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A new species of Cyrtodactylus from Tak Province, Thailand, Cyrtodactylus amphipetraeus sp. nov., is described using an integrative taxonomic analysis based on morphology, color pattern, and the mitochondrial gene NADH dehydrogenase subunit 2 (ND2). The phylogenetic analyses place the new species within the C. sinyineensis group which was previously thought to be endemic to the Salween Basin in southern Myanmar. The phylogeny also places C. inthanon in the C. sinyneensis group which is expanded herein to also include the group’s sister species C. doisuthep. Along with C. amphipetraeus sp. nov., these are the first three species of the C. sinyineensis group to be found outside of Myanmar east of the Tenasserim Mountains. The Tenasserim Mountain region is discussed as an area of cladogeneic turnover.
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ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Accepted by M. Heinicke: 29 Jul. 2020; published: 26 Aug. 2020 179
Zootaxa 4838 (2): 179–209
https://www.mapress.com/j/zt/
Copyright © 2020 Magnolia Press Article
https://doi.org/10.11646/zootaxa.4838.2.2
http://zoobank.org/urn:lsid:zoobank.org:pub:1C7589F7-36DA-41B0-B2AD-23BE22B9F300
A new species Cyrtodactylus Gray (Squamata: Gekkonidae) from western Thailand
and the phylogenetic placement of C. inthanon and C. doisuthep
SIRIWADEE CHOMDEJ1,7#, CHATMONGKON SUWANNAPOOM2,8#, PARINYA PAWANGKHANANT2,9,
WARANEE PRADIT1,10, ROMAN A. NAZAROV3,11, L. LEE GRISMER4* & NIKOLAY A. POYARKOV5,6*
1Research Center in Bioresources for Agriculture, Industry and Medicine, Department of Biology, Faculty of Science, Chiang Mai
University, 239 Huay Kaew Rd. Tambon Su Thep, Chiang Mai 50200, Thailand.
2Division of Fishery, School of Agriculture and Natural Resources, University of Phayao, Phayao, Thailand.
3Zoological Museum, Moscow State University, B. Nikitskaya ul. 2, Moscow 125009, Russia.
4Herpetology Laboratory, Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92515, USA.
lgrismer@lasierra.edu; https://orcid.org/0000-0001-8422-3698
5Faculty of Biology, Department of Vertebrate Zoology, Moscow State University, Moscow, Russia.
n.poyarkov@gmail.com; https://orcid.org/0000-0002-7576-2283
6Laboratory of Tropical Ecology, Joint Russian-Vietnamese Tropical Research and Technological Center, Hanoi, Vietnam
7
siriwadee@yahoo.com; https://orcid.org/0000-0002-2373-8847
8
chatmongkonup@gmail.com; https://orcid.org/0000-0002-3342-1464
9 https://orcid.org/0000-0002-0947-5729
10 https://orcid.org/0000-0002-2369-6405
11
r_nazarov@mail.ru; https://orcid.org/0000-0002-7827-6387
#Authors contributed equally to this work
*Corresponding authors
Abstract
A new species of Cyrtodactylus from Tak Province, Thailand, Cyrtodactylus amphipetraeus sp. nov., is described using
an integrative taxonomic analysis based on morphology, color pattern, and the mitochondrial gene NADH dehydrogenase
subunit 2 (ND2). The phylogenetic analyses place the new species within the C. sinyineensis group which was previously
thought to be endemic to the Salween Basin in southern Myanmar. The phylogeny also places C. inthanon in the C.
sinyneensis group which is expanded herein to also include the group’s sister species C. doisuthep. Along with C.
amphipetraeus sp. nov., these are the first three species of the C. sinyineensis group to be found outside of Myanmar east
of the Tenasserim Mountains. The Tenasserim Mountain region is discussed as an area of cladogeneic turnover.
Key words: Integrative taxonomy, Indochina, Tenasserim Mountains, Bent-toed gecko
Introduction
The rapidly growing number of newly described Indochinese species of the gekkonid genus Cyrtodactylus Gray is a
result of both integrative analyses revealing the existence of “cryptic species” and the discovery of new populations
based on field work (e.g., Grismer et al. 2018a; Murdoch et al. 2019; Nazarov et al. 2012, 2014, 2018; and refer-
ences therein). Accompanying these analyses is a wealth of genetic information enabling authors not only to more
accurately delimit newly discovered species, but to place them into the phylogenetic context of previously described
species. Recent field work in northwestern Thailand recovered a new species from the Tham Sri Fah Cave and the
nearby Tha Ra Rak waterfall, Mae Sot District, Tak Province (Fig. 1). Morphological and molecular data from the
mitochondrial gene NADH dehydrogenase subunit 2 (ND2) indicate that this new species is nested within the C.
sinyieensis group endemic to the Salween Basin in southern Myanmar (sensu Grismer et al. 2018a, 2020a,b). Addi-
tionally, we obtained ND2 data for C. inthanon Kunya, Sumontha, Panitvong, Dongkumfu, Sirisamphan & Pauwels
and C. doisuthep Kunya, Panmongkol, Pauwels, Sumontha, Meewasana, Bunkhwamdi & Dangsri and for the first
time are able to reconstruct their phylogenetic relationships. The newly discovered species from Tak Province is
described herein and we provide a discussion on the importance of phylogenetic information accompanying species
descriptions by using C. inthanon and C. doisuthep as an example.
CHOMDEJ ET AL.
180 · Zootaxa 4838 (2) © 2020 Magnolia Press
Materials and methods
Sampling. Specimens were collected in Mae Sot District, Tak Province, northwestern Thailand, and from Chiang
Mai Province, northern Thailand, by Siriwadee Chomdej, Chatmongkon Suwannapoom, Parinya Pawangkhanant,
and Nikolay A. Poyarkov during several field surveys in 2016 and 2019. The location of the surveyed localities and
the distribution of the C. sinyieensis species group members in Thailand are shown in Figure 1. Geographic coor-
dinates and elevation were obtained using a Garmin GPSMAP 60CSx and recorded in WGS 84 datum. Specimens
were fixed in 10% buffered formalin and later transferred to 70% ethanol. Liver tissues were sampled prior to speci-
men fixation and preserved in 95% ethanol. Specimens and tissues were subsequently deposited in the herpetologi-
cal collections of the School of Agriculture and Natural Resources, University of Phayao (AUP, Phayao, Thailand)
and of the Zoological Museum of Moscow University (ZMMU, Moscow, Russia).
FIGURE 1. Distribution of the species of the Cyrtodactylus sinyineensis group in Thailand and Myanmar.
Specimen collection protocols and animal use were approved by the Institutional Ethical Committee of Animal
Experimentation of the University of Phayao, Phayao, Thailand (certificate number UP-AE61-01-04-0022 issued
to Chatmongkon Suwannapoom) and were strictly complacent with the ethical conditions of the Thailand Animal
Welfare Act. Field work, including collection of animals in the field and specimen exportation, was authorized by
the Institute of Animals for Scientific Purpose Development (IAD), Bangkok, Thailand (permit number U1-04995-
2559, issued to Siriwadee Chomdej).
Species delimitation. The general lineage concept (GLC: de Queiroz 2007) adopted herein proposes that a
species constitutes a population of organisms evolving independently from other such populations owing to a lack
of gene flow. By “independently,” it is meant that new mutations arising in one species cannot spread readily into
another species (Barraclough et al. 2003 and de Queiroz 2007). Integrative studies on the nature and origins of
species are using an increasingly wider range of empirical data to delimit species boundaries (Coyne & Orr 1998;
Fontaneto et al. 2007; Knowles & Carstens 2007; Leaché et al. 2009), rather than relying solely on morphology and
traditional taxonomic methods. Under the GLC implemented herein, molecular phylogenies were used to recover
A NEW CYRTODACTYLUS FROM THAILAND Zootaxa 4838 (2) © 2020 Magnolia Press · 181
monophyletic mitochondrial lineages of individuals (populations) in order to develop initial species-level hypoth-
eses—the grouping stage of Hillis (2019). Discrete color pattern data and univariate and multivariate analyses of
morphological data were then used to search for characters and morphospatial patterns bearing statistically signifi-
cant differences that were consistent with the previous designations of the species-level hypotheses—the construc-
tion of boundaries representing the hypothesis-testing step of Hillis (2019)—thus providing independent diagnoses
to complement the molecular analyses.
Molecular data. The goal of this study was to investigate the taxonomy and phylogenetic relationships of
a newly discovered Thai population of Cyrtodactylus belonging to the C. sinyineensis group from the Salween
Basin and to recover the phylogenetic relationships of C. doisuthep and C. inthanon. To do so, a phylogeny was
constructed using 1534 bp of ND2 and its flanking tRNAs (WANCY region). All other species of the Indochinese
clade of Grismer et. al. (2020b) and C. tibetanus were used as outgroups to root the tree. New sequences not listed
in Grismer et al. (2020b) were deposited in GenBank (Table 1).
TABLE 1. GenBank accession numbers for the newly added specimens of the Cyrtodactylus sinyineensis group used for
the molecular phylogenetic analyses.
Taxon Catalog no. Locality GenBank no.
Cyrtodactylus doisuthep AUP-00774 Doi Suthep Mt., Chiang Mai Province, Thailand
(N 18.808786 E 98.926325)
MT550626
Cyrtodactylus inthanon AUP-00154 Maeyanoi waterfall, Doi Inthanon N.P., Chiang
Mai Province, Thailand (N 18.43968 E 98.59800)
MT550625
Cyrtodactylus amphipetraeus
sp. nov.
ZMMU R-16626 Tha Ra Rak waterfall, Mae Sot District, Tak Prov-
ince, Thailand (N 16.569339 E 98.694566)
MT550630
Cyrtodactylus amphipetraeus
sp. nov.
AUP-00690 Tham Sri Fah Cave, Mae Sot District, Tak Prov-
ince, Thailand (N 16.602162 E 98.712481)
MT550629
Cyrtodactylus amphipetraeus
sp. nov.
AUP-00691 Tham Sri Fah Cave, Mae Sot District, Tak Prov-
ince, Thailand (N 16.602162 E 98.712481)
MT550628
Cyrtodactylus amphipetraeus
sp. nov.
AUP-00698 Tha Ra Rak waterfall, Mae Sot District, Tak Prov-
ince, Thailand (N 16.569339 E 98.694566)
MT550627
For molecular phylogenetic analyses, total genomic DNA was extracted from ethanol-preserved liver tissue
using standard phenol-chloroform—proteinase K (final concentration 1 mg/ml) extraction procedures with con-
sequent isopropanol precipitation following Hillis et al. (1996). 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 Scientific), and consequently adjusted to ca. 100 ng DNA/μL.
The ND2 mitochondrial DNA gene was amplified using a double-stranded Polymerase Chain Reaction (PCR)
under the following conditions: 1.0 µl genomic DNA (~10–30 ng), 1.0 µl light strand primer (10 µM), 1.0 µl heavy
strand primer (10 µM), 1.0 µl dinucleotide pairs (1.0 µM), 2.0 µl 5x buffer (2.0 µM), 1.0 MgCl 10x buffer (1.0 µM),
0.10 µl Taq polymerase (5u/µl), and 7.4 µl H2O. PCR reactions were executed on a Bio-Rad T100™ Thermal Cycler
under the following conditions: initial denaturation at 95˚C for 2 min, followed by a second denaturation at 95˚C
for 35 s, annealing at 48˚C for 35 s, followed by a cycle extension at 72˚C for 35 s, for 31 cycles. All PCR products
were visualized on a 1.0 % agarose electrophoresis gel. Successful PCR products were sent to Evrogen® (Moscow,
Russia) for PCR purification, cycle sequencing, sequencing purification, and sequencing using the same primers
as in the amplification step (Table 2). Sequences were analyzed from both the 3’ and the 5’ ends separately to con-
firm congruence between reads. Forward and reverse sequences were uploaded and edited in GeneiousTM 2019.0.4
(https://www.geneious.com). Following sequence editing we aligned the protein-coding region and the flanking
tRNAs using the MAFTT v7.017 (Katoh & Kuma 2002) plugin under the default settings in GeneiousTM 2019.0.4
(https://www.geneious.com). Mesquite v3.04 (Maddison & Maddison 2015) was used to calculate the correct amino
acid reading frame and to confirm the lack of premature stop codons in the ND2 portion of the DNA fragment.
