Integrative taxonomy reveals six new species of day geckos of the genus Cnemaspis Strauch, 1887 (Reptilia: Squamata: Gekkonidae) from geographically-isolated hill forests in Sri Lanka

Abstract

Six new day gecko species of the genus Cnemaspis Strauch, 1887 are described from geographically isolated forested hills (Bambarabotuwa, Kadugannawa, Kokagala, Kudumbigala, Maragala and Walapane) in Sri Lanka based on analyses of morphological and molecular traits. We provide an updated mtDNA-based genealogy of Sri Lankan Cnemaspis and provide further evidence that diversity of the genus in the island may still be underestimated. The six new Cnemaspis species described herein are small to medium (27 – 40 mm SVL) in size and can be differentiated from all other Sri Lankan congeners by a suite of distinct morphometric, meristic and molecular characteristics. They are recorded from wet, cool, spacious granite caves found within rock outcrops embedded in forests distributed across low and mid-elevations (~ 25 – 600 m) with minimal anthropogenic disturbance. Existing data suggest that each of these geckos have a highly restricted (point endemic) distribution ranges. Further, their area of occurrence, extent of distribution, and relative a abundance appear to be low, thus all these species are categorized as Critically Endangered (CR) under IUCN Red List criteria. With the descriptions of these species, the number of Cnemaspis described from Sri Lanka increases to 32, all of which are endemic to the island. The discovery of these new species highlights the understudied diversity of geckos in isolated hills. Being rupicolous microhabitat specialists with a scansorial mode of life, these species are susceptible to both localized and widespread threats. Therefore, isolated hill forests of Sri Lanka, especially in the intermediate and dry zones, warrant special conservation, habitat protection, indepth research and specific management actions.

Full text

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ISSN 1864-5755 | eISSN 2625-8498 | DOI: 10.26049/VZ69-3-2019-02
69
(3): 247 – 298
2019
© Senckenberg Gesellschaft für Naturforschung, 2019.
Integrative taxonomy reveals six new species of day
geckos of the genus Cnemaspis Strauch, 1887 (Reptilia:
Squamata: Gekkonidae) from geographically-isolated
hill forests in Sri Lanka
S K1,
*, N A. P 2, 3, A  S 4, M
M 5, M B 6, 7, V A. G 2, T S 8,
D G 6, K D.B. U 9 & A M. B 10
1
Nature Explorations and Education Team, No: B-1 / G-6, De Soysapura Flats, Moratuwa 10400, Sri Lanka —
2
Department of Vertebrate
Zoology, Lomonosov Moscow State University, Leninskiye Gory, GSP – 1, Moscow 119991, Russia —
3
Joint Russian-Vietnamese Tropical
Research and Technological Center, 63 Nguyen Van Huyen Road, Nghia Do, Cau Giay, Hanoi, Vietnam —
4
Amphibia and Reptile Research
Organization of Sri Lanka, 15/1, Dolosbage Road, Gampola, Sri Lanka —
5
Victorian Herpetological Society, P.O. Box 4208, Ringwood, VIC
3134, Australia. —
6
Biodiversity Conservation Society, 150/6, Stanly Thilakaratne Mawatha, Nugegoda 10250, Sri Lanka —
7
Central Envi-
ronmental Authority, 104, Denzil Kobbekaduwa Mawatha, Battaramulla 10120, Sri Lanka —
8
Department of Biological Sciences, Bridgewater
State University, Bridgewater, MA, USA —
9
Department of Biological Sciences, Faculty of Applied Sciences, Rajarata University of Sri Lanka,
Mihintale 50300, Sri Lanka —
10
Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, PA 19085, USA — * Corre-
sponding author: suranjan.karu@gmail.com
Submitted March 12, 2019.
Accepted June 20, 2019.
Published online at www.senckenberg.de/vertebrate-zoology on August 15, 2019.
Published in print Q4/2019.
Editor in charge: Raffael Ernst
Abstract
Six new day gecko species of the genus Cnemaspis Strauch, 1887 are described from geographically isolated forested hills (Bambarabotuwa,
Kadugannawa, Kokagala, Kudumbigala, Maragala and Walapane) in Sri Lanka based on analyses of morphological and molecular traits.
We provide an updated mtDNA-based genealogy of Sri Lankan Cnemaspis and provide further evidence that diversity of the genus in the
island may still be underestimated. The six new Cnemaspis species described herein are small to medium (27 – 40 mm SVL) in size and can
be differentiated from all other Sri Lankan congeners by a suite of distinct morphometric, meristic and molecular characteristics. They are
recorded from wet, cool, spacious granite caves found within rock outcrops embedded in forests distributed across low and mid-elevations
(~ 25 – 600 m) with minimal anthropogenic disturbance. Existing data suggest that each of these geckos have a highly restricted (point en-
demic) distribution ranges. Further, their area of occurrence, extent of distribution, and relative a abundance appear to be low, thus all these
species are categorized as Critically Endangered (CR) under IUCN Red List criteria. With the descriptions of these species, the number of
Cnemaspis described from Sri Lanka increases to 32, all of which are endemic to the island. The discovery of these new species highlights
the understudied diversity of geckos in isolated hills. Being rupicolous microhabitat specialists with a scansorial mode of life, these species
are susceptible to both localized and widespread threats. Therefore, isolated hill forests of Sri Lanka, especially in the intermediate and dry
zones, warrant special conservation, habitat protection, indepth research and specic management actions.
Key words
Anthropogenic threats, biogeography, conservation, endangered species, granite caves, microhabitat, natural history, mtDNA, morphology,
point endemic, systematics, wildlife.
Introduction
In recent years, the number of day gecko species rec-
ognized in the Afro-Asian genus Cnemaspis has grown
rapidly, starting from 39 species in 2000 (Das & Bauer,
2000) and now exceeding 150 species (uetz et al., 2019),
making it the third most diverse gecko genus in the Old
World after Cyrtodactylus and Hemidactylus (Grismer et

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
248
al., 2014a; uetz et al., 2019). Despite, the highly conserv-
ative body morphology observed in Cnemaspis species
throughout their broad distribution, molecular phyloge-
netic evidence supports polyphyly, with three independ-
ent derivations of the Cnemaspis-morph in Africa, South
Asia, and Southeast Asia (GamBle et al., 2012; Grismer
et al., 2014a; WooD et al., 2013). Recent additions of new
species have mostly derived from Southeast Asia, espe-
cially Indonesia, Laos, Thailand, and Malaysia (Chan et
al., 2010; Grismer, 2010b; Grismer et al., 2010a, 2014a;
Grismer & Chan, 2010; WooD et al., 2013). A number of
new species have been reported from the Indian mainland
(CyriaC & umesh, 2014; CyriaC et al., 2018; mirza et al.,
2014; sayyeD et al., 2016, 2018; srinivasulu et al., 2015)
as well as from Sri Lanka (BatuWita & uDuGampala,
2017; BatuWita et al., 2019; manamenDra-araChChi et
al., 2007; viDanapathirana et al., 2014; WiCkramasinGhe
& muninDraDasa, 2007; WiCkramasinGhe et al., 2016).
South Asian Cnemaspis are generally diminutive and
slender-bodied; they possess comparatively large, for-
ward and upward-directed eyes with round pupils, broad
attened heads, and widely-splayed limbs bearing elon-
gated, slender digits (DeraniyaGala, 1953; manamenDra-
araChChi et al., 2007). These geckos are mostly rupicol-
ous although a few are arboreal or ground-dwelling. Most
are cryptically patterned, secretive, and either diurnal or
crepuscular in activity period. Microhabitat associations
are mostly restricted to shaded surfaces of rocks, caves,
and trees. The anatomy of members of the genus demon-
strates specialization for a scansorial mode of life with
adaptations for navigating on inclined surfaces. The con-
servative body plan, ecological and behavioural crypsis,
and microhabitat use observed within Cnemaspis have
contributed to the masking of species boundaries and
resulted in taxonomic confusion ( aGarWal et al., 2017;
BatuWita & uDuGampala, 2017; BatuWita et al., 2019;
Bauer et al., 2007; De silva et al., 2019; Das & Bauer,
1998; karunarathna et al., 2019).
Since the resurrection of Cnemaspis by smith (1935),
the South Asian species have undergone much taxonomic
revision (Bauer et al., 2007; manamenDra-araChChi et
al., 2007). Currently, there are 26 species recognized in
Sri Lanka, all of which are endemic to the island (aGar-
Wal et al., 2017; BatuWita & uDuGampala, 2017; Batu-
Wita et al., 2019; De silva et al., 2019; karunarathna et
al., 2019; manamenDra-araChChi et al., 2007; viDanapa-
thirana et al., 2014; WiCkramasinGhe & muninDraDasa,
2007; WiCkramasinGhe et al., 2016). These recent stud-
ies have suggested that the true species richness of Cne-
maspis in Sri Lanka is yet to be documented, as many
areas and habitats remain unexplored (Bauer et al., 2007;
De silva et al., 2019; karunarathna et al., 2019), which
emphasizes the need for continuing taxonomic studies
(aGarWal & karanth, 2015; Bauer et al., 2007). With
this recognition, we conducted eld studies in numerous
unexplored or under-sampled areas of Sri Lanka. Herein,
we report the discovery of six species of Cnemaspis from
three different bioclimatic zones of Sri Lanka that could
not be assigned to known species.
Materials and Methods
Specimens. Museum acronyms follow saBaj pérez
(2015). The type material discussed in this paper is de-
posited in the National Museum of Sri Lanka (NMSL),
Colombo. Specimens were hand caught and were photo-
graphed in life. They were euthanized using halothane and
xed in 10% formaldehyde for two days, washed in run-
ning water and transferred to 70% ethanol for long-term
storage. Tail tips were collected as tissue samples before
xation and were stored in 95% ethanol under cool con-
ditions (10 °C). For comparison, we examined 384 Cne-
maspis specimens (catalogued and uncatalogued) repre-
senting all recognized Sri Lankan species including all
type specimens housed at the National Museum, Sri Lanka
(NMSL), The Natural History Museum, London (BMNH)
and in the privete collection of Anslem de Silva (ADS)
and Aaron Bauer (AMB),, which has been deposited in the
NMSL. Specimens that formerly belonged to the Wildlife
Heritage Trust (WHT) collection and bearing WHT num-
bers are currently deposited in the NMSL, catalogued un-
der their original numbers. Specimens in this study were
collected during a survey on lizards of Sri Lanka under
permit numbers WL/3/2/1/14/12, and WL/3/2/42/18a, b
issued by the Department of Wildlife Conservation and
permit numbers FRC/5, and FRC/6 issued by the For-
est Department of Sri Lanka. Additional information on
morphology and natural history of Sri Lankan Cnemaspis
species was extracted from the relevant literature (aGar-
Wal et al., 2017; BatuWita & uDuGampala, 2017; Batu-
Wita et al., 2019; Bauer et al., 2007; De silva et al., 2019;
karunarathna et al., 2019; manamenDra-araChChi et
al., 2007; viDanapathirana et al., 2014; WiCkramasinGhe
& muninDraDasa, 2007; WiCkramasinGhe et al., 2016).
Assignment of unidentied specimens to species was
based on the presence of shared morphometric and meris-
tic characters (BatuWita & uDuGampala, 2017; BatuWita
et al., 2019; De silva et al., 2019; karunarathna et al.,
2019; manamenDra-araChChi et al., 2007; WiCkramasin-
Ghe & muninDraDasa, 2007).
Morphometric characters. Forty morphometric measure-
ments were taken using a Mitutoyo digital Vernier calli-
per (to the nearest 0.1 mm), and detailed observations of
scales and other structures were made through Leica Wild
M3Z and Leica EZ4 dissecting microscopes. The follow-
ing symmetrical morphometric characters were taken on
the left side of the body: eye diameter (ED), horizontal
diameter of eye ball; orbital diameter (OD), the greatest
diameter of orbit; eye to nostril length (EN), the distance
between anteriormost point of the orbit and the posterior
border of the nostril; snout length (ES), the distance be-
tween anteriormost point of the orbit and the tip of snout;
snout to nostril length (SN), the distance between tip of
snout and the anteriormost point of the nostril; nostril
width (NW), the maximum horizontal width of the nos-
trils; eye to ear distance (EE), the distance between the
posterior border of eye and the anteriormost point of ear

249
VERTEBRATE ZOOLOGY — 69
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opening; snout to axilla distance (SA), the distance be-
tween axilla and tip of snout; ear length (EL), the maxi-
mum length of the ear opening; interorbital width (IO),
the shortest distance between the left and right supracili-
ary scale rows; inter-ear distance (IE) the distance across
the head between the two ear openings; head length (HL),
the distance between posterior edge of mandible and the
tip of the snout; head width (HW), the maximum width
of the head in-between the ears and the orbits; head depth
(HD), the maximum height of the head at the level of the
eye; jaw length (JL), the distance between the tip of snout
and the corner of the mouth; internarial distance (IN), the
smallest distance between the inner margins of nostrils;
snout to ear distance (SED), the distance between the tip
of snout and anteriormost point of the ear; upper-arm
length (UAL), the distance between the axilla and the an-
gle of the elbow; lower-arm length (LAL), the distance
from the elbow to the wrist with palm exed; palm length
(PAL), the distance between the wrist (carpus) and the
tip of longest nger excluding the claw; length of digits
I – V of manus (DLM), the distance between the juncture
of the basal phalanx with the adjacent digit and the tip of
the digit, excluding the claw; snout-vent length (SVL)
the distance between tip of snout and the anterior mar-
gin of vent; trunk length (TRL), the distance between the
axilla and the groin; trunk width (TW), the maximum
width of body; trunk depth (TD), the maximum depth of
body; femur length (FEL), the distance between the groin
and the knee; tibia length (TBL), the distance from the
knee to the heel with ankle exed; heel length (HEL),
the distance between ankle (tarsus) and the tip of longest
toe (excluding the claw) with both foot and tibia exed;
length of pedal digits I – V (DLP), the distance between
the juncture of the basal phalanx with the adjacent digit
and the digit tip, excluding the claw; tail length (TAL),
the distance between the anterior margin of the vent and
the tail tip; tail base depth (TBD), the maximum height
of the tail base; tail base width (TBW), the widest point
of the tail base.
Meristic characters. Thirty one discrete characters were
observed and recorded using Leica Wild M3Z and Leica
EZ4 dissecting microscopes on both the left and the right
side of the body (reported in the form L/R): number of
supralabials (SUP) and infralabials (INF) between the rst
labial scale and the corner of the mouth; number of inter-
orbital scales (INOS), between the left and right supra-
ciliary scale rows; number of postmentals (PM) bounded
by chin scales, 1st infralabial on the left and right and the
mental; number of chin scales (CHS) scales touching
medial edge of infralabials and mental between juncture
of 1st and 2nd infralabials on the left and right; number of
supranasal (SUN) scales between nares; presence of the
postnasal (PON) scales posterior to the naris; presence of
the internasal (INT) scale between supranasals; number of
supraciliary scales (SUS) above the eye; number of scales
between the eye and tympanum (BET) from posterior-
most point of the orbit to anterior-most point of the tym-
panum; number of canthal scales (CAS), number of scales
from posterior-most point of naris to anterior most point
of the orbit; total lamellae on manus I – V (SLM) counted
from rst proximal enlarged scansor greater than twice
width of the largest palm scale, to distalmost lamella at tip
of digits; number of dorsal paravertebral granules (PG)
between pelvic and pectoral limb insertion points along
a straight line immediately left of the vertebral column;
number of midbody scales (MBS) from the centre of mid-
dorsal row diagonally towards the ventral scales; number
of midventral scales (MVS) from the rst scale posterior
to the mental to last scale anterior to the vent; number
of belly scales (BLS) across the venter between the low-
est rows of granular dorsal scales; total lamellae on pes
I – V (SLP), counted from rst proximal enlarged scansor
greater than twice the width of the largest heel scale, to
distalmost lamella at tip of digits; number of precloacal
pores (PCP) anterior to the cloaca; number of femoral
pores (FP) present on the femur; number of non-pored
anterior femoral scales (AFS) counted from distal end of
precloacal pore row to rst femoral pore; number of non-
pored posterior femoral scales (PFS) counted from distal
end of femoral pore row to knee; interfemoral scales (IFS)
number of non-pored scales between innermost femoral
pores on both femurs. In addition, we also evaluated the
texture (smooth or keeled) of the ventral scales, the tex-
ture (homogeneous or heterogeneous) of the dorsal scales,
the number of spinous scales on the anks (FLSP), and
characteristics such as appearance of the caudal scales
(except in specimens with regenerated tails). Coloration
was determined from digital images of living specimens
and also from direct observations in the eld.
Habitat and ecology. New species described herein were
collected during eld surveys conducted in various habi-
tats of Sri Lanka (Fig. 1). During this survey, behavioural
and other aspects of natural history of the focal species
were observed through opportunistic and non-systematic
means. Such observations were done at a minimum dis-
tance of 2 – 3 m away from the focal animals while taking
precautions to avoid disturbances. Eggs were measured
with a Mitutoyo digimatic calliper (Mituyoto Worldwide,
Aurora, Illinois) to the nearest 0.1 mm. The ambient tem-
perature and the substrate temperature were measured
using a standard thermometer and a N19 Q1370 infrared
thermometer (Dick Smith Electronics, Shanghai, China),
respectively. The relative humidity and light intensity
were measured with a QM 1594 multifunction environ-
ment meter (Digitek Instruments Co., Ltd, Hong Kong,
China). To record elevation and georeference species
locations, an eTrex® 10 GPS (Garmin, Johannesburg,
South Africa) was used. Sex was determined by the
presence (male) or absence (female) of precloacal and
femoral pores. The conservation status of the species was
evaluated using the 2001 IUCN Red List Categories and
Criteria version 3.1 (IUCN, 2012). Habitat descriptions
were based on the National Atlas and oristic regions of
Sri Lanka (Gunatileke & Gunatileke, 1990; Survey De-
partment of Sri Lanka, 2007) as well as on direct visual
observations during the survey.

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
250
DNA isolation, PCR amplication and sequencing.
Total genomic DNA was extracted from ethanol-pre-
served tail tissue using standard phenol-chloroform-pro-
teinase K extraction procedures with consequent iso-
propanol precipitation (protocols followed hillis et al.,
1996 and samBrook & DaviD, 2001). The isolated total
genomic DNA was visualized in agarose electrophoresis
in presence of ethidium bromide. The concentration of
total DNA was measured in 1 μl using NanoDrop 2000
(Thermo Scientic, USA), and consequently adjusted to
ca. 100 ng DNA/μL.
To estimate species diversity of Sri Lankan Cne-
maspis and their genealogical realtionships we amplied
a 1041 bp fragment of ND2 mitochondrial gene follow-
ing aGarWal et al. (2017); this gene has been widely
applied in biodiversity surveys and phylogenetic stud-
ies on geckos (e.g. aGarWal et al., 2016; Grismer et al.,
2012, 2014b, 2018; murDoCh et al., 2019; WooD et al.,
2012 and references therein). PCR amplication was per -
formed in 20 μl reactions using ca. 50 ng genomic DNA,
10 nmol of each primer, 15 nMol of each dNTP, 50 nMol
additional MgCl2, Taq PCR buffer (10 mM Tris-HCl,
pH 8.3, 50 mM KCl, 1.1 mM MgCl2 and 0.01% gela-
tine) and 1 U of Taq DNA polymerase. Primers used in
PCR and sequencing followed aGarWal et al. (2017)
and included two forward primers: Metf1, used for
amplification (5’-AAGCTTTCGGGCCCATACC-3’;
maCey et al., 1997), and ND2f17, used for sequenc-
ing (5’-TGACAAAAAATTGCNCC-3’; maCey et al.,
2000), and two reverse primers: CO1R1, used for am-
plication (5’-AGRGTGCCAATGTCTTTGTGRTT-3’;
maCey et al., 1997); and ND2r102, used for sequencing
(5’-CAGCCTAGGTGGGCGATTG-3’; GreenBaum et al.,
2007). The PCR conditions followed aGarWal et al. (2017).
PCR products were loaded onto 1% agarose gels in
presence of ethidium bromide and visualized in agarose
Fig. 1. Currently known distribution of C. butewai sp. nov. (Bambarabotuwa), C. gotaimbarai sp. nov. (Kokagala), C. hitihami sp. nov.
(Maragalakanda), C. kivulegedarai sp. nov. (Walapane), C. kohukumburai sp. nov. (Kadugannawa), and C. nandimithrai sp. nov. (Ku-
dumbigala) and other localities of Sri Lankan Cnemaspis examined in the present study. Since distributions of Cnemaspis clade A and clade
B largely overlap, they are shown separately for each clade. For locality numbers see Table 1. Colors of icons correspond to those in Fig. 2.
Star denotes localities of the new species described herein. Dot in a center of icon indicates the type locality.

251
VERTEBRATE ZOOLOGY — 69
(3) 2019
gel electrophoresis. PCR products were puried using
2 μl of a 1:4 dilution of ExoSapIt (Amersham, UK) per
5 μl of PCR product prior to cycle sequencing. Puried
PCR products were sequenced bidirectionally at the Ge-
netech Sri Lanka Pvt. Ltd. Colombo. The obtained se-
quences were deposited in GenBank under the accession
numbers MK562336 – MK562365 (Tab. 1).
Phylogenetic analyses. The ND2 dataset of aGarWal et
al. (2017) with addition of our newly obtained sequences
was used to examine the matrilineal genealogy of Cne-
maspis in Sri Lanka (summarized in Tab. 1). In total, we
analysed ND2 sequence data for 77 specimens, including
69 samples of ca. 30 species of Sri Lankan Cnemaspis,
ve samples of four Cnemaspis species from southern In-
dia and Indonesia, and three outgroup sequences of other
Gekkonidae representatives which were used to root the
tree.
Nucleotide sequences were initially aligned in
MAFFT v.6 (katoh et al., 2002) with default parame-
ters, and subsequently checked by eye in BioEdit 7.0.5.2
(Hall, 1999) and MEGA 6.0 (tamura et al., 2013) and
slightly adjusted. Mean uncorrected genetic distances
(p-distances) were calculated in MEGA 6.0. MODEL-
TEST v.3.6 (posaDa & CranDall, 1998) was applied to
estimate the optimal evolutionary models for the data
set analysis. The best-tting model for both BI and ML
analyses was the HKY+G model of DNA evolution as
suggested by the Akaike Information Criterion (AIC) for
all three codon partitions of the ND2 gene.
The matrilineal genealogy was inferred using Bayes-
ian inference (BI) and Maximum Likelihood (ML) ap-
proaches. BI was conducted in MrBayes 3.1.2 (ronquist
& huelsenBeCk, 2003); Metropolis-coupled Markov chain
Monte Carlo (MCMCMC) analyses were performed with
one cold chain and three heated chains for twenty million
generations and sampled every 2000 generations. Five
independent MCMCMC iterations were performed and
1000 trees were discarded as burn-in. The convergence
of the iterations was diagnosed by examining the likeli-
hood plots in TRACER v1.6 (ramBaut et al., 2014); the
effective sample sizes (ESS) were all above 200. Nodal
support was assessed by calculating posterior probabili-
ties (BI PP). ML was conducted using the RAxML web
server (http://embnet.vital-it.ch/raxml-bb/; kozlov et al.,
2018). Condence in nodal topology was estimated by
non-parametric bootstrapping (ML BS) with 1000 pseu-
doreplicates (Felsenstein, 1985). Three nodes with BI PP
values > 0.95 and LM BS values ≥ 57% were regarded
as a priori; BI PP value between 0.95 – 0.90 and ML BS
values between 75 – 50% were regarded as tendencies.
Lower values were regarded as indicating essentially un-
resolved nodes (huelsenBeCk & hillis, 1993).
Ordination of morphometric data. All morphometric
measurements of the six new species were normalized to
the snout-vent length (SVL). A principal component anal-
ysis (PCA) was performed on the normalized morpho-
metric measurements via a singular-value decomposition
algorithm. All morphometric variables were centered
and scaled to reach unit variance. Two principal compo-
nents (PCs) were extracted that explained a substantial
proportion of the overall variability of the original vari-
ables. Based on those two PCs, an ordination plot was
produced to visualize the separation of the species. PCA
was performed using R (PCA: function prcomp, base
package; ordination: ggbipplot function, ggbiplot pack -
age) (R Development Core team, 2018).
Results
Phylogenetic analyses
Sequence and statistics. The nal alignment of the ND2
gene partial sequences contained 1041 aligned charac-
ters, of which, 401 sites were conserved and 637 sites
were variable, of which 540 were found to be parsimony-
informative. The transition – transversion bias (R) was
estimated as 3.49. Nucleotide frequencies were 35.29%
(A), 21.71% (T), 32.1% (C), and 10.83% (G) (all data
given for ingroup only).
ND2 gene genealogy. BI and ML analyses generated es-
sentially similar topologies, with relationships varying
only in several poorly supported nodes; most of the nodes
in the tree were well-resolved and supported (Fig. 2). The
BI genealogy (Fig. 2) inferred the following set of phylo-
genetic relationships, which is generally consistent with
the results of aGarWal et al. (2017):
All examined samples of Cnemaspis were clustered
into two major groups, which we indicate as Clade A (po-
dihuna clade of aGarWal et al., 2017) and Clade B (kan-
diana clade of aGarWal et al., 2017). These two groups
are reciprocally monophyletic with high node support
(1.0/100%; hereafter node support values are given for
BI PP/ML BS, respectively). The Clade A includes an un-
described species Cnemaspis sp. 9 from Kerala State, In-
dia (southern part of Western Ghats), which forms a sister
group with respect to other members of the clade A, all
of which occur in Sri Lanka. The Sri Lankan members of
Clade A are clustered in two reciprocally monophyletic
subclades: the species group AI, joining C. phillipsi and
C. scalpensis (sister species; for both of them we ana-
lyzed topotype specimens), C. gemunu and undescribed
candidate species Cnemaspis sp. 7 from Ratnapura Dis-
trict (sister species); the second subclade joins the re-
maining species of the Clade A (species groups AII and
AIII) (Fig. 2). The second subclade of Clade A includes
C. podihuna (AII; Fig. 2), which is a sister lineage with
respect to other species (AIII), genealogical relation-
ships among which are poorly resolved. Cnemaspis po-
dihuna in our analysis comprises four divergent mtDNA
lineages, indicating that C. podihuna is likely a cryptic
species complex. None of the analysed C. cf. podihuna
lineages included material from the type locality of this

