions, it may be able to traverse anthropogenic habitats and plantations. Future
exploration is likely to show that the range of this species is substantially larger than is recorded at present.
Mascaro et al. (2008), show that native animals of a region could survive sometimes even in non-pristine envi-
ronments. Such species may not be new recruits to the environment, but survivors (“relicts”) of a past destruction
of their original habitat. Clearly, this seems to be the case also for some of the Sri Lankan shrub frogs such as P.
schneideri and P. tanu; but given increased risk, such as noted above, that these species face in a non-native habitat
such as tea plantations, management and education of the public could play an important role in their conservation.
Pseudophilautus hankeni and P. schmarda are sister species (Figs. 3) but are separated from each other by sub-
stantial genetic distances. The combined 12s and 16s uncorrected percent genetic distance between P. schamarda
and P. hankeni is 2.43, and for cytochrome-b it is 8.44. The closest species to WHT2667 (P. schneideri) are P. zorro
and P. limbus; uncorrected percent genetic distance for the combined 12s and 16s gene fragment for P. schneideri
and these species are 3.5–4.0; for cytochrome-b it is 15.9–21.8. These genetic distances are well over the 2% cyto-
chrome-b genetic distance that indicates species-level divergence in several groups of mammals (Bradley & Baker
2001). Johns & Avise (1998) also indicate that 90% of the putative sister species across a wide range of vertebrate
taxa showed more than 2% divergence in the cytochrome-b gene, adding confidence to our recognition of these
taxa at the species level. The 16s rRNA gene has been recognized as a suitable barcoding gene for amphibians
(Vences et al. 2005); for instance in Mantellidae, a wide range of inter-species genetic distances, ranging from 1–
16.5%, has been recorded (Vences et al. 2005), and a 3% divergence has been proposed as a species-level threshold
(Fouquet et al. 2007).
Acknowledgements
Prof. Christopher J. Schneider (Boston University) and Prof. James Hanken (Harvard University) are honored,
through naming these two frogs, for their contribution to Sri Lanka’s amphibian conservation through supporting
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research work and training students from Sri Lanka. We are very grateful to Rohan Pethiyagoda for his support in
all aspects of this study, and for reviewing the manuscript. We also thank Suyama Meegaskumbura (University of
Peradeniya, Sri Lanka) for comments that helped significantly to improve the manuscript; Mohomed Bahir and
Sudath Nanayakkara for fieldwork and support at WHT’s Agrapatana field station; Sudesh Batuwita (WHT) for
assistance in the field and hospitality while working in the lowland areas. An Anonymous Reviewer and Miguel
Vences are acknowledged for suggestions that improved the paper significantly. This study was supported finan-
cially by US National Science Foundation (Grant No: 0345885 to CJS and JH) and National Geographic Society
(Grant No: 7612-04 to CJS). We are very grateful to the Forest Department of Sri Lanka for permits to work in their
reserves, enthusiastic support, and accommodation during visits; and the Department of Wildlife Conservation Sri
Lanka for collection and export permits. MM is also thanks the Ziff Environmental Postdoctoral Fellowship
through the Harvard University Center for the Environment (HUCE), which facilitated part of this work.
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