Phylogenetic analyses. Two model-based phylogenetic analyses were employed. A Maximum Likelihood
(ML) analysis was implemented in the IQ-TREE webserver (Nguyen et al. 2015; Trifinopoulos et al. 2016) pre-
ceded by the selection of a substitution model using the Bayesian Information Criterion (BIC) in ModelFinder
(Kalyaanamoorthy et al. 2017), which supported GTR+F+I+Γ4 as the best fit model of evolution for the tRNAs and
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182 · Zootaxa 4838 (2) © 2020 Magnolia Press
TIM3+F+I+Γ4, TN+F+I+Γ4, and TVM+F+I+Γ4 for ND2 codon positions 1–3, respectively. One-thousand boot-
strap pseudoreplicates via the ultrafast bootstrap (UFB; Hoang et al. 2018) approximation algorithm were employed
and nodes having ML UFB values of 95 and above were considered highly supported (Minh et al. 2013) and we
considered with nodes with values of 90–94 as well-supported.
TABLE 2. Primer sequences used in this study for amplification and sequencing the ND2 gene and the flanking tRNAs.
L4437b (Macey et al. 1997) External 5’-AAGCAGTTGGGCCCATACC-3
H5934 (Macey et al. 1997) External 5-AGRGTGCCAATGTCTTTGTGRIT-3
A Bayesian phylogenetic tree was estimated using BEAST version 2.4.6 (Drummond et al. 2012) implemented
in CIPRES (Cyberinfrastructure for Phylogenetic Research; Miller et al. 2010). The input file constructed in BEAU-
ti (Bayesian Evolutionary Analysis Utility) version 2.4.6 was run in BEAST version 2.4.6 (Drummond et al. 2012)
on CIPRES (Cyberinfrastructure for Phylogenetic Research; Miller et al. 2010), using a lognormal relaxed clock
with unlinked site models, linked trees and clock models, and a Yule prior. bModelTest, implemented in BEAST,
was used to numerically integrate over the uncertainty of substitution models while simultaneously estimating
phylogeny using Markov chain Monte Carlo (MCMC). MCMC chains were run for 50,000,000 million generations
and logged every 5000 generations. The BEAST log file was visualized in Tracer v. 1.6.0 (Rambaut et al. 2014) to
ensure effective sample sizes (ESS) were well above 200 for all parameters. A maximum clade credibility tree us-
ing mean heights at the nodes was generated using TreeAnnotator v.1.8.0 with a burnin of 1000 trees (10%). Nodes
with Bayesian posterior probabilities (BPP) of 0.95 and above were considered strongly supported (Huelsenbeck et
al. 2001; Wilcox et al. 2002) and we considered nodes with values of 0.90–0.94 as well-supported. After removing
outgroup taxa, MEGA7 (Kumar et al. 2016) was used to calculate uncorrected pairwise sequence divergence among
the ingroup species.
Morphological analysis. Color notes and digital images were taken from living specimens prior to preserva-
tion. Measurements were taken on the left side of the body when possible to the nearest 0.1 mm using Mitutoyo dial
calipers under a Nikon SMZ 1500 dissecting microscope. Measurements following Grismer et al. (2018a,b) were:
snout-vent length (SVL), taken from the tip of snout to the vent; tail length (TL), taken from the vent to the tip of
the tail, original or regenerated; tail width (TW), taken at the base of the tail immediately posterior to the postcloacal
swelling; forearm length (FL), taken on the dorsal surface from the posterior margin of the elbow while flexed 90°
to the inflection of the flexed wrist; tibia length (TBL), taken on the ventral surface from the posterior surface of
the knee while flexed 90° to the base of the heel; axilla to groin length (AG), taken from the posterior margin of the
forelimb at its insertion point on the body to the anterior margin of the hind limb at its insertion point on the body;
head length (HL), the distance from the posterior margin of the retroarticular process of the lower jaw to the tip of
the snout; head width (HW), measured at the angle of the jaws; head depth (HD), the maximum height of head mea-
sured from the occiput to the throat; eye diameter (ED), the greatest horizontal diameter of the eye-ball; eye-to-ear
distance (EE), measured from the anterior edge of the ear opening to the posterior edge of the eye-ball; eye to snout
distance (ES), measured from anteriormost margin of the eye-ball to the tip of snout; eye-to-nostril distance (EN),
measured from the anterior margin of the eye ball to the posterior margin of the external nares; inter orbital distance
(IO), measured between the anterior edges of the orbit across the frontal bone; ear diameter (EL), the greatest verti-
cal distance of the ear opening; and internarial distance (IN), measured between the nares across the rostrum.
Meristic characters taken were the numbers of supralabial scales (SL) counted from the largest scale immedi-
ately below the middle of the eyeball to the rostral scale and infralabial scales (IL), the large scales counted from
the mental scale to the commissure of the jaw. This count was modified from Grismer et al. (2018b) whose counts
terminated below the middle of the eyeball. These counts were re-taken for all members of the C. sinyineensis group
and used herein in the statistical analyses. Other meristic characters were the number of paravertebral tubercles (PV)
between limb insertions counted in a straight line immediately left of the vertebral column; the number of longitu-
dinal rows of body tubercles (LT) counted transversely across the center of the dorsum from between ventrolateral
folds; the number of longitudinal rows of ventral scales (VS) counted transversely across the center of the abdomen
between ventrolateral folds; the number of expanded subdigital lamellae proximal to the digital inflection on the
fourth toe (ET4) counted from the base of the first phalanx where it contacts the body of the foot to the largest scale
on the digital inflection (see Grismer et al. 2018a: Fig. 3), large continuous scales on the palmar and plantar surfaces
were not counted; the number of small, unmodified subdigital lamellae distal to the digital inflection to the claw
A NEW CYRTODACTYLUS FROM THAILAND Zootaxa 4838 (2) © 2020 Magnolia Press · 183
including the claw sheath on the fourth toe (UT4) counted from the digital inflection to the claw (see Grismer et al.
2018a: Fig. 3); and the total number of subdigital lamellae (TT4) beneath the fourth toe (i.e. ET4 + UT4 = TT4). The
total number of enlarged femoral scales (FS) from each thigh were combined as a single metric. The total number
of femoral pores (FP) in males (i.e., the sum of the number of enlarged pore-bearing femoral scales from each leg
combined as a single metric—not all enlarged femoral scales have pores). The number of enlarged precloacal scales
(PS); the number of precloacal pores in (PP) in males; and the number of rows of post-precloacal scales (PPS) on
the midline between the enlarged precloacal scales and the vent (see Grismer et al. 2018a:Fig. 4); number of dark
body bands (BB) between the occiput and the hind limb insertions not including the sacral or postsacral bands; the
number of light-colored caudal bands on an original tail; the number of dark caudal bands on an original tail; and if
a mature regenerated tail was spotted or not.
Non-meristic morphological characters evaluated were the degree of body tuberculation—weak tuberculation
referring to dorsal body tubercles that are relatively low, small, less densely packed, and weakly keeled whereas
prominent tuberculation refers to tubercles that are larger, higher (raised), and prominently keeled (see Grismer et
al. 2018a: Fig. 6); body tubercles extending past the postcloacal swelling or not (see Grismer et al. 2018a: Fig. 7);
and the relative length-to-width ratio of the transversely expanded, median subcaudal scales and whether or not they
extend onto the lateral surface of the tail (see Grismer et al. 2018a: Fig. 8).
Color pattern characters (see Grismer et al. 2018a: Fig. 5) evaluated were the nuchal loop being continuous from
eye to eye, separated medially into paravertebral halves, bearing an anterior azygous notch or not, and the posterior
border being straight (smooth), sinuous, v-shaped, jagged, or having two posteriorly directed projections; dorsal
body bands bearing paired, paravertebral elements or not; dark dorsal body bands wider than light interspaces, with
or without lightened centers, edged with light-colored tubercles or not, jagged or more regularly shaped (straight or
even-edged); dark markings present or absent in the dorsal interspaces; top of head bearing combinations of dark
diffuse mottling or dark, distinct blotches overlain with a light-colored reticulating network or not; light caudal
bands bearing dark markings or immaculate; light caudal bands encircle tail or not; dark caudal bands wider than
light caudal bands; and regenerated tail bearing a pattern of distinct, dark spots or not.
All statistical analyses were performed using the platform R v 3.2.1 (R Core Team 2018). Analyses of vari-
ance (ANOVA) were conducted on meristic characters with statistically similar variances (i.e. p values ≤ 0.05 in a
Levene’s test) to search for the presence of statistically significant mean differences (p < 0.05) among species across
the data set. Characters bearing statistical differences were subjected to a TukeyHSD test to ascertain which species
pairs differed significantly from each other for those particular characters. Boxplots were generated for discrete
meristic characters in order to visualize the range, mean, median, and degree of differences between pairs of species
bearing statistically different mean values.
Morphospatial positions were subsequently visualized using principal component analysis (PCA) from the
ADEGENET package in R (Jombart et al. 2010) to determine if their positioning was consistent with the putative
species boundaries delimited by the molecular phylogenetic analyses and defined by the univariate analyses (see
below). PCA, implemented using the “prcomp()” command in R, is an indiscriminate analysis plotting the overall
variation among individuals (i.e. data points) while treating each individual independently (i.e. not coercing data
points into pre-defined groups). Meristic data used were SL, IL, PV, LT, VS, ET4, UT4, TT4, FS, and PS. Femoral
and precloacal pore counts were excluded from the PCA due to their absence in females. Because the data were
skewed by values ranging from 6–35, all characters were scaled to their standard deviation prior to analysis in order
to normalize their distribution so as to ensure characters with very large and very low values did not over-leverage
the results owing to intervariable nonlinearity and to ensure the data were analyzed on the basis of correlation not
covariance. Subsequent to the PCA, a discriminant analysis of principle components (DAPC) was performed which
places the individuals of each predefined population inferred from the phylogeny into separate clusters (i.e., plots of
points) bearing the smallest within-group variance that produce linear combinations of centroids having the greatest
between-group variance (i.e. linear distance; Jombart et al. 2010). DAPC relies on standardized data from its own
PCA as a prior step to ensure that variables analyzed are not correlated and number fewer than the sample size.
DAPC principal components with eigenvalues accounting for 90–95% of the variation in the data set were retained
for the DAPC analysis according to the criterion of Jombart et al. (2010).
Museum abbreviations follow Sabaj (2016) except for LSUHC referring to the La Sierra University Herpe-
tological Collection, La Sierra University, Riverside, California, 92505, USA; AUP referring to the Agricultural
University of Phayao, Phayao Province, Thailand; and of the Zoological Museum of Moscow University (ZMMU,
Moscow, Russia).
