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A new insular species of the Cyrtodactylus pulchellus
group (Reptilia, Gekkonidae) from Tarutao Island,
southern Thailand revealed by morphological
and genetic evidence
Korkhwan Termprayoon1, Attapol Rujirawan1, Natee Ampai2,
Perry L. Wood Jr3,4, Anchalee Aowphol1
1Department of Zoology, Faculty of Science, Kasetsart University, Bangkok, 10900 ailand 2Department of
Biology, Faculty of Science, Srinakharinwirot University, Bangkok, 10110 ailand 3Department of Biologi-
cal Sciences and Museum of Natural History, Auburn University, Auburn, AL, USA 4Department of Ecology
and Evolutionary Biology, University of Michigan, Ann Arbor, MI 48109-1085, USA
Corresponding author: Anchalee Aowphol (fsciacl@ku.ac.th)
Academic editor: omas Ziegler|Received 29 August 2021|Accepted 4 October 2021|Published 12 November 2021
http://zoobank.org/FDBBD099-29F0-4521-9B18-AF841BC88F41
Citation: Termprayoon K, Rujirawan A, Ampai N, Wood Jr PL, Aowphol A (2021) A new insular species of the
Cyrtodactylus pulchellus group (Reptilia, Gekkonidae) from Tarutao Island, southern ailand revealed by morphological
and genetic evidence. ZooKeys 1070: 101–134. https://doi.org/10.3897/zookeys.1070.73659
Abstract
e bent-toed geckos of the Cyrtodactylus pulchellus group are widely distributed along the ai-Malay
Peninsula. Although taxonomic and phylogenetic studies of this species group have been continuously
conducted, only some populations from ailand 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 ailand. Cyrtodactylus stellatus
sp. nov. can be distinguished from its congeners by the combination of the following morphological char-
acters: 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 uncor-
rected pairwise divergence of 9.78–12.37%. Cyrtodactylus stellatus sp. nov. is currently only known from
Tarutao Island, ailand. e 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 un-
derstanding of this group and its geographic range.
ZooKeys 1070: 101–134 (2021)
doi: 10.3897/zookeys.1070.73659
https://zookeys.pensoft.net
Copyright Korkhwan Termprayoon et al. This is an open access ar ticle distributed under the terms of the Creative Commons Attribution License
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Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
102
Keywords
Cyrtodactylus astrum, Cyrtodactylus stellatus sp. nov., karst, morphology, phylogeny, taxonomy
Introduction
Bent-toed geckos in the genus Cyrtodactylus Gray, 1827 are geographically widespread
and inhabit lowland (e.g., peat swamps, karst formations, and limestone forests) to
mountainous regions (> 1,500 m a.s.l) of South Asia to Melanesia, ranging from In-
dia, Myanmar, ailand, Vietnam, Cambodia, Malaysia, Java, Papua New Guinea to
northern Australia (Wood et al. 2012; Nielsen and Oliver 2017; Pauwels et al. 2018;
Purkayastha et al. 2020; Riyanto et al. 2020; Grismer et al. 2020a, 2021a, 2021b). is
genus is the most diverse group of gekkotans, comprising 314 nominal species (Uetz et
al. 2021). During the last two decades, the number of new species described in this genus
has signicantly increased with the exploration of unsurveyed karst formations (Luu et
al. 2016; Nazarov et al. 2018; Davis et al. 2019; Grismer et al. 2018, 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 phy-
logenetic analysis to date, and have recognized 31 species groups (Grismer et al. 2021b).
One clade of particular interest is the Cyrtodactylus pulchellus group. is relatively
diverse group is distributed along the ai-Malay Peninsula and has high morphologi-
cal and molecular variation. Cyrtodactylus pulchellus Gray, 1827 was thought to be a
single wide-ranging species across their distributional range, but following an integra-
tive approach many new species have been described (e.g., C. bintangrendah Grismer et
al., 2012, C. langkawiensis Grismer et al., 2012, and C. sharkari Grismer et al., 2014).
is species group has been recovered as monophyletic and currently contains 16 rec-
ognized species, based on multiple phylogenetic studies (Grismer et al. 2012, 2014,
2016; Quah et al. 2019; Wood et al. 2020; Termprayoon et al. 2021). is group is
distributed from the south of the Isthmus of Kra, southern ailand to southern Pen-
insular Malaysia and some of its oshore islands (Grismer and Ahmad 2008; Sumon-
tha et al. 2012; Grismer et al. 2012, 2014, 2016; Quah et al. 2019; Wood et al. 2020;
Termprayoon et al. 2021). During eld surveys, specimens of the C. pulchellus group
were collected from Tarutao Island, Satun Province, southern ailand. Initially, these
specimens were recognized as an insular population of C. astrum Grismer et al., 2012
due to their supercial resemblance in coloration pattern and dorsal tuberculation. A
re-examination of these specimens showed morphological dierences from its other
congeners and mitochondrial DNA sequence data revealed corroborative evidence that
the new population of Cyrtodactylus from Tarutao Island represents a distinct mono-
phyletic lineage and is the sister species to C. astrum from the adjacent mainland.
Based on integrative analyses, we considered this new Cyrtodactylus population from
Tarutao Island as distinct and described it as a new species below.
A new insular species of the Cyrtodactylus pulchellus group 103
Materials and methods
Sampling
Field surveys were conducted on Tarutao Island, Mueang Satun District, Satun Prov-
ince, southern ailand from November 2017 to November 2019 (Fig. 1). Specimens
of the C. pulchellus group were collected from karst forest at night (1900–2200 h).
Ecological data (air temperature and relative humidity) were recorded using a Kestrel
4000 Weather Meter, and habitat use of each specimen was noted. Geographical co-
ordinates and elevation were recorded using a Garmin GPSMAP 64s. For molecular
studies, liver tissue was taken from each euthanized specimen, individually preserved
in 95% ethyl alcohol, and stored at -20 °C. Specimens were initially xed in 10%
formalin and later transferred into 70% ethyl alcohol for permanent storage. Voucher
specimens were deposited in the herpetological collections of the Zoological Museum,
Kasetsart University, ailand (ZMKU). Additional preserved specimens were exam-
ined in the holdings of the ailand Natural History Museum (THNHM), ailand,
and the La Sierra University Herpetological Collection (LSUHC), La Sierra Univer-
sity, Riverside, California, USA.
DNA extraction and PCR amplification
Total genomic DNA was extracted from ethanol-preserved liver tissue of ve Cyr-
todactylus specimens from Tarutao Island (Table 1) using a NucleoSpin Tissue Kit
(Macherey-Nagel GmbH & Co. KG, Germany). A fragment of mitochondrial NADH
dehydrogenase subunit 2 (ND2) gene and its anking tRNAs was amplied using a
double-stand Polymerase Chain Reaction (PCR) under the following conditions: ini-
tial denaturation at 94 °C for 4 min, followed by 35 cycles of denaturation at 94 °C for
30 sec, annealing at 48–52 °C for 30 sec, extension at 72 °C for 1 min 30 sec, and nal
extension at 72 °C for 7 min using the primers Metf6 (5' AAGCTTTCGGGCCCA-
TACC 3'; Macey et al. 1997), and COIH (5' AGRGTGCCAATGTCTTTGTGRTT
3'; Macey et al. 1997). PCR products were puried using NucleoSpin Gel and PCR
Clean-Up kit (Macherey-Nagel GmbH & Co. KG, Germany). Puried products were
sequenced for both strands using the same amplifying primers on an ABI 3730XL
DNA Sequencer (Applied Biosystems, CA, USA). Sequences were visually checked
and aligned in Geneious R11 (Biomatters, Ltd, Auckland, New Zealand). e protein-
coding region of ND2 was translated to amino acids and checked to conrm the lack
of premature stop codons. All sequences were deposited in GenBank under the acces-
sion numbers OK094494–OK094503 (Table 1).
Phylogenetic analyses
Phylogenetic trees were reconstructed using two dierent methods, Maximum Like-
lihood (ML) and Bayesian Inference (BI). e best substitution model for each par-
tition was determined using the Bayesian Information Criterion (BIC) under the
Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
104
Figure 1. Map showing the type locality of Cyrtodactylus stellatus sp. nov. from Tarutao Island, Mueang
Satun District, Satun Province, ailand and the type localities of closely related species, C. astrum,
C.dayangbuntingensis, C. langkawiensis, and C. lekaguli.
greedy search algorithm as implemented in PartitionFinder2 on XSEDE (Lanfear et
al. 2016). e selected models for ML and BI were TIM+G for 1st and 2nd codon posi-
tions of ND2, TVM+I+G for 3rd codon position of ND2 and TRN+I+G for tRNAs.
e ML analysis was performed in IQ-TREE web server v1.6.12 (Trinopoulos et al.