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252
Table 1. Sequences and voucher specimens of Cnemaspis and outgroup taxa used in this study. For sampling localities in Sri Lanka see Fig.
1. Locality codes are given separately for members of clades A (A1 – A16) and B (B1 – B31). Sequences generated in this study are marked
with an asterisk (*); n-dash (—) denotes no data available. (Continues on next page).
No. Genbank A.N. Specimen ID Species Country Locality
1 KY038004 WHT7334 Cnemaspis cf. podihuna Sri Lanka A1 Monaragala District,
Maligawila
2 KY038005 58A Cnemaspis cf. podihuna Sri Lanka A2 Monaragala District,
Kukulagoda
3 KY038006 AMB7449 Cnemaspis cf. podihuna Sri Lanka A3 Anuradhapura District,
Mihintale
4 KY038002 70A Cnemaspis podihuna Sri Lanka A4 Budulla District,
Kuruwekotha
5 KY038003 71A Cnemaspis podihuna Sri Lanka A4 Budulla District,
Kuruwekotha
6 KY037997 AMB7447 Cnemaspis alwisi Sri Lanka A5 Anuradhapura District,
Ritigala
7 MK562336 NMSL2019.05.01 Cnemaspis kohukumburai sp. nov. Sri Lanka A6 Kandy District,
Kadugannawa
8 KY038010 AMB7436 Cnemaspis nilgala Sri Lanka A7 Monaragala District,
Pitakumbura, Serawa
9 KY038009 AMB7418 Cnemaspis nilgala Sri Lanka A7 Monaragala District,
Pitakumbura, Serawa
10 KY038011 47A Cnemaspis sp. 8 Sri Lanka A8 Monaragala District,
Maligathenna
11 KY038012 WHT5918 Cnemaspis hitihami sp. nov. Sri Lanka A9 Monaragala District,
Kumaradola estate
12 MK562337 NMSL2019.06.01 Cnemaspis hitihami sp. nov. Sri Lanka A9 Monaragala District,
Maragala
13 MK562338 NMSL2019.06.02 Cnemaspis hitihami sp. nov. Sri Lanka A9 Monaragala District,
Maragala
14 MK562339 NMSL2019.06.03 Cnemaspis hitihami sp. nov. Sri Lanka A9 Monaragala District,
Maragala
15 KY038007 AA80 Cnemaspis punctata Sri Lanka A10 Matale District, Rattota,
Gammaduwa
16 KY037999 WHT7348 Cnemaspis gemunu Sri Lanka A11 Nuwara Eliya District, Near
Hakgala
17 MK562340 ADS 217 Cnemaspis gemunu Sri Lanka A12 Nuwara Eliya District,
Ohiya
18 MK562341 ADS 216 Cnemaspis gemunu Sri Lanka A12 Nuwara Eliya District,
Ohiya
19 MK562342 ADS 218 Cnemaspis gemunu Sri Lanka A12 Nuwara Eliya District,
Ohiya
20 KY037998 AMB7495 Cnemaspis gemunu Sri Lanka A11 Nuwara Eliya District,
Hakgala
21 KY038000 AMB7507 Cnemaspis sp. 7 Sri Lanka A13 Ratnapura District,
Borangamuwa
22 KY038001 AA81 Cnemaspis phillipsi Sri Lanka A14 Matale District, Rattota,
Gammaduwa
23 MK562343 ADS 220 Cnemaspis phillipsi s.str. Sri Lanka A15 Matale District,
Gammaduwa
24 KY038008 WHT7268 Cnemaspis scalpensis Sri Lanka A16 Kandy District, Gannoruwa
25 MK562344 ADS 219 Cnemaspis scalpensis s.str. Sri Lanka A16 Kandy District, Gannoruwa
26 MK562351 NMSL2019.07.01 Cnemaspis butewai sp. nov. Sri Lanka B1 Ratnapura District,
Bambarabotuwa
27 MK562352 NMSL2019.07.02 Cnemaspis butewai sp. nov. Sri Lanka B1 Ratnapura District,
Bambarabotuwa
28 MK562353 NMSL2019.07.03 Cnemaspis butewai sp. nov. Sri Lanka B1 Ratnapura District,
Bambarabotuwa
29 KY037992 AMB7529 Cnemaspis sp. 3 Sri Lanka B2 Galle District, Haycock
30 MK562354 ADS 205 Cnemaspis pulchra s.str. Sri Lanka B3 Ratnapura District,
Morningside
31 MK562355 ADS 206 Cnemaspis pulchra s.str. Sri Lanka B3 Ratnapura District,
Morningside

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No. Genbank A.N. Specimen ID Species Country Locality
32 MK562356 ADS 207 Cnemaspis pulchra s.str. Sri Lanka B3 Ratnapura District,
Morningside
33 KY037991 AMB7508 Cnemaspis sp. 2 Sri Lanka B4 Ratnapura District,
Masimbula, Godakawela
34 KY037993 AA87 Cnemaspis sp. 4 Sri Lanka B5 Budulla District, Haputale
35 KY037994 AA87B Cnemaspis sp. 4 Sri Lanka B5 Budulla District, Haputale
36 MK562348 NMSL2019.08.01 Cnemaspis kivulegedarai sp. nov. Sri Lanka B6 Nuwara Eliya District,
Walapane
37 MK562349 NMSL2019.08.02 Cnemaspis kivulegedarai sp. nov. Sri Lanka B6 Nuwara Eliya District,
Walapane
38 MK562350 NMSL2019.08.03 Cnemaspis kivulegedarai sp. nov. Sri Lanka B6 Nuwara Eliya District,
Walapane
39 KY037976 WHT7214 Cnemaspis latha Sri Lanka B7 Nuwara Eliya District,
Bandarawela
40 MK562360 ADS 208 Cnemaspis kumarasinghei s.str. Sri Lanka B8 Monaragala District,
Maragala
41 MK562361 ADS 209 Cnemaspis kumarasinghei s.str. Sri Lanka B9 Monaragala District,
Maragala
42 KY037974 AMB7431 Cnemaspis kumarasinghei Sri Lanka B10 Monaragala District,
Rahathankanda (Buttala)
43 MK562358 ADS 210 Cnemaspis kumarasinghei Sri Lanka B11 Badulla District, Udakiruwa
44 MK562359 ADS 212 Cnemaspis kumarasinghei Sri Lanka B11 Badulla District, Udakiruwa
45 KY037975 AA13 Cnemaspis cf. kumarasinghei Sri Lanka B12 Budulla District, Tonacombe
Estate (Namunukula)
46 MK562357 ADS 211 Cnemaspis cf. kumarasinghei Sri Lanka B11 Badulla District, Udakiruwa
47 MK562362 NMSL2019.03.01 Cnemaspis nandimithrai sp. nov. Sri Lanka B13 Ampara District,
Kudumbigala
48 MK562363 NMSL2019.03.02 Cnemaspis nandimithrai sp. nov. Sri Lanka B13 Ampara District,
Kudumbigala
49 MK562364 NMSL2019.04.01 Cnemaspis gotaimbarai sp. nov. Sri Lanka B14 Ampara District, Kokagala
50 KY037984 AA88 Cnemaspis silvula Sri Lanka B15 Galle District, Hiyare forest
reserve
51 KY037990 WHT7331 C. ingerorum Sri Lanka B16 Hambantota District,
Sandagala
52 KY037969 WHT7303 Cnemaspis upendrai Sri Lanka B17 Nuwara Eliya District,
Nanuoya
53 KY037985 WHT7258 Cnemaspis upendrai Sri Lanka B18 Nuwara Eliya District,
Punduloya
54 KY037987 AA12 Cnemaspis upendrai Sri Lanka B19 Nuwara Eliya District, near
Dimbula Junction
55 MK562345 ADS 213 Cnemaspis upendrai Sri Lanka B20 Nuwara Eliya District,
Ramboda
56 KY037988 AMB7488 Cnemaspis upendrai Sri Lanka B21 Kandy District, Helboda,
near Pussellawa
57 KY037986 AA83 Cnemaspis upendrai Sri Lanka B21 Kandy District, Pussellawa
58 KY037979 WHT7261 Cnemaspis pava Sri Lanka B20 Nuwara Eliya District,
Ramboda
59 MK562346 ADS 214 Cnemaspis pava Sri Lanka B20 Nuwara Eliya District,
Ramboda
60 KY037981 AA19 Cnemaspis pava Sri Lanka B22 Nuwara Eliya District,
Ambegamuwa
61 KY037980 AMB7494 Cnemaspis pava Sri Lanka B23 Nuwara Eliya District,
Labookellie
62 KY037983 AMB7505 Cnemaspis samanalensis Sri Lanka B24 Nuwara Eliya District,
Upcot tea estate
63 KY037971 AA57 Cnemaspis kandiana Sri Lanka B25 Kandy District, Gannnoruwa
64 KY037972 AMB7487 Cnemaspis kandiana Sri Lanka B26 Kandy District, Gampola
65 MK562347 ADS 215 Cnemaspis kandiana Sri Lanka B27 Matale District, Rathtota
66 KY037973 AA01 Cnemaspis kandiana Sri Lanka B28 Kandy District,
Loolecondera Tea Estate
Table 1 continued.

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254
species (Lahugala, Eastern Province), thus the identica-
tion as C. podihuna is tentative.
Members of the C. punctata species group (AIII; Fig. 2)
include C. punctata, C. nilgala (listed as Cnemaspis sp.
8 in aGarWal et al., 2017) and its sister undescribed
candidate species Cnemaspis sp. 8 from the Monaragala
District, C. alwisi and its sister species from the Kandy
District, described herein as Cnemaspis kohukumburai
sp. nov., and a new species from the Monaragala Dis-
trict, described herein as Cnemaspis hitihami sp. nov.
(listed as Cnemaspis sp. 10 in aGarWal et al., 2017). The
Clade B shows generally shallower phylogenetic structur-
ing as compared with the Clade A; a number of nodes of
the clade radiation in Sri Lanka remain unresolved. The
basal position in the Clade B is occupied by two unde-
scribed Indian Cnemapsis species (Cnemaspis sp. 5 and
Cnemaspis sp. 6 from Karnataka State) both occurring in
the northern part of the Western Ghats. Cnemaspis sp. 6 is
reconstructed as a sister lineage with respect to all other
members of Clade B, and Cnemaspis sp. 5 is suggested as
a sister lineage of Sri Lankan and Southeast Asian mem-
bers of the clade. The Sumatran species C. modiglianii is
also a member of the kandiana Clade B in full accordance
with the results of Bauer et al. (2007) and aGarWal et al.
(2017); it is reconstructed as a sister lineage with respect
to Sri Lankan radiation of the clade (Fig. 2); monophyly
of the latter receives strong support (0.95/97). Phyloge-
netic relationships among the Sri Lankan species of the
Clade B are essentially unresolved; ve species groups
are supported in ND2 gene genealogy. An undescribed
species, Cnemaspis sp. 1 from the Matara District, occu-
pies an unresolved position in this radiation.
The rst species group BI (0.96/78; Fig. 2) of Clade
B, includes C. kandiana, C. retigalensis, and C. kallima
from the northern part of Central Highlands and Anu-
radhapura District in northern part of Sri Lanka (Fig. 1).
The second species group BII (1.0/100; Fig. 2) of Clade
B comprises C. upendrai, C. pava and C. samanalensis
from the western part of the Central Highlands of Sri
Lanka. The third species group BIII (0.95/78; Fig. 2)
of Clade B, includes taxa from the eastern and southern
parts of Sri Lanka: C. kumarasinghei, C. silvula and sis-
ter species Cnemaspis ingerorum from the Hambantota
District, and two new species from the Ampara District,
which we describe herein as Cnemaspis nandimithrai
sp. nov. and Cnemaspis gotaimbarai sp. nov. The fourth
species group BIV (1.0/100; Fig. 2) of Clade B, includes
C. latha and its sister species, described herein as Cne-
maspis kivulegedarai sp. nov.; both species occur in the
Central Highlands of Sri Lanka (Fig. 1). The fth spe-
cies group BV (1.0/100; Fig. 2) of Clade B, consists of a
number of species from southwestern part of Sri Lanka,
including C. pulchra, three undescribed candidate spe-
cies Cnemaspis sp. 2 (Ratnapura District), Cnemaspis
sp. 3 (Galle District), Cnemaspis sp. 4 (Badulla District),
and a new species from the Ratnapura District which we
describe herein as Cnemaspis butewai sp. nov.
Sequence divergence. The uncorrected p-distances for
the ND2 gene fragment among and within examined Sri
Lankan Cnemaspis species are given in Tab. 2. Intraspe-
cic distances ranged from p = 0% in a number of ex-
amined species to p = 3.5% in the C. kandiana complex
and p = 4.3% in the C. podihuna complex. aGarWal et al.
(2017) applied a conservative threshold of 3.7% uncor-
rected p-distance of ND2 as indicative of putative species-
level divergence. Deep divergence within C. podihuna
and C. kandiana complexes likely indicates an incom-
plete taxonomy of these groups and a more detailed study
including topotype materials on these species is required.
Table 1 continued.
No. Genbank A.N. Specimen ID Species Country Locality
67 KY037982 AMB7448 Cnemaspis retigalensis Sri Lanka B29 Anuradhapura District,
Mihintale
68 KY037970 AA82 Cnemaspis kallima Sri Lanka B30 Matale District, Rattota,
Gammaduwa
69 KY037989 AA17 Cnemaspis sp. 1 Sri Lanka B31 Matara District, Naotunna
(near Thalalla)
70 KY037977 MVZ239314 Cnemaspis modiglianii Indonesia —
Sumatra, Kecematan
Enggano, Pulau Enggano,
near Malakoni
71 KY037978 MVZ239315 Cnemaspis modiglianii Indonesia —
Sumatra, Kecematan
Enggano, Pulau Enggano,
near Malakoni
72 KY038013 SB151 Cnemaspis sp. 9 India — Kerala, Thrissur District,
Athirappilly Falls
73 KY037995 SB048 Cnemaspis sp. 5 India — Karnataka, Kodagu District,
Kumarahalli
74 KY037996 JB239 Cnemaspis sp. 6 India — Karnataka (pet trade)
75 KY038015 2MA29 Urocotyledon inexpectata Seychelles — —
76 MK562365 ADS 198 Gehyra mutilata Sri Lanka — Kandy District,
Kadugannawa
77 KY038014 PL17 Ailuronyx seychellensis Seychelles — —

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The interspecic distances within Sri Lankan Cnemaspis
varied from p = 2.8% (between C. pulchra and Cnemaspis
sp. 3) to p = 29.5% (between C. latha and C. alwisi)
(Tab. 2). The newly discovered lineages of Sri Lankan
Cnemaspis are highly divergent from other congeners
with interspecic distances varying from p = 5.1% (be-
tween Cnemaspis kivulegedarai sp. nov. and C. latha) to
p = 29.4% (between Cnemaspis gotaimbarai sp. nov. and
C. alwisi), and are thus notably higher than the p = 3.7%
divergence proposed by aGarWal et al. (2017) (Tab. 2).
Fig. 2. Bayesian inference tree of Cnemaspis derived from the analysis of 1041 bp of ND2 gene sequences. For voucher specimen infor-
mation and GenBank accession numbers see Table 1. Numbers at tree nodes correspond to BI PP/ML BS support values, respectively;
an asterisk (*) indicates strongly supported nodes (BI PP = 1.0; ML BS > 95%). Outgroup taxa not shown. Colors of clades and locality
numbers correspond to those in Fig. 1. Photos showing the six new species of Cnemaspis described herein taken by Majintha Madawala
and Suranjan Karunarathna.

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
256
Table 2. Uncorrected p-distance (percentage) between the sequences of ND2 mtDNA gene (below the diagonal), estimate errors (above the diagonal) and intraspecic genetic p-distance (on the diagonal) of
Sri Lankan Cnemaspis species included in phylogenetic analyses.
Species 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30
1 C. butewai sp. nov. 1.0 0.6 0.7 0.8 0.9 0.9 0.8 0.8 1.1 0.9 0.9 0.9 0.9 0.8 1.0 0.8 0.9 0.9 0.8 1.4 1.5 1.5 1.5 1.5 1.7 1.4 1.5 1.5 1.6 1.6
2 Cnemaspis sp. 3 4.9 — 0.5 0.6 0.8 0.8 0.7 0.8 0.9 0.9 0.9 0.9 0.8 0.8 0.9 0.8 0.9 0.8 0.7 1.5 1.4 1.5 1.5 1.5 1.7 1.5 1.5 1.5 1.6 1.7
3 C. pulchra 4.6 2.8 0.9 0.6 0.9 0.8 0.8 1.0 1.2 0.9 1.0 1.0 0.8 0.8 0.9 0.8 1.0 0.8 0.8 1.5 1.4 1.5 1.5 1.6 1.7 1.5 1.7 1.7 1.7 1.7
4 Cnemaspis sp. 2 6.1 4.0 3.1 — 0.8 0.7 0.7 0.8 0.9 0.8 0.8 0.9 0.7 0.7 0.8 0.7 0.9 0.8 0.8 1.4 1.5 1.4 1.5 1.5 1.6 1.4 1.5 1.5 1.7 1.7
5 Cnemaspis sp. 4 7.8 7.4 7.1 7.2 0.0 0.7 0.7 0.7 1.0 0.8 0.7 0.9 0.7 0.8 0.8 0.7 0.8 0.7 0.8 1.4 1.4 1.5 1.5 1.5 1.7 1.4 1.5 1.5 1.6 1.6
6 C. kivulegedarai sp. nov. 7.7 7.4 6.6 6.3 6.9 0.0 0.6 0.8 1.0 0.8 0.8 0.8 0.7 0.8 0.8 0.7 0.8 0.7 0.8 1.4 1.5 1.5 1.6 1.5 1.6 1.4 1.4 1.4 1.6 1.6
7 C. latha 8.2 8.1 7.6 7.5 7.4 5.1 — 0.7 1.0 0.8 0.8 0.7 0.7 0.7 0.8 0.7 0.8 0.7 0.7 1.3 1.4 1.5 1.5 1.4 1.5 1.4 1.4 1.3 1.5 1.5
8 C. kumarasinghei 9.9 9.0 9.4 9.0 8.5 8.5 8.9 2.6 0.7 0.7 0.8 0.8 0.7 0.7 0.8 0.7 0.7 0.7 0.7 1.3 1.3 1.4 1.4 1.4 1.5 1.3 1.4 1.3 1.5 1.5
9 C. gotaimbarai sp. nov. 11.2 10.2 10.4 9.9 10.3 10.1 10.1 8.1 — 0.8 0.8 0.8 0.9 0.9 1.0 0.9 1.0 0.9 1.0 1.3 1.3 1.4 1.4 1.4 1.6 1.5 1.5 1.4 1.5 1.6
10 C. nandimithrai sp. nov. 9.4 8.3 8.1 8.2 7.6 7.7 8.1 6.8 7.9 0.7 0.8 0.8 0.8 0.8 0.8 0.8 0.8 0.8 0.7 1.3 1.4 1.4 1.4 1.4 1.5 1.4 1.5 1.4 1.5 1.6
11 C. ingerorum 9.0 8.4 8.4 7.5 7.6 7.2 8.2 8.7 8.6 8.0 — 0.7 0.8 0.8 0.9 0.8 0.9 0.7 0.6 1.4 1.5 1.5 1.5 1.4 1.6 1.5 1.5 1.4 1.6 1.7
12 C. silvula 10.0 8.9 8.9 8.2 8.5 7.8 8.5 8.3 9.0 7.5 5.9 — 0.8 0.8 0.8 0.7 0.8 0.7 0.7 1.4 1.4 1.4 1.5 1.4 1.6 1.4 1.5 1.4 1.6 1.6
13 C. upendrai 8.0 8.0 7.4 7.2 7.8 6.5 7.3 7.7 9.8 8.4 7.6 7.9 0.3 0.6 0.7 0.6 0.8 0.7 0.8 1.4 1.4 1.4 1.5 1.4 1.6 1.3 1.4 1.3 1.6 1.6
14 C. pava 7.5 7.9 7.5 8.0 7.8 6.7 6.7 7.9 9.8 8.2 7.1 8.2 3.7 1.4 0.7 0.6 0.7 0.7 0.8 1.4 1.5 1.5 1.5 1.5 1.6 1.4 1.4 1.5 1.6 1.6
15 C. samanalensis 9.4 9.2 8.7 8.9 8.7 7.8 8.3 8.7 10.8 9.0 9.0 8.7 5.6 5.8 — 0.7 0.9 0.8 0.8 1.4 1.4 1.5 1.5 1.4 1.6 1.3 1.4 1.4 1.6 1.6
16 C. kandiana 8.5 8.3 7.6 8.3 8.0 6.8 7.9 8.3 9.9 7.7 7.9 8.1 6.1 5.9 7.7 3.5 0.7 0.5 0.7 1.4 1.4 1.4 1.4 1.4 1.6 1.3 1.4 1.4 1.5 1.6
17 C. retigalensis 9.7 8.9 8.8 8.8 8.0 7.1 8.8 8.1 10.6 8.5 8.7 8.7 7.4 7.4 8.5 6.5 — 0.7 0.7 1.4 1.4 1.4 1.5 1.5 1.6 1.4 1.5 1.5 1.6 1.6
18 C. kallima 7.8 7.6 6.7 7.4 7.1 5.8 6.5 7.8 9.3 7.3 7.1 8.1 5.7 5.6 6.5 5.4 5.9 — 0.7 1.4 1.4 1.4 1.5 1.4 1.6 1.3 1.4 1.4 1.5 1.6
19 Cnemaspis sp. 1 7.3 7.7 7.0 7.7 6.5 6.7 7.1 7.8 9.6 7.6 6.7 7.3 5.9 6.3 7.0 7.0 7.5 5.4 — 1.4 1.5 1.5 1.5 1.5 1.7 1.3 1.4 1.3 1.6 1.6
20 C. hitihami sp. nov. 27.4 26.9 25.6 26.9 26.6 26.8 27.1 27.4 27.6 27.0 27.1 26.9 25.5 26.3 26.8 25.7 26.5 26.2 25.9 0.2 1.0 1.1 0.9 1.0 1.2 1.1 1.2 1.2 1.2 1.3
21 C. nilgala 27.7 27.7 26.2 27.8 26.8 26.7 27.4 27.4 27.7 27.0 27.9 27.4 26.1 26.7 27.0 26.5 27.8 27.5 26.8 12.4 0.0 0.9 1.0 1.0 1.0 1.0 1.2 1.2 1.2 1.2
22 Cnemaspis sp. 8 28.3 28.1 26.7 27.5 27.2 26.9 27.4 27.3 28.3 27.0 27.8 27.0 26.0 26.6 26.9 26.2 27.6 26.6 26.3 11.9 7.5 — 1.1 1.1 1.0 1.1 1.2 1.3 1.3 1.3
23 C. alwisi 29.4 28.9 28.1 29.1 28.5 28.3 29.5 28.6 29.4 28.8 29.2 29.1 27.2 28.3 28.5 27.9 28.7 28.2 27.5 12.0 13.0 12.2 — 1.0 1.2 1.1 1.2 1.2 1.1 1.1
24 C. kohukumburai sp. nov. 29.1 28.4 28.2 28.9 28.3 28.1 28.5 28.1 29.0 28.3 29.2 29.1 27.5 28.6 28.3 28.0 28.5 27.8 27.7 13.7 14.9 14.3 11.9 — 1.2 1.1 1.4 1.4 1.3 1.3
25 C. punctata 29.4 28.6 27.7 28.2 28.7 28.3 29.0 28.9 29.3 28.1 28.8 29.1 27.0 27.9 29.0 28.5 29.5 28.2 28.0 13.9 13.4 13.1 13.1 14.9 — 1.2 1.3 1.3 1.3 1.3
26 C. podihuna 27.1 26.8 25.6 26.6 26.7 26.0 26.7 27.5 28.9 26.8 26.8 27.0 26.4 26.9 27.0 26.5 27.3 26.0 25.6 16.7 16.8 16.1 17.0 17.6 18.0 4.3 1.1 1.2 1.2 1.2
27 C. gemunu 27.3 26.9 25.8 26.4 26.0 26.3 26.8 26.9 28.2 27.5 27.4 27.6 25.7 26.8 27.1 26.2 27.2 25.9 26.0 18.8 19.5 18.9 18.3 19.3 20.5 19.8 0.4 0.7 0.9 0.9
28 Cnemaspis sp. 7 25.8 25.3 24.3 25.0 24.6 24.4 24.8 24.8 25.9 25.7 25.5 25.8 23.9 25.0 24.9 24.5 25.1 24.0 23.8 16.9 17.8 18.0 17.4 18.3 19.0 18.5 6.8 — 1.0 1.0
29 C. phillipsi 27.7 27.4 26.6 27.2 26.5 26.4 27.2 27.2 27.9 27.6 27.4 27.3 25.9 26.9 27.3 26.3 27.6 26.6 26.3 18.7 17.5 17.6 17.8 19.5 19.0 18.3 10.7 9.7 2.2 0.7
30 C. scalpensis 27.7 27.3 26.6 27.2 26.6 26.6 27.2 27.2 28.2 27.4 27.7 27.7 26.3 27.4 27.9 26.4 27.8 27.3 26.6 18.5 17.9 17.9 18.0 19.9 19.6 18.8 10.6 10.3 4.5 0.2

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Taxonomy
The results of our phylogenetic analyses of ND2 mtDNA
gene fragment are largely concordant with the previously
published trees of Bauer et al. (2007) and aGarWal et al.
(2017). In addition to the data reported in aGarWal et al.
(2017) our sampling adds six more previously unknown
mtDNA lineages of Sri Lankan Cnemaspis, ve of which
correspond to undescribed candidate species. With addi-
tion of C. pulchra for which our study presents the rst
DNA data, C. nilgala and C. ingerorum, originally re-
ported as Cnemaspis sp. 8 and Cnemaspis sp. 2 in the
work by aGarWal et al. (2017) and recently described by
karunarathna et al. (2019) and BatuWita et al. (2019),
respectively, our genealogy includes 18 nominal species
of Sri Lankan Cnemaspis and additionaly reports on 12
currently undescribed candidate species. As we demon-
strate below, six of them show signicant morphological
differences from all other congeners and can be easily
distinguished from currently recognized Cnemaspis spe-
cies occurring in Sri Lanka. Their divergence in ND2
gene fragment from all congeners for which comparable
genetic data are available is greater than 5.2%. These ge-
netic distances are consistent with observed morphologi-
cal differentiation and are greater than the proposed spe-
cies-level divergence threshold of 3.7% of substitutions
in the ND2 gene. These data suggest that these six line-
ages of Cnemaspis from Sri Lanka represent currently
undescribed species new to science which we describe
below.
Morphophonemic analyses
The PCA produced two PCs that collectively explained
65.8% of the overall variability of the morphometric
measurements of the six new species (Fig. 3). The PC 1
explained 40.8 % of the variability while PC 2 accounted
for 25% of the variability. The six new species showed
substantial separation in the ordination space based on
the morphometric data.
Systematics
Cnemaspis nandimithrai sp. nov.
ZooBank urn:lsid:zoobank.org:act:76DAB508-4201-4FB4-B05F-
5AB9BAD6C5D0
Nandimithras’ day gecko (English)
Nandimithrage diva-seri hoona (Sinhala)
Figs. 4 – 6; Table 3.
Fig. 3. Principal component analysis (PCA) on morphometric variables of the six new species. The ellipses represent 95% condent inter-
vals around the centroids.