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184 · Zootaxa 4838 (2) © 2020 Magnolia Press
Results
The ML and BEAST analyses recovered trees with identical topologies that corroborate the relationships of the
Cyrtodactylus sinyineensis group recovered by Grismer et al. (2020b). The C. sinyineensis group is composed of
two major sister clades (Fig. 2) plus one additional species (see below). Clade 1 contains C. inthanon which is the
sister species of the Burmese species C. welpyanensis Grismer, Wood, Thura, Zin, Quah, Murdoch, Grismer, Lin,
Kyaw & Lwin and these two species are most closely related to the Burmese sister species C. sinyineensis and C.
taungwineensis. Clade 2 is composed of six Burmese species that are endemic to the Salween Basin and the two new
Thai populations from Tak Province which are recovered as the sister species to a monophyletic group composed of
C. aequalis Bauer 2003, C. bayinnyiensis Grismer, Wood Jr, Thura, Quah, Murdoch, Grismer, Herr, Lin & Kyaw,
and C. dattkyaikensis Grismer, Wood Jr, Thura, Quah, Oaks & Lin.. Collectively, these species are most closely
related to C. naungkayaingensis Grismer, Wood Jr, Thura, Quah, Grismer, Herr, Lin & Kyaw. The sister species
C. chaunghanakwaensis Grismer, Wood Jr, Thura, Quah, Murdoch, Grismer, Herr, Lin & Kyaw and C. dammath-
etensis Grismer, Wood Jr, Thura, Zin, Quah, Murdoch, Grismer, Lin, Kyaw & Lwin compose the sister group to the
remaining species of clade 2. The analyses recovered C. doisuthep as the sister species to (clade 1 + clade 2) with
moderate maximum likelihood support but weak Bayesian support (0.62/90). Given its phylogenetic relationships
and geographic proximity to the species of clades 1 and 2, we here include the new lineage from Tak Province as
part of the C. sinyieensis group.
The PCA of clade 2 recovered the Tak Province population as morphologically distinct from all other species in
the clade (Fig. 3A). Although the Tak Province population overlaps with C. chaunghanakwaensis, C. naungkayain-
gensis, and C. dammethetensis along PC1 it plots separately from them. It is well separated from C. aequalis, C.
dattkyaikensis, and C. bayinnyiensis along PC1 and from C. bayinnyensis and C. dammathetensis along PC2. PCs
1–3 account for 71.2% of the variation in the data set (Table 3). PC1 accounts for 42.7% and loads most heavily
for toe lamellae and femoral scales; PC2 accounts for 17.2% and loads most heavily for longitudinal rows of body
tubercles; and PC3 accounts for 11.8% of the variation and loads most heavily for precloacal scales (Fig. 3C). The
first seven PCs were retained in the DAPC which accounted for 96.6% of the variation in the data set and recovered
the Tha Pha Pum population as separate from all populations except for C. naugkayaingensis along the first and
second linear discriminants (LD; Fig. 3B).
FIGURE 2. BEAST maximum clade credibility tree of the Cyrtodactylus sinyineensis group based on ND2 and its flanking
tRNAs.
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TABLE 3. Summary statistics and principal component analysis scores for the species of the Cyrtodactylus sinyineensis group. Abbreviations are listed in the Materials and
methods.
PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8 PC9 PC10
Standard deviation 2.06753 1.31119 1.08729 0.86649 0.84105 0.69452 0.60995 0.53498 0.47408 0.01961
Proportion of Variance 0.42747 0.17192 0.11822 0.07508 0.07074 0.04824 0.0372 0.02862 0.02248 4.00E-05
Cumulative Proportion 0.42747 0.59939 0.71761 0.79269 0.86343 0.91166 0.94887 0.97749 0.99996 1
Eigenvalue 4.27469 1.71922 1.18219 0.75081 0.70736 0.48235 0.37204 0.28620 0.22476 0.00038
SL 0.31773 0.32691 -0.27856 -0.35452 0.15735 -0.25256 0.44613 -0.37341 -0.40234 -0.00182
IL 0.32263 0.22057 -0.37948 -0.49445 -0.05815 0.04984 -0.21699 0.29344 0.56542 0.00383
PV 0.14158 0.46889 0.40486 0.16791 -0.60734 0.03499 0.30918 -0.14686 0.28416 0.00240
LT -0.24737 0.52520 -0.02820 0.17391 0.03586 -0.54899 -0.17713 0.50447 -0.20828 -0.00204
VS 0.29310 -0.36373 0.18578 -0.15357 -0.34973 -0.61301 -0.41368 -0.22590 -0.05838 -0.00539
ET4 -0.36795 -0.30528 0.07664 -0.23720 0.04555 -0.38510 0.53591 0.13346 0.34560 0.36668
UT4 -0.39348 0.11853 -0.11872 -0.35620 -0.41189 0.26094 -0.24682 -0.10454 -0.32229 0.52601
TT4 -0.44377 -0.06223 -0.04228 -0.36058 -0.26038 0.00283 0.08982 -0.00244 -0.05229 -0.76732
FS 0.37102 -0.28707 0.03767 -0.06651 -0.31127 0.15868 0.29216 0.63889 -0.39808 0.00339
PS 0.05490 0.15150 0.74566 -0.47982 0.37813 0.11726 -0.11932 0.10216 -0.07986 0.00362
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FIGURE 3. A. PCA of the species of clade 1 of the Cyrtodactylus sinyeensis group based on meristic characters. B. DAPC of
same. C. Histograms of the factor loadings of the characters contributing the most to the variation along PC1, PC2, and PC3.
The ANOVA and subsequent TukeyHSD tests indicated that the Tak Province population bore 1–5 statistically
significant mean character differences from all other species in clade 2 except for C. naungkayaingensis (Table 4,
Fig. 4). However, they are not sister species (Fig. 2) and differ by a 7.0% uncorrected pairwise sequence divergence.
Furthermore, the jagged dorsal bands of C. naungkayaingensis are not wider in the interspaces whereas the broken
to hour glass-shaped dorsal bands of lizards of the Tak Province population usually are wider and the lizards of the
Tak Province population have a maximum SVL of 93.4 mm and a green iris whereas C. naungkayaingensis has a
66.9 mm maximum SVL and a reddish iris. Based on the results of the phylogenetic, multivariate, and univariate
analyses and genetic distance, we consider the Tak Province population from Thailand to be a diagnosable indepen-
dent lineage on its own phylogenetic trajectory and as such hypothesize it to be a new species belonging to the C.
sinyineensis species group which is described below.
A NEW CYRTODACTYLUS FROM THAILAND Zootaxa 4838 (2) © 2020 Magnolia Press · 187
TABLE 4. Character list of species bearing statistically significant mean differences from Cyrtodactylus amphipetraeus
sp. nov. based on ANOVA and subsequent TukeyHSD tests showing the upper and lower ranges of the confidence inter-
vals, ANOVA F statistics, and the difference between mean values.
mean difference lower range upper range p adjusted ANOVA
Supralabials (SL) F = 14.76
aequalis 0.878787879 0.243217077 1.514358681 0.001433869
bayinnyiensis 1 0.057296156 1.942703844 0.030442842
Infralabilas (IL) F = 21.09
chaunghanakwaensis -0.958333333 -1.527299203 -0.389367463 5.04E-05
Paravertebral tubercles (PV) F = 37.97
aequalis 2.272727273 1.02675132 3.518703226 9.58E-06
bayinnyiensis 8.666666667 6.818585671 10.51474766 0
chaunghanakwaensis 2.125 0.829929728 3.420070272 8.35E-05
dammathetensis 3 0.791120722 5.208879278 0.00184912
Longitudinal rows of dorsal tuber-
cles (LT)
F = 15.2
aequalis -2.656565657 -4.233120848 -1.080010465 5.00E-05
chaunghanakwaensis -1.652777778 -3.291452871 -0.014102684 0.04669276
dammathetensis 3.555555556 0.760621925 6.350489186 0.004398933
Ventral scales (VS) F = 9.45
aequalis 3.787878788 2.17349512 5.402262456 1.32E-08
bayinnyiensis 2.733333333 0.33881539 5.127851277 0.015075513
chaunghanakwaensis 2.958333333 1.280339234 4.636327433 1.99E-05
dattkyaikensis 3 0.138003648 5.861996352 0.033811365
Expanded toe lamellae (ET4) F = 27.28
aequalis -1.666666667 -2.337807604 -0.99552573 2.15E-09
bayinnyiensis -2 -2.99546288 -1.00453712 9.82E-07
dammathetensis -1.666666667 -2.856472381 -0.476860952 0.001185945
dattkyaikensis -2.333333333 -3.523139048 -1.143527619 1.79E-06
Unexpanded Toe Lamellae (UT4) F = 24.44
aequalis -3.111111111 -4.079417364 -2.142804859 0
bayinnyiensis -1.977777778 -3.414008051 -0.541547505 0.001520244
chaunghanakwaensis -1.152777778 -2.159237525 -0.146318031 0.014557173
dattkyaikensis -1.777777778 -3.49440129 -0.061154266 0.037516985
Total toe lamellae (TTL) F = 63.07
aequalis -4.777777778 -5.773662986 -3.78189257 0
bayinnyiensis -3.977777778 -5.454914252 -2.500641303 5.19E-11
chaunghanakwaensis -1.277777778 -2.312903154 -0.242652401 0.006347698
dammathetensis -2.777777778 -4.543293552 -1.012262004 0.000180252
dattkyaikensis -4.111111111 -5.876626885 -2.345595337 1.67E-08
Femoral scales (FS) F = 24.40
aequalis 6.151515152 4.157159924 8.145870379 0
bayinnyiensis 4.866666667 1.908559823 7.824773511 7.94E-05
chaunghanakwaensis 3.708333333 1.635395921 5.781270745 1.46E-05
dattkyaikensis 5.666666667 2.131052737 9.202280596 0.000129186
Precloacal scales (PS) F = 168
bayinnyiensis 1.8 0.270475017 3.329524983 0.010922307
dammathetensis 2.333333333 0.505201315 4.161465352 0.004220516
CHOMDEJ ET AL.
188 · Zootaxa 4838 (2) © 2020 Magnolia Press
dattkyaikensis -16 -17.82813202 -14.17186798 0
FIGURE 4. Boxplot comparisons of discrete meristic characters among the species of clade 2 of the Cyrtodactylus sinyinensis
group. Light blue circle is the mean and the black horizontal bars is the median. Asterisks denote species bearing statistically
significant mean differences from C. amphipetraeus sp. nov.
Taxonomy
The Tak Province population can be placed in the Cyrtodactylus sinyineensis group (as revised and expanded here-
in) by having 9–10 supralabials; 7–9 infralabials; raised, moderately to strongly keeled, dorsal body tubercles that
extend beyond base of tail; 34–38 paravertebral tubercles; 17–20 longitudinal rows of body tubercles; 28–30 ventral
scales; seven expanded subdigital fourth toe lamellae; 11 or 12 unmodified fourth toe subdigital lamellae; 18 or 19
total subdigital lamellae; continuous enlarged femoral and precloacal scales; 27–34 similarly sized enlarged femoral
scales; 10–12 femoral and 7–9 precloacal pore-bearing scales in males that do not form a continuous row; 8–11
enlarged precloacal scales; three post-precloacal scale rows; transverse, median subcaudal scales usually twice as
wide as long, usually extending onto lateral surface of tail; no anterior, azygous notch in nuchal loop; approximately
4–7 variably shaped body bands; anterodorsal margins of thighs and brachia darkly pigmented; 10–12 light caudal
bands; 11–13 dark caudal bands; and maximum SVL 93.4 mm.
A NEW CYRTODACTYLUS FROM THAILAND Zootaxa 4838 (2) © 2020 Magnolia Press · 189
Cyrtodactylus amphipetraeus sp. nov.
Tak Bent-toed Gecko
(Figs. 5, 6)
Holotype. Adult male AUP-00696 was collected on August 3, 2019, at 21:00 hrs by S. Chomdej, C. Suwannapoom
and P. Pawangkhanant from limestone rocks at the entrance to the Tham Sri Fah Cave, Mae Sot District, Tak Prov-
ince, Thailand (16.602162°N, 98.712481°E WGS; 710 m in elevation).
Paratypes. Nine specimens, including the paratypes AUP-00688–90 (three adult males), and AUP-00691–93
and AUP-0097 (four adult females) bear the same collecting data as the holotype; AUP-00698, an adult male, was
collected on August 3, 2019, at 20:00 hrs by S. Chomdej, C. Suwannapoom and P. Pawangkhanant from granite
rocks near the Tha Ra Rak waterfall, Mae Sot District, Tak Province, Thailand (16.569339° N, 98.694566° E WGS;
610 m in elevation); ZMMU R-16626 (field No. NAP-06637) was collected on November 13, 2016, at 20:00 hrs
by N.A. Poyarkov and P. Pawangkhanant from granite rocks near the Tha Ra Rak waterfall, Mae Sot District, Tak
Province, Thailand (16.569339° N, 98.694566° E WGS; 610 m in elevation).