2016) with 1,000 bootstrap replicates using ultrafast bootstrap approximation (Minh
et al. 2013). e BI analysis was performed in MrBayes 3.2.6 on XSEDE (Ronquist
et al. 2012) using the CIPRES Science Gateway v3.3 (Miller et al. 2010). Two simul-
taneous runs were performed with four chains per run, three hot and one cold under
the default settings. e analysis was run for 10,000,000 generations and sampled
every 1,000 generations from the Markov chain Monte Carlo (MCMC), with the
rst 25% of each run discarded as burn-in. Stationarity and the eective sample
sizes (ESS) for all parameters were assessed in Tracer v1.7.1. (Rambaut et al. 2018).
Nodes with ultrafast bootstrap support (UFB) of ≥ 95 and Bayesian posterior prob-
abilities (BPP) of ≥ 0.95 were considered to be strongly supported (Huelsenbeck and
Ronquist 2001; Wilcox et al. 2002; Minh et al. 2013). Intraspecic and interspecic
uncorrected pairwise genetic divergences (p-distance) were calculated in MEGA X
10.0.5 using the pairwise deletion option for the treatment of gaps and missing data
in the dataset (Kumar et al. 2018).
A new insular species of the Cyrtodactylus pulchellus group 105
Table 1. Samples used in the molecular analyses, including their GenBank accession number (ND2),
voucher number and locality. WM = West Malaysia; TH = ailand.
SpeciesLocalityMuseum No. GenBank
Accession
No.
Reference
Outgroup
Agamura persica Pakistan, Baluchistan Province, Makran District, Gwadar
division
FMNH 247474 JX440515 Wood et al. (2012)
Hemidactylus frenatus Unknow NC 00155 JX519468 Grismer et al. (2012)
Tropiocolotes steudneri captive JB 28 JX440520 Wood et al. (2012)
C. elok WM, Pahang, Fraser’s Hill, e Gap LSUHC 6471 JQ889180 Wood et al. (2012)
C. hontreensis Vietnam, Kien Giang Province, Kien Hai District, Hon
Tre Island
LSUHC 8583 JX440539 Wood et al. (2012)
C. intermedius TH, Chantaburi Province, Khao Khitchakut District LSUHC 9513 JX519469 Grismer et al. (2012)
TH, Chantaburi Province, Khao Khitchakut District LSUHC 9514 JX519470 Grismer et al. (2012)
C. interdigitalis Lao, Khammouan Province, Nakai District FMNH 255454 JQ889181 Wood et al. (2012)
Cyrtodactylus sp. TH, Loei, Phu Rua FMNH 265806 JX519471 Grismer et al. (2012)
Ingroup
C. astrum WM, Perlis, Gua Kelam LSUHC 8806 JX519481 Grismer et al. (2012)
WM, Perlis, Gua Kelam LSUHC 8807 JX519478 Grismer et al. (2012)
WM, Perlis, Gua Kelam LSUHC 8808 JX519479 Grismer et al. (2012)
WM, Perlis, Gua Kelam LSUHC 8809 JX519480 Grismer et al. (2012)
WM, Perlis, Kuala Perlis LSUHC 8815 JX519482 Grismer et al. (2012)
WM, Perlis, Kuala Perlis LSUHC 8816
(paratype)
JX519483 Grismer et al. (2012)
WM, Perlis, Perlis State Park LSUHC 9215 JX519473 Grismer et al. (2012)
WM, Perlis, Perlis State Park, Gua Wang Burma LSUHC 9962 JX519475 Grismer et al. (2012)
WM, Perlis, Perlis State Park, Gua Wang Burma LSUHC 9986 JX519476 Grismer et al. (2012)
WM, Perlis, Perlis State Park, Gua Wang Burma LSUHC 9987 JX519477 Grismer et al. (2012)
WM, Perlis, Wang Kelian LSUHC 10023 JX519474 Grismer et al. (2012)
WM, Perlis, Wang Kelian LSUHC 10024 JX519472 Grismer et al. (2012)
C.
australotitiwangsaensis
WM, Pahang, Fraser’s Hill LSUHC 8086 JX519486 Grismer et al. (2012)
WM, Pahang, Fraser’s Hill LSUHC 8087 JX519485 Grismer et al. (2012)
WM, Pahang, Genting Highlands LSUHC 6637
(holotype)
JX519484 Grismer et al. (2012)
C. bintangrendah WM, Kedah, Bukit Mertajam LSUHC 10331
(paratype)
MN125076 Quah et al. (2019)
WM, Kedah, Bukit Mertajam LSUHC 10519 MN125077 Quah et al. (2019)
WM, Kedah, Bukit Mertajam LSUHC 10520
(paratype)
MN125078 Quah et al. (2019)
WM, Kedah, Bukit Palang LSUHC 9984 JX519487 Grismer et al. (2012)
C. bintangtinggi WM, Perak, Bukit Larut LSUHC 8862 JX519493 Grismer et al. (2012)
WM, Perak, Bukit Larut LSUHC 9006
(paratype)
JX519494 Grismer et al. (2012)
C. dayangbuntingensis WM, Kedah, Dayang Bunting Island LSUHC 14353 MN125090 Quah et al. (2019)
WM, Kedah, Dayang Bunting Island LSUHC 14354 MN125091 Quah et al. (2019)
WM, Kedah, Dayang Bunting Island LSUHC 14355 MN125092 Quah et al. (2019)
C. evanquahi WM, Kedah, Gunung Baling BYU 53435
(holotype)
MN586889 Wood et al. (2020)
WM, Kedah, Gunung Baling BYU 53436
(paratype)
MN586890 Wood et al. (2020)
WM, Kedah, Gunung Baling BYU 53437
(paratype)
MN586891 Wood et al. (2020)
C. hidupselamanya WM, Kelantan, Felda Chiku 7 LSUHC 12161
(paratype)
KX011415 Grismer et al. (2016)
WM, Kelantan, Felda Chiku 7 LSUHC 12162
(paratype)
KX011416 Grismer et al. (2016)
WM, Kelantan, Felda Chiku 7 LSUHC 12163
(holotype)
KX011417 Grismer et al. (2016)
WM, Kelantan, Felda Chiku 7 LSUHC 12173
(paratype)
KX011420 Grismer et al. (2016)
Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
106
SpeciesLocalityMuseum No. GenBank
Accession
No.
Reference
C. jelawangensis WM, Gunung Stong, Kelantan LSUHC 11060
(paratype)
KJ659850 Grismer et al. (2014)
WM, Gunung Stong, Kelantan LSUHC 11062
(holotype)
KJ659852 Grismer et al. (2014)
WM, Kelantan, Gunung Stong LSUHC 11061
(paratype)
KJ659851 Grismer et al. (2014)
C. langkawiensis WM, Kedah, Pulau Langkawi, Wat Wanaram LSUHC 9120 JX519502 Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Wat Wanaram LSUHC 9122 JX519501 Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Wat Wanaram LSUHC 9123 JX519500 Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Wat Wanaram LSUHC 9124
(paratype)
JX519499 Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Wat Wanaram LSUHC 9125 JX519496 Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Wat Wanaram LSUHC 9434 JX519498 Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Wat Wanaram LSUHC 9435 JX519495 Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Wat Wanaram LSUHC 9437 JX519497 Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Wat Wanaram LSUHC 14347 MN125093 Quah et al. (2019)
WM, Kedah, Pulau Langkawi, Wat Wanaram LSUHC 14348 MN125094 Quah et al. (2019)
C. lekaguli TH, Phang-nga Province, Mueang Phang-nga District ZMKU R 00720 KX011425 Grismer et al. (2016)
TH, Phang-nga Province, Mueang Phang-nga District ZMKU R 00721 KX011426 Grismer et al. (2016)
TH, Phang-nga Province, Mueang Phang-nga District ZMKU R 00722 KX011427 Grismer et al. (2016)
TH, Phang-nga Province, Mueang Phang-nga District ZMKU R 00723 KX011428 Grismer et al. (2016)
TH, Trang Province, Na Yong District ZMKU R 00918 OK094494 is study
TH, Trang Province, Na Yong District ZMKU R 00919 OK094495 is study
TH, Trang Province, Na Yong District ZMKU R 00920 OK094496 is study
TH, Trang Province, Na Yong District ZMKU R 00921 OK094497 is study
TH, Trang Province, Na Yong District ZMKU R 00922 OK094498 is study
C. lenggongensis WM, Perak, Lenggong Valley LSUHC 9974
(holotype)
JX519490 Grismer et al. (2012)
WM, Perak, Lenggong Valley LSUHC 9975
(paratype)
JX519488 Grismer et al. (2012)
WM, Perak, Lenggong Valley LSUHC 9976
(paratype)
JX519489 Grismer et al. (2012)
WM, Perak, Lenggong Valley LSUHC 9977
(paratype)
JX519491 Grismer et al. (2012)
C. macrotuberculatus TH, Phuket Province, Kathu District, Kathu Waterfall ZMKU R 00890 MW809301 Termprayoon et al.
(2021)
TH, Phuket Province, Kathu District, Kathu Waterfall ZMKU R 00891 MW809302 Termprayoon et al.