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
258
Holotype. NMSL.2019.03.01, adult male, 27.9 mm SVL (Fig. 4),
collected from granite cave in Kudumbigala, Kumana, Ampara
District, Eastern Province, Sri Lanka (6.667519° N, 81.747839° E,
WGS1984; elevation 28 m; around 10.00 hrs) on 15 August 2018
by Suranjan Karunarathna and Anslem de Silva.
Paratypes. NMSL.2019.03.02, adult male, 31.7 mm SVL, and
NMSL.2019.03.03, adult female, 29.7 mm SVL, collected from
granite cave in Kudumbigala, Kumana, Ampara District, Eastern
Province, Sri Lanka (6.658708° N, 81.753736° E, WGS1984; el-
evation 31 m; around 11.00 hrs), on 15 August 2018 by Suranjan
Karunarathna and Anslem de Silva.
Diagnosis. Cnemaspis nandimithrai sp. nov., can be
readily distinguished from its Sri Lankan congeners by a
combination of the following morphometric and meristic
characteristics as well as color pattern: maximum SVL
31.7 mm; dorsum with homogeneous, smooth granular
scales; 2/2 supranasals, 2 internasals and 1/1 postnasal
present; 3 enlarged postmentals; postmentals bounded
by 5 chin scales; chin, gular, pectoral and abdominal
scales smooth, subimbricate; 25 – 27 belly scales across
midbody; 3 – 4 weakly developed tubercles on posterior
ank; 95 – 99 paravertebral granules linearly arranged;
2 – 4 precloacal pores, 2 – 4 femoral pores on each side
in males, separated by 9 – 11 unpored anterior femoral
scales, 5 – 7 unpored posterior femoral scales; 108 – 112
ventral scales; 87 – 89 midbody scales; subcaudals
smooth, median row comprising an irregular series of
oval shaped, small scales; 5 – 6 supralabials; 6 infralabi-
als; 12 – 13 total lamellae on fourth digit of manus, and
19 total lamellae on fourth digit of pes.
Comparisons with other species. Among species of the
C. kandiana clade sensu aGarWal et al. (2017), Cne-
maspis nandimithrai sp. nov. differs by having smooth
ventral scales versus keeled scales in C. pava manamen-
Dra-araChChi, BatuWita & pethiyaGoDa, 2007, C. pul-
chra manamenDra-araChChi, BatuWita & pethiyaGoDa,
2007, C. samanalensis WiCkramasinGhe & muninDraDa-
sa, 2007, C. silvula manamenDra-araChChi, BatuWita
& pethiyaGoDa, 2007, C. tropidogaster (BoulenGer,
1885), and C. upendrai manamenDra-araChChi, Batu-
Wita & pethiyaGoDa, 2007, and also in having smooth
gular scales versus keeled in C. amith manamenDra-
araChChi, BatuWita & pethiyaGoDa, 2007; it can also
be diagnosed from C. ingerorum BatuWita, aGarWal
& Bauer, 2019, C. kallima manamenDra-araChChi,
BatuWita & pethiyaGoDa, 2007, C. kandiana (kelaart,
1852), C. menikay manamenDra-araChChi, BatuWita &
pethiyaGoDa, 2007, and C. retigalensis WiCkramasinGhe
& muninDraDasa, 2007 by having homogeneous (ver-
sus heterogeneous) dorsal scales; the new species differs
from C. kumarasinghei WiCkramasinGhe & muninDra-
Dasa, 2007, and C. latha manamenDra-araChChi, Batu-
Wita & pethiyaGoDa, 2007 by having a greater number of
belly scales (25 – 27 versus 17 – 21 and 13 – 15, respec-
tively), having a lower number of supralabial scales (5 – 6
versus 7 – 8 and 7 – 8, respectively), and having greater
number of paravertebral granules (95 – 99 versus 61 – 68
and 72 – 79, respectively); further it can be differentiated
from C. kumarasinghei by having a lower number of
ventral scales (108 – 112 versus 120 – 134).
Among species of the C. podihuna clade sensu aGar-
Wal et al. (2017), the new species differs from C. al-
wisi WiCkramasinGhe & muninDraDasa, 2007, C. ge-
munu Bauer, De silva, GreenBaum & jaCkman, 2007,
C. godagedarai De silva, Bauer, Botejue & karunar-
athna, 2019, C. kandambyi BatuWita & uDuGampala,
2017, C. molligodai WiCkramasinGhe & muninDraDasa,
2007, C. nilgala karunarathna, Bauer, De silva, sur-
asinGhe, somaratna, maDaWala, GaBaDaGe, Botejue,
henkanaththeGeDara & ukuWela, 2019, C. phillipsi
manamenDra-araChChi, BatuWita & pethiyaGoDa,
2007, C. podihuna DeraniyaGala, 1944, C. punctata
manamenDra-araChChi, BatuWita & pethiyaGoDa,
2007, C. rajakarunai WiCkramasinGhe, viDanapathi-
rana & rathnayake, 2016, C. rammalensis viDanapa-
thirana, rajeev, WiCkramasinGhe, FernanDo & WiCk-
ramasinGhe, 2014, and C. scalpensis (FerGuson, 1877)
by having smaller and more irregularly shaped subcaudal
scales (versus clearly enlarged hexagonal or subhex-
agonal scales), and by the presence (versus absence) of
precloacal pores except in C. kandambyi, C. molligo-
dai and C. podihuna, which can be differentiated from
C. nandimithrai sp. nov. by having a greater number of
belly scales (23 – 25 versus 16 – 17, 15 – 19, and 13 – 20,
respectively).
Description of Holotype. An adult male, 27.9 mm SVL.
Body slender, relatively short (TRL 40.8% of SVL).
Head relatively large (HL 30.4% of SVL, HL 74.5% of
TRL), narrow (HW 16.3% of SVL, HW 53.7% of HL),
depressed (HD 11.6% of SVL, HD 38.2% of HL) and
distinct from neck. Snout relatively long (ES 78.2% of
HW, ES 41.9% of HL), more than twice length of eye
diameter (ED 46.9% of ES), more than half length of jaw
(ES 70.5% of JL), snout slightly concave in lateral view;
eye relatively small (ED 19.7% of HL), larger than the
ear (EL 16.2% of ED), pupil rounded; orbit length less
than eye to ear distance (OD 92.9% of EE) and less than
length of digit IV of manus (OD 72.2% of DLM IV); su-
praocular ridges neither prominent or absent; ear opening
very small (EL 3.2% of HL), deep, taller than wide, larg-
er than nostrils; single row of scales separate orbit from
supralabials; interorbital distance is narrow (IO 89.0% of
ES), shorter than head length (IO 37.3% of HL); eye to
nostril distance greater than the eye to ear distance (EN
128.6% of EE).
Dorsal surface of trunk with smooth homogeneous
granules, 94 in paravertebral row; 108 smooth midven-
tral scales; 89 midbody scales; 4/3 weakly developed tu-
bercles on the anks; ventrolateral scales not enlarged;
granules on snout very smooth, larger than those on
interorbital and occipital regions; canthus rostralis not
pronounced, 12/12 smoothly rounded scales from eye to
nostril; scales of the interorbital region ovaland smooth;
tubercles present on the sides of the neck but absent
around the ear; ear opening vertically oval, slanting from
anterodorsal to posteroventral, 18/17 scales between an-

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VERTEBRATE ZOOLOGY — 69
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terior margin of the ear opening and the posterior margin
of the eye. Supralabials 6/6, infralabials 6/6, becoming
smaller towards the gape. Rostral scale wider than long,
partially divided (75%) by a median groove, in contact
with rst supralabial. Nostrils separated by 2/2 enlarged
supranasals, 2 internasals and 1/1 postnasal; no enlarged
scales behind the supranasals. Nostrils oval, dorsolater-
ally orientated, not in contact with rst supralabials.
Mental subrhomboidal in shape, as wide as long, pos-
teriorly in contact with 3 enlarged postmentals (smaller
than mental, and larger than chin scales); postmentals in
contact and bordered posteriorly by ve unkeeled chin
scales (smaller than nostrils), in contact with the 1st in-
fralabial; ventral scales smaller than chin scales. Smooth,
rounded, juxtaposed scales on the chin and the gular re-
gion; pectoral and abdominal scales smooth, subimbri-
Fig. 4. Close-ups of Cnemaspis nandimithrai sp. nov. male holotype (NMSL.2019.03.01) (A) dorsal head, (B) lateral head, (C) ventral
head, (D) homogeneous dorsal scales, (E) scaleson lateral surface of trunk, (F) smooth ventral scales, (g) cloacal characters with precloacal
pores and femoral pores (H) subdigital lamellae on manus, (I) subdigital lamellae on pes, (J) dorsal scalation of tail, (K) lateral side of tail,
and (l) oval shaped subcaudals (Photos: Suranjan Karunarathna).

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
260
cate to imbricate towards precloacal region, abdominal
scales slightly larger than dorsals; 25 belly scales across
venter; smooth scales around vent and base of tail, sub-
imbricate; 4 precloacal pores; 4/3 femoral pores; 11/11
unpored anterior femoral scales between pores on each
side; 5/5 enlarged posterior femoral scales. Original tail
of holotype longer than the snout-vent length (TAL
121.3% of SVL); hemipenial bulge greatly swollen
(TBW 2.9 mm), heterogeneous scales on the dorsal as-
pect of the tail directed backwards, spine-like tubercles
present at base of tail; tail with 4 – 5 enlarged attened
obtuse scales forming whorls; a small post-cloacal spur
on each side, dorsoventrally attened and narrow; sub-
caudals small, smooth, oval, arranged inamedian series.
Forelimbs moderately short, slender (LAL 14.0% of
SVL, UAL 14.1% of SVL); hind limbs long, tibia short-
er than the femur (TBL 18.2% of SVL, FEL 20.1% of
SVL). Anterior and posterior surfaces of upper arm with
keeled and less imbricate scales, scales of the anterior
surface twice as largre as those of other faces of limb;
dorsal and anterior surfaces of lower arm with keeled
and less imbricate scales, ventral and posterior surfaces
Table 3. Morphometric and Meristic data of holotype and two paratypes of Cnemaspis nandimithrai sp. nov. from Kudumbigala, Ampara
District, Sri Lanka (Abbreviations: L – left, R – right, M – male, F – female).
Measurements
(mm)
NMSL
2019.03.01
NMSL
2019.03.02
NMSL
2019.03.03 Counts
NMSL
2019.03.01
NMSL
2019.03.02
NMSL
2019.03.03
Holotype
(M)
Paratype
(M)
Paratype
(F)
Holotype
(M)
Paratype
(M)
Paratype
(F)
ED 1.7 1.7 1.8 FLSP (L/R) 4/3 4/4 4/4
OD 2.2 2.2 2.3 SUP (L/R) 6/6 5/6 6/5
EN 3.1 3.2 3.1 INF (L/R) 6/6 6/6 6/6
ES 3.6 2.5 3.6 INOS 31 33 34
SN 0.9 0.9 1.1 PM 3 3 3
NW 0.2 0.1 0.1 CHS 5 5 5
EE 2.4 2.6 2.5 SUN (L/R) 2 2 2
SA 12.8 13.9 14.5 PON (L/R) 1 1 1
EL 0.3 0.6 0.5 INT 2 2 2
IO 3.2 3.4 3.4 SUS (L/R) 14/15 14/13 12/14
IE 3.5 3.7 3.7 BET (L/R) 18/17 18/18 19/18
HL 8.5 9.1 8.8 CAS (L/R) 12/12 12/12 12/12
HW 4.6 4.8 4.9 TLM (i) (L/R) 10/10 9/10 9/9
HD 3.2 3.3 3.4 TLM (ii) (L/R) 15/15 15/15 14/14
JL 5.1 5.5 5.4 TLM (iii) (L/R) 17/17 18/17 18/18
IN 1.2 1.1 1.1 TLM (iv) (L/R) 13/13 12/12 12/12
SED 8.0 8.1 8.3 TLM (v) (L/R) 10/10 10/10 11/11
UAL 3.9 4.1 4.5 PG 95 96 99
LAL 3.9 3.9 4.1 MBS 89 87 87
PAL 2.8 2.9 3.1 MVS 108 111 112
DLM (i) 1.5 1.5 1.6 BLS 25 25 27
DLM (ii) 2.1 2.2 1.9 TLP (i) (L/R) 8/8 8/8 9/9
DLM (iii) 2.8 2.8 2.9 TLP (ii) (L/R) 13/13 14/13 14/14
DLM (iv) 3.1 3.1 3.2 TLP (iii) (L/R) 18/18 17/18 17/17
DLM (v) 2.4 2.4 2.5 TLP (iv) (L/R) 19/19 19/19 19/19
SVL 27.9 31.7 29.7 TLP (v) (L/R) 16/16 16/16 17/17
TRL 11.4 13.8 12.2 PCP 4 2 —
TW 4.7 4.8 4.9 FP (L/R) 4/3 2/2 —
TD 2.6 2.7 2.9 AFS (L/R) 11/11 9/10 —
FEL 5.6 5.7 5.9 PFS (L/R) 5/5 7/6 —
TBL 5.1 5.2 5.5
HEL 5.2 5.2 5.4
DLP (i) 1.4 1.5 1.3
DLP (ii) 2.1 2.1 2.0
DLP (iii) 3.3 3.3 3.4
DLP (iv) 3.6 3.7 3.7
DLP (v) 2.7 2.8 2.8
TAL 33.9 34.8 36.2
TBW 2.9 3.0 2.6
TBD 2.3 2.3 2.1

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VERTEBRATE ZOOLOGY — 69
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with unkeeled imbricate scales. Scales on dorsal surface
of the femur smooth and granular, less imbricate scales
on anterior, posterior and ventral surfaces, scales on an-
terior and ventral surfaces twice the size of those of other
faces of limbs. Dorsal, posterior and ventral surfaces of
tibia with smooth, less imbricate scales, anterior surface
with keeled subimbricate scales, scales of the ventral sur-
face twice as large as those of the other faces of the limb.
Dorsal and ventral scales on manus and pes with keeled
granules; dorsal surfaces of digits with granular scales.
Digits elongate and slender with inected distal phalan-
geal joint, all bearing slightly recurved claws. Subdigital
lamellae entire (except divided at rst interphalangeal
joint), unnotched; total lamellae on manus (left/right):
digit I, 10/10, digit II, 15/15, digit III, 17/17, digit IV,
13/13, digit V, 10/10; total lamellae on pes (left/right):
digit I, 8/8, digit II, 13/13, digit III, 18/18, digit IV, 19/19,
digit V, 16/16; interdigital webbing absent; length of dig-
its of manus (left): I (1.5 mm), II (2.1 mm), V (2.4 mm),
III (2.8 mm), IV (3.1 mm); length of digits of pes (left):
I (1.4 mm), II (2.1 mm), V (2.7 mm), III (3.3 mm), IV
(3.6 mm).
Variation of the type series. The SVL of adult speci-
mens in the type series (n = 3) ranges from 27.9 to 31.7
mm; supralabials 5 – 6; interorbital scales 31 – 34; supra-
ciliaries 12 – 15; scales from eye to tympanum 17 – 19;
total lamellae on digit I of the manus 9 – 10, lamellae on
digit II of manus 14 – 15, lamellae on digit III of manus
17 – 18, lamellae on digit IV of manus 12 – 13, lamellae
on digit V of manus 10 – 11; total lamellae on digit I of pes
8 – 9, lamellae on digit II of pes 13 – 14, lamellae on digit
III of pes 17 – 18, lamellae on digit V of pes 16 – 17; ven-
tral scales 108 – 112; belly scales across venter 25 – 27;
midbody scales 87 – 89; paravertebral granules 95 – 99;
precloacal pores in males 2 – 4; femoral pores in males
2 – 4; unpored anterior femoral scales in males 9 – 11, and
unpored posterior femoral scales in males 5 – 7.
Color of living specimens. The color of the head, body
and limbs on the dorsal side generally varies from light
grey to light brown, 5 irregular black bands on the dor-
sum of the body (Fig. 5); an oblique black line is present
between the eye and the nostrils on either side; 2 straight,
dark brown postorbital stripes on each side extend from
the eyes posteroventrally, and a dark longitudinal line is
present on the occipital area; there is a narrow subtrangle
shaped, brown patch on the occipital area with scattered
cream white spots; 4 distinct, irregular cream spots on
the lateral surfaces of the trunk; tail grey-white on the
dorsum, with 3 faded brown cross-bands present in the
rst half; the pupil is circular and black with surround-
ing red and white margins, with supraciliaries scales be-
ing yellowish; supralabials and infralabaials are cream
colored with black spots; mid-gular scales are dark yel-
low without dark spots; pectoral, abdominal, cloacal and
subcaudal scales are immaculate white; dorsal of limbs
have brown patches; manus and pes with cream white
and black cross stripe arrangement.
Color of preserved specimens. Dorsally dark brown
with pale colored, pale irregular brown bands and white
irregular lines are distinct; ventral surface completely
cream colored with some scales on thigh, tail base and
arms with dark brown margins.
Etymology. The specic epithet is an eponym Latinized
(nandimithrai) in the masculine genitive singular, hon-
ouring ‘Nandimithra Yodaya’ (a gladiator, one of the ten
giant warriors of King Dutugemunu’s army) for his val-
iant services to his king and his contribution to rebuilding
the Kudumbigala monastery (granite caves, especially
Maha Sudharshana Lena), later gifted to the Buddhist
Monks.
Habitat and ecology. Kudumbigala Sanctuary (
6.658114°
–
6.681811° N and 81.726844° – 81.762328° E; altitude
ranges from 12 to 67 m) is dominated by a tropical dry
mixed semi-evergreen forest with a Manilkara-dominant
oristic community (Gunatileke & Gunatileke, 1990),
and is approximately 800 ha in size, situated in the low-
land dry zone (Eastern Province, Ampara District) of
eastern Sri Lanka. The mean annual rainfall varies be-
tween 1,000 and 1,500 mm, received mainly during the
northeast monsoon (November – February). The mean
annual temperature of the area is 29.8 – 31.4 ºC. The Ku-
dumbigala monastic complex was built in 246 BC as a
refuge for Buddhist monks and includes over 200 caves
(Fig. 6). Our surveys suggest that C. nandimithrai sp.
nov. was rare in this locality given the low encounter
rate of 7 (± 0.1) geckos per man-hour. After surveying
a total area of 50 ha, we only found this species in four
distinct locations within the type locality. Microhabitat
associations of this species were restricted to rock out-
crops and the interior of granite caves in forested areas,
and are found up to a height of up to 8 m on vertical
surfaces. These microhabitats were well-shaded (light
intensity 0 – 643 Lux), relatively moist (relative humi di-
ty: 71 – 83%), cool and shady (ambient temeratur: 30.5 –
32.2 °C, rock-surface temperature: 26.6 – 28.4 °C, cano py
cover: 60 – 75%). The geckos were active during daylight
hours (0800 – 1700 h) and, when disturbed, sought ref-
uge in the rock crevices. The new species was sympatric
(at both the local habitat and microsite scales) with two
other micro-endemic cave-dwelling geckos (Calodac-
tylodes illingworthorum and Hemidactylus hunae) and
other generalist species (Cnemaspis podihuna, Gehyra
mutilata, Hemidactylus depressus, H. frenatus, H. trie-
drus, H. parvimaculatus). Eggs were observed in granite
rock crevices, typically laid in pairs. The eggs were pure
white in color almost spherical in shape (mean diameter
4.8 ± 0.02 mm), with a slightly attened side attached to
the rocky substrate. So far, this species has only been re-
corded from rock outcrops and granite caves at the type
locality, many of which have been human occupied since
the stoneage until the colonial period. It is likely that this
species is distributed throughout the Kudumbigala and
nearby habitats, but occupancy is restricted to granite
caves and rock outcrops.

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
262
Fig. 5. Cnemaspis nandimithrai sp. nov. male holotype (NMSL.2019.03.01) in life in-situ (A) dorsolateral view of the full body displaying
color pattern, and showing ectoparasites in red, (B) ventral aspect depicting gular and abdomen color (Photos: Nimantha Abeyarathna).
Conservation status. Application of the IUCN Red List
criteria indicates that C. nandimithrai sp. nov. is Critical-
ly Endangered (CR) due to having an area of occupancy
(AOO) < 10 km2 (four locations, 0.1 km2 in total assum-
ing a 100 m radius around the georeferenced location)
and an extent of occurrence (EOO) < 100 km2 (2.2 km2)
in the lower elevations of Eastern Province. The low
numbers we recorded in our survey also implies that the
species is rare, thus, warrants a higher conservation sta-
tus [Applicable criteria is B2-b (iii)].
Phylogenetic position. A member of C. kandiana Clade
B; the new species belongs to C. kumarasinghei species
group (BIII) (Fig. 2), within which it is most closely re-
lated to the new species C. gotaimbarai sp. nov. (p-dis-
tance 7.9%), C. ingerorum (p-distance 8.0%) and C. ku-
marasinghei (p-distance 6.8%) (see Tab. 2).
Geographical distance. Cnemaspis nandimithrai sp. nov.
most closely resembles C. gotaimbarai sp. nov., C. in-
gerorum and C. kumarasinghei the type localities of
which are ~48 km (Maragala in Monaragala District),
~ 102 km (Kokagala in Ampara District) and ~ 61 km
(San da gala in Hambantota District) straight line distance
from Ku dum bigala in Panama.
Cnemaspis gotaimbarai sp. nov.
ZooBank urn:lsid:zoobank.org:act:54D8F176-3105-479A-9474-
2951D8654309
Gotaimbaras’ day gecko (English)
Gotaimbarage diva-seri hoona (Sinhala)
Figs. 7 – 9; Table 4.
Holotype. NMSL.2019.04.01, adult male, 32.9 mm SVL (Fig. 7),
collected from granite cave in Kokagala, Padiyatalawa in Ampara
District, Eastern Province, Sri Lanka (7.439517° N, 81.207967° E,
WGS1984; elevation 292 m; around 14.00 hrs) on 22 October 2018
by Suranjan Karunarathna and Anslem de Silva.
Paratypes. NMSL.2019.04.02, adult male, 32.1 mm SVL, and
NMSL.2019.04.03, adult female, 33.7 mm SVL, collected from
granite caves in Kokagala, Padiyatalawain Ampara District, East-
ern Province, Sri Lanka (7.413086° N, 81.210161°E, WGS1984;
elevation 306 m; around 12.00 hrs) on 22 October 2018 by Suran-
jan Karunarathna and Anslem de Silva.
Diagnosis. Cnemaspis gotaimbarai sp. nov., can be
readily distinguished from its Sri Lankan congeners by
a combination of the following morphological and mer-
istic characteristics and color pattern: maximum SVL
33.7 mm; dorsum with homogeneous, smooth granular
scales; 2/2 supranasals, 1 internasal and 2/2 postnasals
present; 2 enlarged postmentals; postmentals bounded

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VERTEBRATE ZOOLOGY — 69
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by 5 chin scales; chin, gular, pectoral and abdominal
scales smooth, subimbricate; 23 – 25 belly scales across
venter; 5 – 6 weakly developed tubercles on posterior
ank; 117 – 121 paravertebral granules linearly arranged;
2 – 4 precloacal pores, 3 femoral pores on each side in
males, on each side separated by 10 – 13 unpored anterior
femoral scales, 1 – 5 unpored posterior femoral scales;
129 – 138 ventral scales; 72 – 79 midbody scales; subcau-
dals smooth, median row tiny, in an irregular series of
subrhomboidal scales; 7 – 8 supralabials; 8 – 9 infralabi-
als; 16 – 17 total lamellae on fourth digit of manus, and
19 – 20 total lamellae on fourth digit of pes.
Comparisons with other species. The new species,
C. go taimbarai sp. nov. differs from species of the
C. kan diana clade sensu aGarWal et al. (2017) in having
smooth ventral scales versus keeled in C. pava, C. pul-
chra, C. samanalensis, C. silvula, C. tropidogaster, and
C. upen drai, and in also having smooth gular scales
versus keeled gular scales in C. amith. It can be diag-
nosed from C. in gerorum, C. kallima, C. kandiana, C. me-
nikay, and C. reti galensis by having homogeneous (ver-
sus heterogeneous) dorsal scales. The new species differs
from C. ku ma rasinghei and C. latha by having a greater
number of belly scales (23 – 25 versus 17 – 21 and 13 – 15,
respectively), having greater number of paravertebral
granules (117 – 121 versus 61 – 68 and 72 – 79, respective-
ly), and also having greater number of interorbital scales
(34 – 36 versus 24 – 26 and 22 – 25, respectively). Further,
it can be differentiated from C. nandimithrai sp. nov. by
having more supralabial scales (7 – 8 versus 5 – 6), more
paravertebral granules (117 – 121 versus 95 – 99), and
more ventral scales (129 – 138 versus 108 – 112).
The new species also clearly differs from the fol-
lowing species of the C. podihuna clade sensu aGarWal
et al. (2017): C. alwisi, C. gemunu, C. godagedarai, C. kan-
dambyi, C. molligodai, C. nilgala, C. phillipsi, C. po-
di huna, C. punctata, C. rajakarunai, C. rammalensis,
and C. scalpensis by having small and irregular shaped
subcaudal scales (versus clearly enlarged hexagonal or
subhexagonal scales), and by the presence (versus ab-
sence) of precloacal pores, except in C. kandambyi,
C. molligodai and C. podihuna, which differ from C. go-
taimbarai sp. nov. by having a lower number of belly
scales (16 – 17, 15 – 19 and 13 – 20 versus 23 – 25).
Description of Holotype. An adult male, 32.9 mm SVL.
Body slender, relatively long (TRL 44.2% of SVL).
Fig. 6. General habitat of Cnemaspis nandimithrai sp. nov. at Kudumbigala isolated forest hill, Ampara District, Sri Lanka (A) complete
view of the hill, (B) archaeological monuments on rock outcrops, (C) prehistorical-occupied granite caves with inscription, and evidence
of remains of early humans (Photos: Majintha Madawala).