Diagnosis. Cyrtodactylus amphipetraeus sp. nov. differs from all species in the C. sinyineensis group by hav-
ing the combination of nine supralabials; seven infralabials; 34–38 paravertebral tubercles; 17–20 longitudinal rows
of dorsal tubercles; 28–30 ventral scales ventral scales; seven expanded subdigital lamellae on the fourth toe; 11
or 12 unmodified subdigital lamellae on the fourth toe; 18 or 19 total subdigital lamellae on the fourth toe; 27–34
enlarged femoral scales; a total of 10–12 pore-bearing femoral scales in males; 8–11 enlarged precloacal scales; 7–9
pore-bearing precloacal scales in males; three rows of enlarged post-precloacal scales; approximately 4–7 broken
to hour glass shaped dorsal body bands; 10–12 light-colored caudal bands (n=2); 11–13 dark-colored caudal bands
(n=2); raised and strongly keeled dorsal tubercles that extend beyond base of tail; enlarged femoral and precloacal
scales nearly the same size and continuous; pore-bearing femoral and precloacal scales not continuous; medial sub-
caudals two to three times wider than long and extending onto lateral side of tail; iris green; nuchal loop lacking an
anterior azygous notch, and bearing a jagged posterior border; dorsal bands bearing paravertebral elements, gener-
ally equal in width than interspaces, bearing lightened centers, edged with white tubercles; dark markings in dorsal
interspaces; light caudal bands in adults bearing dark-colored markings; light-colored caudal bands not encircling
tail; and mature regenerated tail not spotted (Table 5).
Description of holotype. Adult male SVL 81.3 mm (Fig. 5); head moderate in length (HL/SVL 0.30), wide
(HW/HL 0.69), flat (HD/HL 0.47), distinct from neck, triangular in dorsal profile; lores inflated, prefrontal region
concave, canthus rostralis rounded; snout elongate (ES/HL 0.39), rounded in dorsal profile, broad in lateral profile;
eye large (ED/HL 0.25); ear opening oval (EL/HL 0.10); eye to ear distance greater than diameter of eye; rostral
rectangular, partially divided dorsally, bordered posteriorly by supranasals and one internasal, laterally by first su-
pralabials; external nares bordered anteriorly by rostral, dorsally by supranasals, posteriorly by smaller postnasals,
and ventrally by first supralabials; 9(R,L) rectangular supralabials extending to below midpoint of eye; 7(R,L)
infralabials tapering posteriorly to commissure of jaw; scales of rostrum and lores slightly raised, larger than granu-
lar scales on top of head and occiput; scales on top of head and occiput intermixed with small tubercles; dorsal
superciliaries weakly pointed and directed posteriorly; mental triangular, bordered laterally by first infralabials and
posteriorly by large left and right trapezoidal postmentals which contact medially for 50% of their length posterior
to mental; one row of enlarged chinshields, outermost row bordering first five infralabials; gular and throat scales
granular, grading posteriorly into larger, subimbricate pectoral and ventral scales.
Body relatively short (AG/SVL 0.42) with well-defined ventrolateral folds; dorsal scales small, raised and in-
terspersed with large, raised, semi-regularly arranged, strongly keeled tubercles; tubercles extend from top of head
onto base of tail just beyond the postcloacal swelling; tubercles on nape smaller than those on body; 35 paravertebral
tubercles; approximately 18 longitudinal rows of dorsal tubercles; 28 flat, subimbricate, ventral scales larger than
dorsal scales; 11 enlarged precloacal scales; eight pore-bearing precloacal scales separated on the midline by one
poreless scale; three rows of large, post-precloacal scales; and no deep precloacal groove or depression.
Forelimbs moderate in stature, relatively short (FL/SVL 0.17); slightly raised, juxtaposed scales of forearm
larger than those on body, intermixed with large tubercles; palmar scales slightly raised, juxtaposed; digits well-
developed, relatively long, inflected at basal, interphalangeal joints; digits narrow distal to inflections; widened
proximal subdigital lamellae do extend onto palm; slight webbing at base of digit; claws well-developed, sheathed
by a dorsal and ventral scale at base; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.15),
covered dorsally by small, raised, juxtaposed scales intermixed with large pointed tubercles and bearing flat, slightly
CHOMDEJ ET AL.
190 · Zootaxa 4838 (2) © 2020 Magnolia Press
larger imbricate scales anteriorly; ventral femoral scales flat, imbricate, much larger than dorsals; one row of 13/14
(R,L) enlarged femoral scales and enlarged precloacal scales continuous; enlarged femoral scales nearly equal in
size; small, postfemoral scales form an abrupt union with larger, flat ventral scales on posteroventral margin of
thigh; six (R,L) pore-bearing femoral scales not continuous with pore-bearing precloacal scales; subtibial scales
flat, imbricate; plantar scales raised; digits relatively long, well-developed, inflected at basal, interphalangeal joints;
seven (R,L) transversely expanded subdigital lamellae on fourth toe proximal to joint inflection that do not extend
onto sole, 11 (R,L) unmodified subdigital lamellae distal to inflection; and claws well-developed, base of claw
sheathed by a dorsal and ventral scale.
Tail complete, original, gracile in proportions, 115.1 mm in length, 9.9 mm in width at base, tapering to a point,
TL/SVL (1.42); dorsal scales of tail flat, forming indistinct whorls; median row of transversely expanded subcaudal
scales three times as wide as long, extending onto lateral subcaudal region; three enlarged postcloacal tubercles at
base of tail on hemipenal swellings; and postcloacal scales large, flat.
Color pattern (Fig. 5, Fig. 6A). Dorsal ground color of head, body, limbs, and tail yellowish-brown; top of
head and rostrum nearly unicolor, bearing areas of slightly darker, diffuse irregularly shaped markings; nuchal loop
jagged posteriorly, divided medially, bearing two posterior projections; triangular occipital band composed of three
dark oval-shaped blotches; approximately five dark hour glass shaped body bands with paravertebral elements
bearing lightened centers and edged with whitish tubercles extend from the shoulder to the presacral region; lighter
colored interspaces between bands bear darker markings; whitish tubercles scattered on flanks; sacral and postsacral
bands continue onto the tail to form 13 black caudal bands that are wider than the 12 light-colored caudal bands;
light-colored caudal bands bear dark markings and do not encircle tail; limbs bear distinct, dark-colored irregularly
shaped markings; gular scales bearing only two or three black stipples; black stippling in throat, pectoral region, and
anterior portion of belly more dense; anterior one-half of subcaudal region light-colored, posterior one-half bearing
faint whitish mottling.
Variation. The dorsal color pattern in the type series is highly variable. The body bands are often so irregularly
shaped it is difficult to accurately assess how many there are. Paratypes AUP-00688, AUP-00692–93, and AUP-
00698 have well-defined generally oval-shaped paravertebral components to their dorsal bands that are separated
on the midline of the body (Fig. 6B,E). Whereas the banding pattern of the holotype and paratypes AUP-00689–91,
00697–98 are generally more irregularly shaped (Fig. 6). Paratypes AUP-00689, 00691–93, 00698 and ZMMU
R-16626 have partially regenerated tails (Fig. 6). Additional variation in meristic and mensural characters are pre-
sented in Table 6.
Distribution. Cyrtodactylus amphipetraeus sp. nov. is known only from the type locality at Tham Sri Fah
Cave, Mae Sot District, environs of Mae Sot, Tak Province, western Thailand; and Tha Ra Rak Waterfall located
approximately 4 km to the southwest from the cave (Fig. 1).
Etymology. The specific epithet amphipetraeus is a Latinized adjective in nominative singular, derived from
Greek amphi or ἀμφί (meaning of both kinds) and petra or πέτρα (for rock). The species name is given in reference
to the remarkable natural history of this species which inhabits both limestone and granite rocks. The recommended
vernacular name in English is Tak Bent-toed Gecko; in Thai is Tuk kai tak.
Natural history. The type locality—Tham Sri Fah Cave (or Blue Cave), is located in Mae Sot District of Tak
Province, and represents a karstic cave on the northern slope of a small limestone hill with a Buddhist temple at
the entrance (Fig. 7). The hill runs northeast to southwest reaching 730 m in elevation and is covered with sparse
secondary vegetation and is approximately 500 m in length and approximately 20 m in its widest central point.
Hill slopes are jagged, bearing numerous karstic alcoves, deep cracks and crevices, creating suitable microhabitat
for Cyrtodatcylus. This small limestone area is surrounded by agricultural landscapes. At the type locality several
specimens were seen at night after 20:00 hrs on the cave walls near the entrance and on rocks near cracks and holes
in karst. The species is quite weary and would rapidly retreat into crevices and holes when disturbed by light. Ap-
proximately 4 km to the southwest from the Tham Sri Fah Cave is the Tha Ra Rak waterfall, formed by ca. 3–5
m wide river flowing within highly disturbed secondary evergreen forest. The valley of this river is covered with
large granite boulders, where the new species was observed at night sitting on granite rocks or hiding among them.
Only few specimens were spotted and just two of them collected. Remarkably, specimens recorded on granite rocks
had distinctive dorsal pattern forming a more blotched (Fig. 6B,E) rather than banded (Fig. 6A) appearance. Cyrto-
dactylus amphipetraeus sp. nov. is expected to occur in several nearby limestone hills within the Mae Sot District.
Reproductive biology, diet and predators of the new species remain unknown.
A NEW CYRTODACTYLUS FROM THAILAND Zootaxa 4838 (2) © 2020 Magnolia Press · 191
TABLE 5. Summary statistics and diagnostic characters of the species of the Cyrtodactylus sinyineensis species group. SD = standard deviation, n = sample size, and / indicates
data unavailable.
clade2 clade 1
amphipetraeus sp. nov.
aequalis
bayinnyiensis
chaunghanakwaensis
dammathetensis
dattkyaikensis
naungkayaingensis
inthanon
sinyineensis
taungwineensis
welpyanensis
doisuthep
supralabial scales (SL)
mean (±SD) 9.0 (0.00) 8.1 (0.46) 8.0 (0.84) 9.4 (0.71) 9.0 (0.00) 8.3 (0.58) 8.3 (0.58) 11.0
(1.00)
9.7 (0.57) 9.1
(7–10)
8.3
(0.6)
11.0 (1.0)
Range 9 7–9 7–9 8–11 9 8 or 9 8 or 9 10–12 9 or 10 7–10 8 or 9 10–12
n 10 37 5 24 3 3 3 3 3 14 3 3
infralabial scales (IL)
Mean (±SD) 7.0 (0.00) 6.8 (0.53) 6.2 (0.44) 7.9 (0.50) 7.7 (0.58) 6.3 (0.57) 7.0 (0.00) 9.3 (0.58) 8 7.00
(0.39)
7.0
(0.00)
9.5 (1.5)
Range 7 6–8 6 or 7 7–9 7 or 8 6 or 7 7 9 or 10 8 6–8 7 8–11
n 10 37 5 24 3 3 3 3 3 14 3 3
paravertebral tubercles (PV)
Mean (±SD) 35.7
(0.50)
32.0
(1.64)
26.0
(0.71)
32.5
(0.98)
31.7
(1.15)
34. 0
(0.00)
34.3
(0.58)
/ 33.7 30.3
(0.84)
31.7
(1.5)
/
Range 34–38 29–35 25–27 31–36 31–33 33–35 34 or 35 / 33–35 29–32 30–33 /
n 10 37 5 24 3 3 3 / 3 14 3 /
longitudunal rows of body
tubercles (LT)
Mean (±SD) 17.9
(0.78)
20.6
(1.42)
17.4
(01.14)
19.5
(1.47)
14.3
(1.15)
19.0
(1.00)
17.3
(1.54)
19. 0
(1.00)
15 18.7
(0.61)
16.0
(0.00)
19.7 (0.58)
Range 17–20 18–23 16–19 17–22 13–15 18–20 16–18 18–20 15 18–20 16 19 or 20
n 10 37 5 24 3 3 3 3 3 14 3 3
ventral scales (VS)
......continued on the next page
CHOMDEJ ET AL.