(2021)
TH, Phuket Province, alang District, ep Krasatti ZMKU R 00894 MW809305 Termprayoon et al.
(2021)
TH, Phuket Province, alang District, ep Krasatti ZMKU R 00895 MW809306 Termprayoon et al.
(2021)
TH, Phuket Province, alang District, ep Krasatti ZMKU R 00896 MW809307 Termprayoon et al.
(2021)
TH, Satun Province, Mueang Satun District, Adang Island ZMKU R 00875 MW809295 Termprayoon et al.
(2021)
TH, Satun Province, Mueang Satun District, Rawi Island ZMKU R 00883 MW809299 Termprayoon et al.
(2021)
TH, Satun Province, Mueang Satun District, Rawi Island ZMKU R 00887 MW809300 Termprayoon et al.
(2021)
TH, Songkhla Province, Hat Yai District, ung Tam Sao ZMKU R 00876 MW809296 Termprayoon et al.
(2021)
TH, Songkhla Province, Hat Yai District, ung Tam Sao ZMKU R 00877 MW809297 Termprayoon et al.
(2021)
TH, Songkhla Province, Hat Yai District, ung Tam Sao ZMKU R 00878 MW809298 Termprayoon et al.
(2021)
WM, Kedah, Hutan Lipur Sungai Tupah LSUHC 9671 JX519510 Grismer et al. (2012)
WM, Kedah, Hutan Lipur Sungai Tupah LSUHC 9672 JX519511 Grismer et al. (2012)
WM, Kedah, Hutan Lipur Sungai Tupah LSUHC 9693 JX519517 Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Gunung Machinchang LSUHC 9448 JX519507 Grismer et al. (2012)
A new insular species of the Cyrtodactylus pulchellus group 107
SpeciesLocalityMuseum No. GenBank
Accession
No.
Reference
C. macrotuberculatus WM, Kedah, Pulau Langkawi, Gunung Raya LSUHC 9428 JX519506 Grismer et al. (2012)
WM, Kedah, Pulau Langkawi, Lubuk Sembilang LSUHC 6829 JX519505 Grismer et al. (2012)
WM, Perlis, Bukit Chabang LSUHC 10037 JX519519 Grismer et al. (2012)
WM, Perlis, Bukit Chabang LSUHC 10038 JX519518 Grismer et al. (2012)
C. pulchellus WM, Penang, Pulau Pinang, Empangan Air Itam LSUHC 6668 JX519523 Grismer et al. (2012)
WM, Penang, Pulau Pinang, Moongate Trail LSUHC 6727 JX519526 Grismer et al. (2012)
WM, Penang, Pulau Pinang, Moongate Trail LSUHC 6728 JX519525 Grismer et al. (2012)
WM, Penang, Pulau Pinang, Moongate Trail LSUHC 6729 JX519528 Grismer et al. (2012)
C. sharkari WM, Pahang, Merapoh, Gua Gunting LSUHC 11022
(holotype)
KJ659853 Grismer et al. (2014)
Cyrtodactylus stellatus
sp. nov.
TH, Satun Province, Mueang Satun District, Tarutao Island ZMKU R 00903
(paratype)
OK094499 is study
TH, Satun Province, Mueang Satun District, Tarutao Island ZMKU R 00905
(holotype)
OK094500 is study
TH, Satun Province, Mueang Satun District, Tarutao Island ZMKU R 00906
(paratype)
OK094501 is study
TH, Satun Province, Mueang Satun District, Tarutao Island ZMKU R 00907
(paratype)
OK094502 is study
TH, Satun Province, Mueang Satun District, Tarutao Island ZMKU R 00908
(paratype)
OK094503 is study
C. timur WM, Gunung Tebu, Terengganu LSUHC 10886 KJ659854 Grismer et al. (2014)
WM, Gunung Tebu, Terengganu LSUHC 11183
(paratype)
KJ659855 Grismer et al. (2014)
WM, Gunung Tebu, Terengganu LSUHC 11184
(paratype)
KJ659856 Grismer et al. (2014)
WM, Gunung Tebu, Terengganu LSUHC 11185
(paratype)
KJ659857 Grismer et al. (2014)
C. trilatofasciatus WM, Pahang, Cameron Highlands LSUHC 10064 JX519529 Grismer et al. (2012)
WM, Pahang, Cameron Highlands LSUHC 10065 JX519530 Grismer et al. (2012)
WM, Pahang, Cameron Highlands LSUHC 10066 JX519531 Grismer et al. (2012)
Morphology
e morphological characters and their denition used in this study were modied
from previous studies of the C. pulchellus group (Grismer and Ahmad 2008; Grismer
et al. 2012, 2014, 2016; Quah et al. 2019; Wood et al. 2020), and abbreviations are
derived from Grismer et al. (2018, 2020c). All mensural characters were taken with
digital calipers to the nearest 0.01 mm on the left side, while scale counts were made
on both sides when possible. Scalation and external morphology were evaluated under
a Nikon SMZ745 dissecting microscope. Measurement and meristic characters are
shown in Table 2, and external morphological characters evaluated are described below.
External morphological characters examined in the C. pulchellus group were the
degree of body tuberculation, weak tuberculation referring to dorsal body tubercles
that are low and rounded whereas prominent tuberculation refer to tubercles that are
raise and keeled; the presence or absence of tubercles on the dorsal and ventral surface
of the forearms; the presence or absence of tubercles in the gular region, throat, and
ventrolateral body folds; the width of the dark body bands relative to the width of the
interspace between the bands; the presence or absence of dark pigmentation infused in
the white caudal bands of adults; the presence or absence of a precloacal depression or
groove; the presence or absence of scattered white/yellow tubercles on the dorsum; and
Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
108
the presence or absence of white tail tip on the posterior portion of the original tail in
hatchlings and juveniles. Color pattern characteristics were taken from digital images
of live specimens in both sexes and of all possible age classes prior to preservation.
Statistical analyses
All analyses were performed using the base statistical software in R v3.6.1 (R Core Team
2019). To eliminate bias of sexual dimorphism, adult males and females were analyzed
separately. Morphological analyses were run on 15 mensural characters. Tail length
(TL) was not included due to their dierent condition (e.g., original, regenerated,
and broken). All measurements of each species were size-adjusted in order to remove
potential eects of allometry using the following allometric equation: Xadj = log[X ±
β(SVL ± SVLmean)], where Xadj = adjusted value; X = measured value; β = unstandard-
ized regression coecient for each OTU; SVL = measured snout–vent length; SVLmean
Table 2. Measurement and meristic characters used in this study, with abbreviations and explanations.
Abbreviations Characters
Measurement
SVL Snout–vent length, taken from the tip of snout to the vent
TW Tail width, taken at the base of the tail immediately posterior to the postcloacal swelling
TL Tail length, taken from vent to the tip of the tail, original or regenerated
FL Forearm length, taken from the posterior margin of the elbow while exed 90º to the inection of the exed wrist
TBL Tibia length, taken from the posterior surface of the knee while exed 90º to the base of the heel
AG Axilla to groin length, 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
HL Head length, the distance from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout
HW Head width, measured at the angle of the jaws
HD Head depth, the maximum height of head from the occiput to the throat
ED Eye diameter, the greatest horizontal diameter of the eyeball
EE Eye to ear distance, measured from the anterior edge of the ear opening to the posterior edge of the eyeball
ES Eye to snout distance, measured from anterior most margin of the eyeball to the tip of snout
EN Eye to nostril distance, measured from the anterior margin of the eyeball to the posterior margin of the external nares
IO Inter orbital distance, measured between the anterior edges of the orbit
EL Ear length, the greatest vertical distance of the ear opening
IN Internarial distance, measured between the nares across the rostrum
Meristic
SL Supralabial scales, counted from the largest scale immediately posterior to the dorsal inection of the posterior portion of the
upper jaw to the rostral scale
IL Infralabial scales, counted from the largest scale immediately posterior to the dorsal inection of the posterior portion of the
upper jaw to the mental scale
PVT e number of paravertebral tubercles between limb insertions, counted in a straight line immediately left or right of the
vertebral column
LRT e number of longitudinal rows of body tubercles, counted transversely across the center of the dorsum from one
ventrolateral fold to the other
VS e number of longitudinal rows of ventral scales, counted transversely across the center of the abdomen from one
ventrolateral fold to the other
4TL e number of subdigital lamellae beneath the fourth toe, counted from the base of the rst phalanx to the claw
FPP e total number of precloacal and femoral pores in male (i.e., the sum of the number of femoral and precloacal scales bearing
pores combined as a single meristic referred to as the femoroprecloacal pores)
PP e number of precloacal pores in female
BB e number of dark body bands between limb insertions
DCB e number of dark caudal bands on the original tail
A new insular species of the Cyrtodactylus pulchellus group 109
= overall average SVL of each OTU (orpe 1975, 1983; Turan 1999; Lleonart et al.