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
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Head relatively large (HL 30.2% of SVL, HL 68.4% of
TRL), narrow (HW 15.6% of SVL, HW 51.6% of HL),
depressed (HD 10.1% of SVL, HD 33.4% of HL) and
distinct from neck. Snout relatively long (ES 76.2% of
HW, ES 39.3% of HL), more than twice eye diameter
(ED 46.3% of ES), more than half length of jaw (ES
63.0% of JL), snout slightly concave in lateral view; eye
relatively small (ED 18.2% of HL), larger than the ear
(EL 24.3% of ED), pupil rounded; orbit length less than
eye to ear distance (OD 80.1% of EE) and of length of
IV digit of manus (OD 63.9% of DLM IV); supraocular
ridges are moderately developed; ear opening very small
(EL 4.4% of HL), deep, taller than wide, larger than nos-
trils; 3 rows of scales separate orbit from supralabials; in-
Fig. 7. Close-ups of Cnemaspis gotaimbarai sp. nov. male holotype (NMSL.2019.04.01) (A) dorsal head, (B) lateral head, (C) ventral
head, (D) homogeneous dorsal scales, (E) scaleson lateral surface of trunk, (F) smooth ventral scales, (G) cloacal characters with precloacal
pores and femoral pores (H) subdigital lamellae on manus, (I) subdigital lamellae on pes, (J) dorsal scalation of tail, (K) lateral side of tail,
and (L) oval shaped subcaudals (Photos: Suranjan Karunarathna).

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VERTEBRATE ZOOLOGY — 69
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Fig. 8. Cnemaspis gotaimbarai sp. nov. male holotype (NMSL.2019.04.01) in life in-situ (A) dorsolateral view of the full body, and (B)
dorsal view depicting color pattern (Photos: Majintha Madawala).
terorbital distance is broader than orbit length (IO 87.5%
of ES), shorter than head length (IO 34.4% of HL); eye
to nostril distance slightly greater than eye to ear distance
(EN 114.5% of EE).
Dorsal surface of the trunk with unkeeled homogene-
ous granules, 117 paravertebral granules; 132 midventral
scales, smooth; 79 midbody scales; 5/5 weakly devel-
oped tubercles on the anks; ventrolateral scales slightly
enlarged; granules on snout somewhat rough, larger than
those on interorbital and occipital regions; canthus ros-
tralis not pronounced,12/12 keeled round scales from
eye to nostril; scales of the interorbital region circular
and fairly keeled; tubercles present both on the sides of
the neck and around the ear; ear opening vertically oval,
slanting from anterodorsal to posteroventral, 17/17 scales
between anterior margin of the ear opening and the poste-
rior margin of the eye. Supralabials 7/8, infralabials 8/6,
becoming smaller towards the gape. Rostral scale wider
than long, partially divided (60%) by a median groove,
contact with rst supralabial. Nostrils separated by 2/2
enlarged supranasals with 1 internasal; no enlarged
scales behind the supranasals. Nostrils oval, dorsolater-
ally orientated, not in contact with rst supralabials; 2/2
postnasals, smooth, larger than nostrils, partially in con-
tact with rst supralabial.
Mental sub-triangular in shape, as wide as long, poste-
riorly in contact with 2 enlarged postmentals (smaller than
mental, and lager than chin scales); postmentals in con-
tact and bordered posteriorly with 5 unkeeled chin scales
(smaller than nostrils), in contact with the 1st infralabial;
ventral scales smaller than posterior postmentals. Smooth,
rounded, juxtaposed scales on the chin and the gular re-
gion; pectoral and abdominal scales smooth, subimbricate
to imbricate towards precloacal region, abdominal scales
slightly larger than dorsals; 25 belly scales across venter;
smooth scales around vent and base of tail, subimbricate;
4 precloacal pores; 3/3 femoral pores; 10/11 unpored an-
terior femoral scales in between pores on each side; 4/5
enlarged posterior femoral scales. Regenerated tail of
holotype longer than the snout-vent length (TAL 123.6%

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266
of SVL); Hemipenial bulge greatly swollen (TBW 3.1
mm), heterogeneous scales on the dorsal aspect of the tail
directed backwards, spine-like tubercles present at the
base of tail; tail with 4 – 5 enlarged attened obtuse scales
forming whorls; a large, blunt post-cloacal spur on each
side, dorso-ventrally attened and narrow; subcaudals
smooth, subrhomboidal, arranged in a median series.
Forelimbs moderately short, slender (LAL 17.7% of
SVL, UAL 13.8% of SVL); hind limbs long, tibia short-
er than the femur (TBL 17.1% of SVL, FEL 18.6% of
SVL). Anterior and posterior surfaces of upper arm with
keeled and less imbricate scales, scales of the anterior
surface twice as large as those of the other parts; dor-
sal and anterior surfaces of lower arm with keeled and
less imbricate scales, ventral and posterior surfaces with
unkeeled imbricate scales those are imbricate. Scales on
dorsal surface of femur smooth, rounded and juxtaposed,
those on anterior surface well keeled, posterior and ven-
tral surfaces smooth and imbricate, scales on the ventral
surface twice the size ofthose of other aspects. Dorsal,
Table 4. Morphometric and Meristic data of holotype and two paratypes of Cnemaspis gotaimbarai sp. nov. from Kokagala, Ampara Dis-
trict, Sri Lanka (Abbreviations: L – left, R – right, M – male, F – female).
Measurements
(mm)
NMSL
2019.04.01
NMSL
2019.04.02
NMSL
2019.04.03 Counts
NMSL
2019.04.01
NMSL
2019.04.02
NMSL
2019.04.03
Holotype
(M)
Paratype
(M)
Paratype
(F)
Holotype
(M)
Paratype
(M)
Paratype
(F)
ED 1.8 1.7 1.6 FLSP (L/R) 5/5 5/6 6/6
OD 2.1 2.0 2.0 SUP (L/R) 7/8 8/8 8/8
EN 2.9 2.9 3.0 INF (L/R) 8/8 8/9 8/8
ES 3.9 3.8 3.7 INOS 35 34 36
SN 0.7 0.7 0.8 PM 2 2 2
NW 0.2 0.2 0.2 CHS 5 5 5
EE 2.6 2.5 2.4 SUN (L/R) 2 2 2
SA 15.6 15.5 15.4 PON (L/R) 2 2 2
EL 0.4 0.4 0.4 INT 1 1 1
IO 3.4 3.3 3.2 SUS (L/R) 15/16 15/15 16/16
IE 3.7 3.6 3.5 BET (L/R) 17/17 18/17 18/18
HL 9.9 9.8 9.7 CAS (L/R) 12/12 12/11 12/12
HW 5.1 5.1 4.9 TLM (i) (L/R) 13/13 12/13 12/12
HD 3.3 3.3 3.1 TLM (ii) (L/R) 15/15 15/15 14/14
JL 6.2 6.1 5.9 TLM (iii) (L/R) 18/18 18/17 17/17
IN 1.2 1.2 1.2 TLM (iv) (L/R) 17/17 17/17 16/17
SED 8.5 8.4 8.2 TLM (v) (L/R) 15/14 15/15 15/15
UAL 4.5 4.4 4.3 PG 117 121 119
LAL 5.8 5.7 5.6 MBS 79 72 76
PAL 3.3 3.3 3.1 MVS 132 138 129
DLM (i) 1.4 1.4 1.2 BLS 25 23 23
DLM (ii) 1.9 1.9 1.8 TLP (i) (L/R) 9/9 9/9 8/8
DLM (iii) 2.6 2.6 2.4 TLP (ii) (L/R) 13/13 12/13 12/12
DLM (iv) 3.2 3.1 2.9 TLP (iii) (L/R) 14/14 14/15 15/15
DLM (v) 2.3 2.2 2.1 TLP (iv) (L/R) 19/19 19/20 20/19
SVL 32.9 32.1 33.7 TLP (v) (L/R) 21/21 22-22 21/22
TRL 14.5 14.4 14.9 PCP 4 2 —
TW 4.9 4.9 5.1 FP (L/R) 3/3 3/3 —
TD 3.3 3.2 3.3 AFS (L/R) 11/10 12/13 —
FEL 6.1 6.0 5.9 PFS (L/R) 4/5 2/1 —
TBL 5.6 5.5 5.4
HEL 4.3 4.3 4.2
DLP (i) 1.2 1.2 1.2
DLP (ii) 1.9 1.9 1.7
DLP (iii) 3.3 3.3 3.1
DLP (iv) 3.4 3.4 3.3
DLP (v) 2.7 2.7 2.6
TAL 40.7 39.3 36.7
TBW 3.1 3.1 2.7
TBD 2.6 2.5 2.7

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VERTEBRATE ZOOLOGY — 69
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anterior and posterior surfaces of tibia with keeled and
less imbricate scales, ventral surface with smooth and
subimbricate scales, scales of the ventral surface twice
as large as those on other aspects. Dorsal and ventral
surfaces of manus and pes with keeled granules; dorsal
surfaces of digits with granular scales. Digits elongate
and slender with inected distal phalanges joint, all bear-
ing slightly recurved claws. Subdigital lamellae entire
(except divided at rst interphalangeal joint), unnotched;
total lamellae on manus (left/right): digit I, 13/13, digit
II, 15/15, digit III, 18/18, digit IV, 17/17, digit V, 15/14;
total lamellae on pes (left/right): digit I, 9/9, digit II,
13/13, digit III, 14/14, digit IV, 19/19, digit V, 21/21; in-
terdigital webbing absent; length of manual digits (left):
I (1.4 mm), II (1.9 mm), V (2.3 mm), III (2.6 mm), IV
(3.2 mm); length of pedal digits (left): I (1.2 mm), II (1.9
mm), V (2.7 mm), III (3.3 mm), IV (3.4 mm).
Variation of the type series. The SVL of adult speci-
mens in the type series (n = 3) size ranges from 32.1 to
23.7 mm; supralabials 7 – 8, and infralabials 8 – 9; in-
terorbital scales 34 – 36; supraciliaries 15 – 16; scales
from eye to tympanum 17 – 18; scales from eye to nostril
11 – 12; total lamellae on digit I of the manus 12 – 13, la-
mellae on digit II of manus 14 – 15, lamellae on digit III
of manus 17 – 18, lamellae on digit IV of manus 16 – 17,
lamellae on digit V of manus 14 – 15; total lamellae on
digit I of pes 8 – 9, lamellae on digit II of pes 12 – 13,
lamellae on digit III of pes 14 – 15, lamellae on digit IV
of pes 19 – 20, lamellae on digit V of pes 21 – 22; ven-
tral scales 129 – 138; belly scales across venter 23 – 25;
midbody scales 72 – 79, and paravertebral granules
117 – 121; precloacal pores in males 2 – 4, anterior femo-
ral scales in males 10 – 13, posterior femoral scales in
males 1 – 5.
Color of living specimens. The color of the dorsum of
head, body and limbs generally varies from light brown
to light grey with 4 – 5 irregular dirty white cloud-like
blotches (Fig. 8); an oblique black line is present along
the canthal line, 2 straight, dark brown and grey postorbi-
tal stripes (each side) extend from eyes posteroventrally,
and a longitudinal golden line with dark margins present
in the occipital area; there is a narrow and faded human-
face-like brown patch on the occipital area with scattered
cream white spots; 3 – 4 distinct irregular cream white
spots on the lateral surfaces of the body; tail brown-
black on dorsum, with 6 – 7 faded white cross-bands; the
pupil is circular and golden with surrounding red and
white margins, with supraciliaries scales being yellow-
ish; supralabials and infralabaials are cream white and
black spotted; mid gular scales are yellow with tiny dark
spots; pectoral, abdominal, cloacal and subcaudal scales
are white with dark spots; dorsum of limbs with white
and brown patches; manus and pes with cream white and
black stripe arrangement.
Color of preserved specimens. Dorsally dark brown
with pale colored, pale irregular blotches and much dark
irregular lines are distinct; ventral surface completely
cream colored with some scales on thigh, tail base and
arms with dark brown margins.
Etymology. The specic epithet is an eponym Latinized
(gotaimbarai) in the masculine genitive singular, hon-
ouring ‘Gotaimbara Yodaya’ (a gladiator, one of the ten
giant warriors of King Dutugemunu’s army) for his val-
iant services to his king and country.
Habitat and ecology. Kokagala Forest Reserv
(7.400000° – 7.450000° N and 81.200000° – 81.233333° E;
altitude ranges from 285 – 650 m) is a tropical dry mixed
semi-evergreen forest with a Vatica-dominant oristic
community (Gunatileke & Gunatileke, 1990). It is ap-
proximately 1000 ha in size and islocated in the lowland
mid-peneplains of the dry bioclimatic zone of Eastern Sri
Lanka (Eastern Province, Ampara District). The mean
annual rainfall varies between 1,500 and 2,500 mm, re-
ceived mainly during the northeast monsoon (Novem-
ber – February). The mean annual temperature of the
area is 29.6 – 30.5 ºC. Kokagala is rich in granite caves
with over 100 having been documented. Cnemaspis go-
taimbarai sp. nov. could be a rare species as we only
found 9 (± 0.2) geckos per man-hour after surveying ap-
proximately a total area of 35 ha with 5 conrmed loca-
tions. This species was restricted to rock outcrops and
granite caves in forested areas, and was found up to 10
m in height on vertical surfaces (Fig. 9). The microhabi-
tats of C. gotaimbarai sp. nov. were poorly illuminated
(light intensity: 0 – 821 Lux), relatively moist (relative
humidity: 69 – 85%), cool and dry (ambient temperature:
30.2 – 31.8 °C, substrate temperature: 27.2 – 28.7 °C, can-
opy cover: 70 – 85%). The geckos were active during day-
light hours (0800 – 1700 h) and, when disturbed, sought
refuge in rocky crevices. The new species was sympatric
(at both local habitat and the microsite scale) with several
other micro-endemic cave-dwelling geckos (Calodac ty-
lodes illingworthorum and Hemidactylus hunae) as well
as more generalist species (Cnemaspis cf. po dihuna, Ge-
hyra mutilata, Hemidactlus depressus, H. fre na tus, H.
triedrus, H. parvimaculatus). No eggs were found.
Conservation status. Application of the IUCN Red List
criteria indicates that C. gotaimbarai sp. nov. is Critical-
ly Endangered (CR) due to having an area of occupancy
(AOO) <10 km2 (ve locations, 0.2 km2 in total, assum-
ing a 100 m radius around the georeferenced locations)
and an extent of occurrence (EOO) <100 km2 (3.5 km2)
in the mid-elevations of the Eastern Province [Applicable
criteria is B2-b (iii)].
Phylogenetic position. Cnemaspis gotaimbarai sp. nov.
is a member of C. kandiana Clade B; the new species be-
longs to C. kumarasinghei species group (BIII) (Fig. 2),
within which it is most closely related to C. ingerorum
(p-distance 8.6%), C. kumarasinghei (p-distance 8.1%)
and the new species Cnemaspis nandimithrai sp. nov. (p-
distance 7.9%) (see Tab. 2).

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
268
Geographical distance. Cnemaspis gotaimbarai sp. nov.
most closely resembles C. ingerorum, C. kumarasinghei
and C. nandimithrai sp. nov morphologically, the type
localities of which is separated by 121 km (Sandagala in
Hambantota District), ~55 km (Maragala in Monaragala
District) and ~ 102 km (Kudumbigala in Ampra District),
straight line distance from Kokagala in Padiyatalawa.
Cnemaspis kohukumburai sp. nov.
ZooBank urn:lsid:zoobank.org:act:B0F2073D-A77F-43DA-85E3-
27464BD2CF64
Kohukumbures’ day gecko (English)
Kohukumburege diva-seri hoona (Sinhala)
Figs. 10 – 12; Table 5.
Holotype. NMSL.2019.05.01, adult male, 33.9 mm SVL (Fig. 10),
collected from a granite tunnel in Kadugannawa, Kandy Dis-
trict, Central Province, Sri Lanka (7.251800° N, 80.509378° E,
WGS1984; elevation 427 m; around 09.00 hrs) on 12 October 2018
by Suranjan Karunarathna and Anslem de Silva.
Paratypes. NMSL.2019.05.02, adult male, 32.5 mm SVL, and
NMSL.2019.05.03, adult female, 31.0 mm SVL collected from
a granite tunnel in Kadugannawa, Kandy District, Central Prov-
ince, Sri Lanka (7.251658° N, 80.512561° E, WGS1984; eleva-
tion 435 m; around 10.00 hrs), on 12 October 2018 by Suranjan
Karunarathna and Anslem de Silva.
Diagnosis. Cnemaspis kohukumburai sp. nov. can be
readily distinguished from its Sri Lankan congeners by a
combination of the following morphological and meristic
characteristics and color pattern: maximum SVL 33.9 mm;
dorsum with homogeneous, smooth granular scales; 2/2
supranasals, 1 internasal and 1/1 postnasal present; 3 en-
larged postmentals; postmentals bounded by 5 chin scales;
chin and gular scales smooth, granular, juxta posed; pecto-
ral and abdominal scales smooth, subimbricate; 7 – 8 well
developed tubercles on posterior ank; 150 – 159 paraver-
tebral granules linearly arranged; 23 belly scales across
the venter; precloacal pores absent in males, 6 – 9 femo-
ral pores on each side in males separated by 25 unpored
femoro-precloacal scales, 1 – 2 unpored posterior femoral
scales; 131 – 134 ventral scales; 81 – 88 midbody scales;
subcaudals smooth, subhexagonal, enlarged, subequal,
forming aregular median row; 8 – 9 supra labials; 7 – 8 in-
fralabials; 21 – 22 total lamellae on digit IV of manus, and
23 – 25 total lamellae on digit IV of pes.
Fig. 9. General habitat of Cnemaspis gotaimbarai sp. nov. at Kokagala isolated forest hill, Ampara District, Sri Lanka (A) complete view
of the hill and corn plantation, (B) granite boulders along the stream, (C) abandoned wattle and daub house in lower margin (Photos:
Madhava Botejue and Majintha Madawala).

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Comparisons with other species. The new species dif-
fers from species of the C. podihuna clade sensu aGar-
Wal et al. (2017) as follows: from C. kandambyi, C. mol-
ligodai and C. podihuna by the absence (versus pres-
ence) of precloacal pores; from C. gemunu, C. godage-
darai, C. phillipsi, C. rammalensis and C. scalpensis
by the presence of fewer femoral pores (6 – 9 versus
11 – 14, 12 – 13, 15 – 16, 15 and 13 – 15, respectively); it
further differs from C. gemunu and C. scalpensis by the
presence of a greater number of belly scales (23 versus
13 – 16 and 17 – 19, respectively), from C. rammalensis
by the presence of fewer ventral scales (131 – 134 versus
186 – 207), and from C. phillipsi by the presence of more
total lamellae on digit IV of pes (23 – 25 versus 17 – 19).
Fig. 10. Close-ups of Cnemaspis kohukumburai sp. nov. male holotype (NMSL.2019.05.01) (A) dorsal head, (B) lateral head, (C) ventral
head, (D) homogeneous dorsal scales, (E) scales on lateral surface of trunk, (F) smooth ventral scales, (G) cloacal characters with precloa-
cal pores and femoral pores (H) subdigital lamellae on manus, (I) subdigital lamellae on pes, (J) dorsal scalation of tail, (K) lateral side of
tail, and (L) oval shaped subcaudals (Photos: Suranjan Karunarathna).

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270
It can be diagnosed from C. alwisi and C. rajakarunai
by the presence of fewer ventral scales (131 – 134 ver-
sus 145 – 153 and 146 – 186, respectively), and by the
presence of more unpored femoro-precloacal scales (25
versus 18 – 19 and 20 – 22, respectively). It can be differ-
entiated from C. punctata by the greater number of para-
vertebral granules (150 – 159 versus 83 – 91), by the pres-
ence of more total lamellae on digit IV of manus (21 – 22
versus 17 – 18), and by fewer well-developed tubercles
on the posterior ank (7 – 8 versus 11 – 13). Further, it
can be distinguished from C. nilgala by the presence of a
greater number of belly scales (23 versus 17 – 19) across
the midbody, and by the presence of more total lamellae
on digit IV of pes (23 – 25 versus 17 – 19).
Cnemaspis kohukumburai sp. nov. also clearly dif-
fers by the absence of precloacal pores and the presence
of clearly enlarged, hexagonal or subhexagonal subcau-
dal scales from the following species of the C. kandiana
clade sensu aGarWal et al. (2017): C. amith, C. gotaim-
barai sp. nov., C. ingerorum, C. kallima, C. kandiana,
C. kumarasinghei, C. latha, C. menikay, C. nandimithrai
sp. nov., C. pava, C. pulchra, C. retigalensis, C. samana-
lensis, C. silvula, C. tropidogaster and C. upendrai.
Description of Holotype. An adult male, 33.9 mm SVL.
Body slender, relatively short (TRL 39.3% of SVL).
Head relatively large (HL 30.9% of SVL, HL 78.6% of
TRL), narrow (HW 19.0% of SVL, HW 61.6% of HL),
depressed (HD 6.6% of SVL, HD 21.4% of HL) and
distinct from neck. Snout relatively short (ES 61.0% of
HW, ES 37.6% of HL), more than twice eye diameter
(ED 47.7% of ES), more than half length of jaw (ES
63.2% of JL), snout slightly concave in lateral view;
eye relatively small (ED 17.9% of HL), larger than the
ear (EL 23.9% of ED), pupil rounded; orbit length less
than eye to ear distance (OD 75.2% of EE) and nearly
half ofthe length of digit IV of the manus (OD 51.5% of
DLM IV); supraocular ridges not prominent; ear open-
ing very small (EL 4.3% of HL), deep, taller than wide,
larger than nostrils; 2 rows of scales separate orbit from
supralabials; interorbital distanceis narrow (IO 78.9% of
ES), shorter than head length (IO 29.6% of HL); eye to
nostril distance little greater than the eye to ear distance
(EN 103.2% of EE).
Dorsal surface of the trunk with smooth, small ho-
mogeneous granules, 150 paravertebral granules; 131
smooth midventral scales; 88 midbody scales; 8/8 well
developed tubercles on the anks; ventrolateral scales
slightly enlarged; granules on snout smooth and fairly
raised, larger than those on interorbital and occipital re-
gions; canthus rostralis not pronounced, 15/15 smooth
round scales from eye to nostril; scales of the interorbi-
tal region oval and smooth; 3/3 tubercles present on the
sides of the neck and around the ear; ear opening verti-
cally oval, slanting from anterodorsal to posteroventral,
25/26 scales between anterior margin of the ear opening
and the posterior margin of the eye. Supralabials 8/9 in-
fralabials 8/8, becoming smaller towards the gape. Ros-
tral scale wider than long, partially divided (80%) by a
median groove, in contact with rst supralabial. Nostrils
separated by 2/2 enlarged supranasals with 1 internasal
and 1/1 postnasal; no enlarged scales behind the suprana-
sals. Nostrils oval, dorsolaterally orientated, not in con-
tact with rst supralabials.
Mental sub-triangular, as wide as long, posteriorly
in contact with three enlarged postmentals (smaller than
mental, and larger than chin scales); postmentals in con-
tact and bordered posteriorly by ve smooth chin scales
(smaller than nostrils), in contact only with the 1st infral-
abials; ventral scales smaller than chin scales. Smooth,
rounded, juxtaposed scales on chin and gular region;
pectoral and abdominal scales smooth, subimbricate to-
wards precloacal region, abdominal scales larger than
dorsals; 23 belly scales across venter; smooth, subim-
bricate scales around vent and base of tail; 8/9 femoral
pores; 25 unpored femoro-precloacal scales; 2/1 small
posterior femoral scales. Original tail of holotype longer
than snout-vent length (TAL 121.5% of SVL); Hemipe-
nial bulge greatly swollen (TBW 3.2 mm), homogeneous
scales on the dorsal aspect of the tail directed backwards,
3/3 spine-like tubercles present at base of tail, latter very
smooth; tail with 3 – 4 enlarged attened obtuse scales
forming whorls; a small, blunt post-cloacal spur on each
side, dorso-ventrally attened and narrow; smooth sub-
caudals are arranged into a median series of clearly en-
larged, hexagonal or subhexagonal scales.
Forelimbs moderately short, slender (LAL 15.2% of
SVL, UAL 19.2% of SVL); hind limbs long, tibia shorter
than the femur (TBL 21.6% of SVL, FEL 26.2% of SVL).
Dorsal, anterior, ventral and posterior surfaces of upper
arm with smooth scales, those on anterior surface twice-
as large as those on other faces of limb; dorsal, anterior,
ventral and posterior surfaces of lower arm with smooth
scales, anterior surface twice as large as those of the other
parts; scales on dorsal surface of the femur smooth and
granular, less imbricate scales on the anterior, posterior
and ventral surfaces, scales on the anterior surface is twice
the size ofthose of the other parts. Dorsal, anterior, poste-
rior and ventral surfaces of tibia with smooth scales; both
anterior and posterior surfaces of limbs are with smooth
granules, scales of the ventral surface twice as large as
those of the other parts. Dorsal and ventral scales on the
manus and the pes smooth, granular; dorsal surfaces of
digits with granular scales. Digits elongate and slender
with inected distal phalanges joint, all bearing slightly
recurved claws. Subdigital lamellae entire (except divid-
ed at rst interphalangial joint), unnotched; total lamellae
on manus (left/right): digit I, 12/12, digit II, 18/19, digit
III, 20/19, digit IV, 22/22, digit V, 24/24; total lamellae
on pes (left/right): digit I, 12/12, digit II, 18/18, digit III,
20/19, digit IV, 24/25, digit V, 22/23; interdigital webbing
absent; length of digits of manus (left): I (2.5 mm), II (3.3
mm), III (3.6 mm), V (3.6 mm), IV (4.1 mm); length of
digits of pes (left): I (1.8 mm), II (3.7 mm), III (4.1 mm),
V (4.2 mm), IV (4.9 mm).
Variation of the type series. The SVL of adult specimens
in the type series (n = 3) ranges from 31.0 to 33.9 mm;