192 · Zootaxa 4838 (2) © 2020 Magnolia Press
TABLE 5. (Continued)
clade2 clade 1
amphipetraeus sp. nov.
aequalis
bayinnyiensis
chaunghanakwaensis
dammathetensis
dattkyaikensis
naungkayaingensis
inthanon
sinyineensis
taungwineensis
welpyanensis
doisuthep
Mean (±SD) 28.3
(0.71)
24.5
(1.68)
25.6
(1.52)
25.4
(1.10)
26.7
(1.53)
25.3
(0.58)
27.0
(0.00)
30.7
(2.89)
28 (1.0) 32.3
(2.02)
29.3
(1.20)
32.0 (3.00)
Range 28–30 22–31 24–28 23–27 25–28 25 or 26 27 29–34 27–29 30–36 28–30 29–35
n 10 37 5 24 3 3 3 3 3 14 3 3
expanded 4th toe lamellae
(ET4)
Mean (±SD) 7.0 (0.00) 8.5 (0.77) 9.0 (0.00) 7.1 (0.61) 8.7 (0.58) 9.3 (0.58) 7. 0
(0.00)
/ 9 7.4 (0.50) 8.0
(0.00)
/
Range 7 7–10 9 6–9 8 or 9 9 or 10 7 / 8–10 7 or 8 8 /
n 10 37 5 24 3 3 3 / 3 14 3 /
unmodified 4th toe lamellae
(UT4)
Mean (±SD) 11.2
(0.44)
14.4
(1.01)
13.3
(0.50)
12.4
(0.92)
12.3
(0.58)
13.0
(0.00)
11.3
(0.58)
/ 11.3
(0.58)
12.1
(0.62)
12.0
(1.0)
/
Range 11 or 12 13–17 13 or 14 11–14 12 or 13 13 11 or 12 / 11 or 12 11–13 11–13 /
n 10 37 5 24 3 3 3 / 3 14 3 /
total 4th toe lamellae (TT4)
Mean (±SD) 18.2
(0.44)
22.9
(0.94)
22.3
(0.50)
19.5
(1.06)
21.0
(0.00)
23.3
(0.58)
18.3
(0.58)
/ 20.3 (1.6) 19.4
(0.85)
20.0
(1.0)
/
Range 18 or 19 21–25 22 or 23 18–21 21 22 or 23 18 or 19 / 19–21 18–21 19–21 /
n 10 37 5 24 3 3 3 / 3 14 3 /
enlarged femoral scales (FS)
......continued on the next page
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TABLE 5. (Continued)
clade2 clade 1
amphipetraeus sp. nov.
aequalis
bayinnyiensis
chaunghanakwaensis
dammathetensis
dattkyaikensis
naungkayaingensis
inthanon
sinyineensis
taungwineensis
welpyanensis
doisuthep
Mean (±SD) 31.2
(2.06)
25.4
(1.85)
26.8
(1.79)
28.0
(1.06)
33.0
(2.65)
26.0 (1.7) 31.7
(2.89)
/ 26.3
(0.58)
24.1
(3.83)
30.3
(0.58)
/
Range 28–34 22–30 24–28 27–32 31–36 25–28 30–35 / 26 or 27 18–30 30–31 /
n 10 37 5 24 3 3 3 / 3 14 3 /
femoral pores (FP)
Mean (±SD) 11.4
(0.89)
14.4
(2.94)
15.0
(0.00)
28.6
(1.51)
36.0
(0.00)
25.0
(0.00)
11.0
(0.00)
5.0 (1.00) 16 16.3
(3.15)
20.0
(0.00)
5.0 (0.00)
Range 10–12 10–19 15 27–32 36 25 11 4–6 16 13–22 20 0
n 571912231721
enlarged precloacal scales (PS)
Mean (±SD) 10.0
(1.22)
9.1 (0.77) 8.2 (0.45) 10.2
(0.70)
8.3 (0.58) 8.3 (0.58) 9.7
(0.2.08)
/ 11(1.0) 8.6 (0.84) 12.0
(1.0)
/
Range 8–11 7–10 8 or 9 9–12 8 or 9 8 or 9 8–12 / 10–12 7–10 11–13 /
n 10 37 5 24 3 3 3 / 3 14 3 /
precloacal pores (PP) 5 (0.00) 6 (0.00)
Mean (±SD) 8.0 (1.0) 7.4 (2.07) 9.0 (0.00) 9.9 (0.60) 9.0 (0.00) 7.0 (1.41) 3.5 (0.5) 5.0 5 6.6 (0.79) 7.5
(0.5)
6.0
Range 7–9 5–10 9 9–11 9 6–8 3 or 4 5 5 6–8 7 or 8 6
n 571912211721
post-precloacal scale rows
(PPS)
......continued on the next page
CHOMDEJ ET AL.
194 · Zootaxa 4838 (2) © 2020 Magnolia Press
TABLE 5. (Continued)
clade2 clade 1
amphipetraeus sp. nov.
aequalis
bayinnyiensis
chaunghanakwaensis
dammathetensis
dattkyaikensis
naungkayaingensis
inthanon
sinyineensis
taungwineensis
welpyanensis
doisuthep
Mean (±SD) 3.0 (0.00) 3.0 (0.00) 2.4 (0.54) 3.0 0.21) 3.0 (0.00) 3.0 (0.00) 3. 0
(0.00)
3.0 (0.00) 3 3.0 (0.00) 3.0
(0.00)
3.0 (0.00)
Range 3 3 2 or 3 2 or 3 3 3 3 3 3 3 3 3
n 10 37 5 24 3 3 3 3 3 3 3 3
body bands (BB)
Mean (±SD) 5.5 5.3 (1.00) 5.2 (0.83) 5.92
(0.41)
5.0 (0.00) 6.0 (0.00) 4.0 (0.00) 6.0 (0.00) 5.5 (0.71) 4.1 (0.26) 6.0
(0.00)
8.7 (1.53)
Range 4–7
(0.94)
5–7 4–6 5–7 5 6 4 6 5 or 6 4 or 5 6 7–10
n 10 12 5 24 3 3 3 3 2 14 3 3
light caudal bands (LCB)
Mean (±SD) 11.0
(1.00)
9.6 (0.52) 9.3 (2.08) 12.3
(1.37)
10.3
(0.71)
8.0 (1.41) 11.0
(0.00)
12.0
(0.00)
9 12.1
(0.60)
9.0
(0.00)
12.0 (0.00)
Range 10–12 9 to 10 7–11 10–14 10 or 11 7–9 11 12 9 11–13 9 12
n 283632121911
dark caudal bands (DCB)
Mean (±SD) 12.0 (1.0) 9.5 (0.76) 10.3
(2.08)
12.7
(1.37)
10.7
(0.71)
8.5 (0.71) 10.0
(0.00)
11.0
(0.00)
9 12.0
(0.71)
10.0
(0.00)
11.0 (0.00)
Range 11–13 9–11 8–12 11–15 10 or 11 8 or 9 10 11 9 11–13 10 11
n 283632121911
Morphology
......continued on the next page
A NEW CYRTODACTYLUS FROM THAILAND Zootaxa 4838 (2) © 2020 Magnolia Press · 195
TABLE 5. (Continued)
clade2 clade 1
amphipetraeus sp. nov.
aequalis
bayinnyiensis
chaunghanakwaensis
dammathetensis
dattkyaikensis
naungkayaingensis
inthanon
sinyineensis
taungwineensis
welpyanensis
doisuthep
Body tubercles low, weakly
keeled
no no yes/no yes no no no no no no no no
body tubercles raised, moder-
ately to strongly keeked
yes yes yes/no no yes yes yes yes yes yes yes yes
tubercles extend beyond base
of tail
yes yes yes yes yes yes yes yes yes yes no yes
enlarged femoral and precloa-
cal scales continuous
yes yes yes yes yes yes yes yes yes variable yes yes
pore-bearing femoral and pre-
cloacal scales continuous
no no no yes yes no no yes no no no yes
enlarged proximal femoral
scales ~1/2 size of distal
femorals
no no no no yes no no no no no no no
medial subcaudals 2 or 3 times
wider than long
yes yes yes yes yes yes yes yes yes yes yes yes
medial subcaudals extend onto
lateral surface of tail
no no yes yes yes no yes no no no no no
Color Pattern
color of iris green reddish reddish green green reddish reddish green reddish reddish green reddish
nuchal loop divided medially variable variable no no variable yes no no no no no no
nuchal loop with anterior
azygous notch
no no no no no no no no no no no no
......continued on the next page
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196 · Zootaxa 4838 (2) © 2020 Magnolia Press
TABLE 5. (Continued)
clade2 clade 1
amphipetraeus sp. nov.
aequalis
bayinnyiensis
chaunghanakwaensis
dammathetensis
dattkyaikensis
naungkayaingensis
inthanon
sinyineensis
taungwineensis
welpyanensis
doisuthep
posterior border of nuchal loop jagged smooth jagged jagged jagged protract-
ed
jagged jagged jagged protract-
ed
jagged jagged
band on nape variable yes yes yes yes no variable yes yes no yes yes
dorsal banding with paraverte-
brtal elements
yes yes variable yes yes no yes no no no no no
dorsal bands wider than inter-
spaces
usually equal to
wider
yes/no yes yes yes no yes yes yes yes yes
dorsal bands bearing lightened
centers
yes no yes/no variable no yes yes yes weak yes no anterior
edge
dorsal bands edged with light-
colored tubercles
yes partly no variable no partly no yes yes partly no posteriorly
shape of dorsal bands broken
to hour
glass
highly
variable
jagged jagged jagged zig-zag jagged zig-zag jagged zig-zag jagged straight
dark markings in dorsal inter-
spaces in adults
yes yes yes yes yes yes yes yes yes yes yes yes
ventrolateral body fold whitish no no no no faintly no no no no no yes no
top of head diffusely mottled,
blotched, or patternless
variable mottled mottled
(adult)
mottled mottled mottled mottled mottled mottled mottled mottled no
light reticulum on top of head no no no no no no no no no no no yes
anterodorsal margin of thighs
darkly pigmented
yes yes yes yes yes yes yes yes yes yes yes yes
......continued on the next page
A NEW CYRTODACTYLUS FROM THAILAND Zootaxa 4838 (2) © 2020 Magnolia Press · 197
TABLE 5. (Continued)
clade2 clade 1
amphipetraeus sp. nov.
aequalis
bayinnyiensis
chaunghanakwaensis
dammathetensis
dattkyaikensis
naungkayaingensis
inthanon
sinyineensis
taungwineensis
welpyanensis
doisuthep
anterodorsal margin of brachia
darkly pigmented
yes yes yes yes yes yes yes yes yes yes yes yes
light caudal bands bearing dark
markings in adults
yes no no yes no yes yes yes yes yes yes yes
light caudal bands encircle tail no variable no no no no no no no no no no
dark caudal bands wider than
light caudal bands
yes yes yes yes yes yes yes yes yes yes yes yes
mature regenerated tail spotted no no yes no / no no no no no no no
maximum SVL (mm) 93.4 87.0 84.1 76.3 69.3 83.0 66.9 87.3 91.6 82.0 70.6 90.5
CHOMDEJ ET AL.
198 · Zootaxa 4838 (2) © 2020 Magnolia Press
FIGURE 5. Holotype of adult male Cyrtodactylus amphipetraeus sp. nov. (AUP-00696). A. Dorsal view of body showing
nuchal loop, dorsal band, and caudal patterns (in preservative). B. Lateral view of head showing greenish coloration of iris. C.