2000)—implemented through the R package GroupStruct (Chan and Grismer 2021).
Morphological measurements of C. astrum, C. dayangbuntingensis Quah et al., 2019,
C. langkawiensis and C. lekaguli Grismer et al., 2012 were obtained from their original
descriptions (Grismer et al. 2012; Quah et al. 2019). Additional preserved specimens
of C. astrum (from Malaysia) and C. lekaguli (topotypes) were examined and included
in the analyses (Appendix I). Morphometric adjustments were conducted separately
on each species and then concatenated into a single data frame to ensure there was no
interspecic conation of variation (Reist 1985; McCoy et al. 2006). Specimens were
assigned into ve groups (= species) based on phylogenetic analyses which are Cyrto-
dactylus Tarutao Island samples (N = 5 males, 5 females), C. astrum (N = 5 males, 3 fe-
males), C. dayangbuntingensis (N = 2 males), C. langkawiensis (N = 2 males, 4 females),
and C. lekaguli (N = 7 males, 9 females).
Principal components analysis (PCA) was performed on size-adjusted data for each
sex using FactoMineR package (Lê et al. 2008) and were visualized with the R pack-
age ggplot2 (Wickham 2016). For univariate analysis, Shapiro-Wilk test was used to
evaluate data to meet normality assumptions (p ≥ 0.05) and Levene’s test for testing for
equality of variance (p ≥ 0.05). Morphological dierences were compared using Analy-
sis of variance (ANOVA) or Kruskal-Wallis test. ANOVA was conducted on normally
distributed data with homogeneous variances and were subjected to Tukey HSD post
hoc tests (Tukey’s test) to determine which characters had statistically dierent mean
values for which pairs of species if ANOVA had a p-value of less than 0.05. Kruskal-
Wallis test was performed on non-normally distributed data and followed by a post
hoc Dunn’s multiple comparison (Dunn’s test). Due to limited sample sizes, C. dayang-
buntingensis (N = 2 males) and males of C. langkawiensis (N = 2 males) were excluded
from the univariate analysis.
Results
Phylogenetic relationships
e aligned matrix contained 1,429 mtDNA characters from 93 individuals of the
C. pulchellus group and nine individuals of outgroup species (Table 1). e standard
deviation of split frequencies among the two simultaneous BI runs was 0.002676, and
the ESS values of all parameters were greater than or equal to 2,494.4. e maximum
likelihood value of the best ML tree was lnL = -15,115.412.
e topologies of ML and BI analyses were largely concordant. e ML and BI
analyses recovered the C. pulchellus group as monophyletic with strong support (≥ 95
UFB, ≥ 0.95 BPP) which is comprised of two major clades referred to as Clades A and
B (Figs 2, 3). e Cyrtodactylus specimens from Tarutao Island represented a strongly
supported monophyletic group (≥ 95 UFB, ≥ 0.95 BPP) within Clade A containing
Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
110
Figure 2. e Maximum Likelihood tree of the Cyrtodactylus pulchellus group based on 1,429 bp of the
ND2 gene and anking tRNAs. Support values on branches are ultrafast bootstrap (UFB). Black circles
represent nodes strongly supported (UFB ≥ 95).
A new insular species of the Cyrtodactylus pulchellus group 111
Figure 3. e Bayesian consensus tree of the Cyrtodactylus pulchellus group based on 1,429 bp of the
ND2 gene and anking tRNAs. Support values on branches are Bayesian posterior probabilities (BPP).
Black circles represent nodes strongly supported (BPP ≥ 0.95).
Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
112
C. astrum, C. dayangbuntingensis, C. langkawiensis, and C. lekaguli. e Tarutao Island
samples were weakly recovered as a sister species to C. astrum from the adjacent Pen-
insular Malaysian mainland (64 UFB, 0.82 BPP). Clade B is composed of all other
species including C. australotitiwangsaensis Grismer et al., 2012, C. bintangrendah, C.
bintangtinggi Grismer et al., 2012, C. evanquahi Wood et al., 2020, C. hidupselamanya
Grismer et al., 2016, C. jelawangensis Grismer et al., 2014, C. lenggongensis Grismer et
al., 2016, C. macrotuberculatus Grismer and Ahmad, 2008, C. pulchellus, C. sharkari,
C. timur Grismer et al., 2014 and C. trilatofasciatus Grismer et al., 2012. Uncorrected
pairwise genetic divergences (p-distance) range from 0.00–1.17% within the Tarutao
Island specimens and 8.46–12.37% between the Tarutao Island specimens and other
species in Clade A (Table 3).
Morphology
e PCA was conducted on members from Clade A. e plots on the rst two PC axes
showed that the Tarutao Island specimens are clustered separately from other species in
both sexes (Fig. 4). In male, the rst two principal components explained 66.95% of
the morphological variation (Table 4). e rst principal component (PC1) accounted
for 45.88% of the variation and was heavily loaded on FLadj, TBLadj, HWadj, HDadj, EE-
adj, ESadj, and ENadj; and the PC2 accounted for 21.07% of the variation and was heavily
loaded on TWadj, AGadj, IOadj, and ELadj. PC analysis of females accounted for 56.74%
of the variation in rst two components. e PC1 accounted for 34.81% of the vari-
ation and was heavily loaded on TBLadj, HWadj, EEadj, ESadj, and ENadj; and the PC2
accounted for 21.93% of the variation and was heavily loaded on TWadj, IOadj and INadj.
e univariate analyses (ANOVA or Kruskal-Wallis test) were signicantly dif-
ferent (p < 0.05) in most morphological characters among the members of Clade A
(except C. dayangbuntingensis). In the comparison of adult males, the Tarutao Island
population was signicantly dierent from C. astrum and C. lekaguli in twelve mor-
phological characters (ANOVA or Kruskal-Wallis test, p < 0.001–0.006) except AGadj,
EDadj and INadj (ANOVA or Kruskal-Wallis test, p = 0.051–0.122). Subsequent Tukey’s
test or Dunn’s test demonstrated that Tarutao Island population was signicantly dif-
ferent from C. astrum in SVLadj, FLadj, TBLadj, HLadj, HWadj, HDadj, EEadj, ESadj, and
ENadj; and C. lekaguli in SVLadj, TWadj, FLadj, HLadj, HWadj, HDadj, EEadj, ESadj, IOadj,
and ELadj. In adult females, the Tarutao Island population was signicantly dierent
Table 3. Percentage uncorrected pairwise genetic divergence (p-distances) of Cyrtodactylus stellatus sp.
nov. and closely related species (Clade A) calculated from 1,429 base pairs of mitochondrial ND2 gene
and anking tRNAs.
Species N1 2 3 4 5
1Cyrtodactylus stellatus sp. nov. 5 0.48 (0.00–1.17)
2C. astrum 12 10.50 (9.78–12.37) 1.37 (0.00–2.97)
3C. dayangbuntingensis 3 9.90 (9.56–10.88) 9.86 (9.51–11.21) 0.14 (0.07–0.22)
4C. langkawiensis 10 10.49 (9.86–11.69) 10.19 (9.71–11.59) 7.62 (7.39–7.83) 0.42 (0.00–0.69)
5C. lekaguli 9 9.33 (8.46–10.80) 9.94 (8.98–11.77) 8.58 (8.00–9.59) 9.39 (8.42–10.54) 2.30 (0.00–4.27)
A new insular species of the Cyrtodactylus pulchellus group 113
Figure 4. Plots of the rst two principal components of Cyrtodactylus stellatus sp. nov. and the closely
related species in Clade A based on adjusted mensural characters of A males and B females. e letters in
the scatter plots refer to holotype (= H), paratype (= P), and topotype (= T).
Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
114
from C. astrum, C. langkawiensis and C. lekaguli in nine characters (ANOVA, p <
0.001–0.007) except SVLadj, TWadj, AGadj, HLadj, EDadj and ELadj (ANOVA or Kruskal-
Wallis test, p = 0.052–0.631). Subsequent Tukey’s test revealed that the Tarutao Island
population was signicantly dierent from C. astrum in FLadj, TBLadj, HWadj, HDadj,
EEadj, ESadj, and ENadj; C. langkawiensis in HWadj, ESadj, ENadj, and INadj; and C. lekaguli
in HDadj, IOadj, and INadj. Summary pairwise results (Tukey’s test or Dunn’s test) of sig-
nicant dierences in morphological characters for adult males and females of Clade A
are shown in Table 5. Additional dierences in meristic characters and coloration are
discussed in the comparison sections.