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VERTEBRATE ZOOLOGY — 69
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number of supralabials 8 – 9, and infralabials 7 – 8; inter-
orbital scales 38 – 39; supraciliaries 13 – 15; scales from
eye to tympanum 24 – 26; total lamellae on digit I of the
manus 11 – 12, lamellae on digit II of manus 17 – 19, la-
mellae on digit III of manus 19 – 20, lamellae on digit IV
of manus 21 – 22, lamellae on digit V of manus 23 – 24;
total lamellae on digit II of pes 17 – 18, lamellae on digit
III of pes19 – 20, lamellae on digit IV of pes 23 – 25, la-
mellae on digit V of pes 21 – 23; ventral scales 131 – 134,
midbody scales 81 – 88; paravertebral granules 150 – 159;
femoral pores in males 6 – 9, and unpored posterior femo-
ral scales in males 1 – 2
Color of living specimens. The dorsum of the head, body
and limbs light brown and light grey; 5 irregular cloud-
like black bands on the dorsum of the body; 2 oblique
black and white lines are present between the eye and the
nostrils on either side (Fig. 11); a straight, dark brown
postorbital stripe extends from each eye posteroventrally
across the ear, and broad dark spots are present in the oc-
cipital area; there is a narrow ‘I’ shaped, black patch on
the occipital area with scattered small, irregular cream-
white spots surrounding it. Tail dorsum is brownish, with
9 faded grey cross-bands; the pupil is circular and black
with surrounding orange and white margins, with grey-
ish supraciliaries scales; supralabials and infralabaials
light brown with black spots; mid-gular scales dirty white
without dark spots; pectoral, abdominal, cloacal and sub-
caudal scales are cream without dark spots; dorsum of
limbs with brown patches; manus and pes with with an ar-
rangement of black and cream white stripe on the dorsum.
Color of preserved specimens. Dorsally dark brown
with pale colored, 5 irregular cloud-like pale bands clear
Fig. 11. Cnemaspis kohukumburai sp. nov. (A) female paratype (NMSL.2019.05.03) in life in-situ (B) male holotype (NMSL.2019.05.01)
in life in-situ depicting dorsal color pattern (Photos: Majintha Madawala).

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
272
on the dorsum of the body; dark and narrow ‘I’ shaped
patch on the occipital area; ventral surface completely
whitewith some scales on thighs, tail base and arms with
dark brown margins.
Etymology. The specic epithet is an eponym Latinized
(kohukumburai) in the masculine genitive singular, hon-
ouring the Sri Lankan warrior ‘Kohukumbure Walauwe
Rate Rala’ for his valiant feats in the Great Rebellion of
1817 – 1818 which was initiated in Uva-Wellassa.
Habitat and ecology. The Kadugannawa area
(7.232356° – 7.261469° N and 80.497639° – 80.540933° E;
altitude range from 354 – 567 m) is a tropical wet wet-
ever green forest (Gunatileke & Gunatileke, 1990) mix-
ed with a mosaic of anthropogenic and anthropocentric
habitats, and is approximately 600 ha in size, situated in
the low-country wet zone of central Sri Lanka (Central
Province, Kandy District). The mean annual rainfall var-
ies between 3,000 and 4,500 mm, received mostly via the
southwest monsoon (May – September). The mean annual
Table 5. Morphometric and Meristic data of holotype and two paratypes of Cnemaspis kohukumburai sp. nov. from Kadugannawa, Kandy
District, Sri Lanka (Abbreviations: L – left, R – right, M – male, F – female).
Measurements
(mm)
NMSL
2019.05.01
NMSL
2019.05.02
NMSL
2019.05.03 Counts
NMSL
2019.05.01
NMSL
2019.05.02
NMSL
2019.05.03
Holotype
(M)
Paratype
(M)
Paratype
(F)
Holotype
(M)
Paratype
(M)
Paratype
(F)
ED 1.9 1.8 1.9 FLSP (L/R) 8/8 8/7 7/7
OD 2.1 2.1 2.1 SUP (L/R) 8/9 8/8 8/8
EN 2.9 2.9 2.9 INF (L/R) 8/8 7/8 7/7
ES 3.9 3.9 3.8 INOS 38 38 39
SN 0.8 0.8 0.8 PM 3 3 3
NW 0.2 0.2 0.2 CHS 5 5 5
EE 2.8 2.8 2.6 SUN (L/R) 2 2 2
SA 16.9 17.1 17.0 PON (L/R) 1 1 1
EL 0.5 0.5 0.5 INT 1 1 1
IO 3.1 3.1 3.2 SUS (L/R) 14/15 15/15 13/13
IE 3.3 3.5 3.5 BET (L/R) 25/26 24/24 24/25
HL 10.5 11.2 11.0 CAS (L/R) 15/15 14/14 15/14
HW 6.5 6.5 6.3 TLM (i) (L/R) 12/12 11/11 11/11
HD 2.3 2.3 2.4 TLM (ii) (L/R) 18/19 18/18 18/17
JL 6.2 6.3 6.0 TLM (iii) (L/R) 20/19 20/20 19/19
IN 1.3 1.3 1.3 TLM (iv) (L/R) 22/22 21/21 21/21
SED 10.3 10.4 10.3 TLM (v) (L/R) 24/24 23/24 23/23
UAL 6.5 6.4 6.4 PG 150 154 159
LAL 5.2 5.2 5.1 MBS 88 83 81
PAL 6.1 6.1 5.9 MVS 131 134 134
DLM (i) 2.5 2.5 2.5 BLS 23 23 23
DLM (ii) 3.3 3.3 3.4 TLP (i) (L/R) 12/12 12/12 12/12
DLM (iii) 3.6 3.6 3.5 TLP (ii) (L/R) 18/18 17/17 17/18
DLM (iv) 4.1 4.1 4.2 TLP (iii) (L/R) 20/19 19/19 19/19
DLM (v) 3.6 3.6 3.7 TLP (iv) (L/R) 24/25 24/24 23/23
SVL 33.9 32.5 31.0 TLP (v) (L/R) 22/23 23/23 21/21
TRL 13.3 13.6 14.1 FP (L/R) 8/9 7/6 —
TW 5.4 5.4 5.3 PFS (L/R) 2/1 2/2 —
TD 3.8 3.8 3.7 IFS 25 25 —
FEL 8.9 8.6 8.3
TBL 7.3 7.3 7.1
HEL 7.8 7.6 7.3
DLP (i) 1.8 1.8 1.8
DLP (ii) 3.7 3.7 3.7
DLP (iii) 4.1 4.1 4.2
DLP (iv) 4.9 4.9 4.8
DLP (v) 4.2 4.2 4.2
TAL 41.2 41.4 42.0
TBW 3.2 3.2 3.2
TBD 2.7 2.7 2.5

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VERTEBRATE ZOOLOGY — 69
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temperature of the area is 25.2 – 27.5ºC; C. kohukumburai
sp. nov. is a rare species as 2 (± 1) geckos per man-hour
were found after surveying a total area of 25 ha with
three conrmed locations. This species was restricted
to rock outcrops and granite caves in forested areas, and
was found at heights of up to 5 m on vertical surfaces
(Fig. 12). These microhabitats were poorly illuminated
(light intensity: 0 – 686 Lux), relatively moist (relative
humidity: 73 – 89%), cool and wet (ambient temperature:
29.8 – 31.2°C, rock surface temperature: 26.2 – 27.8°C,
canopy cover: 70 – 90%). The geckos were active during
the day time (0800 – 1700 h) and when disturbed, sought
refuge in rocky crevices. The new species was sympatric
(at local habitat scale) with several other geckos (Cne-
maspis kandiana, Gehyra mutilata, Hemidactylus depres-
sus, H. frenatus, H. parvimaculatus, Hemiphyllodactylus
typus). Eggs likely belonging to C. kohukumburai were
found in crevices, typically laid in pairs communally. The
eggs were pure white in color almost spherical in shape
(mean diameter 4.9±0.02 mm; n = 22), with a slightly at-
tened side attached to the rocky substrate. This species
was recorded from granite caves, tunnel and rock out-
crops located in both relatively undisturbed forests and
less modied habitats in the Kadugannawa area.
Conservation status. Application of the IUCN Red List
criteria indicates that C. kohukumburai sp. nov. is Criti-
cally Endangered (CR) due to having an area of occu-
pancy (AOO) <10km2 (three locations, 0.1 km2 in total,
assuming a 100 m radius around the georeferenced lo-
cations) and an extent of occurrence (EOO) <100 km2
(0.01 km2) in the lower elevations of Central Province
[Applicable criteria is B2-b (iii)].
Phylogenetic position. A member of C. podihuna Clade
A; the new species belongs to the species group AIII
(Fig. 2), within which it is reconstructed as a sister spe-
cies of C. alwisi (p-distance 11.9%) and C. punctata (p-
distance 14.9%) (see Tab. 2).
Geographical distance. Cnemaspis kohukumburai sp.
nov. most closely resembles C. alwisi and C. rajakaru-
nai. The type localities of these species are separated by
~ 35 km and ~ 40 km straight line distance (Dolukanda
in Kurunegala and Salgala in Kegalle, respectively) from
Kadugannawa in Kandy.
Fig. 12. General habitat of Cnemaspis kohukumburai sp. nov. at Kadugannawa isolated forest hill, Kandy District, Sri Lanka (A) complete
view of the mountain range from top of the hill, (B) the granite tunnel in type locality along the tar road, (B) communal egg laying site
(Photos: Majintha Madawala and Kalangi Rodrigo).

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
274
Cnemaspis hitihami sp. nov.
ZooBank urn:lsid:zoobank.org:act:8DC0B6FC-C1F5-46FF-83C6-
8FBF33563F5C
Hitihamis’ day gecko (English)
Hitihamige diva-seri hoona (Sinhala)
Figs. 13 – 15; Table 6.
Holotype. NMSL.2019.06.01, adult male, 41.7 mm SVL (Fig. 13),
collected from a granite cave in Kumaradola, Maragalakanda,
Monaragala District, Uva Province, Sri Lanka (6.875428° N,
81.357289° E, WGS1984; elevation 421 m; around 16.00 hrs)
on 18 September 2018 by Suranjan Karunarathna and Anslem de
Silva.
Paratypes. NMSL.2019.06.02, adult male, 38.8 mm SVL, and
NMSL.2019.06.03, adult female, 39.5 mm SVL, collected from a
granite cave in Kumaradola, Maragalakanda, Monaragala District,
Uva Province, Sri Lanka (6.879725° N, 81.364986° E, WGS1984;
elevation 437 m; around 17.00 hrs) on 18 September 2018 by
Suranjan Karunarathna and Anslem de Silva.
Diagnosis. Cnemaspis hitihami sp. nov., can be readily
distinguished from its Sri Lankan congeners by a combi-
nation of the following morphological and meristic char-
acteristics and color pattern: maximum SVL 41.7 mm;
dorsum with homogeneous, smooth granular scales; 2/2
supranasals, 1 internasal and 1/1 postnasal present; 3 en-
larged postmentals; postmentals bounded by 5 chin scales;
chin and gular scales smooth. granular, juxtaposed; pecto-
ral and abdominal scales smooth, subimbricate; 21 belly
scales across the venter; 4 – 5 well developed tubercles on
posterior ank; 143 – 149 paravertebral granules linearly
arranged; precloacal pores absent in males, 5 – 10 femoral
pores in males on each side separated by 24 – 26 unpored
femoro-precloacal scales, 5 – 7 unpored posterior femoral
scales; 132 – 135 ventral scales; 96 – 99 midbody scales;
enlarged subcaudals smooth, subequal, subhexagonal, ar-
ranged in a regular median row; 8 – 9 supralabials; 7 – 9
infralabials; 18 – 19 total lamellae on digit IV of manus,
and 21 – 22 total lamellae on digit IV of pes.
Comparisons with other species. The new species,
C. hitihami sp. nov. differs from all members of the
C. podihuna clade sensu aGarWal et al. (2017): from
C. kandambyi, C. molligodai and C. podihuna it differs
by the absence (versus presence) of precloacal pores.
The new species differs from C. godagedarai, C. phil-
lipsi, C. rammalensis and C. scalpensis in having fewer
femoral pores (5 – 10 versus 12 – 13, 15 – 16, 15 and
13 – 15, respectively). It also differs from C. godage-
darai by its greater number of paravertebral granules
(143 – 149 versus 101 – 106), from C. phillipsi by its
greater number of total lamellae on digit IV of the pes
(21 – 22 versus 17 – 19), from C. rammalensis by fewer
belly scales (21 versus 25 – 28) across the midbody, and
from C. scalpensis by the presence of more belly scales
(21 versus 17 – 19). It can be diagnosed from C. alwisi
and C. rajakarunai by lower number of ventral scales
(132 – 135 versus 145 – 153 and 146 – 186, resperc-
tively), and by its greater number of unpored femoro-
precloacal scales (24 – 26 versus 18 – 19 and 20 – 22,
respectively). It can be differentiated from C. gemunu
and C. nilgala by its greater number of belly scales (21
versus 13 – 16 and 17 – 19) and by a greater number to-
tal lamellae on digit IV of pes (21 – 22 versus 18 – 19
and 17 – 18, respectively). The new species also differs
from C. punctata by in having more midbody scales
(96 – 99 versus 71 – 78), by greater number of paraver-
tebral granules (143 – 149 versus 83 – 91), and by fewer
well developed tubercles on posterior ank (4 – 5 versus
11 – 13). It can be diagnosed from C. kohukumburai sp.
nov. by a lower number of interorbital scales (30 – 33
versus 37 – 39), and by more total lamellae on digit IV
of pes (21 – 22 versus 18 – 19).
The new species clearly differs from C. amith, C. go-
taimbarai sp. nov., C. ingerorum, C. kallima, C. kandia na,
C. kumarasinghei, C. latha, C. menikay, C. nandimi th -
rai sp. nov., C. pava, C. pulchra, C. retigalensis, C. sa-
ma nalensis, C. silvula, C. tropidogaster and C. upend-
rai, members of the C. kandiana clade sensu aGarWal
et al. (2017), by the absence of precloacal pores and the
presence of clearly enlarged, hexagonal or subhexagonal
subcaudal scales.
Description of Holotype. An adult male, 41.7 mm SVL.
Body slender, relatively long (TRL 42.6% of SVL).
Head relatively small (HL 27.2% of SVL, HL 63.9% of
TRL), narrow (HW 16.6% of SVL, HW 61.1% of HL),
depressed (HD 8.1% of SVL, HD 29.9% of HL) and dis-
tinct from neck. Snout relatively short (ES 67.1% of HW,
ES 41.0% of HL), more than twice eye diameter (ED
41.4% of ES), more than half length of jaw (ES 59.4%
of JL), snout slightly concave in lateral view; eye rela-
tively small (ED 17.0% of HL), larger than the ear (EL
37.3% of ED), pupil rounded; orbit length greater than
eye to ear distance (OD 118.5% of EE) and more than
half of the length of IV digit of manus (OD 76.7% of
DLM IV); supraocular ridges not prominent; ear open-
ing very small (EL 6.3% of HL), deep, taller than wide,
larger than nostrils; 2 rows of scales separate orbit from
supralabials; interorbital distance is broad (IO 100.6% of
ES), shorter than head length (IO 41.3% of HL); eye to
nostril distance little greater than the eye to ear distance
(EN 103.6% of EE).
Dorsal surface of the trunk with small, smooth, ho-
mogeneous granules; 145 paravertebral granules; 132
smooth midventral scales; 96 midbody scales; 5/5 well
developed tubercles on the anks; ventrolateral scales not
enlarged; granules on snout smooth and fairly raised, larg-
er than those on interorbital and occipital regions; canthus
rostralis not pronounced, 18/19 smooth round scales from
eye to nostril; scales of the interorbital region oval and
smooth; 2/2 tubercles present on the sides of the neck and
around the ear; ear opening vertically oval, slanting from
anterodorsal to posteroventral, 18/18 scales between ante-
rior margin of the ear opening and posterior margin of the
eye. Supralabials 9/9 infralabials 8/9, becoming smaller
towards the gape. Rostral scale wider than long, partially
divided (50%) by a median groove, in contact with rst
supralabial. Nostrils separated by 2/2 enlarged suprana-

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VERTEBRATE ZOOLOGY — 69
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sals with 1 internasal and 1/1 postnasal; no enlarged scales
behind the supranasals. Nostrils oval, dorsolaterally ori-
entated, not in contact with rst supralabials.
Mental sub-triangular in shape, as wide as long,
posteriorly in contact with three enlarged postmentals
(smaller than mental, and lager than chin scales); post-
mentals in contact and bordered posteriorly by ve
smooth chin scales (smaller than nostrils), in contact with
the 1st infralabials; ventral scales smaller than chin scales.
Smooth, rounded, juxtaposed scales present on chin and
gular region; pectoral and abdominal scales smooth, sub-
imbricate towards precloacal region, abdominal scales
Fig. 13. Close-ups of Cnemaspis hitihami sp. nov. male holotype (NMSL.2019.06.01) (A) dorsal head, (B) lateral head, (C) ventral head,
(D) homogeneous dorsal scales, (E) scales on lateral surface of trunk, (F) smooth ventral scales, (G) cloacal characters with precloacal
pores and femoral pores (H) subdigital lamellae on manus, (I) subdigital lamellae on pes, (J) dorsal scalation of tail, (K) lateral side of tail,
and (L) oval shaped subcaudals (Photos: Suranjan Karunarathna).

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
276
larger than dorsals; 21 belly scales across venter; smooth,
subimbricate scales around vent and base of tail; 5/8
femoral pores; 24 unpored femoro-precloacal scales; 6/5
small posterior femoral scales. Original tail of holotype
longer than snout-vent length (TAL 129.7% of SVL);
Hemipenial bulge greatly swollen (TBW 4.4 mm), ho-
mogeneous scales on the dorsal aspect of the tail directed
backwards, 3/3 spine-like tubercles present at the base of
tail, latter very smooth; tail with 5 – 7 enlarged attened
obtuse scales forming whorls; a small, blunt post-cloacal
spur on each side, dorsoventrally attened and narrow;
smooth, enlarged hexagonal or subhexagonal subcaudals
arranged in a median series.
Forelimbs moderately short, slender (LAL 17.8% of
SVL, UAL 16.7% of SVL); hind limbs long, tibia quite
shorter than the femur (TBL 21.4% of SVL, FEL 22.6%
of SVL). Dorsal, anterior, ventral and posterior surfaces
of upper arm with smooth scales, those on anterior sur-
face twice as large as those on other faces of limb. Dorsal,
anterior, ventral and posterior surfaces of lower arm with
Table 6. Morphometric and Meristic data of holotype and two paratypes of Cnemaspis hitihami sp. nov. from Maragalakanda, Monaragala
District, Sri Lanka (Abbreviations: L – left, R – right, M – male, F – female).
Measurements
(mm)
NMSL
2019.06.01
NMSL
2019.06.02
NMSL
2019.06.03 Counts
NMSL
2019.06.01
NMSL
2019.06.02
NMSL
2019.06.03
Holotype
(M)
Paratype
(M)
Paratype
(F)
Holotype
(M)
Paratype
(M)
Paratype
(F)
ED 1.9 1.9 1.9 FLSP (L/R) 5/5 5/4 4/5
OD 3.3 3.3 3.1 SUP (L/R) 9/9 8/8 8/9
EN 2.9 3.0 2.9 INF (L/R) 8/9 8/7 9/7
ES 4.7 4.8 5.4 INOS 31 33 31
SN 1.7 1.6 1.5 PM 3 3 3
NW 0.2 0.2 0.3 CHS 5 5 5
EE 2.8 2.7 2.7 SUN (L/R) 2 2 2
SA 18.8 18.0 18.6 PON (L/R) 1 1 1
EL 0.7 0.8 0.7 INT 1 1 1
IO 4.7 4.6 4.6 SUS (L/R) 15/15 15/16 14/14
IE 4.9 4.9 4.8 BET (L/R) 18/18 19/19 20/19
HL 11.4 10.8 10.5 CAS (L/R) 18/19 18/18 19/19
HW 6.9 6.7 6.5 TLM (i) (L/R) 12/12 12/13 12/12
HD 3.4 3.1 3.0 TLM (ii) (L/R) 15/14 15/15 15/15
JL 7.9 7.7 6.9 TLM (iii) (L/R) 15/15 16/16 15/16
IN 1.8 1.8 1.7 TLM (iv) (L/R) 18/18 18/18 18/19
SED 10.4 10.0 9.9 TLM (v) (L/R) 17/16 16/16 17/17
UAL 7.0 6.6 6.3 PG 145 143 149
LAL 7.4 6.7 7.1 MBS 96 97 99
PAL 5.9 5.7 5.9 MVS 132 133 135
DLM (i) 2.8 2.8 2.7 BLS 21 21 21
DLM (ii) 3.5 3.5 3.3 TLP (i) (L/R) 13/13 12/13 13/13
DLM (iii) 3.7 3.6 3.6 TLP (ii) (L/R) 16/16 16/16 16/16
DLM (iv) 4.3 4.1 4.1 TLP (iii) (L/R) 19/18 18/18 18/18
DLM (v) 3.8 3.6 3.6 TLP (iv) (L/R) 21/21 21/22 22/22
SVL 41.7 38.8 39.5 TLP (v) (L/R) 18/19 18/18 19/19
TRL 17.8 15.6 16.5 FP (L/R) 5/8 10/9 —
TW 7.8 7.9 7.3 PFS (L/R) 6/5 7/7 —
TD 3.9 3.9 4.3 IFS 24 26 —
FEL 9.4 8.4 8.7
TBL 8.9 7.5 8.6
HEL 7.3 7.1 8.2
DLP (i) 1.9 1.8 1.8
DLP (ii) 3.9 3.7 3.8
DLP (iii) 4.2 4.2 4.0
DLP (iv) 4.8 4.7 4.6
DLP (v) 4.3 4.1 4.1
TAL 54.1 45.9 44.1
TBW 4.4 4.3 3.9
TBD 3.3 3.3 2.9

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VERTEBRATE ZOOLOGY — 69
(3) 2019
smooth scales, those on anterior surface twice as large as
those of the other faces of limb; scales on dorsal surface
of femur smooth and granular, less imbricate on anterior,
posterior and ventral surfaces, scales on the anterior sur-
face twice the size of those on other parts. Dorsal, ante-
rior, posterior and ventral surfaces of tibia with smooth
scales; both anterior and posterior surfaces of limbs with
smooth granules, scales of the ventral surface twice as
large as those of the other surfaces. Dorsal and ventral
scales on the manus and the pes with smooth granules;
dorsal surfaces of digits also with granular scales. Dig-
its elongate and slender with an inected distal phalan-
ges joint, all bearing slightly recurved claws. Subdigital
lamellae entire (except divided at rst interphalangial
joint), unnotched; total lamellae on manus (left/right):
digit I, 12/12, digit II, 15/14, digit III, 15/15, digit IV,
18/18, digit V, 17/16; total lamellae on pes (left/right):
digit I, 13/13, digit II, 16/16, digit III, 19/18, digit IV,
21/21 digit V, 18/19; interdigital webbing absent; length
of digits of manus (left): I (2.8 mm), II (3.5 mm), III (3.7
mm), V (3.8 mm), IV (4.3 mm); length of digits of pes
(left): I (1.9 mm), II (3.9 mm), III (4.2 mm), V (4.3 mm),
IV (4.8 mm).
Variation of the type series. The SVL of adult speci-
mens in the type series (n = 4) ranges from 38.6 to 41.7
mm; supralabials 8 – 9; infralabials 7 – 9; interorbital
scales 30 – 33; supraciliaries 14 – 16; scales from naris to
orbit 18 – 19; scales from eye to tympanum 18 – 20; total
lamellae on digit I of the manus 12 – 13, lamellae on digit
II of manus 14 – 15, lamellae on digit III of manus 15 – 16,
lamellae on digit IV of manus 18 – 19, lamellae on digit V
of manus 16 – 17; total lamellae on digit I of pes 12 – 13,
lamellae on digit II of pes 15 – 16, lamellae on digit III of
pes 18 – 19, lamellae on digit IV of pes 21 – 22, lamellae
on digit V of pes 18 – 19; spine-like tubercles on ank
4 – 5; ventral scales 131 – 135, belly scales across venter
21, midbody scales 95 – 99, and paravertebral granules
143 – 149; femoral pores in males 7 – 11, unpored femo-
ro-precloacal scales in males 24 – 26, and unpored poste-
rior femoral scales in males 5 – 7.
Fig. 14. Cnemaspis hitihami sp. nov. male holotype (NMSL.2019.06.01) in life in-situ (A) dorsolateral view of the full body, and (B)
ventral aspect depicting gular and abdomen color (Photos: Majintha Madawala).