Gular region showing mental, postmental, and chin scales arrangement. D. Dorsal view of top of head. E. Precloacal and femoral
region showing scale sizes and pore arrangements, and plantar view of feet showing subdigital lamellae morphology. Photos by
C. Suwannapoom.
Comparisons. Cyrtodactylus amphipetraeus sp. nov. (n=10) differs from various combinations of all other
species except C. naungkayaingensis in clade 2 of the C. sinyineensis group in having statistically different mean
values across 1–5 of the nine scale characters. These differences are summarized in Tables 4 and 5 and visualized
A NEW CYRTODACTYLUS FROM THAILAND Zootaxa 4838 (2) © 2020 Magnolia Press · 199
TABLE 6. Meristic, mensural, and color pattern data from the type series of Cyrtodactylus amphipetraeus sp. nov. R = right, L = left, and / = data unobtainable or not applica-
ble.
Character AUP-00688
paratype
AUP-00689
paratype
AUP-00690
paratype
AUP-00691
paratype
AUP-00692
paratype
Sex M M M F F
Supralabials 9 9 9 9 9
Infralabials 7 7 7 7 7
Body tubercles low, weakly keeled no no no no no
Body tubercles raised, moderately to strongly keeled yes yes yes yes yes
Paravertebral tubercles 34 35 35 35 35
Longitudinal rows of body tubercles 17 18 19 19 18
Tubercles extend beyond base of tail yes yes yes yes yes
Ventral scales 28 28 30 28 29
Expanded subdigital lamellae on 4th toe 7 7 7 7 7
Unmodified subdigital lamellae on 4th toe 11 11 12 12 11
Total subdigital lamellae on 4th toe 18 18 19 19 18
Enlarged femoral scales (R/L) R16L15 R17L16 R15L14 17RL 15RL
Total femoral scales 31 33 29 34 30
Femoral pores (R/L) 5/5 6/6 6/6 / /
Total femoral pores in males 10 12 12 / /
Enlarged precolacal scales 10 10 11 10 8
Precloacal pores 9 7 9 / /
Post-precloacal scale rows 3 3 3 3 3
Enlarged femoral and precloacal scales continuous yes yes yes yes yes
Pore-bearing femoral and precloacal scales continuous no no no / /
Enlarged proximal femoral scales ~1/2 size of distal femorals no no no no no
Medial subcaudals 2 or 3 times wider than long yes yes yes yes yes
Medial subcaudals extend onto lateral surface of tail yes yes yes yes yes
Nuchal loop divided medially partly partly no yes no
2 posterior projections from nuchal loop yes yes yes yes yes
Nuchal loop with anterior azygous notch no no no no no
......continued on the next page
CHOMDEJ ET AL.
200 · Zootaxa 4838 (2) © 2020 Magnolia Press
TABLE 6. (Continued)
Character AUP-00688
paratype
AUP-00689
paratype
AUP-00690
paratype
AUP-00691
paratype
AUP-00692
paratype
Triangular marking anterior to nuchal loop no no no no no
Posterior border of nuchal loop jagged jagged jagged jagged jagged
Band on nape yes yes no yes yes
Dorsal banding with paravertebral elements yes yes yes yes yes
Number of body bands ~6 ~6 4 ~7 ~6
Dorsal body bands wider than interspaces equal equal equal yes yes
Dorsal body bands with lightened centers yes yes yes yes yes
Dorsal bands edged with white tubercles yes yes yes yes yes
Shape of dorsal bands hour glass hour glass hour glass hour glass broken
Dark markings in dorsal interspaces yes yes yes yes yes
Ventrolateral fold whitish no no no no no
Top of head diffusely mottled, blotched, or patternless blotched blotched diffusely mottled diffusely mottled diffusely mottled
Light-colored reticulum on top of head no no no no no
Anterodorsal margin of thighs darkly pigmented yes yes yes yes yes
Anterodorsal margin of brachia darkly pigmented yes yes yes yes yes
White caudal bands with dark markings yes / yes yes yes
White caudal bands encircle tail no / no no no
Number of light caudal bands / / / / /
Number of dark caudal bands / / / / /
Dark caudal bands wider than light caudal bands yes / yes yes yes
Mature regenerated tail spotted no no no no no
SVL 75 84.7 93.4 89.7 87.3
TL 95.2 98.2 124.7 91.8 106.65
TW 9.4 10.7 11.1 9.8 9.6
FL 12.9 15.2 15.6 15.5 12.3
TBL 13.1 13.3 14.7 13.5 13.0
AG 27.3 34.9 40 34.3 37
......continued on the next page
A NEW CYRTODACTYLUS FROM THAILAND Zootaxa 4838 (2) © 2020 Magnolia Press · 201
TABLE 6. (Continued)
Character AUP-00688
paratype
AUP-00689
paratype
AUP-00690
paratype
AUP-00691
paratype
AUP-00692
paratype
HL 21.8 24.9 27 25.7 24.5
HW 15 17.2 19.1 18.5 17.8
HD 9.7 11.1 11.9 11.9 10.7
ED 5.3 6.2 6.1 6.4 6.1
EE 6.4 7.8 7.9 7.3 7.2
ES 9.1 10.1 10.9 10 9.6
EN 6.9 8 9.2 8 8.4
IO 6.8 7.9 7.9 7.8 7.4
EL 1.9 2.5 2.1 2.2 1.7
IN 2.6 3.4 3.3 3.3 3
......continued on the next page
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202 · Zootaxa 4838 (2) © 2020 Magnolia Press
TABLE 6. (Continued)
Character AUP-00693
paratype
AUP-00696
holotype
AUP-00697
paratype
AUP-00698
paratype
ZMMU
R-16626
Sex F M F M F
Supralabials 9 9 9 9 10
Infralabials 7 7 7 7 9
Body tubercles low, weakly keeled no no no no no
Body tubercles raised, moderately to strongly keeled yes yes yes yes yes
Paravertebral tubercles 34 35 34 35 38
Longitudinal rows of body tubercles 17 18 17 18 20
Tubercles extend beyond base of tail yes yes yes yes yes
Ventral scales 28 28 28 28 29
Expanded subdigital lamellae on 4th toe 7 7 7 7 7
Unmodified subdigital lamellae on 4th toe 11 11 11 11 11
Total subdigital lamellae on 4th toe 18 18 18 18 18
Enlarged femoral scales (R/L) 15RL 13R14L 15RL 17RL 14RL
Total femoral scales 30 34 30 34 28
Femoral pores (R/L) / 6/6 / 6/5 /
Total femoral pores in males / 12 / 11 /
Enlarged precolacal scales 8 11 11 11 8
Precloacal pores / 8 / 7 /
Post-precloacal scale rows 3 3 3 3 3
Enlarged femoral and precloacal scales continuous yes yes yes yes yes
Pore-bearing femoral and precloacal scales continuous / no / no /
Enlarged proximal femoral scales ~1/2 size of distal femorals no no no no no
Medial subcaudals 2 or 3 times wider than long yes yes yes yes reg.tail.
......continued on the next page
A NEW CYRTODACTYLUS FROM THAILAND Zootaxa 4838 (2) © 2020 Magnolia Press · 203
TABLE 6. (Continued)
Character AUP-00693
paratype
AUP-00696
holotype
AUP-00697
paratype
AUP-00698
paratype
ZMMU
R-16626
Medial subcaudals extend onto lateral surface of tail yes yes yes yes reg.tail.
Nuchal loop divided medially partly yes yes no partly
2 posterior projections from nuchal loop yes yes yes yes yes
Nuchal loop with anterior azygous notch no no no no no
Triangular marking anterior to nuchal loop no no no no no
Posterior border of nuchal loop jagged jagged jagged jagged jagged
Band on nape yes yes yes yes no
Dorsal banding with paravertebral elements yes yes yes yes yes
Number of body bands ~6 ~5 ~5 ~4.5 ~7
Dorsal body bands wider than interspaces equal equal no equal equal
Dorsal body bands with lightened centers yes yes yes yes yes
Dorsal bands edged with white tubercles yes yes yes yes yes
Shape of dorsal bands broken hour glass broken broken irregular
Dark markings in dorsal interspaces yes yes yes yes yes
Ventrolateral fold whitish no no no no no
Top of head diffusely mottled, blotched, or patternless nearly patternless nearly patternless nearly patternless nearly patternless diffusely mottled
Light-colored reticulum on top of head no no no no no
Anterodorsal margin of thighs darkly pigmented yes yes yes yes yes
Anterodorsal margin of brachia darkly pigmented yes yes yes yes yes
White caudal bands with dark markings yes yes yes / /
White caudal bands encircle tail no no no / /
Number of light caudal bands / 12 10 /
Number of dark caudal bands / 13 11 / /
......continued on the next page
CHOMDEJ ET AL.
204 · Zootaxa 4838 (2) © 2020 Magnolia Press
TABLE 6. (Continued)
Character AUP-00693
paratype
AUP-00696
holotype
AUP-00697
paratype
AUP-00698
paratype
ZMMU
R-16626
Dark caudal bands wider than light caudal bands yes yes yes / /
Mature regenerated tail spotted no / / no no
SVL 72.2 81.3 74.9 82.8 89
TL 96.7 115.1 94.4 95.8 74 reg
TW 8.3 9.9 7.8 10.6 7.3
FL 12.2 13.6 13.2 15.3 13.9
TBL 11.1 12.6 11.4 13.1 11.4
AG 28.3 34.3 32.9 33.3 38.6
HL 21.5 24.2 22 25.9 23.8
HW 14.6 16.7 14.3 18.7 16.1
HD 8.9 11.3 8.7 11.1 9.75
ED 5 6 5.2 6.1 6.1
EE 5.7 7 6.3 7.8 7.2
ES 8.5 9.4 8.8 10.6 9.8
EN 7.1 7 7 8.4 7.6
IO 6.7 6.1 5.6 8 7.8
EL 1.1 2.3 2 1.9 2.2
IN 2.6 2.9 2.8 3.6 3.0
A NEW CYRTODACTYLUS FROM THAILAND Zootaxa 4838 (2) © 2020 Magnolia Press · 205
in Figure 4 and do not need to be written out here. Cyrtodactylus amphipetraeus sp. nov. is not the sister species of
C. naungkayaingensis (Fig. 2) and differs from it by a 7.0% uncorrected pairwise sequence divergence. As noted
above, the jagged dorsal bands of C. naungkayaingensis are not wider than the interspaces whereas the broken to
hour glass-shaped dorsal bands of C. amphipetraeus sp. nov. are wider than the interspaces and it has a maximum
SVL of 93.4 mm and a green iris whereas C. naungkayaingensis has a maximum SVL 66.9 mm and a reddish iris.
Additional differences between C. amphipetraeus sp. nov., C. doisuthep, and the species of clade 1 are listed in
Table 5.
FIGURE 6. Variation in dorsal pattern in Cyrtodactylus amphiptraeus sp. nov. in life. A. Adult holotype male (AUP-00696)
from Tham Sri Fah Cave, Pop Pra District, Tak Province, Thailand. B. Adult paratype male (AUP-00698) from Tha Ra Rak
waterfall, Pop Pra District, Tak Province, Thailand. C. General view of the adult paratype female (ZMMU R-16626) from Tha
Ra Rak waterfall, Pop Pra District, Tak Province, Thailand. D. Close-up of the head of the same specimen showing green iris.
E. Adult male (not collected) hiding in crevice near Tha Ra Rak waterfall, Pop Pra District, Tak Province, Thailand. F. Juvenile
specimen (not collected) from Tha Ra Rak waterfall, Pop Pra District, Tak Province, Thailand. Photos by C. Suwannapoom
(A–B), N.A. Poyarkov (C–D), and P. Pawangkhanant (E–F).