Table 4. Summary statistics and factor loadings of the rst three principal components (PC) of 15 men-
sural characters for males and females of Cyrtodactylus stellatus sp. nov. and its closely related species includ-
ing C. astrum, C. dayangbuntingensis, C. langkawiensis, and C. lekaguli. Bold texts indicate high loadings.
Characters Males Females
PC1 PC2 PC3 PC1 PC2 PC3
SVLadj 0.660 -0.284 0.293 0.374 -0.097 -0.127
TWadj 0.436 0.795 -0.073 0.380 0.845 0.194
FLadj 0.855 -0.228 -0.051 0.693 0.111 0.003
TBLadj 0.951 0.098 -0.032 0.881 0.181 0.102
AGadj -0.261 -0.728 0.364 -0.030 0.157 0.824
HLadj 0.459 -0.508 -0.560 0.532 -0.194 0.537
HWadj 0.943 -0.168 0.177 0.785 -0.350 -0.197
HDadj 0.838 -0.325 0.188 0.526 -0.647 0.170
EDadj 0.552 -0.195 -0.491 0.137 -0.086 0.612
EEadj 0.829 -0.171 0.194 0.755 -0.410 0.266
ESadj 0.932 0.113 0.054 0.900 -0.078 -0.271
ENadj 0.875 0.246 0.142 0.922 0.122 -0.263
IOadj 0.143 0.850 0.435 0.361 0.854 0.033
ELadj 0.303 0.770 -0.417 0.343 0.690 -0.298
INadj -0.105 0.093 0.560 -0.089 0.719 0.238
Eigenvalue 6.882 3.161 1.587 5.222 3.289 1.835
Percentage of variance 45.879 21.073 10.581 34.813 21.928 12.235
Cumulative proportion 45.879 66.952 77.533 34.813 56.741 68.976
Table 5. Summary pairwise results of statistically signicant characters (Tukey’s test; p < 0.05) from 15
mensural characters for males and females of Cyrtodactylus stellatus sp. nov. and closely related species
(Clade A). Abbreviations are listed in Table 2. Key: * tested by Dunn’s test; M = male; F = female.
Cyrtodactylus stellatus sp. nov. C. astrum C. langkawiensis
M F M F M F
Cyrtodactylus
stellatus sp. nov.
M – – – – – –
F – – – – – –
C. astrum M SVL, FL, TBL, HL,
HW, HD, EE, ES*, EN
– – – – –
F – FL, TBL, HW,
HD, EE, ES, EN
– – – –
C. langkawiensis M – – – – – –
F – HW, ES, EN, IN – HW, IN – –
C. lekaguli M SVL, TW, FL, HL, HW,
HD, EE, ES*, IO, EL
– TW, FL, TBL, HW,
EN, IO, EL
– – –
F – HD, IO, IN – FL, TBL, HW, ES,
EN, IO, EL
– ES
A new insular species of the Cyrtodactylus pulchellus group 115
Taxonomic hypotheses
Cyrtodactylus samples from Tarutao Island, Mueang Satun District, Satun Province
diered from its congeners in mtDNA, morphometrics and morphological compari-
sons. ese corroborated lines of evidence provide sucient support to warrant them
specic species status and is described as new below.
Taxonomy
Cyrtodactylus stellatus sp. nov.
http://zoobank.org/F2AF3CB9-F0FE-4749-9785-F57C7CAC021C
Figures 5–11
Stellar Bent-toed Gecko
Holotype. Adult male (ZMKU R 00905, Figs 5–7) collected from ailand, Satun
Province, Mueang Satun District, Tarutao National Park, Tarutao Island, Pha (= cli)
Toe Boo (6°42.185'N; 99°38.895'E; 2 m a.s.l.), on 11 March 2019 by Korkhwan Term-
prayoon, Anchalee Aowphol, Attapol Rujirawan, Natee Ampai and Siriporn Yodthong.
Paratypes (Figs 8–9). Two adult males (ZMKU R 00906–00907) and two adult
females (ZMKU R 00908–00909), same data as holotype. One adult female (ZMKU
R 00913) same data as holotype except collected on 12 May 2019. One adult male
(ZMKU R 00903) and two adult females (ZMKU R 00899–00900), same data as
holotype, except collected on 5 November 2017 by Korkhwan Termprayoon, Attapol
Rujirawan, Natee Ampai, and Siriporn Yodthong. One adult male (ZMKU R 00915)
collected from ailand, Satun Province, Mueang Satun District, Tarutao National
Park, Tarutao Island, Tarutao Outcrop (6°41.617'N; 99°38.796'E; 3 m a.s.l.) on 12
March 2019 by Korkhwan Termprayoon, Anchalee Aowphol, Attapol Rujirawan, Na-
tee Ampai and Siriporn Yodthong.
Referred specimens. ZMKU R 00901 (immature male) and ZMKU R 00902
(immature female) same data as holotype except collected on 5 November 2017 by
Korkhwan Termprayoon, Attapol Rujirawan, Natee Ampai, and Siriporn Yodthong.
ZMKU R 00904 (immature male) same data as holotype, except collected on 5 April
2018. ZMKU R 00910–00911 (two immature males) and ZMKU R 00912 (imma-
ture female) same data as holotype. ZMKU R 00914 (immature female) same data
as holotype except collected on 12 May 2019. ZMKU R 00916 (immature male)
and ZMKU R 00917 (juvenile) collected from ailand, Satun Province, Mueang
Satun District, Tarutao National Park, Tarutao Island, Tarutao Outcrop (6°41.617'N;
99°38.796'E; 3 m a.s.l.) on 12 March 2019 by Korkhwan Termprayoon, Anchalee
Aowphol, Attapol Rujirawan, Natee Ampai and Siriporn Yodthong.
Diagnosis. Cyrtodactylus stellatus sp. nov. can be distinguished from all other spe-
cies of the C. pulchellus group by the combination of the following characters: (1) SVL
86.3–95.9 mm in adult males, 86.6–96.1 mm in adult females; (2) 12–15 supralabial
and 10–13 infralabial scales; (3) weak tuberculation on body; (4) no tubercles on ven-
Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
116
tral surfaces of forelimbs, gular region, or in ventrolateral body folds; (5) 32–47 para-
vertebral tubercles; (6) 19–23 longitudinal rows of dorsal tubercles; (7) 32–40 rows of
ventral scales; (8) 20–23 subdigital lamellae on the fourth toe; (9) 24–29 femoropre-
cloacal pores in adult males; (10) precloacal pores present in adult females; (11) deep
precloacal groove in males; (12) dorsum bearing a scattered pattern of white tubercles;
(13) four dark dorsal body bands; (14) 10–12 dark caudal bands on original tail; (15)
white caudal bands in adults heavily infused with dark pigmentation; and (16) poste-
rior portion of tail in hatchlings and juveniles white.
Description of holotype. Adult male SVL 94.2 mm; head large, moderate in
length (HL/SVL 0.29) and wide (HW/HL 0.61), somewhat attened (HD/HL 0.38),
distinct from neck, and triangular in dorsal prole; lores concave anteriorly, inated
posteriorly; frontal and prefrontal regions deeply concave; canthus rostralis rounded
anteriorly; snout elongate (ES/HL 0.39), rounded in dorsal prole, laterally constrict-
ed; eye large (ED/HL 0.25); ear opening elliptical, moderate in size (EL/HL 0.09),
obliquely oriented; eye to ear distance slightly greater than diameter of eye; rostral rec-
tangular, divided dorsally by an inverted Y-shaped furrow, bordered posteriorly by left
and right supranasals and internasal, bordered laterally by rst supralabials; external
nares bordered anteriorly by rostral, dorsally by a large anterior supranasal, posteriorly
by two postnasals, ventrally by rst supralabial; 13/14 (left/right) rectangular suprala-
bials extending to just beyond upturn of labial margin, tapering abruptly below mid-
point of eye; second supralabial slightly larger than rst; 11/11 infralabials tapering in
size posteriorly; scales of rostrum and lores slightly raised, larger than granular scales on
top of head and occiput, those on posterior portion of canthus rostralis slightly larger;
scales on occiput intermixed with small tubercles; large, boney frontal ridges bordering
orbit conuent with boney, V-shaped, transverse, parietal ridge; dorsal superciliaries
elongate, smooth, largest anteriorly; mental triangular, bordered laterally by rst infral-
abials and posteriorly by left and right, trapezoidal postmentals which contact medially
for 50% of their length; one row of slightly enlarged, elongate sublabials extending
posteriorly to the seventh (left) and fth (right) infralabials; small, granular, gular scales
grading posteriorly into larger, at, smooth, imbricate, pectoral and ventral scales.