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278
Color of living specimens. The dorsum of the head, body
and limbs generally varies from golden brown to dark
brown mixed with light grey; 5 – 7 small, irregular black
paravertebral spots present; a narrow, short black longitu-
dinal line on the occipital area with scattered cream-white
spots;a single row of white spots present along vertebral
line (Fig. 14). Tail dorsum cinnamon brown, with 8 – 12
faded grey cross-bands; pupil circular and black with sur-
rounding orange and white margins, with supraciliaries
scales cinnamon brown; supralabials and infralabaials
light brown with black spots; chin and gular scales bright
yellow, without dark spots; pectoral, abdominal, cloacal
and subcaudal scales cream without dark spots; dorsum
of limbs have brown patches; manus and pes intermixed
with black and cream white stripe arrangement.
Color of preserved specimens. Dorsally dark brown
with pale colored, paravertebral spots much dark and
with pale vertebral spot line; ventral surface completely
dirty white, with some scales on thighs, tail base and
arms with dark brown margins.
Etymology. The specic epithet is an eponym Latinized
(hitihami) in the masculine genitive singular, honouring
the Sri Lankan warrior ‘Meegahapitiye Walauwe Hiti-
hami Mudiyanse Rate Rala’ (a long-range archer with
precise aim) for his valiant feats in the Great Rebellion of
1817 – 1818, which was initiated in Uva-Wellassa.
Habitat and ecology. The Maragalakanda area
(
6.842525°– 6.922392
° N and
81.336822°– 81.413719
° E;
altitude ranges from 354 – 750 m) is characterized as trop-
ical dry mixed semi-evergreen forest (Gunatileke & Gu-
natileke, 1990) mixed with anthropogenic habitats and
rubber plantations. It is approximately 4,100 ha in size,
situated in the lowland intermediate bioclimatic zone of
eastern Sri Lanka (Uva Province, Monaragala District).
The mean annual rainfall varies between 1,500 and 2,500
mm, received mostly during the northeast monsoon sea-
son (November – February). The mean annual tempera-
ture of the area is 26.8 – 28.9°C. Cnemaspis hitihami sp.
nov. appeared to be very common, at least at the type
locality, as we recorded 21 (± 0.2) geckos per man-hour
after surveying a total area of 25 ha with 7 conrmed lo-
cations. This species was restricted to rock outcrops and
granite caves in forested areas and inside active rubber
plantations, and reached heights of up to 6 m on vertical
surfaces (Fig. 15). These habitats were poorly illuminat-
ed (light intensity: 0 – 753 Lux), relatively moist (relative
humidity: 68 – 84%), cool and wet (ambient temperature:
Fig. 15. General habitat of Cnemaspis hitihami sp. nov. at Maragala isolated forest hill, Monaragala District, Sri Lanka (A) a complete
view of the granite hill, (B) rock outcrop with rubber plantation, (C) granite cave habitat on roadside (Photos: Majintha Madawala).

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VERTEBRATE ZOOLOGY — 69
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29.8 – 32.3°C, rock surface temperature: 25.8 – 27.5°C,
canopy cover: 75 – 90%). The geckos were active during
the day time (0800 – 1700 h) and when disturbed, sought
refuge in the crevices of rocky caves or jump on to the
forest oor and retreated under leaitter. The new species
was sympatric (at both local habitat and the microsite
scales) with two other micro-endemics (Calodactylodes
illingworthorum and Hemidactylus hunae) and generalist
gecko species (Cnemaspis kumarasinghei, C. podihuna,
Gehyra mutilata, Hemidactylus depressus, H. frenatus,
H. leschenaultii, H. parvimaculatus and H. triedrus).
Eggs were observed in granite rock crevices, typically
laid communally in pairs. The eggs were pure white, al-
most spherical in shape (mean diameter 5.1 ± 0.02 mm;
n = 16), with a slightly attened side attached to the rocky
substrate. This species was recorded from granite caves,
rock outcrops in both undisturbed and anthropogenic
habitats around Maragalakanda.
Conservation status. Application of the IUCN Red List
criteria indicates that C. hitihami sp. nov. is Critically
Endangered (CR) due to having an area of occupancy
(AOO) < 10 km2 (seven locations, 0.2 km2 in total assum-
ing a 100 m radius around the georeferenced locations)
and an extent of occurrence (EOO) < 100 km2 (2.3 km2)
in the lower elevations of Uva Province [Applicable cri-
teria is B2-b (iii)].
Phylogenetic position. A member of C. podihuna Clade
A; the new species belongs to species group AIII (Fig. 2),
within which it is reconstructed as a sister species of the
clade comprising C. nilgala (p-distance 12.4%) + unde-
scribed candidate species Cnemaspis sp. 8 (p-distance
11.9%) (see Tab. 2).
Geographic distance. Cnemaspis hitihami sp. nov. most
closely resembles C. nilgala and undescribed candidate
species Cnemaspis sp. 8. The type localities of these spe-
cies are separated by an straight line distance of ~25 km
(Nilgala in Bibila) and 38 km (Rahathankanda in Buttala)
from Maragala in Monaragala.
Cnemaspis butewai sp. nov.
ZooBank urn:lsid:zoobank.org:act:6E8FB8CC-6F2C-4E75-B312-
B300FF29CE38
Butewes’ day gecko (English)
Butewege diva-seri hoona (Singhala)
Figs. 16 – 18; Table 7.
Holotype. NMSL.2019.07.01, adult male, 31.7 mm SVL (Fig. 16),
collected from a large granite cave in Bambarabotuwa, Opanayake,
Ratnapura District, Sabaragamuwa Province, Sri Lanka (6.630364°
N, 80.628925° E, WGS1984; elevation 445 m; around 12.00 hrs)
on 26 November 2018 by Suranjan Karunarathna and Anslem de
Silva.
Paratypes. NMSL.2019.07.02, adult female, 27.2 mm SVL, and
NMSL.2019.07.03, Adult male, 31.8 mm SVL, collected from a
large granite cave in Bambarabotuwa, Opanayake, Ratnapura Dis-
trict, Sabaragamuwa Province, Sri Lanka (6.646919° N, 80.653892°
E, WGS1984; elevation 458 m; around 11.00 hrs) on 26 November
2018 by Suranjan Karunarathna and Anslem de Silva.
Diagnosis. Cnemaspis butewai sp. nov., can be readily
distinguished from its Sri Lankan congeners by a combi-
nation of the following morphological and meristic char-
acteristics: maximum SVL 31.8 mm; dorsum with het-
erogeneous, smooth, granular scales; 1 – 2 internasals, 2/2
supranasals and 1 – 2 postnasal present; 37 – 39 interorbital
scales; 15 – 17 supraciliaries, 10 – 11 canthal scales, 23 – 26
eye to tympanum scales; 3 enlarged postmentals; post-
mentals bounded by 5 – 6 chin scales; gular scales keeled,
chin, pectoral and abdominal scales smooth, subimbricate;
23 – 25 belly scales across the venter; 5 – 6 weakly devel-
oped tubercles on posterior ank; 134 – 138 paravertebral
granules linearly arranged; 3 – 5 precloacal pores, 5 femo-
ral pores on each side in males separated by 9 – 10 unpored
anterior femoral scales, 2 – 3 unpored posterior femoral
scales; 125 – 128 ventral scales; 92 – 98 midbody scales;
subcaudals smooth, median row comprising a regular se-
ries of small, diamond shaped scales; 8 supralabials; 7 – 8
infralabials; 16 total lamellae on fourth digit of manus, and
17 – 18 total lamellae on fourth digit of pes.
Comparisons with other species. Among species of
the C. kandiana clade sensu aGarWal et al. (2017)
C. butewai sp. nov. differs by having heterogeneous
(versus homogeneous) dorsal scales from C. amith,
C. gotaimbarai sp. nov., C. kumarasinghei, C. latha,
and C. nandimithrai sp. nov.; it can also be diagnosed
from C. ingerorum, C. pava, C. pulchra, C. samanalen-
sis, C. silvula, C. tropidogaster and C. upendrai by hav-
ing smooth (versus keeled) pectoral and gular scales. The
new species differs from C. kandiana and C. retigalensis
in having more belly scales (23 – 25 versus 19 – 20 and
16 – 20, respectively), more femoral pores (5 versus 3 – 4
and 3 – 4), and in having more precloacal pores (3 – 5 ver-
sus 2 – 4 and 1, respectively). It differs from C. menikay
in having more femoral pores (5 versus 3 – 4), and more
precloacal pores (3 – 5 versus 1 – 2) and from C. kallima
by having fewer tubercles on the posterior ank (5 – 6
versus 12 – 15), and keeled (versus smooth) gular scales.
The new species clearly differs from the species of
the C. podihuna clade sensu aGarWal et al. (2017): from
C. alwisi, C. gemunu, C. godagedarai, C. hitihami sp.
nov., C. kohukumburai sp. nov., C. nilgala, C. phillipsi,
C. punctata, C. rajakarunai, C. rammalensis and C. scal-
pensis by the presence (versus absence) of precloacal
pores, and by the absence (versus presence) of clearly en-
larged, hexagonal or subhexagonal subcaudal scales from
C. kandambyi, C. molligodai and C. podihuna it differ
by having heterogeneous (versus homogeneous) dorsal
scales and by the absence of clearly enlarged, hexa gonal
or subhexagonal subcaudal scales.
Description of Holotype. An adult male, 31.7 mm SVL.
Body slender, relatively short (TRL 34.1% of SVL).

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
280
Head relatively large (HL 29.0% of SVL, HL 84.9% of
TRL), narrow (HW 16.6% of SVL, HW 57.3% of HL),
depressed (HD 8.9% of SVL, HD 30.7% of HL) and dis-
tinct from neck. Snout relatively long (ES 71.2% of HW,
ES 40.8% of HL), more than twice eye diameter (ED
49.9% of ES), more than half length of jaw (ES 60.2%
of JL), snout slightly concave in lateral view; eye rela-
tively small (ED 20.3% of HL), larger than the ear (EL
38.0% of ED), pupil rounded; orbit length greater than
eye to ear distance (OD 111.4% of EE) and greater than
to the length of IV digit of manus (OD 111.8% of DLM
IV); supraocular ridges not prominent; ear opening very
small (EL 7.7% of HL), deep, taller than wide, larger than
nostrils; single row of scales separate orbit from suprala-
Fig. 16. Close-ups of Cnemaspis butewai sp. nov. male holotype (NMSL.2019.07.01) (A) dorsal head, (B) lateral head, (C) ventral head,
(D) homogeneous dorsal scales, (E) scales on lateral surface of trunk, (F) smooth ventral scales, (G) cloacal characters with precloacal
pores and femoral pores (H) subdigital lamellae on manus, (I) subdigital lamellae on pes, (J) dorsal scalation of tail, (K) lateral side of tail,
and (L) oval shaped subcaudals (Photos: Suranjan Karunarathna).

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VERTEBRATE ZOOLOGY — 69
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bials; interorbital distance less narrow than snout length
(IO 96.5% of ES), shoter than head length (IO 39.3%
of HL); eye to nostril distance little less than eye to ear
distance (EN 93.8% of EE).
Dorsal surface of the trunk with heterogeneous tri-
hedral granules; 134 paravertebral granules; 128 smooth
midventral scales; 96 midbody scales; 5/6 weakly devel-
oped tubercles on the anks; ventrolateral scales not en-
larged; granules on snout very smooth, larger than those
on interorbital and occipital regions; canthus rostralis
not pronounced, 10/10 smoothly oval scales from eye to
nostril; scales of the interorbital region oval and smooth;
blunt tubercles on the sides of neck, but absent around
ear; ear opening vertically oval, slanting from antero-
dorsal to posteroventral, 24/26 scales between anterior
margin of ear opening and posterior margin of the eye.
Supralabials 8/8, infralabials 8/8, becoming smaller to-
wards the gape. Rostral scale wider than long, partially
divided (80%) by a median groove, in contact with rst
supralabial. Nostrils separated by 2/2 enlarged suprana-
Table 7. Morphometric and Meristic data of holotype and two paratypes of Cnemaspis butewai sp. nov. from Bambarabotuwa, Ratnapura
District, Sri Lanka (Abbreviations: L – left, R – right, M – male, F – female).
Measurements
(mm)
NMSL
2019.07.01
NMSL
2019.07.02
NMSL
2019.07.03 Counts
NMSL
2019.07.01
NMSL
2019.07.02
NMSL
2019.07.03
Holotype
(M)
Paratype
(M)
Paratype
(F)
Holotype
(M)
Paratype
(M)
Paratype
(F)
ED 1.9 1.8 1.8 FLSP (L/R) 5/6 5/5 6/6
OD 3.2 3.3 3.3 SUP (L/R) 8/8 8/8 8/8
EN 2.7 2.8 2.7 INF (L/R) 8/8 7/8 8/7
ES 3.8 3.8 3.4 INOS 39 38 37
SN 1.6 1.7 1.6 PM 3 3 3
NW 0.2 0.2 0.2 CHS 5 5 6
EE 2.9 2.9 2.5 SUN (L/R) 2 2 2
SA 14.6 14.9 12.2 PON (L/R) 1 2 1
EL 0.7 0.7 0.8 INT 2 2 1
IO 3.6 3.7 3.8 SUS (L/R) 16/15 15/15 17/15
IE 3.8 3.9 4.0 BET (L/R) 24/26 25/24 23/25
HL 9.2 9.3 8.5 CAS (L/R) 10/10 11/10 10/10
HW 5.3 5.4 5.0 TLM (i) (L/R) 8/8 8/9 8/8
HD 2.8 2.9 2.4 TLM (ii) (L/R) 12/12 12/12 13/12
JL 6.2 6.3 6.0 TLM (iii) (L/R) 16/16 14/14 15/15
IN 1.6 1.7 1.6 TLM (iv) (L/R) 16/16 16/16 16/16
SED 8.7 8.7 8.5 TLM (v) (L/R) 14/14 14/15 14/14
UAL 5.9 5.9 5.5 PG 134 135 138
LAL 5.4 5.5 5.2 MBS 96 98 92
PAL 3.5 3.5 3.1 MVS 128 126 125
DLM (i) 1.4 1.5 1.4 BLS 23 25 23
DLM (ii) 1.8 1.8 1.7 TLP (i) (L/R) 9/9 8/8 8/8
DLM (iii) 2.4 2.3 2.4 TLP (ii) (L/R) 13/13 13/13 12/13
DLM (iv) 2.9 2.8 3.0 TLP (iii) (L/R) 15/16 15/15 15/15
DLM (v) 2.3 2.4 2.4 TLP (iv) (L/R) 18/18 18/17 18/18
SVL 31.7 31.8 27.2 TLP (v) (L/R) 17/17 17/17 17/17
TRL 10.8 10.9 9.8 PCP 3 5 —
TW 5.4 5.4 5.2 FP (L/R) 5/5 5/5 —
TD 3.4 3.4 3.1 AFS (L/R) 10/10 10/9 —
FEL 5.9 5.9 5.7 PFS (L/R) 2/2 3/3 —
TBL 5.6 5.7 5.3
HEL 5.2 5.3 5.1
DLP (i) 1.4 1.5 1.5
DLP (ii) 3.1 3.2 3.2
DLP (iii) 3.6 3.5 3.4
DLP (iv) 4.1 4.1 4.2
DLP (v) 3.9 3.8 3.9
TAL 36.8 37.9 32.1
TBW 3.7 3.9 3.4
TBD 3.1 3.2 2.9

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
282
sals with 2 internasals and 1/1 postnasal; no enlarged
scales behind the supranasals. Nostrils oval, dorsolater-
ally orientated; not in contact with rst supralabials.
Mental subtriangular, as wide as long, posteriorly in
contact with three enlarged postmentals (smaller than
mental, and lager than chin scales); postmentals not in
contact and bordered posteriorly by 5 unkeeled chin
scales (smaller than nostrils), in contact with only 1st in-
fralabials; ventral scales smaller than chin scales. Chin
scales smooth, rounded, juxtaposed; gular scales keeled;
pectoral and abdominal scales smooth, subimbricate to
imbricate towards precloacal region, abdominal scales
slightly larger than dorsals; 23 belly scales across venter;
scales around vent and base of tail smooth, subimbricate;
3 precloacal pores; 5/5 femoral pores; 10/10 unpored ante-
rior femoral scales; 2/2 enlarged posterior femoral scales.
Regenerated tail of holotype longer than the snout-vent
length (TAL 116.0% of SVL); hemipenial bulge greatly
swollen (TBW 3.7 mm). Heterogeneous scales on dorsum
of tail directed backwards, spine-like tubercles present at
base of tail; tail with 4 – 5 small attened obtuse scales
forming whorls; a large blunt post-cloacal spur on each
side, dorso-ventrally attened and narrow; subcaudals
smooth, diamond-shaped, tiny, arranged in an irregular
median series.
Forelimbs moderately short, slender (LAL 17.1% of
SVL, UAL 18.5% of SVL); hind limbs long, tibia quite
shorter than the femur (TBL 17.8% of SVL, FEL 18.6%
of SVL). Anterior and posterior surfaces of upper arm
with keeled and less imbricate scales, scales of anterior
surface twice as large as those of the other parts; An-
terior, dorsal and posterior surfaces of lower arm with
keeled and less imbricate scales, ventral surface with
smooth, imbricate scales. Scales on dorsal surface of fe-
mur smooth and granular, less imbricate, keeled scales
on the anterior and posterior surfaces, scales on the an-
terior and ventral surfaces twice the size of those on the
other aspects of limb. Dorsal, anterior and posterior sur-
faces of tibia with keeled and less imbricate scales, ven-
tral surface with smooth, subimbricate scales, scales of
the ventral surface twice as large as those of other parts.
Manus and pes with keeled granules dorsally and ventral-
ly; dorsal surfaces of digits with granular scales. Digits
elongate and slender with inected distal phalanges joint,
all bearing slightly recurved claws. Subdigital lamellae
entire (except divided at rst interphalangial joint), un-
notched; total lamellae on manus (left/right): digit I, 8/8,
digit II, 12/12, digit III, 16/16, digit IV, 16/16, digit V,
14/14; total lamellae on pes (left/right): digit I, 9/9, digit
II, 13/13, digit III, 15/16, digit IV, 18/18, digit V, 17/17;
interdigital webbing absent; length of digits of manus
(left): I (1.4 mm), II (1.8 mm), V (2.3 mm), III (2.4 mm),
IV (2.9 mm); length of digits of pes (left): I (1.4 mm), II
(3.1 mm), III (3.6 mm), V (3.9 mm), IV (4.1 mm).
Variation of the type series. The SVL of adult speci-
mens in the type series (n = 3) ranges from 27.2 to 31.8
mm; infralabials 7 – 8; interorbital scales 37 – 39; chin
scales 5 – 6; postnasals 1 – 2, internasals 1 – 2; canthal
scales 10 – 11; supraciliaries above the eye 15 – 17; scales
from eye to tympanum 23 – 26; tubercles on posterior
ank 5 – 6; total lamellae on digit I of the manus 8 – 9,
lamellae on digit II of manus 12 – 13, lamellae on digit III
of manus 14 – 16, lamellae on digit V of manus 14 – 15;
total lamellae on digit I of pes 8 – 9, lamellae on digit II
of pes 12 – 13, lamellae on digit III of pes 15 – 16, lamel-
lae on digit IV of pes 17 – 18; precloacal pores in males
3 – 5, unpored anterior femorals in males 9 – 10, unpored
posterior femoral scales in males 2 – 3; ventral scales
125 – 128, paravertebral granules 134 – 138, midbody
scales 92 – 98, and belly scales across venter 23 – 25.
Color of living specimens. Dorsum of head, body and
limbs varies from light brown to golden yellow, 5 – 6 fad-
ed irregular ‘W’-shaped, dark cross bands on the trunk;
an oblique black line between the eye and the nostrils on
either side, 2 straight, faded brown postorbital stripes ex-
tend from eyes posteroventrally, and a dark line is present
in the occipital area (Fig. 17); tail grey-white dorsally,
with 10 – 12 faded brown cross-bands; pupil is circular
and black with the surrounding orange and white mar-
gins, with supraciliaries scales being yellowish; suprala-
bials yellowish, infralabaials greyish dusted with black;
snout yellowish; mid-gular scales cream with dark spots;
pectoral, abdominal, cloacal and subcaudal scales white,
without spotting; dorsumof limbs with brown patches;
manus and pes with black and cream white stripe ar-
rangement.
Color of preserved specimens. Dark brown dorsum with
pale colored, ‘W’-shaped irregular pale bands distinct;
ventral surface dirty white with some scales on throat,
thigh, tail base and arms with dark brown margins.
Etymology. The specic epithet is an eponym Latinized
(butewai) in the masculine genitive singular, honouring
the Sri Lankan warrior ‘Butewe Rate Rala’ for his valiant
feats in the Great Rebellion of 1817 – 1818, which was
initiated in Uva-Wellassa.
Habitat and ecology. Bambarabotuwa Forest Reserve
(6.623506°–6.674431° N and 80.588214°–80.643431°E;
altitude range from 380 – 850 m) is a Dipterocarpus-
dominated (Mahogany mixed) tropical wet-evergreen
rainforest (Gunatileke & Gunatileke, 1990), appro xi -
mately 3,500 ha in size, situated in the wet zone of
south-central Sri Lanka (Sabaragamuwa Province, Rat-
napura District). The mean annual rainfall varies be-
tween 3,000 – 4,000 mm, received mainly during the
southwest monsoon (May – September). The mean an-
nual temperature of the area is 28.5 – 30.7°C ºC. Many
granite caves and rock ledges are found at this locality
(Fig. 18). Cnemaspis butewai sp. nov. is very common,
at least at this locality with 6 conrmed locations. We
found 24 (± 0.2) geckos per man-hour after surveying
a total area of 45 ha. This species was restricted to rock
outcrops and granite caves in forested areas, and reached
heights of 10 – 15 m on vertical surfaces. These micro-

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habitats were poorly illuminated (light intensity: 0 – 578
Lux), relatively moist (relative humidity: 73 – 92%),
cool and wetty (ambient temperature: 29.7 – 30.4°C,
rock surface temperature: 25.2 – 27.9°C, canopy cover:
70 – 90%). The geckos were active during the day time
(900 – 1600 h) and, when disturbed, sought refuge in the
crevices of the rocky caves and ledges. The new spe-
cies was sympatric (at both local habitat and the micro-
site scale) with several other geckos (Cyrtodactylus sp.,
Gehyra mutilata, Hemidactylus depressus, H. frenatus,
H. parvima culatus). Eggs were observed in granite crev-
ices, typically laid in pairs. The eggs were pure white
and almost spherical in shape (mean diameter 4.7 ±0.01
mm (n = 19)), with a slightly attened side attached to
the rocky substrate.
Conservation status. Application of the IUCN Red List
criteria indicates that C. butewai sp. nov. is Critically
Endangered (CR) due to having an area of occupancy
(AOO) <10km2 (six locations, 0.2 km2 in total assuming
a 100 m radius around the georeferenced location) and
an extent of occurrence (EOO) <100 km2 (3.8 km2) at the
lower elevations of Sabaragamuwa Province [Applicable
criteria is B2-b (iii)].
Phylogenetic position. A member of C. kandiana Clade
B; the new species belongs to C. pulchra species group
(BV) (Fig. 2), within which it is most closely related to
C. pulchra (p-distance 4.6%) and the undescribed can-
didate species Cnemaspis sp. 3 (p-distance 4.9%) (see
Tab. 2).
Gegraphic distance. Cnemaspis butewai sp. nov. most
closely resembles C. kandiana and C. menikay morpho-
logically. The type localities of these species are sepa-
rated by ~ 70 km (Kandy area) and ~ 85 km (Allauwa
in Kegalle) straight line distance from Bambarabotuwa
in Opa naka.
Fig. 17. Cnemaspis butewai sp. nov. male paratype (NMSL.2019.07.02) in life in-situ (A) dorsolateral aspect showing irregular colors on
labials and trunk, and (B) dorsal view of the full body depicting color pattern (Photos: Majintha Madawala).