CHOMDEJ ET AL.
206 · Zootaxa 4838 (2) © 2020 Magnolia Press
FIGURE 7. Karstic habitat of Cyrtodactylus amphipetraeus sp. nov. at the type locality of Tha Pha Pum, Tham Sri Fah Cave,
Tak Province, Thailand.
Discussion
Prior to this study, there were 36 species of Cyrtodactylus Gray known from Thailand which is comparable to that
of neighboring Myanmar (45 species), Laos (23 species), Vietnam (43 species), and Peninsular Malaysia (33 spe-
cies) but far exceeds that of China (five species) and Cambodia (eight species) (Uetz et al. 2020). However, in stark
contrast to the neighboring countries, the phylogenetic relationships of 44% (16) of the Thai species are unknown
because most of the recent species descriptions did not include molecular data (Bauer et al. 2009, 2010; Chan-ard
& Makchai, 2011; Kunya et al. 2014, 2015; Pauwels & Sumontha 2014; Pauwels et al. 2014a,b, 2016; Sumontha et
al., 2014, 2015), thus precluding any downstream comparative analyses (see Grismer et al. 2020a). To this end, we
add C. inthanon and C. doisuthep and lower that value to 38%.
The Cyrtodactylus sinyinensis group was constructed based on the descriptions of C. sinyineensis, C. dammath-
etensis, and C. sinyineensis and the inclusion of C. aequalis—all endemic to the Salween Basin of southern Myan-
mar west of the Tenasserim Mountains. Grismer et al. (2018a,b) noted that the other newly constructed Burmese
species groups were most closely related to lineages outside of Myanmar east of the Tenasserim Mountains and
their uplift from 21–16 mya was hypothesized to be the source of divergence between the western Burmese species
groups and the eastern Indochinese lineages (see Grismer et al. 2018b:Fig. 2). Now, however, the placement of east-
ern species C. doisuthep as the sister lineage to the remaining members of the C. sineyieensis group and the eastern
species C. tigroides as the sister lineage to all of them, would suggest the C. sineyineensis group may have evolved
east of the Tenasserim Mountains and recently invaded Myanmar. Complicating the issue further is the phylogenetic
placement of the eastern species C. inthanon and C. amphipetraeus sp. nov. nested among the western species of C.
sinyeensis group—something that does not occur within the other Burmese species groups—indicating there were
at least two independent cladogeneic events associated with the Tenasserim Mountains much later than the earlier
events proposed by Grismer et al. (2018b).
These new biogeographic developments arise from the expanded phylogeny which now includes Cyrtodac-
tylus doisuthep, C. inthanon, and C. amphipetraus sp. nov., underscoring the importance of phylogeny to any
downstream analyses. To try to propose a new biogeographic scenario at this juncture would be pointless with the
phylogenetic relationships of so many Thai species still unknown. We are in the process of acquiring tissue samples
for these species and stress herein that in this age of molecular genetics and biodiversity crises, collection of tissue
samples and application of molecular methods is crucial for taxonomic practice in studies of herpetofaunal diver-
A NEW CYRTODACTYLUS FROM THAILAND Zootaxa 4838 (2) © 2020 Magnolia Press · 207
sity in Southeast Asia (Smith et al. 2008; Murphy et al. 2013). Not only is this paramount for any comparative or
biogeographic analyses, it is now becoming a fundamental cornerstone of biodiversity conservation (Shaffer et al.
2015).
Acknowledgements
We are grateful to the Laboratory Animal Research Center, University of Phayao and The Institute of Animal for Sci-
entific Purposes Development (IAD), Thailand for the permission to do field work. Specimen collection protocols
and animal use were approved by the Institutional Ethical Committee of Animal Experimentation of the University
of Phayao, Phayao, Thailand (certificate number UP-AE61-01-04-0022 issued to Chatmongkon Suwannapoom).
Fieldwork, including collection of animals in the field and specimen exportation, was authorized by the Institute
of Animals for Scientific Purpose Development (IAD), Bangkok, Thailand (permit number U1-04995-2559, is-
sued to Siriwadee Chomdej). We thank Kanokwan Yimyoo, Thiti Ruengsuwan, Kawin Jiaranaisakul, Akkrachai
Aksornneam for help and support, and Evgeniy S. Popov for useful suggestions in the field of Latin and Ancient
Greek grammar. We are grateful to Anna S. Dubrovskaya and Platon V. Yushchenko for assistance in the lab. We
express our sincere gratitude to two anonymous reviewers for their useful suggestions on the earlier version of the
manuscript. Fieldwork, specimen collection, morphological examination, molecular phylogenetic analyses and data
analyses for this paper were conducted with the financial support of the Russian Science Foundation (RSF grant
No. 19-14-00050 to Nikolay A. Poyarkov). This work was partially financially supported by the Thailand Science
Research and Innovation (TSRI) (DBG6180025) with additional support from Chiang Mai University; specimen
collection and data analysis were also partially supported by the grants of the Unit of Excellence 2020 on Biodiver-
sity and Natural Resources Management, University of Phayao (No. UoE63005) and Plant Genetic Conservation
Project under the Royal Initiative of Her Royal Highness Princess Maha Chakri Sirindhorn, University of Phayao
(RD61017) to Chatmongkon Suwannapoom. The research was carried out within the frameworks of Russian State
projects AAAA-A16116021660077-3 and АААА-А17-117030310017-8.
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... In the north, are C. doisuthep Kunya, Panmongkol, Pauwels, Sumontha, Meewasana, Bunkhwamdi & Dangsri at Doi Suthep and C. inthanon Kunya, Sumontha, Panitvong, Dongkumfu, Sirisamphan & Pauwels from Doi Inthanon from Chiang Mai Province. Approximately 240 km to the south in the Tenasserim Mountains, is the newly described C. amphipetraeus Chomdej, Suwannapoom, Pawangkhanant, Pradit, Nazarov, Grismer & Poyarkov from Tak Province (Chomdej et al. 2020). An integrative taxonomic analysis (Chomdej et al. 2020) places these three upland species in the C. sinyneeneis group (sec. ...
... Approximately 240 km to the south in the Tenasserim Mountains, is the newly described C. amphipetraeus Chomdej, Suwannapoom, Pawangkhanant, Pradit, Nazarov, Grismer & Poyarkov from Tak Province (Chomdej et al. 2020). An integrative taxonomic analysis (Chomdej et al. 2020) places these three upland species in the C. sinyneeneis group (sec. Gris-mer et al. 2018a,b;2020a) that was previously considered to be endemic to a karstic archipelago in the lowlands of the Salween Basin of Myanmar. ...
... This new population lies approximately 60 km to the southwest of C. inthanon of the eastern branch of the range. Phylogenetic analyses using the mitochondrial gene NADH dehydrogenase subunit 2 (ND2) and its flanking tRNAs place this new population in clade 1 of the C. sinyineensis group (Chomdej et al. 2020). However, based on a combination of scale and color pattern characters, morphometrics, and molecular divergences, it cannot be ascribed to any known species of that group. ...
Article
A new gekkonid lizard, Cyrtodactylus maelanoi sp. nov., from Mae Hong Son Province of the Thai Highlands is described using an integrative taxonomic analysis based on morphology, color pattern, and the mitochondrial gene NADH dehydrogenase subunit 2 (ND2) and its flanking tRNAs. Phylogenetic analyses place the new species within clade 1 of the C. sinyineensis group and as the sister species to C. inthanon with an uncorrected pairwise sequence divergence of 3.9%. Collection data gathered in the field indicate that C. maelanoi sp. nov. is a habitat generalist. Reconstruction of the ancestral habitat preference for the C. sinyineensis group by way of stochasitc character mapping (SCM) indicates that karstic environments were the ancestral condition out of which the general habitat preference of the ancestor of C. maelanoi sp. nov. and C. inthanon and that of C. amphipetreaus and C. doisuthep evolved three times independently. Additionally, SCM demonstrated that the evolution of a granitic habitat preference from a karst-adapted ancestor happened in C. aequalis. The discovery of a new upland species in the Thai Highlands brings into focus the understudied nature of the mountain systems of western Thailand and the need for their continued exploration and conservation.
... Grismer et al. 2018b) ranges from the uplands of northwestern Thailand to the lowland karstic archipelagos of the Salween Basin in southeastern Myanmar (Grismer et al. 2018b(Grismer et al. , 2020c and is composed of 12 nominal species (Figs 3, 24, 33). The group is divisible into two strongly supported clades (Chomdej et al. 2020;Fig. 3). ...
... am phipetraeus) margins of the Salween Basin (Figs. 3,24). All species in this group are narrow-range endemics (Grismer et al. 2018b(Grismer et al. , 2020dChomdej et al. 2020) and more species belonging to this group are expected to be discovered when the vast number of small, isolated, karstic hills in the Salween Basin and the many unsurveyed mountain tops in western Thailand are explored. ...
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The gekkonid genus Cyrtodactylus is the third most speciose vertebrate genus in the world, containing well over 300 species that collectively range from South Asia to Melanesia across some of the most diverse landscapes and imperiled habitats on the planet. A genus-wide phylogeny of the group has never been presented because researchers working on different groups were using different genetic markers to construct phylogenies that could not be integrated. We present here Maximum likelihood and Bayesian inference mitochondrial and mito-nuclear phylogenies incorporating of 310 species that include dozens of species that had never been included in a genus-wide analysis. Based on the mitochondrial phylogeny, we partition Cyrtodactylus into 31 well-supported monophyletic species groups which, if used as recommended herein, will increase the information content of future integrative taxonomic analyses that continue to add new species to this genus at an ever-increasing annual rate. Data presented here reiterate the outcome of several previous studies indicating that Cyrtodactylus comprises an unprecedented number of narrow-range endemics restricted to single mountain tops, small islands, or karst formations that still remain unprotected. This phylogeny can provide a platform for various comparative ecological studies that can be integrated with conservation management programs across the broad diversity of landscapes and habitats occupied by this genus. Additionally, these data indicate that the true number of Cyrtodactylus remains substantially underrepresented.
... Grismer et al. 2018b) ranges from the uplands of northwestern Thailand to the lowland karstic archipelagos of the Salween Basin in southeastern Myanmar (Grismer et al. 2018b(Grismer et al. , 2020c and is composed of 12 nominal species (Figs 3, 24, 33). The group is divisible into two strongly supported clades (Chomdej et al. 2020;Fig. 3). ...
... am phipetraeus) margins of the Salween Basin (Figs. 3,24). All species in this group are narrow-range endemics (Grismer et al. 2018b(Grismer et al. , 2020dChomdej et al. 2020) and more species belonging to this group are expected to be discovered when the vast number of small, isolated, karstic hills in the Salween Basin and the many unsurveyed mountain tops in western Thailand are explored. ...
Article
Full-text available
The gekkonid genus Cyrtodactylus is the third most speciose vertebrate genus in the world, containing well over 300 species that collectively range from South Asia to Melanesia across some of the most diverse landscapes and imperiled habitats on the planet. A genus-wide phylogeny of the group has never been presented because researchers working on different groups were using different genetic markers to construct phylogenies that could not be integrated. We present here Maximum likelihood and Bayesian inference mitochondrial and mito-nuclear phylogenies incorporating of 310 species that include dozens of species that had never been included in a genus-wide analysis. Based on the mitochondrial phylogeny, we partition Cyrtodactylus into 31 well-supported monophyletic species groups which, if used as recommended herein, will increase the information content of future integrative taxonomic analyses that continue to add new species to this genus at an ever-increasing annual rate. Data presented here reiterate the outcome of several previous studies indicating that Cyrtodactylus comprises an unprecedented number of narrow-range endemics restricted to single mountain tops, small islands, or karst formations that still remain unprotected. This phylogeny can provide a platform for various comparative ecological studies that can be integrated with conservation management programs across the broad diversity of landscapes and habitats occupied by this genus. Additionally, these data indicate that the true number of Cyrtodactylus remains substantially underrepresented.