Body relatively short (AG/SVL 0.46) with well-dened, non-tuberculate, vent-
rolateral folds; dorsal scales small, granular, interspersed with low, regularly arranged,
weakly keeled tubercles, smaller intervening tubercles occasionally present; tubercles
extend from occiput to caudal constriction, absent from regenerated portion of tail;
tubercles on occiput and nape relatively small, those on body largest; approximately 21
longitudinal rows of tubercles at midbody; 36 paravertebral tubercles; 33 at imbricate
ventral scales between ventrolateral body folds; ventral scales larger than dorsal scales;
precloacal scales large, smooth; deep precloacal groove.
Forelimbs moderate in stature, relatively short (FL/SVL 0.16); scales on dorsal sur-
faces of forelimbs granular intermixed with larger tubercles; scales of ventral surface
of forearm at, subimbricate, tubercles absent; palmar scales small, weakly rounded;
digits well-developed, inected at basal, interphalangeal joints; subdigital lamellae rec-
A new insular species of the Cyrtodactylus pulchellus group 117
Figure 5. Adult male holotype of Cyrtodactylus stellatus sp. nov. (ZMKU R 00905) from Tarutao Island,
Satun Province. A specimen in life and immediately before preservative: B dorsal and C ventral views.
Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
118
Figure 6. Head of the holotype of Cyrtodactylus stellatus sp. nov. (ZMKU R 00905): A lateral view of the
left side B dorsal view, and C ventral view.
tangular proximal to joint inection, only slightly expanded distal to inection; digits
narrower distal to joints; claws well-developed, sheathed by a dorsal and ventral scale;
the fth digit broken on left forearm; hind limbs more robust than forelimbs, moder-
ate in length (TBL/SVL 0.19), larger tubercles on dorsal surface of legs separated by
smaller juxtaposed scales; ventral scales of thigh at, smooth, imbricate, larger than
dorsal granular scales; ventral, tibial scales at, smooth, imbricate; a single row of 34
enlarged femoroprecloacal scales extend nearly from knee to knee through precloacal
region where they are continuous with enlarged, pore-bearing precloacal scales; 27 sep-
arated pore-bearing femoroprecloacal scales (Fig. 10A), forming an inverted T bearing
a deep, precloacal groove; six pore-bearing scales bordering groove (three on each side
A new insular species of the Cyrtodactylus pulchellus group 119
Figure 7. Male holotype of Cyrtodactylus stellatus sp. nov. (ZMKU R 00905) in preservation. A dorsal
and B ventral views C tuberculation on dorsum, and D ventral view of left foot.
Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
120
of groove); postfemoral scales immediately posterior to enlarged scale row small, nearly
granular, forming an abrupt union with postfemoral scales on posteroventral margin
of thigh; plantar scales weakly rounded to at; digits well developed, inected at basal,
interphalangeal joints; subdigital lamellae proximal to joint inection rectangular, only
slightly expanded distal to inection; digits narrower distal to joints; claws well-devel-
oped, sheathed by a dorsal and ventral scale; 21/22 subdigital lamellae on the 4th toe.
Tail 94.8 mm in length, completely regenerated, 9.2 mm in width at base, taper-
ing to a point; regenerated tail covered with small, smooth, rectangular scales dorsally;
base of tail bearing hemipenial swellings; one row of 4/4 medium-sized postcloacal
tubercles on each hemipenial swelling; postcloacal scales smooth, at, large, imbricate.
Coloration in life (Fig. 5). Dorsal ground color of head, body, and limbs light-
brownish grey; a wide, dark-brown nuchal band bordered anteriorly and posteriorly by
thin, creamy-white lines bearing tubercles that extend from the posterior margin of one
eye to the posterior margin of other eye; the color of nuchal band and creamy-white
lines is faded above left ear opening; four dark-brown body bands between nuchal
loop and hind limb insertions that are also bordered anteriorly and posteriorly by thin,
creamy-white lines bearing tubercles, rst band terminates at shoulders, second and
third bands terminate just dorsal of ventrolateral folds, the fourth band terminates at
Figure 8. Male paratypes of Cyrtodactylus stellatus sp. nov. in preservation. A dorsal and B ventral views;
from left to right: ZMKU R 00903, ZMKU R 00906, ZMKU R 00907, and ZMKU R 00915.
A new insular species of the Cyrtodactylus pulchellus group 121
Figure 9. Female paratypes of Cyrtodactylus stellatus sp. nov. in preservation. A dorsal and B ventral
views; from left to right: ZMKU R 00899, ZMKU R 00900, ZMKU R 00908, ZMKU R 00909, and
ZMKU R 00913.
femurs; dark body bands slightly larger than light-colored interspaces; creamy-white
to light-yellow tubercles scattered on dorsal surfaces of nape, body, and limbs; one ad-
ditional dark-brown band posterior to hind limbs; light-brown regenerated tail, bear-
ing yellowish pigment on some scales; ventral surfaces of head smudged with brown;
abdomen and limbs beige, with slightly darker, lateral regions.
Coloration in preservative (Figs 6, 7). e overall color pattern of head, body,
limbs, and tail similar to that in life with some fading. Ground color of head, body,
limbs, and dorsum light-brown; dark body bands lighter than in life. Colored tubercu-
lation on dorsum fade to o-white. Tan colored on the ventral surface.
Variation. Cyrtodactylus stellatus sp. nov. usually varies in coloration and banding
pattern (Figs 8–11; Tables 6, 7). All specimens possess a clear dark-brown nuchal band
which is less clearly dened in ZMKU R 00903 and the holotype. In adult females,
precloacal pores are present but they lack the precloacal groove (Fig. 10B). Four speci-
mens (ZMKU R 00903, ZMKU R 00907, ZMKU R 00911, and ZMKU R 00913)
have prominent light-yellow tubercles scattered on the dorsum and limbs. Male para-
type (ZMKU R 00907) has continuous pore-bearing femoroprecloacal scales. Original
Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
122
tails (ZMKU R 00899, ZMKU R 00901–00902, ZMKU R 00910, ZMKU R 00912,
and ZMKU R 00916) have 10–12 dark caudal bands (Fig. 11A, B), light bands dif-
fused with dark pigment in adults (immaculate in immature and juvenile), subcaudal
scales transversely enlarged and shallow caudal furrows. Male paratypes have a single
row of 3–4L/2–4R postcloacal tubercles on each hemipenial swelling except ZMKU
R 00907 which has two rows of 4/5 on each side. is character in female paratypes is
very small, and a single row of 2–4/2–4 on each side at the base of tail.
In life, the juvenile (ZMKU R 00917; SVL 43.1 mm) had a body pattern similar
to the adults but with less prominent tuberculation, brownish yellow ground color of
body, dark body bands are bordered by yellow lines, some bearing tubercles, the origi-
nal tail has approximately 10 dark caudal bands, the posterior portion of tail is white,
and light caudal bands are immaculate (Fig. 11C).
Figure 10. Precloacal region of Cyrtodactylus stellatus sp. nov. showing A precloacal depression with
pore-bearing femoroprecloacal scales in male holotype (ZMKU R 00905), and B pore-bearing precloacal
scales with lacking depression in female paratype (ZMKU R 00900). Red arrows show pore-bearing scales.
A new insular species of the Cyrtodactylus pulchellus group 123
Figure 11. Variation of Cyrtodactylus stellatus sp. nov. A adult female ZMKU R 00899 having 11 dark
caudal bands on the original tail and white caudal bands infused with dark pigmentation B immature
female ZMKU R 00902 (eld number AA 05272) having 12 dark caudal bands on the original tail with
immaculate white caudal bands, and C juvenile ZMKU R 00917 having light-yellow color on the body
and bearing white tail tip.
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124
Distribution. Cyrtodactylus stellatus sp. nov. is currently known only from Tarutao
Island, Satun Province, ailand (Figs 1, 12A).
Natural history. All specimens of C. stellatus sp. nov. were collected from a karst
forest at night (1950–2100 h) with temperatures between 27.1–32.2 °C and relative
humidity between 71.4–93.0%. e specimens were found on karst walls, within karst
crevices and on nearby karst boulders. Some specimens occurred on tree trunks or
Table 6. Descriptive measurement (millimeters), meristic (left/right) and color pattern characters of the
type series of Cyrtodactylus stellatus sp. nov. Key: H = holotype; P = paratype; M = male; F = female; / = data
unavailable or unapplicable; b = broken; r = regenerated. Morphological abbreviations are dened in Table 2.