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Cnemaspis kivulegedarai sp. nov.
ZooBank urn:lsid:zoobank.org:act:EE63DFF2-250D-47E4-ACE7-
DFE5EDD93E02
Kivulegedaras’ day gecko (English)
Kivulegedarage diva-seri hoona (Singhala)
Figs. 19 – 21; Table 8.
Holotype. NMSL.2019.08.01, adult male, 28.5 mm SVL (Fig. 19),
collected from a large granite wall in Keer thibandarapura, Walapa-
ne, Nuwara Eliya District, Central Province, Sri Lanka (7.127933°
N, 80.867364° E, WGS 1984; elevation 539 m; around 14.00 hrs)
on 17 June 2018 by Suranjan Karunarathna and Anslem de Silva.
Paratypes. NMSL.2019.08.02, adult female, 31.2 mm SVL, and
NMSL.2019.08.03, adult female, 29.8 mm SVL, collected from a
large granite wall in Keerthibandarapura, Walapane, Nuwara Eliya
District, Central Province, Sri Lanka (7.140061° N, 80.872467° E,
WGS 1984; elevation 562 m; around 15.00 hrs) on 17 June 2018 by
Suranjan Karunarathna and Anslem de Silva.
Diagnosis. Cnemaspis kivulegedarai sp. nov., can be
readily distinguished from its Sri Lankan congeners by
a combination of the following morphological and mer-
istic characteristics: maximum SVL 31.2 mm; dorsum
with heterogeneous, smooth, granular scales; 1 interna-
sal, 2/2 supranasals and 1/1 postnasal present; 30 – 34 in-
terorbital scales; 15 – 17 supraciliaries, 12 canthal scales,
17 – 21 eye to tympanum scales; 3 enlarged postmentals;
postmentals bounded by 5 chin scales; chin with smooth
granules; gular, pectoral and abdominal scales smooth,
subimbricate; 19 belly scales across the venter; 4 – 5
weakly developed tubercles on posterior ank; 131 – 133
paravertebral granules linearly arranged; 2 precloacal
pores, 4 – 5 femoral pores on each side in males separat-
ed by 9 – 10 unpored anterior femoral scales, 2 unpored
posterior femoral scales; 109 – 114 ventral scales; 69 –
76 midbody scales; subcaudals smooth, diamond shaped,
in irregular series forming a narrow median row; 7 su-
pralabials; 6 – 7 infralabials; 14 total lamellae on fourth
digit of manus, and 15 total lamellae on fourth digit of
pes.
Comparisons with other species. Among species of the
C. kandiana clade sensu aGarWal et al. (2017) C. ki vu-
legedarai sp. nov. differs from C. amith, C. gotaimbarai
sp. nov., C. kumarasinghei, C. latha and C. nandimithrai
sp. nov. by having heterogeneous (versus homogeneous)
dorsal scales. It also diagnosed from C. pava, C. pul-
chra, C. samanalensis, C. silvula, C. tropidogaster and
C. upendrai by having smooth (versus keeled) pectoral
and ventral scales, from C. butewai sp. nov., C. kandi-
Fig. 18. General habitat of Cnemaspis butewai sp. nov. at Bambarabotuwa isolated forest hill, Ratnapura District, Sri Lanka (A) complete
view of the wet forest, (B) thick, cool and shady forest, (C) moss covered granite cave wall microhabitat (Photos: Majintha Madawala).

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ana, C. menikay and C. retigalensis by having smooth
(versus keeled) chin and gular scales; from C. kallima by
having fewer ventral scales (109 – 114 versus 131 – 138),
and fewer total lamellae on digit IV of pes (15 versus
18 – 20); and from C. ingerorum by having more ventral
scales (109 – 114 versus 88 – 95), and fewer total lamellae
on digit IV of pes (15 versus 17 – 18).
The new species, C. kivulegedarai sp. nov. also
clearly differs from the species of the C. podihuna clade
sensu aGarWal et al. (2017). From C. alwisi, C. gemunu,
C. godagedarai, C. hitihami sp. nov., C. kohukumburai
sp. nov., C. nilgala, C. phillipsi, C. punctata, C. raja-
karunai, C. rammalensis, and C. scalpensis it differs
by the presence (versus absence) of precloacal pores
Fig. 19. Close-ups of Cnemaspis kivulegedarai sp. nov. male holotype (NMSL.2019.08.01) (A) dorsal head, (B) lateral head, (C) ventral
head, (D) homogeneous dorsal scales, (E) scales on lateral surface of trunk, (F) smooth ventral scales, (G) cloacal characters with precloa-
cal pores and femoral pores (H) subdigital lamellae on manus, (I) subdigital lamellae on pes, (J) dorsal scalation of tail, (K) lateral side of
tail, and (L) oval shaped subcaudals (Photos: Suranjan Karunarathna).

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and also by the absence of clearly enlarged, hexagonal
or subhexagonal subcaudal scales. From C. kandambyi,
C. molligodai and C. podihuna it differs by having heter-
ogeneous (versus homogeneous) dorsal scales and by the
absence of clearly enlarged, hexagonal or subhexagonal-
shaped subcaudal scales.
Description of Holotype. An adult male, 28.5 mm SVL.
Body slender, relatively short (TRL 36.8% of SVL).
Head relatively large (HL 32.0% of SVL, HL 86.8% of
TRL), narrow (HW 17.3% of SVL, HW 54.2% of HL),
depressed (HD 8.9% of SVL, HD 27.7% of HL) and
distinct from neck. Snout relatively long (ES 75.7% of
HW, ES 41.0% of HL), more than twice length of eye
diameter (ED 49.1% of ES), more than half length of jaw
(ES 65.2% of JL), snout slightly concave in lateral view;
eye relatively small (ED 20.1% of HL), twice as large as
the ear (EL 48.1% of ED), pupil rounded; orbit length
greater than eye to ear distance (OD 133.2% of EE) and
also greater than to the length of IV digit of manus (OD
101.9% of DLM IV); supraocular ridges not prominent;
ear opening very small (EL 9.7% of HL), deep, taller
than wide, larger than nostrils; single row of scales sepa-
rate orbit from supralabials; interorbital distance equal
to snout length (IO 100.3% of ES), head length twise a
long than interorbital distance (IO 41.1% of HL); eye to
nostril distance little greater than eye to ear distance (EN
105.4% of EE).
Dorsal surface of the trunk mixed with conical het-
erogeneous granules, 131 paravertebral granules; 109
smooth midventral scales; 69 midbody scales; 4/5 weakly
developed tubercles on the anks; ventrolateral scales
not enlarged; granules on snout strongly keeled, larger
than those on interorbital and occipital regions; canthus
rostralis not pronounced, 12/12 smoothly rounded scales
from eye to nostril; scales of interorbital region oval and
smooth; tubercles present on the sides of the neck, and
absent around the ear; ear opening vertically oval, slanting
from anterodorsal to posteroventral, 17/18 scales between
anterior margin of the ear opening and the posterior mar-
gin of the eye. Supralabials 7/7, infralabials 6/6, becoming
smaller towards the gape. Rostral scale wider than long,
partially divided (75%) by a median groove, in contact
with rst supralabial. Nostrils separated by 2/2 enlarged
supranasals with 1 internasal and 1/1 postnasal; no en-
larged scales behind the supranasals. Nostrils oval, dorso-
laterally orientated, not in contact with rst supralabials.
Mental subtriangular, as wide as long, posteriorly in
contact with 3 enlarged postmentals (smaller than mental,
and lager than chin scales); postmentals in contact and
bordered posteriorly by 5 smooth chin scales (smaller
than nostrils), in contact only with 1st infralabials; ventral
scales smaller than chin scales. Smooth, rounded, juxta-
posed scales on the chin and gular region; pectoral and
abdominal scales smooth, subimbricate to imbricate to-
wards precloacal region, abdominal scales slightly larger
than dorsals; 19 belly scales across venter; scales around
vent and base of tail smooth, subimbricate; 2 precloacal
pores; 4/5 femoral pores; 10/9 unpored anterior femoral
scales; 2/2 enlarged posterior femoral scales. Regener-
ated tail of holotype longer than the snout-vent length
(TAL 135.5% of SVL); hemipenial bulge greatly swol-
len (TBW 2.8 mm), heterogeneous scales on tail dorsum
directed backwards, spine-like tubercles present at the
base of tail; tail with 4 – 5 enlarged attened obtuse scales
forming whorls; a large post-cloacal spur on each side,
dorsoventrally attened and narrow; median subcaudals
smooth, irregular and oval.
Forelimbs moderately short, slender (LAL 15% of
SVL, UAL 16.6% of SVL); hind limbs long, tibia quite
shorter than the femur (TBL 19.6% of SVL, FEL 19.7%
of SVL). Anterior and posterior surfaces of upper arm
with keeled and less imbricate scales, scales of the an-
terior surface twice as large as those of the other parts,
scales of dorsal and ventral surfaces smooth. Dorsal, an-
terior and posterior surfaces of lower arm with keeled
and less imbricate scales, ventral surface with smooth,
imbricate scales. Scales on dorsal and posterior surfaces
of the femur smooth and granular, anterior surface with
keeled and subimbricate scales; scales of ventral surface
smooth and less imbricate. Dorsal, anterior and posterior
surfaces of tibia with keeled subimbricate scales, and
ventral scales smooth, subimbricate; scales of ventral
surface twice as large as those on other aspects. Dorsal
and ventral scales ofmanus and pes with keeled gran-
ules; dorsal surfaces of digits with granular scales. Digits
elongate and slender with inected distal phalanges joint,
all bearing slightly recurved claws. Subdigital lamellae
entire (except divided at rst interphalangial joint), un-
notched; total lamellae on manus (left/right): digit I, 8/9,
digit II, 11/11, digit III, 13/13, digit IV, 14/14, digit V,
11/11; total lamellae on pes (left/right): digit I, 8/9, digit
II, 11/11, digit III, 13/13, digit IV, 15/15, digit V, 12/12;
interdigital webbing absent; length of digits of manus
(left): I (1.7 mm), II (1.8 mm), V (2.6 mm), III (2.7 mm),
IV (3.2 mm); length of digits of pes (left): I (1.6 mm), II
(3.2 mm), III (3.7 mm), V (3.9 mm), IV (4.1).
Variation of the type series. The SVL of adult speci-
mens in the type series (n = 3) ranges from 28.5 to 31.2
mm; interorbital scales 30 – 34; supraciliaries above the
eye 15 – 17; scales from eye to tympanum 17 – 21; total
lamellae on digit I of the manus 8 – 9, lamellae on digit
III of manus 12 – 13; total lamellae on digit I of pes 8 – 9,
lamellae on digit II of pes 10 – 11, lamellae on digit III
of pes 13 – 14, lamellae on digit V of pes 11 – 12; para-
vertebral granules 131 – 133, ventral scales 109 – 114,
midbody scales 69 – 76; femoral pores in males 4 – 5, and
unpored anterior femoral scales in males 4 – 5.
Color of living specimens. Dorsum of head, body and
limbs generally orange-brown; marked with 5 faded, ir-
regular brown cross-markings on the body dorsum and
7 – 8 cream vertebral blotches; an oblique black line is
present between the eye and the nostrils on either side,
two straight, dark brown postorbital stripes extend from
eyes posteroventrally (Fig. 20). Tail grey-brown on
dor sum, with 9 – 11 irregular ‘W’-shaped faded brown

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cross-bands present; pupil circular and black with the
surrounding ring is gold and orange, with supraciliary
scales light brownish; supralabials yellowish dusted with
black; infralabaials greyish dusted with black; mid-gular
scales greyish white dusted with black; pectoral, abdomi-
nal, cloacal and subcaudal scales white without tiny dark
spots; dorsum of limbs with irregular brown patches and
lines; dorsum of manus and pes with black and cream
white stripes alternating.
Color of preserved specimens. Dorsum dark brown with
grey, with indistinct irregular brown markings; vertebral
blotches dirty white; venterdirty white with some gular,
abdominal, thigh, tail base and forelimb scales with dark
brown margins.
Etymology. The specic epithet is an eponym Latinized
(kivulegedarai) in the masculine genitive singular, hon-
ouring ‘‘Kivulegedara Mohottala’ (a warrior) for his val-
iant feats in the Great Rebellion of 1817 – 1818, which
was initiated in Uva-Wellassa.
Habitat and ecology. The Keerthibandarapura area
(7.138467°– 7.156942° N and 80.856094°– 80.880067°
E; altitude range from 500 – 750m) is characterized as a
tropical wet-evergreen forest interspaced with savannah
(Gunatileke & Gunatileke, 1990), approximately 900 ha
in size, located in the submontane intermediate zone of
central Sri Lanka (Central Province, Nuwara Eliya Dis-
trict). The mean annual rainfall is 3,000 – 4,000 mm,
received mainly during the southwest monsoon
(May – September) season. The mean annual tempera-
ture of the area is 22.6 – 25.2 ºC. Cnemaspis kivulege-
darai sp. nov. was very commonat this locality, as we
found 31 (±0.3) geckos per man-hour while surveying a
total surveyed area of 60 ha with 4 conrmed locations.
Fig. 20. Cnemaspis kivulegedarai sp. nov. male holotype (NMSL.2019.08.01) in life in-situ (A) dorsolateral view of the full body, and (B)
dorsal view depicting the pale vertebral line (Photos: Majintha Madawala).

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288
This species was restricted to rock outcrops and granite
caves in forested areas, and was found at heights of 6
m along vertical surfaces (Fig. 21). These habitats were
poorly illuminated (0 – 471 Lux), relatively moist (rela-
tive humidity: 76 – 89%), and cool (ambient temperature:
28.9 – 30.1°C, rock surface temperature: 25.8 – 27.4°C,
canopy cover: 65 – 80%). The geckos were diurnally ac-
tive (0800 – 1700 h) and, when disturbed, sought refuge
in the crevices of rocky caves. The new species was sym-
patric (at both local habitat and microsite scales) with
several other geckos (Gehyra mutilata, Hemidactylus
depressus, H. frenatus, H. parvimaculatus). Eggs were
observed on eadge of granite rock walls, typically laid
in pairs (rarely trias) in communal nests. The eggs were
pure white and almost spherical in shape (mean diameter
4.7 ± 0.02 mm; n = 12), with a slightly attened side at-
tached to the rocky substrate.
Table 8. Morphometric and Meristic data of holotype and two paratypes of Cnemaspis kivulegedarai sp. nov. from Walapane, Nuwara
Eliya District, Sri Lanka (Abbreviations: L – left, R – right, M – male, F – female).
Measurements
(mm)
NMSL
2019.08.01
NMSL
2019.08.02
NMSL
2019.08.03 Counts
NMSL
2019.08.01
NMSL
2019.08.02
NMSL
2019.08.03
Holotype
(M)
Paratype
(F)
Paratype
(F)
Holotype
(M)
Paratype
(F)
Paratype
(F)
ED 1.8 1.8 1.7 FLSP (L/R) 4/5 4/4 5/5
OD 3.2 3.1 3.1 SUP (L/R) 7/7 7/7 7/7
EN 2.5 2.6 2.6 INF (L/R) 6/6 6/7 6/6
ES 3.7 3.8 3.9 INOS 32 30 34
SN 1.3 1.4 1.4 PM 3 3 3
NW 0.2 0.1 0.2 CHS 5 5 5
EE 2.4 2.5 2.5 SUN (L/R) 2 2 2
SA 14.1 14.2 14.3 PON (L/R) 1 1 1
EL 0.9 0.8 0.6 INT 1 1 1
IO 3.7 3.7 3.7 SUS (L/R) 16/17 16/16 15/16
IE 3.9 3.9 3.9 BET (L/R) 18/17 19/21 18/17
HL 9.1 8.9 9.0 CAS (L/R) 12/12 12/12 12/12
HW 4.9 4.2 4.6 TLM (i) (L/R) 8/9 9/9 9/9
HD 2.5 2.6 2.6 TLM (ii) (L/R) 11/11 11/11 11/11
JL 5.7 5.7 5.5 TLM (iii) (L/R) 13/13 12/13 13/13
IN 1.7 1.5 1.6 TLM (iv) (L/R) 14/14 14/14 14/14
SED 8.3 8.1 8.2 TLM (v) (L/R) 11/11 11/11 11 /11
UAL 4.7 4.6 4.6 PG 131 133 132
LAL 4.3 4.1 4.2 MBS 69 76 71
PAL 3.1 3.2 3.2 MVS 109 114 111
DLM (i) 1.7 1.5 1.6 BLS 19 19 19
DLM (ii) 1.8 1.9 1.9 TLP (i) (L/R) 8/9 8/8 9/9
DLM (iii) 2.7 2.7 2.9 TLP (ii) (L/R) 11/11 11/11 11/10
DLM (iv) 3.2 3.1 3.2 TLP (iii) (L/R) 13/13 14/14 13/13
DLM (v) 2.6 2.5 2.5 TLP (iv) (L/R) 15/15 15/15 15/15
SVL 28.5 31.2 29.8 TLP (v) (L/R) 12/12 11/12 12/12
TRL 10.5 10.5 10.6 PCP 2 — —
TW 5.7 5.9 5.9 FP (L/R) 4/5 — —
TD 3.6 3.5 3.6 AFS (L/R) 10/9 — —
FEL 5.6 5.4 5.5 PFS (L/R) 2/2 — —
TBL 5.6 5.4 5.5
HEL 4.0 3.8 3.8
DLP (i) 1.6 1.5 1.6
DLP (ii) 3.2 3.3 3.2
DLP (iii) 3.7 3.7 3.6
DLP (iv) 4.1 4.2 4.2
DLP (v) 3.9 3.9 3.7
TAL 38.6 38.6 39.0
TBW 2.8 2.6 2.7
TBD 2.4 2.1 2.2

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Conservation status. Application of the IUCN Red List
criteria indicates that C. kivulegedarai sp. nov. is Criti-
cally Endangered (CR) due to having an area of occu-
pancy (AOO) < 10 km2 (four locations, 0.1 km2 in total
assuming a 100 m radius around the georeferenced lo-
cation) and an extent of occurrence (EOO) < 100 km2
(2.3 km2) in the mid-elevations of Central Province [Ap-
plicable criteria is B2-b (iii)].
Phylogenetic position. A member of C. kandiana Clade
B; the new species belongs to the C. latha species group
(BIV) (Fig. 2), within which it is reconstructed as a sister
species of C. latha (p-distance 5.1%) (see Tab. 2).
Geographic distance. Cnemaspis kivulegedarai sp. nov.
most closely resembles C. latha, the type locality of which
is ~ 32 km (Bandarawela in Badulla District) straight line
distance from Keerthibandarapura in Walapane.
Discussion
Sri Lanka is recognized as one of the global hotspots for
herpetofaunal diversity as well as a local center for en-
demism (Bossuyt et al., 2004; meeGaskumBura et al.,
2002). The addition of six more endemic gecko species
to Sri Lankan reptile checklist underpines that the island
is a centre of reptilian diversity and endemism (amar-
asinGhe et al., 2014, 2015; BatuWita, 2016; BatuWita
& uDuGampala, 2017; BatuWita et al., 2019; De silva
et al., 2019; karunarathna et al., 2019; WiCkramasin-
Ghe, 2016; WiCkramasinGhe et al., 2017). Cnemaspis
now comprises 32 species in Sri Lanka, bringing the
total number of geckos recorded in the country to 54.
Among Sri Lankan gekkonids, 44 (81%) species are en-
demic to the island, most of which are restricted to the
wet zone (> 2,000 mm of annual average precipitation).
Of them, 20 (37%) are Critically Endangered, 9 (16%)
are Endagered, 5 (9%) are Vulnerable and 4 (7%) are
Data Decient. However, as our study demonstrated, Sri
Lanka’s Cnemaspis diversity is not limited to the south-
western lowlands or to the central massif, but is scattered
throughout multiple bio-climatic regions and oristic
Fig. 21. General habitat of Cnemaspis kivulegedarai sp. nov. at Walapane isolated forest hill, Nuwara Eliya District, Sri Lanka (A) com-
plete view of the water shed area in the mountain, (B) rock outcrop area in the type locality, (C) communal egg depositional site (Photos:
Majintha Madawala).

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290
Table 9. Key characters of currently known 32 Cnemaspis species in Sri Lanka (Abbreviations: MM – Millimeters, SVL – Maximum Snout to vent length, SUB – Subcaudals, SUP – Supralabials, INF –
Infralabials, PG – Paravertebral, IFS – Interfemoral scales, FLSP – Flank spines, PCP – Precloacal pores, FP – Femoral pores, HET – Heterogeneous, HOM – Homogeneous, KD – Keeled, SM – Smooth).
Species SVL
(mm) Dorsal Gular Pectoral Abdomen SUB SUP INF Ventrals Belly Midbody PG IFS FLSP PCP FP Lamellae
4th nger
Lamellae
4th toe
Cnemaspis kandiana clade
Group (1)
C. pava 32.4 HET KD KD KD Small 7 – 8 6 – 7 139 – 145 22 – 25 64 – 75 83 – 98 – 9 – 11 2 – 4 4 – 5 16 – 17 18 – 19
C. pulchra 34.2 HET KD KD KD Small 7 – 8 7 – 8 120 – 135 24 – 27 67 – 73 94 – 103 – 5 – 7 3 – 4 4 – 6 15 – 17 17 – 20
C. samanalensis 37.5 HET KD KD KD Small 8 – 10 8 – 9 128 – 144 19 – 20 61 – 67 64 – 72 – 5 – 6 3 – 4 3 – 5 16 – 17 18 – 20
C. silvula 28.6 HET KD KD KD Small 7 – 8 7 – 8 132 – 139 19 – 21 73 – 81 102 – 113 – 10 – 15 3 – 4 4 – 5 15 – 16 18 – 19
C. tropidogaster 31.7 HET KD KD KD Small 7 – 8 7 – 8 132 – 146 21 – 25 92 – 98 99 – 106 – 5 – 7 3 – 4 4 – 5 16 – 17 18 – 19
C. upendrai 35.2 HET KD KD KD Small 7 – 8 7 – 8 112 – 128 16 – 25 69 – 74 97 – 102 – 13 – 15 2 – 3 4 – 5 17 – 18 17 – 21
Group (2)
C. ingerorum 26.9 HET SM SM SM Small 7 7 88 – 95 17 – 21 62 – 69 93 – 101 – 7 – 9 2 5 13 – 16 17 – 18
C. kivulegedarai sp. nov. 31.2 HET SM SM SM Small 7 6 – 7 109 – 114 19 69 – 76 131 – 133 – 4 – 5 24 – 5 14 15
C. kallima 35.1 HET SM SM SM Small 7 – 8 7 – 8 131 – 138 19 – 23 67 – 74 99 – 107 – 12 – 15 3 – 4 4 – 5 16 – 18 18 – 20
C. butewai sp. nov. 31.8 HET KD SM SM Small 8 7 – 8 125 – 128 23 – 25 92 – 98 134 – 138 – 5 – 6 3 – 5 5 16 17 – 18
C. kandiana 34.6 HET KD SM SM Small 8 – 9 7 – 8 119 – 138 19 – 20 68 – 75 86 – 99 – 5 – 7 2 – 4 3 – 4 12 – 14 18 – 20
C. menikay 28.0 HET KD SM SM Small 7 – 9 7 – 8 124 – 138 20 – 26 71 – 79 83 – 98 – 13 – 15 1 – 2 3 – 4 14 – 15 15 – 17
C. retigalensis 30.8 HET KD SM SM Small 7 – 8 7 – 8 121 – 128 16 – 20 69 – 77 82 – 86 – 4 – 5 13 – 4 14 – 15 16 – 20
Group (3)
C. amith 33.0 HOM KD SM SM Small 7 – 8 7123 – 131 19 – 21 67 – 74 79 – 84 – 4 – 5 3 3 16 – 17 18 – 19
Group (4)
C. gotaimbarai sp. nov. 33.7 HOM SM SM SM Small 7 – 8 8 – 9 129 – 138 23 – 25 72 – 79 117 – 121 – 5 – 6 2 – 4 316 – 17 19 – 20
C. kumarasinghei 31.6 HOM SM SM SM Small 7 – 8 7 – 8 120 – 134 17 – 21 87 – 94 61 – 68 – 7 – 9 2 – 3 3 – 5 15 – 16 16 – 18
C. latha 30.4 HOM SM SM SM Small 7 – 8 7 – 8 109 – 115 13 – 15 69 – 73 72 – 79 – 5 – 7 2 – 3 4 – 5 15 – 17 17 – 18
C. nandimithrai sp. nov. 31.7 HOM SM SM SM Small 5 – 6 6108 – 112 25 – 27 87 – 89 95 – 99 – 3 – 4 2 – 4 2 – 4 12 – 13 19 – 20
Cnemaspis podihuna clade
Group (1)
C. alwisi 40.4 HOM SM SM SM Enlarged 8 – 10 7 – 9 145 – 153 27 – 31 71 – 78 89 – 97 18 – 19 4 – 5 – 7 – 9 15 – 17 17 – 21
C. godagedarai 35.5 HOM SM SM SM Enlarged 7 – 8 7 – 8 133 – 137 21 – 23 98 – 102 101 – 106 85 – 6 – 12 – 13 17 – 18 20 – 21
C. gemunu 34.0 HOM SM SM SM Enlarged 8 – 10 7 – 8 112 – 118 13 – 16 74 – 87 79 – 93 10 – 12 7 – 8 – 11 – 14 15 – 17 18 – 19
C. hitihami sp. nov. 41.7 HOM SM SM SM Enlarged 8 – 9 7 – 9 132 – 135 21 96 – 99 143 – 149 24 – 26 4 – 5 – 5 – 10 18 – 19 21 – 22
C. kohukumburai sp.
nov. 34.5 HOM SM SM SM Enlarged 8 – 9 7 – 8 131 – 134 23 81 – 88 150 – 159 25 7 – 8 – 6 – 9 21 – 22 23 – 25
C. nilgala 32.9 HOM SM SM SM Enlarged 7 – 8 6 – 7 122 – 129 17 – 19 71 – 78 179 – 187 14 – 15 3 – 4 – 7 – 9 17 17 – 18