... brings the total number of recognized Ansonia species to 38 (Frost 2022), and the total number of Ansonia species known from Thailand to nine . Suwannapoom et al. (2020Suwannapoom et al. ( , 2021 recently discussed the importance of the entire Tenasserim Mountain region for its notable recent discoveries of endemic amphibians and reptiles (Matsui 2006;Sumontha et al. 2012Sumontha et al. , 2017Wilkinson et al. 2012;Connette et al. 2017;Grismer et al. 2016Grismer et al. , 2020aGrismer et al. , 2020bGrismer et al. , 2020cMatsui et al. 2018;Pawangkhanant et al. 2018;Suwannapoom et al. 2018;Lee et al. 2019;Chomdej et al. 2020Chomdej et al. , 2021Poyarkov et al. 2020Poyarkov et al. , 2021. Additional exploration followed by integrative taxonomic analyses will continue to increase our understanding of this region's exceptional herpetofaunal diversity and endemism and will provide the foundation for sciencebased conservation management programs. ...
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An integrative taxonomic analysis confirmed the new species status of a recently discovered upland population of Ansonia from Thongsong, Thongsong District, Nakhon Si Thammarat Province, Thailand from the sky island archipelago south of the Isthmus of Kra. Ansonia infernalis sp. nov. is a member of the Thai-Burmese clade within the more inclusive Thai-Malaya Peninsula clade of western and southern Thailand. It is separated from all other species of Ansonia by a unique combination of morphometric and discrete morphological and color pattern characteristics and is the sister species of a clade of eight other species found north of the Isthmus of Kra. Ansonia infernalis sp. nov. is the newest member of a long list of range-restricted endemics from the sky island archipelago of the Thai-Malay Peninsula and continues to underscore the unexplored nature of this region and its need for conservation.
... The genus Cyrtodactylus is monophyletic and currently contains 330 recognized species (Uetz et al. 2022) within 32 monophyletic species groups that have been delimited based on molecular data (Grismer et al. 2022). Most of the known species diversity is in mainland Southeast Asia Grismer et al. 2018bGrismer et al. , 2021bUetz et al. 2022), including Thailand, which is home to 39 species or nearly 9% of the described diversity (e.g., Chomdej et al. 2020;Grismer et al. 2020;Termprayoon et al. 2021;Uetz et al. 2022). Although the number of recognized species in the genus has rapidly increased in recent years, the true species diversity of the genus is still underestimated, and many known molecular lineages await formal description as species (Brennan et al. 2017;Chomdej et al. 2021;Grismer et al. 2021b). ...
Article
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Cyrtodactylus monilatus sp. nov. is described from Si Sawat District, Kanchanaburi Province, in western Thailand. The new species superficially resembles C. zebraicus Taylor, 1962 from southern Thailand. However, differences between the new species from C. zebraicus and other congeners were supported by an integrative taxonomic analysis of molecular and morphological data. Phylogenetic analyses based on the mitochondrial NADH dehydrogenase subunit 2 (ND2) gene showed that the new species is a member of the C. oldhami group and closely related to Cyrtodactylus sp. MT468911 from Thong Pha Phum National Park, Thong Pha Phum District, Kanchanaburi Province. Uncorrected pairwise genetic divergences ( p -distances) between the new species and its congeners, including C. zebraicus , ranged from 7.7–17.7%. Cyrtodactylus monilatus sp. nov. can also be distinguished from all members of the C. oldhami group by having a unique combination of morphological characters, including a snout to vent length of 53.7–63.3 mm in adult males and 58.6–75.8 mm in adult females; 22–34 paravertebral tubercles; 34–42 ventral scales; 30–39 enlarged contiguous femoroprecloacal scales; femoral pores and precloacal pores absent in both sexes; four or five rows of postprecloacal scales; enlarged median subcaudal scales absent; weak ventrolateral folds present; 4–7 rows of paired, paravertebral, dark-brown blotches edged in yellow or yellowish white; and two rows of small, diffuse, yellow or yellowish white spots on flanks. The new species occurs in a narrow range of forest at mid to low elevations associated with karst landscapes in the Tenasserim mountain range.
... This topology contradicts several earlier studies on the taxonomy of the group (e.g., Guo et al., 2011;Wang et al., 2020), but generally agrees with the recent multilocus phylogenetic study by 2013). Not only the phylogenetic hypothesis is crucial for any comparative or biogeographic analyses, it now also became a keystone of biodiversity conservation (Shaffer et al., 2015;Chomdej et al., 2020). Phylogenetic studies on the remaining species of Asthenodipsas are required to fully resolve the taxonomy of the genus; furthermore, additional taxon and gene sampling will likely enhance the phylogenetic resolution on the level of the subfamily Pareinae and might lead to discovery of additional new lineages and species. ...
Article
Full-text available
Slug-eating snakes of the subfamily Pareinae are an insufficiently studied group of snakes specialized in feeding on terrestrial mollusks. Currently Pareinae encompass three genera with 34 species distributed across the Oriental biogeographic region. Despite the recent significant progress in understanding of Pareinae diversity, the subfamily remains taxonomically challenging. Here we present an updated phylogeny of the subfamily with a comprehensive taxon sampling including 30 currently recognized Pareinae species and several previously unknown candidate species and lineages. Phylogenetic analyses of mtDNA and nuDNA data supported the monophyly of the three genera Asthenodipsas , Aplopeltura , and Pareas . Within both Asthenodipsas and Pareas our analyses recovered deep differentiation with each genus being represented by two morphologically diagnosable clades, which we treat as subgenera. We further apply an integrative taxonomic approach, including analyses of molecular and morphological data, along with examination of available type materials, to address the longstanding taxonomic questions of the subgenus Pareas , and reveal the high level of hidden diversity of these snakes in Indochina. We restrict the distribution of P. carinatus to southern Southeast Asia, and recognize two subspecies within it, including one new subspecies proposed for the populations from Thailand and Myanmar. We further revalidate P. berdmorei , synonymize P. menglaensis with P. berdmorei , and recognize three subspecies within this taxon, including the new subspecies erected for the populations from Laos and Vietnam. Furthermore, we describe two new species of Pareas from Vietnam: one belonging to the P. carinatus group from southern Vietnam, and a new member of the P. nuchalis group from the central Vietnam. We provide new data on P. temporalis , and report on a significant range extension for P. nuchalis . Our phylogeny, along with molecular clock and ancestral area analyses, reveal a complex diversification pattern of Pareinae involving a high degree of sympatry of widespread and endemic species. Our analyses support the “upstream” colonization hypothesis and, thus, the Pareinae appears to have originated in Sundaland during the middle Eocene and then colonized mainland Asia in early Oligocene. Sundaland and Eastern Indochina appear to have played the key roles as the centers of Pareinae diversification. Our results reveal that both vicariance and dispersal are responsible for current distribution patterns of Pareinae, with tectonic movements, orogeny and paleoclimatic shifts being the probable drivers of diversification. Our study brings the total number of Pareidae species to 41 and further highlights the importance of comprehensive taxonomic revisions not only for the better understanding of biodiversity and its evolution, but also for the elaboration of adequate conservation actions.
... During the last two decades, the number of new species described in this genus has significantly increased with the exploration of unsurveyed karst formations (Luu et al. 2016;Nazarov et al. 2018;Davis et al. 2019;Grismer et al. 2018Grismer et al. , 2020b. Moreover, genetic data has become a useful tool for taxonomic studies, revealing hidden diversity within the genus (Murdoch et al. 2019;Chomdej et al. 2020;Neang et al. 2020;Riyanto et al. 2020;Kamei and Mahony 2021;Liu and Rao 2021). Recent molecular studies have further supported the monophyly of this genus based on the most complete phylogenetic analysis to date, and have recognized 31 species groups (Grismer et al. 2021b). ...
Article
Full-text available
The bent-toed geckos of the Cyrtodactylus pulchellus group are widely distributed along the Thai-Malay Peninsula. Although taxonomic and phylogenetic studies of this species group have been continuously conducted, only some populations from Thailand have been included, resulting in hidden diversity within this group. In this study, we used morphological and molecular data to clarify the taxonomic status and describe a new population from Tarutao Island, Satun Province, southern Thailand. Cyrtodactylus stellatus sp. nov. can be distinguished from its congeners by the combination of the following morphological characters: body size; tuberculation; number of dark body bands, ventral scales, and femoroprecloacal pores in males; presence of precloacal pores in females; and scattered pattern on dorsum. Phylogenetic analyses of the mitochondrial ND2 gene recovered the new species as the sister species to C. astrum , with an uncorrected pairwise divergence of 9.78–12.37%. Cyrtodactylus stellatus sp. nov. is currently only known from Tarutao Island, Thailand. The discovery of this species suggests that the diversity within the C. pulchellus group remains underestimated and future exploration of unsurveyed areas are needed to further the understanding of this group and its geographic range.
... The first 25% of the sampled trees was discarded as burn-in after the standard deviation of split frequencies of the two runs reached a value of less than 0.01, and then the remaining trees were used to create a 50% majority-rule consensus tree and to estimate Bayesian posterior probabilities (BPP). Nodes with BPP of 95 and above were considered strongly supported (Huelsenbeck et al. 2001;Wilcox et al. 2002;Alfaro et al. 2003) and nodes with values of 90-94 as well supported (Chomdej et al. 2020). Maximum Likelihood (ML) analysis was performed in RaxmlGUI v. 1.5 (Silvestro and Michalak 2012), and nodal support was estimated by 1,000 rapid bootstrap replicates. ...
Article
Full-text available
A new species of Cyrtodactylus is described on the basis of five specimens collected from the karst formations of Zhenkang County, Yunnan Province, China. Cyrtodactylus zhenkangensis sp. nov. is recognized by having a unique combination of morphological characters, the most diagnostic being: 12–15 enlarged femoral scales on each thigh; 2–5 femoral pores on each thigh in males, 0–3 pitted scales on each thigh in females; eight or nine precloacal pores in a continuous row or separated by one poreless scale in males, 7–9 pitted scales in females; subcaudals enlarged, arranged alternately as single and double on anterior and mostly single at middle and posterior; dorsal surface of head with obvious reticulations. Phylogenetic analyses show that the new species is a member of the C. wayakonei species group and a sister taxon to a clade consisting of C. wayakonei and C. martini based on Maximum Likelihood analyses and Bayesian Inference and differs from its congeners by at least 12.0% genetic divergence in a fragment of the COI gene.
Article
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The integrated results of morphological and molecular phylogenetic analyses confirmed the new species status of a recently discovered population of Ansonia from Suan Phueng District, Ratchaburi Province, Thailand. Ansonia karen sp. nov. is separated from all other species of Ansonia by a unique combination of mensural, discrete morphological, and color pattern characteristics and is the sister species of A. thinthinae from Tanintharyi Division, Myanmar. This discovery fills a geographic hiatus of 350 km between it and A. kraensis from Ranong Province, Thailand. Ansonia karen sp. nov. is the newest member of a long list of range-restricted endemics having been recently discovered in the northern Tenasserim Mountain region of western Thailand and continues to underscore the unexplored nature of this region and its need for conservation.
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We present the latest version of the Molecular Evolutionary Genetics Analysis (MEGA) software, which contains many sophisticated methods and tools for phylogenomics and phylomedicine. In this major upgrade, MEGA has been optimized for use on 64-bit computing systems for analyzing bigger datasets. Researchers can now explore and analyze tens of thousands of sequences in MEGA. The new version also provides an advanced wizard for building timetrees and includes a new functionality to automatically predict gene duplication events in gene family trees. The 64-bit MEGA is made available in two interfaces: graphical and command line. The graphical user interface (GUI) is a native Microsoft Windows application that can also be used on Mac OSX. The command line MEGA is available as native applications for Windows, Linux, and Mac OSX. They are intended for use in high-throughput and scripted analysis. Both versions are available from www.megasoftware.net free of charge.
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