ZMKU R
00905
ZMKU R
00903
ZMKU R
00906
ZMKU R
00907
ZMKU R
00915
ZMKU R
00899
ZMKU R
00900
ZMKU R
00908
ZMKU R
00909
ZMKU R
00913
HPPPPPPPPP
Sex M M M M M F F F F F
SVL 94.2 95.9 94.6 87.9 86.3 96.1 93.6 94.8 90.4 86.6
TL 94.8r 96.8r 92.3r 69.0r 18.5b 124.4 70.5r 107.5r 81.7r 102.3r
TW 9.2 9.7 9.2 8.4 7.5 7.6 9.1 9.1 10.3 7.1
FL 15.4 15.3 15.1 14.4 14.1 14.9 14.5 15.9 15.6 14.8
TBL 18.1 18.3 17.8 17.4 17.6 17.7 17.3 18.1 17.8 16.5
AG 43.3 43.7 43.9 44.4 39.6 46.0 44.5 44.5 45.4 39.9
HL 27.6 26.3 27.0 25.2 24.8 26.5 26.9 27.3 26.4 25.7
HW 16.7 16.3 17.3 15.4 14.7 16.4 16.8 17.2 15.7 15.2
HD 10.4 9.6 10.8 9.2 8.7 10.4 10.2 10.1 9.4 9.2
ED 6.8 6.7 6.7 5.9 5.8 6.4 6.8 6.6 6.6 5.6
EE 7.1 7.6 7.2 6.3 6.2 6.9 7.0 7.0 6.7 6.8
ES 10.8 10.7 10.9 9.8 9.8 10.9 10.8 10.7 10.4 10.1
EN 8.5 8.3 8.3 7.8 7.5 8.2 8.3 8.5 8.1 7.9
IO 5.9 6.2 6.6 5.8 5.5 6.0 6.4 6.1 5.7 6.2
EL 2.4 2.5 2.2 2.0 2.0 2.0 2.8 2.7 2.1 2.3
IN 2.9 3.2 3.5 3.1 3.0 3.8 3.3 3.2 3.2 2.6
SL 13/14 14/13 15/12 13/14 13/12 13/13 13/13 13/12 12/12 13/13
IL 11/11 10/10 11/12 12/12 11/11 11/11 12/11 12/12 10/11 11/11
PVT 36 35 41 38 43 38 40 40 40 47
LRT 21 20 21 19 22 20 21 19 22 22
VS 33 37 36 35 36 37 39 37 37 36
4TL 21/22 21/21 21/21 21/20 23/22 22/22 21/21 22/23 22/20 20/20
FPP in adult males 27 25 24 29 27 / / / / /
PP in adult females / / / / / 15 12 14 11 11
BB 4444444444
DCB / / / / / 11 / / / /
Body band/ interspace ratio 1.12 1.20 1.04 1.10 1.68 1.07 1.03 1.23 1.06 0.92
Precloacal groove Deep Deep Deep Deep Deep Absent Absent Absent Absent Absent
Femoroprecloacal pores continuous No No No Yes No /////
Tuberculation Weak Weak Weak Weak Weak Weak Weak Weak Weak Weak
Tubercles on ventral surface of
forelimb
No No No No No No No No No No
Tubercles in gular region No No No No No No No No No No
Ventrolateral fold tuberculate No No No No No No No No No No
Dorsum bearing scattered pattern of
white tubercles
Yes Ye s Yes Ye s Yes Ye s Yes Ye s Yes Ye s
Hatchlings/ juveniles with white
tail tip
/ / / / / / / / / /
Adult posterior caudal region white / No ///No /No / /
White caudal bands in adults
immaculate
/No ///No /No /No
A new insular species of the Cyrtodactylus pulchellus group 125
Table 7. Descriptive meristic (left/right) and color pattern characters of the referred specimens of Cyrto-
dactylus stellatus sp. nov. Key: RF = referred specimen; IM-M = immature male; IM-F = immature female;
J = juvenile; / = data unavailable or unapplicable. Morphological abbreviations are dened in Table 2.
ZMKU R
00901
ZMKU R
00902
ZMKU R
00904
ZMKU R
00910
ZMKU R
00911
ZMKU R
00912
ZMKU R
00914
ZMKU R
00916
ZMKU R
00917
RF RF RF RF RF RF RF RF RF
Age IM-M IM-F IM-M IM-M IM-M IM-F IM-F IM-M J
SVL 77.4 68.4 72.5 82.5 81.8 73.8 81.9 79.9 43.1
SL 12/12 13/13 12/12 12/13 14/14 14/14 13/13 13/14 /
IL 10/11 11/12 10/11 11/11 10/12 13/11 10/11 10/11 /
PVT 38 38 32 41 41 41 42 40 /
LRT 20 21 19 22 19 19 22 23 /
VS 34 40 34 37 32 37 38 37 /
4TL 22/21 22/22 21/20 21/21 20/21 23/23 22/21 21/22 /
FFP in adult males / / / / / / / / /
PP in adult females / / / / / / / / /
BB 444444444
DCB 11 12 / 11 / 10 / 11 /
Body band/ interspace ratio 1.35 1.36 1.40 1.09 0.99 1.44 1.22 1.39 /
Precloacal groove / / / / / / / / /
Tuberculation Weak Weak Weak Weak Weak Weak Weak Weak Weak
Tubercles on ventral surface of forelimb No No No No No No No No No
Tubercles in gular region No No No No No No No No No
Ventrolateral fold tuberculate No No No No No No No No No
Dorsum bearing scattered pattern of
white tubercles
Yes Ye s Yes Ye s Yes Ye s Yes Ye s No
Hatchlings/juveniles with white tail tip No No No No No No No No Yes
Adult posterior caudal region white / / / / / / / / /
White caudal bands in adults
immaculate
/ / / / / / / / /
vines near the karst formations (Fig. 12). e holotype was found on a karst wall
approximately 1 m above the ground within karst forest. Eight specimens (ZMKU
R 00900, ZMKU R 00906, ZMKU R 00908, ZMKU R 00911–00912, ZMKU R
00913, and ZMKU R 00915–00916) were found on karst walls from 0.5–3.0 m above
the ground. ZMKU R 00907, ZMKU R 00910, and ZMKU R 00914 were found in
karst crevices. ree specimens (ZMKU R 00901, ZMKU R 00903, and ZMKU R
00909) were found on karst boulders. Four specimens (ZMKU R 00899, ZMKU R
00902, ZMKU R 00904, and ZMKU R 00917) were perched on vegetation near karst
walls or karst boulders.
Two gravid females (ZMKU R 00899–00900) were collected in November 2017
and contained two eggs (externally visible). e juvenile was found on a vine in May
2019. Cyrtodactylus stellatus sp. nov. appears to be nocturnal and sympatric with two
other gekkonids, Gehyra mutilata Wiegmann, 1834 and the diurnal species Cnemaspis
tarutaoensis Ampai et al., 2019.
Etymology. e specic epithet stellatus is Latin word, meaning starry or starred,
and refers to scattered pattern of light-colored tubercles on dorsum and limbs. e
name corresponds with the sister taxon C. astrum that shared similar diagnostic char-
acter (scattered light-colored tubercles pattern on dorsum).
Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
126
Comparison. Cyrtodactylus stellatus sp. nov. can be distinguished from other spe-
cies in the C. pulchellus group by having a combination of weak tuberculation on the
body; no tubercles on ventral surface of forelimbs, gular region, or in ventrolateral
body folds; 19–23 longitudinal tubercle rows; 32–40 ventral scales; 20–23 subdigital
lamellae on the fourth toe; 24–29 femorprecloacal pores in males; deep precloacal
Figure 12. Habitat of Cyrtodactylus stellatus sp. nov. at the type locality, Tarutao Island, Satun Province,
ailand. A Pha Toe Boo karst formation B karst microhabitat structure and C vegetation (vine) used by
a juvenile in karst habitat.
A new insular species of the Cyrtodactylus pulchellus group 127
Table 8. Diagnostic characters of Cyrtodactylus stellatus sp. nov. and its related species within the C. pulchellus group. W = weak; P = prominent; / = data unavailable.
Some information was collected from the following literature (Grismer et al. 2012, 2014, 2016; Quah et al. 2019, Wood et al. 2020, and Termprayoon et al. 2021).