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Species SVL
(mm) Dorsal Gular Pectoral Abdomen SUB SUP INF Ventrals Belly Midbody PG IFS FLSP PCP FP Lamellae
4th nger
Lamellae
4th toe
C. phillipsi 36.6 HOM SM SM SM Enlarged 8 – 9 8 – 9 128 – 143 18 – 25 76 – 91 86 – 93 11 – 14 4 – 6 – 15 – 16 16 – 19 17 – 19
C. punctata 39.9 HOM SM SM SM Enlarged 7 – 10 7 – 9 129 – 137 20 – 29 71 – 78 83 – 91 25 – 27 11 – 13 – 5 – 7 17 – 18 17 – 23
C. rajakarunai 40.2 HOM SM SM SM Enlarged 8 – 9 9 – 11 146 – 186 26 – 29 69 – 74 81 – 85 20 – 22 5 – 6 – 7 – 8 16 – 20 19 – 22
C. rammalensis 53.8 HOM SM SM SM Enlarged 8 – 10 8 – 9 186 – 207 25 – 28 119 – 131 94 – 96 19 – 24 4 – 5 – 14 – 16 22 – 23 22 – 23
C. scalpensis 36.6 HOM SM SM SM Enlarged 7 – 9 7 – 8 120 – 131 17 – 19 81 – 89 102 – 112 8 – 12 9 – 11 – 13 – 15 17 – 18 19 – 21
Group (2)
C. kandambyi 23.6 HOM SM SM SM Enlarged 7 – 8 7 – 8 128 – 137 16 – 17 71 – 77 85 – 92 – 4 – 5 3 – 4 5 – 6 11 – 14 19 – 20
C. molligodai 29.0 HOM SM SM SM Enlarged 8 – 10 7 – 8 127 – 135 15 – 19 73 – 82 76 – 83 – 5 – 7 4 – 5 8 – 9 15 – 18 19 – 23
C. podihuna 24.7 HOM SM SM SM Enlarged 7 – 9 6 – 8 111 – 118 15 – 19 79 – 83 102 – 106 – 4 – 6 3 – 4 3 – 6 14 – 15 18 – 19
Table 9 continued.
regions, which suggests intricate biogeographic patterns possibly due to
multiple colonizations from the Indian mainland rather than a singular
event of insular radiation (aGarWal et al., 2017).
Since the early 2000s, there has been a surge of taxonomic research
on the faunal diversity of Sri Lanka, mostly targeting vertebrates (Batu-
Wita & pethiyaGoDa, 2007; Bossuyt et al., 2004, 2005; manamenDra-
araChChi & pethiyaGoDa, 2005; manamenDra-araChChi et al., 2007;
meeGaskumBura et al., 2002, 2007; pethiyaGoDa et al., 2006, 2008, 2012)
leading to the discovery and description of several new species across
numerous habitats. For example, the number of day geckos of the ge-
nus Cnemaspis recognised by DeraniyaGala (1953) has undergone over
a eight-fold increase (from 4 to 32 species) since this taxonomic renais-
sance (BatuWita et al., 2019; De silva et al., 2019; karunarathna et al.,
2019). Molecular phylogenetic analyses have indicated two distinct Sri
Lankan clades, namely: C. kandiana and C. podihuna (aGarWal et al.,
2017). Furthermore, among recent taxonomic and systematic research
investigations, the rediscovery of purportedly extinct species, ‘C. amith’
and another species categorized as Data Decient, ‘C. tropidogaster’, are
noteworthy (amarasinGhe et al., 2016). Use of molecular phylogenet-
ics, detailed elucidation of morphological character states as well as their
polarity, greater access to remote locations and wilderness, and enhanced
knowledge on historical geology and geography have contributed to the
taxonomic advances not only of Cnemaspis, but also among other reptiles
(aGarWal & karanth, 2015; Bauer et al., 2007; Grismer et al., 2014a;
karunarathna et al., 2016). Thus, continuation of faunal surveys and
detailed examination of morphological as well as genetic diagnostic fea-
tures is critical in revealing the true Cnemaspis diversity in Sri Lanka. We
strongly recommend that such studies focus on isolated hills, smaller for-
ests, rock outcrops, and granite caves, including historical tunnel systems
(De silva et al., 2019).
Among the previously recognized species of Cnemaspis, 14 belong
to C. kandiana clade: C. amith, C. ingerorum, C. kallima, C. kandiana,
C. ku marasinghei, C. latha, C. menikay, C. pava, C. pulchra, C. retiga len-
sis, C. samanalensis, C. silvula, C. tropidogaster and C. upendrai, which
are characterized by small and irregularly shaped subcaudal scales. Out of
six new species, we assign four to this clade on this basis (C. butewai sp.
nov., C. gotaimbarai sp. nov., C. kivulegedarai sp. nov. and C. nandim-
ithrai sp. nov.). Interestingly, only four species of this clade are distrib-
uted at high elevations (above 900 m): C. pulchra (Rakwana Highlands,
southwest of central highlands) and C. samanalensis, C. upendrai and
C. latha (Central Highlands), whereas the rest are low-elevation taxa
(manamenDra-araChChi et al., 2007; WiCkramasinGhe & muniDraDasa,
2007). Further, only two species of the C. kandiana clade (C. ingerorum
and C. retigalensis) are known from the dry zone and three species (Cne-
maspis kallima, Cnemaspis kumarasinghei and Cnemaspis latha) are
known from the intermediate zone. Two new species (C. gotaimbarai sp.
nov. and C. nandimithrai sp. nov.) represent additions to the day-geckos
of the dry zone, and one new species (C. kivulegedarai sp. nov.) from the
intermediate zone. These are morphologically quite similar to C. kumar-
asinghei but each taxon occurs in a geographically isolated mountains
(Kokagala, Kudumbigala and Maragala, respectively), with the straight
line distances between these three locations are no further than 45 km.
Due to notable morphological similarities and the close geographic prox-
imity of the habitats, these three species are closely related and form a
clade in our phylogenetic analyses (see Fig. 2).
Tweleve of the previously recognized species of Sri Lankan Cnemaspis
belong to the C. podihuna clade, which includes C. alwisi, C. gemunu,
C. godagedarai, C. kandambyi, C. molligodai, C. nilgala, C. phillipsi,
C. podihuna, C. punctata, C. rajakarunai, C. rammalensis and C. scal-

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
292
Table 10. Geographic distribution and habitat features of six new Cnemaspis species from Sri Lanka (Abbreviation: m – meter, ha – hectare, mm – milimeters, Lux – light intensity, CR – critically endangered).
Species Bioclimatic
area
District Coordinates Eleva -
tion
Area size Forest
type
Micro-
habitat
Annual
Rainfall
Annual
temperature
Ambient
temperature
Substrate
temperature
Lux Relative
humidity
Canopy
cover
Conser-
vation
status
N E
C. nandimithrai
sp. nov. Dry zone Ampara
6.667519 81.747839 28 m
600 ha
Dry mixed
Semie-
vergreen
Granite cave
1000 –
1500 mm 29.8 –31.4 °C
32.2 °C 26.6 °C 643 77% 70%
CR
6.658708 81.753736 31 m Granite cave 30.5 °C 27.2 °C 532 80% 65%
6.669422 81.750561 25 m Granite cave 30.9 °C 27.7 °C 621 83% 60%
6.645103 81.725086 22 m Granite cave 31.4 °C 28.4 °C 494 71% 75%
C. gotaimbarai
sp. nov. Dry zone Ampara
7.439517 81.207967 292 m
1000 ha
Dry mixed
Semie-
vergreen
Granite cave
1500 –
2000 mm 29.6 –30.5 °C
30.2 °C 28.3 °C 752 72% 85%
CR
7.413086 81.210161 306 m Granite cave 31.5 °C 28.7 °C 821 85% 75%
7.417322 81.226047 327 m Granite cave 31.8 °C 28.2 °C 784 82% 70%
7.421439 81.223567 297 m Granite cave 30.9 °C 27.2 °C 731 69 % 80%
7.408897 81.223928 294 m Granite cave 31.5 °C 28.6 °C 815 76% 70%
C. kohukumburai
sp. nov. Wet zone Kandy
7.251800 80.509378 427 m
600 ha
Tropical
wet
ever-
green
Granite wall
3000 –
4500 mm 25.2 –27.5 °C
30.7 °C 26.2 °C 686 76% 90%
CR7.251658 80.512561 435 m Granite wall 31.2 °C 27.8 °C 547 73% 75%
7.252525 80.510281 413 m Granite cave 29.8 °C 27.3 °C 593 89% 85%
C. hitihami
sp. nov.
Intermediate
zone Monaragala
6.875428 81.357289 421 m
4100 ha
Dry mixed
Semie-
vergreen
Granite cave
1500 –
2000 mm 26.8 –28.9 °C
31.5 °C 27.3 °C 693 79% 90%
CR
6.879725 81.364986 437 m Granite cave 30.8 °C 26.8 °C 521 68% 85%
6.893278 81.369394 512 m Granite cave 29.8 °C 27.1 °C 733 73% 75%
6.895650 81.373047 487 m Granite cave 31.2 °C 27.5 °C 577 84% 80%
6.889972 81.372575 392 m Granite wall 31.9 °C 26.6 °C 684 80% 85%
6.904797 81.370878 465 m Granite cave 32.3 °C 27.2 °C 753 77% 75%
6.911967 81.373014 547 m Granite cave 30.5 °C 25.9 °C 522 82% 80%
C. butewai
sp. nov. Wet zone Ratnapura
6.630364 80.628925 445 m
3500 ha
Tropical
wet
evergreen
Granite cave
3000-
4000 mm 28.5–30.7 °C
30.1 °C 26.8 °C 551 84% 75%
CR
6.646919 80.653892 458 m Granite cave 29.9 °C 25.2 °C 495 81% 70%
6.640125 80.666367 396 m Granite cave 30.4 °C 26.2 °C 507 90% 70%
6.638986 80.659875 514 m Granite cave 29.7 °C 27.4 °C 438 88% 85%
6.648331 80.651686 482 m Granite wall 29.8 °C 25.9 °C 578 92% 90%
6.650092 80.644100 417 mm Granite cave 30.4 °C 27.9 °C 515 79% 70%
C. kivulegedarai
sp. nov.
Intermediate
zone Nuwara Eliya
7.127933 80.867364 539 m
900 ha
Tropical
wet
evergreen
Granite wall
3000-
4000 mm 22.6–25.2 °C
30.1 °C 25.8 °C 394 76% 70%
CR
7.140061 80.872467 562 m Granite wall 29.6 °C 26.1 °C 471 89% 65%
7.148364 80.866878 522 m Granite cave 29.9 °C 27.4 °C 452 81% 75%
7.157342 80.853992 587 m Granite cave 28.9 °C 26.7 °C 426 79% 80%

293
VERTEBRATE ZOOLOGY — 69
(3) 2019
pensis – characterized by clearly enlarged, hexagonal or
subhexagonal subcaudal scales. We assign two of the
new species (C. hitihami sp. nov., and C. kohukum-
burai sp. nov.) to the C. podihuna clade based on this
feature. Only four species in this clade are distributed at
high elevation (above 900 m) namely: C. gemunu (Cen-
tral Highlands), C. godagedarai (Rakwana Highlands),
C. phillipsi and C. punctata (Dumbara Highlands of the
Knuckles Range, northeast of central highlands), whereas
the rest are low elevation taxa (manamenDra-araChChi
et al., 2007; WiCkramasinGhe & muniDraDasa, 2007).
Further, only single species of the C. podihuna clade
(i.e., C. podi huna) are known from the dry zone, and
six species (i.e., Cnemaspis alwisi, Cnemaspis kandam-
byi, Cnemaspis nilgala, Cnemaspis phillipsi, Cnemaspis
punctata and Cnemaspis rammalensis) are known from
the intermediate zone – whereas the rest are wet zone
species. One new species from podihuna clade (C. hiti-
hami sp. nov.) is also from the intermediate zone. Two of
the new species described here, C. hitihami sp. nov. and
C. kohukumburai sp. nov., are morphologically similar
to C. alwisi, C. nilgala and C. rajakarunai but all are re-
stricted to geographically isolated mountains (Maragala,
Kadugannawa, Dolukanda, Nilgala and Salgala, respec-
tively), and the minimum straight line distance between
any two of these localities exceeds 35 km. In our phylo-
genetic analyses, C. hitihami sp. nov. and C. kohukum-
burai sp. nov. form a clade with C. alwisi, C. nilgala and
C. punctata (see Fig. 2). This situation presents a case
similar to that of C. kandiana clade where morphologi-
cal similarities and geographical proximity together with
mtDNA-based genealogy suggest common descent.
Our phylogenetic analyses uncovered six previously
unknown mtDNA lineages within Sri Lankan Cnemaspis,
suggesting that our understanding of the within-genus di-
versity in the island is still far from complete. With the
exception of C. hitihami sp. nov., which was included in
the analysis of aGarWal et al. (2017) as Cnemaspis sp. 10,
the other ve new species described in the present paper
are completely new and have not been included in previ-
ous phylogenies of the genus. At least eight other mtDNA
lineages of Cnemaspis from Sri Lanka are recognized as
putative candidate species and further morphological and
molecular studies are required to evaluate their taxonomic
status. Moreover, our analyses suggest that at least within
the C. podihuna and C. kandiana species complexes mo-
lecular differentiation among mtDNA lineages approach-
es species level. Within the C. podihuna complex, four
mtDNA lineages were revealed; three of them occur in
the southeastern part of Sri Lanka (localities A1, A2, A4,
see Fig. 1), while one highly divergent (p = 4.3%) line-
age tentatively identied as C. cf. podihuna is reported
from northern part of the country (Anuradhapura, locality
A3, see Fig. 1). The observed level of divergence suggest
species status for C. cf. podihuna from the Anuradhapura
District, however we do not here revise the taxonomy of
this species complex pending examination and phyloge-
netic assessment of topotype specimens of C. podihuna
(type locality – Lahugala, Eastern Province, Sri Lanka).
Within the C. kandiana species complex, we report the
presence of three moderately divergent mtDNA lineages
(p = 3.5%) occurring in a comparatively narrow area; fur-
ther molecular and morphological studies are required to
evaluate their distribution and taxonomic status.
Our morphological observations indicate the pres-
ence of four morphologically distinct subgroups within
the C. kandiana clade: subgroup (I) is distinguished by
the presence of heterogeneous dorsal scales and smooth
ventral scales; subgroup (II) is characterized by the pres-
ence of heterogeneous dorsal scales and keeled ventral
scales; subgroup (III) may be distinguished by the pres-
ence of homogeneous dorsal scales and smooth ventral
scales, and subgroup (IV) can be recognized by the
presence of homogeneous dorsal scales and keeled gu-
lar scales (Tab. 3 – 8). Further, we nd morphological
evidence for the presence of two subgroups within the
C. podihuna clade: subgroup (I) characterized by the
presence of homogeneous dorsal scales with precloacal
pores and femoral pores, and subgroup (II) distinguisha-
ble by the presence of homogeneous dorsal scales and the
absence of precloacal pores (Tab. 9). It is interesting to
note that Calodactylodes illingworthorum and Hemidac-
tylus hunae are sympatric with Cnemaspis in caves in the
intermediate and dry zones of Sri Lanka (karunarathna
& amarasinGhe, 2011; karunarathna & kumarasinGhe,
2011), especially in Uva Province, whereas Cyrtodacty-
lus spp. are sympatric with Cnemaspis in caves in the
intermediate and wet zones. Eventhough these species
share the same habitat, the specic use of microhabitats
differ with time of the day. Cnemaspis, being diurnal,
prefer the outer edges of cave walls and illuminated parts
of the caves during daytime, whereas other gecko species
prefer darker parts of the cave. During night time these
preferences are reversed.
The habitats of Cnemaspis species, including the
type localities, are undergoing habitat conversion, and
are threatened by localized human disturbances such as
encroachments primarily for tea, crop-farming, and hu-
man settlements. Other adverse anthropogenic impacts,
such as unplanned infrastructure development like road
construction as well as granite mining, untimely forest
res and logging are further exacerbating the imperil-
ment of these species and degradation of their habi-
tats (see GaBaDaGe et al., 2018; karunarathna et al.,
2016). Being rupicolous microhabitat specialists with
a scansorialmode of life, these species are suscepti-
ble to both localized and widespread threats. Although
saxicolous at the microhabitat scale, their presence is
notably limited to caves and rock outcrops in forests,
woodlands, and savannahs. As such, these habitat spe-
cialists can be vulnerable to adversities of edge effects.
Sri Lanka’s conservation polices should be reformed to
protect both smaller habitat patches and taxonomically
cryptic species that are overlooked in current legislation
(see amarasinGhe et al., 2015, 2016; karunarathna et
al., 2017a, b). Unlike some other reptile lineages, Cne-
maspis is genetically highly diverse and, as such, wild-
life policy reforms should include conservation within

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
294
species level, including conservation of genetically dis-
tinct populations and subclades (zieGler et al., 2016).
Further, our study highlights the importance of granite
caves and rock outcrops, which are shared with many
other unique species assemblies, as critical habitats for
this endemic reptile clade. Thus, we recommend protec-
tion of such cryptic habitats and their surrounding land-
scape as keystone ecosystems.
Moreover, a number of Cnemaspis species have re-
stricted ranges, limited dispersal capabilities, and niche
specialization; thus, most are point endemics (also see
aGarWal et al., 2017; BatuWita et al., 2019; Bauer et al.,
2007; De silva et al., 2019; karunarathna et al., 2019).
The overall ecoregional environmental gradients stem-
ming from habitat heterogeneity, geographic and geo-
logical complexity, and climatic variations have formed
biogeographic barriers leading to the isolation and sub-
sequent allopatric speciation of these diminutive species
(De silva et al., 2019; WiCkramasinGhe & somaWeera,
2002). These ndings suggest the possible speciation of
Cnemaspis in geographically isolated mountains with
granite caves, rock outcrops, and favorable environmen-
tal conditions, and the number of species is predicted
to increase to more than 50 (e.g., aGarWal et al. 2017;
BatuWita et al. 2019; De silva et al., 2019; karunar-
athna et al., 2019). As evidenced by their microende-
mism, this genus may have also underdone peripatric and
parapatric speciation as well. However, these hypotheses
need testing in future studies that integrate phylogenetic
afcinites with historical biogeography.
Acknowledgements
We thank Chandana Suriyabandara (Director General), Laxmen
Peiris (Deputy Director), research committee, and the eld staff of
the Department of Wildlife Conservation – DWC (WL/3/2/1/14/12,
and WL/3/2/42/18a, b) and Conservator General of Forest Depart-
ment – FD (FRC/5, and FRC/6) for granting permission and for
eld staff providing help during the eld surveys. Nanda Wick-
ramasinghe, Sanuja Kasthuriarachchi, Lankani Somaratne, Chan-
drika Munasinghe, Rasika Dasanayake, Ravindra Wickramanay-
ake, P. Gunasiri at National Museum of Sri Lanka (NMSL) assisted
while we were examining collections under their care. Nirmala
Perera, Angelo Hettige, Nimantha Aberathna, Nadeesh Gamage,
Mahesh De Silva, Hasantha Wijethunga, Udaya Chanaka, Gayan
Edirisinghe, Niranjan Karunarathna, Kawmini Karunarathna, D.M.
Kararunarathna, Rashmini Karunarathna, Thesanya Karunarathna,
Tharaka Kusuminda, Chamara Amarasinghe, Evgeniy S. Popov
(BIN RAS) and Alina V. Alexandrova (MSU) provided valu-
able assistance. This work was supported, in part, Nagao Natural
Environment Foundation (2018-20) grant to SK, by the Russian
Science Foundation grant No. 19-14-00050 to NAP, and United
States National Science Foundation grants DEB 1555968 and EF
1241885 (subaward 13-0632) to AMB. Finally, we would like to
thank Thomas Ziegler and Thasun Amarasinghe for constructive
criticism of an earlier draft, and anonymous reviewers for useful
comments that helped signicantly improve this paper.
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Appendix
Comparative materials examined:
Cnemaspis alwisi: NMSL 2004.9.1 (holotype), NMSL 2004.9.2
(paratype), NMSL 2004.9.3 (paratype), WHT 5918, WHT
6518, WHT 6519, WHT 7336, WHT 7337, WHT 7338, WHT
7343, WHT 7344, WHT 7345, WHT 7346.
C. amith: BMNH 63.3.19.1066A (holotype), BMNH 63.3.19.1066B
(paratype), BMNH 63.3.19.1066C (paratype).
C. gemunu: AMB 7495 (holotype), AMB 7507 (paratype), WHT
7221, WHT 7347, WHT 7348, NMSL 2006.11.01, NMSL
2006. 11.02, NMSL 2006.11.03, NMSL 2006.11.04.
C. godagedarai: NMSL 2019.09.01 (holotype), NMSL 2019.16.01
(paratype), NMSL 2019.16.02 (paratype).
C. ingerorum: WHT 7332 (holotype), WHT 7330 (paratype) WHT
7331 (paratype).
C. kallima: WHT 7245 (holotype), WHT 7222 (paratype), WHT
7227 (paratype), WHT 7228 (paratype), WHT 7229 (para-
type), WHT 7230(paratype), WHT 7239 (paratype), WHT
7249 (paratype), WHT 7251 (paratype), WHT 7252 (para-
type), WHT 7253 (paratype), WHT 7254 (paratype), WHT
7255 (paratype).
C. kandambyi: WHT 9466 (holotype), WHT 9467 (paratype).
C. kandiana: BMNH 53.4.1.1 (lectotype), BMNH 80.2.2.119A
(paralectotype), BMNH 80.2.2.119B (paralectotype), BMNH
80.2.2.119C (paralectotype), WHT 7212, WHT 7213, WHT
7267, WHT 7305, WHT 7307, WHT 7308, WHT 7310, WHT
7313, WHT 7319, WHT 7322.
C. kumarasinghei: NMSL 20061301 (holotype), NMSL 20061302
(paratype).
C. latha: WHT 7214 (holotype).
C. menikay: WHT 7219 (holotype), WHT 7218 (paratype), WHT
7349 (paratype).
C. molligodai: NMSL 2006.14.01 (holotype), NMSL 2006.14.02-5
(paratype), NMSL 2006.14.03 (paratype), NMSL 2006.14.04
(paratype), NMSL 2006.14.05 (paratype).
C. nilgala: NMSL 2018.07.01 (holotype), NMSL 2018.06.01 (pa-
ratype), NMSL 2018.06.02 (paratype), NMSL 2018.06.03 (pa-
ra type).
C. pava: WHT 7286 (holotype), WHT 7281 (paratype), WHT 7282
(paratype), WHT 7283 (paratype), WHT 7285 (paratype),
WHT 7288 (paratype), WHT 7289 (paratype), WHT 7290
(paratype), WHT 7291 (paratype), WHT 7292 (paratype),
WHT 7293 (paratype), WHT 7294 (paratype), WHT 7295
(paratype), WHT 7296 (paratype), WHT 7297 (paratype),
WHT 7298 (paratype), WHT 7299 (paratype), WHT 7300
(paratype), WHT 7301 (paratype), WHT 7302 (paratype).
C. phillipsi: WHT 7248 (holotype), WHT 7236 (paratype); WHT
7237 (paratype); WHT 7238 (paratype).
C. podihuna: BMNH 1946.8.1.20 (holotype), NMSL 20061002,
NMSL 20061003, NMSL 20061004.
C. pulchra: WHT 7023 (holotype), WHT 1573a (paratype), WHT
7011 (paratype), WHT 7021 (paratype), WHT 7022 (paratype).
C. punctata: WHT 7256 (holotype), WHT 7223 (paratype), WHT
7226 (paratype), WHT 7243 (paratype), WHT 7244 (paratype).
C. rajakarunai: NMSL 2016.07.01 (holotype), DWC 2016.05.01
(paratype), DWC 2016.05.02 (paratype).

K, S. et al.: Integrative taxonomy reveals six new species of Cnemaspis from Sri Lanka
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C. rammalensis: NMSL 2013.25.01 (holotype), DWC 2013.05.001.
C. retigalensis: NMSL 20061201 (holotype), NMSL 20061202 (pa-
ra type), NMSL 20061203 (paratype), NMSL 20061204 (pa ra-
type).
C. samanalensis: NMSL 2006.15.01 (holotype), NMSL 2006.15.02
(paratype), NMSL 2006.15.03 (paratype), NMSL 2006.15.04
(paratype), NMSL 2006.15.05 (paratype).
C. scalpensis: NMSL 2004.1.1 (neotype), NMSL 2004.2.1, NMSL
2004.3.1, NMSL 2004.4.1, WHT 7265, WHT 7268, WHT
7269, WHT 7274, WHT 7275, WHT 7276, WHT 7320.
C. silvula: WHT 7208 (holotype), WHT 7206 (paratype), WHT
7207 (paratype), WHT 7209 (paratype), WHT 7210 (paratype),
WHT 7216 (paratype), WHT 7217 (paratype), WHT 7018,
WHT 7027, WHT 7202, WHT 7203, WHT 7220, WHT 7354,
WHT 7333.
C. tropidogater: BMNH 71.12.14.49 (lectotype), NMSL 5152,
NMSL 5151, NMSL 5159, NMSL 5157, NMSL 5970, NMSL
5974.
C. upendrai: WHT 7189 (holotype), WHT 7184 (paratype), WHT
7187 (paratype), WHT 7188 (paratype), WHT 7181 (para-
type), WHT 7182 (paratype), WHT 7183 (paratype), WHT
7185 (paratype), WHT 7190 (paratype), WHT 7191 (para-
type), WHT 7192 (paratype), WHT 7193 (paratype), WHT
7194 (paratype), WHT 7195 (paratype), WHT 7196 (paratype),
WHT 7197 (paratype), WHT 7260 (paratype).
ZooBank Registration
at www.zoobank.org
Present article: The LSID for this publication is as follows:
urn:lsid:zoobank.org:pub:1AF1F94A-1091-4F8A-AACF-
B6A574A4D6EA.