Clade A Clade B
stellatus sp. nov.
astrum
dayangbuntingensis
langkawiensis
lekaguli
australotitiwangsaensis
bintangrendah
bintangtinggi
evanquahi
hidupselamanya
jelawangensis
lenggongensis
macrotuberculatus
pulchellus
sharkari
timur
trilatofasciatus
Sample size 10 13 3 10 16 12 6 14 3 14 4 4 39 13 1 5 6
Maximum SVL 96.1 108.3 99.0 99.8 108.3 120.1 114.4 111.1 96.0 102.7 119.8 103.1 117.9 114.1 100.1 120.5 122.2
SL 12–15 10–12 12–14 9–12 10–12 9–12 8–12 9–13 9 or 10 9–11 10–12 10 or 11 9–12 11 10–12 9–12 9–13
IL 10–13 9–12 10–11 8–10 9–11 9–13 8 or 10 8–11 9 or 10 8–10 8–10 8–10 9–11 10 8–11 8–11 7–11
PVT 32–47 40–57 35–36 34–44 30–50 37–45 36–44 31–42 31–34 33–43 38–43 36–41 36–42 31 34–38 39–48 34–49
LRT 19–23 20–29 20–22 21–25 20–25 22–30 22–25 21–26 18–23 19–24 23–25 22–25 19–27 22–26 24 21–24 23–27
VS 32–40 31–46 36–39 38–43 30–43 32–40 31–39 36–40 29–33 26–33 31–36 32 or 33 17–28 29–34 41 31–40 33–36
4TL 20–23 20–24 21–23 19–21 20–25 21–25 21–24 21–24 22–23 19–24 21–24 20–23 19–23 21–26 24 21–25 22–27
FPP in adult males 24–29 28–38 26–29 30 30–40 39–45 41–46 37–41 32–36 17–22 36 39–41 28–42 33–39 46 21 or 22 41–46
PP in adult females 11–15 Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent Absent
No. of body bands 4 4 4 4 or 5 4 or 5 3(1) or 4 4 3(1) or 4 6 or 7 4 4 4 or 5 3–4 4 4 4 3
Body band/ interspace ratio 0.92–
1.68
0.98–
2.07
0.75 0.75–
1.00
0.86–
2.00
1.00–
2.00
1.00–
1.25
1.00–
1.25
0.82–
1.10
1.00–
1.25
1.00–
1.50
0.50–
1.25
0.95–
1.74
0.75–
1.25
1.75 1.00–
1.25
2.00–
2.75
DCB 10–12 13 or 14 >7 11–16 12–14 7–8 8 or 9 8–10 9–11 8–10 10 14 7–10 8–10 7 8–10 6–7
Precloacal groove in males Deep Deep Deep Deep Deep Deep Deep Deep Shallow Deep Deep Deep Deep Deep Shallow Deep Deep
Femoroprecloacal pores continuous Both Ye s Yes Ye s Yes Ye s Yes Ye s Yes Ye s Yes Ye s Yes Ye s Yes Ye s Yes
Tuberculation W W W W W P P P P W P W P P W W P
Tubercles on ventral surface of forelimb No No No No No No No No No No Yes No Ye s No No No No
Tubercles in gular region No No No No No No No No No No No No Yes No No No No
Ventrolateral fold tuberculate No No No No No No Yes No No No No No Ye s No No No No
Dorsum bearing scattered pattern of white
tubercles
Yes Ye s Yes No No No No No No No No No No No No No No
Hatchlings/juveniles with white tail tip Yes Yes Yes Yes Yes No No No Yes Yes Yes /No No /No No
Adult posterior caudal region white No No No No No No No No Yes Yes No No No No No No No
White caudal bands in adults immaculate No No No No No Yes Ye s Yes No Ye s No Yes No Ye s Yes Ye s Yes
Korkhwan Termprayoon et al. / ZooKeys 1070: 101–134 (2021)
128
groove in males; 11–15 precloacal pores in females; scattered pattern of white, cream
or light-yellow tubercles on dorsum; 10–12 dark caudal bands on original tail; white
caudal bands on original tail infused with dark pigmentation in adults; and juveniles
with white tail tip. Additional comparisons between C. stellatus sp. nov. and other spe-
cies in the C. pulchellus group are in Table 8.
Based on phylogenetic tree, C. stellatus sp. nov. is embedded in Clade A along with
C. astrum, C. dayangbuntingensis, C. langkawiensis, and C. lekaguli. It can be distin-
guished from all four species by having smaller maximum SVL of 96.1 mm (vs. 108.3
mm in C. astrum, 99.0 mm in C. dayangbuntingensis, 99.8 mm in C. langkawiensis, and
108.3 in C. lekaguli); 24–29 femoroprecloacal pores in males (vs. 28–38 in C. astrum,
30 in C. langkawiensis, and 30–40 in C. lekaguli); 11–15 precloacal pores in females (vs.
absent in C. astrum, C. dayangbuntingensis, C. langkawiensis, and C. lekaguli); scattered
pattern of white, cream or light-yellow tubercles on dorsum (vs. absent in C. langkawien-
sis, and C. lekaguli); the ratio of dark body bands to the light color interspaces 0.92–1.68
(vs. 0.75 in C. dayangbuntingensis); 10–12 dark caudal bands (vs. 13 or 14 in C. astrum).
Discussion
e discovery of C. stellatus sp. nov. brings the total number of species in the C. pulchellus
group to 17, of which four have been reported from ailand. is new species is only
known from karst habitats on Tarutao Island and seems to have a narrow geographic
distribution (endemic to Tarutao Island). Molecular analyses recovered it as the sister
taxon to C. astrum and is closely related to C. dayangbuntingensis, C. langkawiensis, and
C. lekaguli. Although C. stellatus sp. nov. showed a similar morphological pattern to its
sister species, morphological analyses and comparisons of meristic characters revealed
that this new species is clearly dierent from its congeners species of Cyrtodactylus.
Among Cyrtodactylus, most useful diagnostic characters are associated with the femoral
and precloacal pores (Harvey et al. 2015). ese characters are easily detected in males,
but those in females are supercial and only found in some species (e.g., C. marmoratus
Gray, 1831; C. psarops Harvey et al., 2015; C. sworderi Smith, 1925). We found dif-
ferences in pore-bearing scales between C. stellatus sp. nov. and other species in the C.
pulchellus group, that proved to be useful in distinguishing among species. Members
of the C. pulchellus group mostly possess a continuous series of enlarged, pore-bearing
femoroprecloacal scales in males, but C. stellatus sp. nov. presents a discontinuous row
of femoroprecloacal pores except one individual (ZMKU R 00907), which has a con-
tinuous series. Moreover, the presence of precloacal pores were found in females of C.
stellatus sp. nov., which has not been reported in the other species (Grismer et al. 2012;
Quah et al. 2019; Wood et al. 2020, Termprayoon et al. 2021).
In addition, we found that the reported sampling localities of C. lekaguli (ZMKU
R 00720–00723) were incorrectly stated as “ailand, Changwat Province, Takua Pa
District, Phangnga” in previous studies (i.e., Grismer et al. 2016; Quah et al. 2019;
A new insular species of the Cyrtodactylus pulchellus group 129
Wood et al. 2020, Termprayoon et al. 2021). erefore, we corrected the sampling lo-
calities to “ailand, Phang-nga Province, Mueang Phang-nga District” (see Table 1).
e discovery of this new species suggests that undiscovered species of the C. pulchel-
lus group may still occur in southern ailand where there are still numerous unexplored
karst areas. Additional surveys are needed to determine the extent of the geographic
range of C. stellatus sp. nov. and the C. pulchellus group in as a whole in the region.
Acknowledgments
is research was supported by the Center of Excellence on Biodiversity (BDC), Of-
ce of Higher Education Commission (BDC-PG4-160022). Research protocol was
approved by the Institutional Animal Care and Use Committee of Faculty of Science,
Kasetsart University (project number ACKU61-SCI-006). KT was supported by a
grant from the Faculty of Science, Kasetsart University (50th Anniversary of Faculty
of Science). AR and AA were supported by Kasetsart University Research and Devel-
opment Institute (KURDI), the Department of Zoology, and International SciKU
Branding (ISB), Faculty of Science, Kasetsart University. NA was supported by Sri-
nakharinwirot University Research Grant. e Department of National Parks, Wild-
life and Plant Conservation, ailand for providing permission to conduct the re-
search in Tarutao National Park and Khao Banthat Wildlife Sanctuary. Kanchanapan
Kamhang (Tarutao National Park) and Bamrungrat Ploydam (Khao Banthat Wild-
life Sanctuary) for facilitating the eldwork. L. Lee Grismer (La Sierra University),
Wachara Sanguansombat and Sunchai Makchai (ailand Natural History Museum)
made specimens in their care available for study. Siriporn Yodthong assisted with
eldwork. is paper is contribution number 950 of the Auburn University Museum
of Natural History. We thank L. Lee Grismer and Evan S.H. Quah for providing their
suggestions, which greatly improved the manuscript.
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Appendix I
Specimens examined
Cyrtodactylus astrum: Peninsular Malaysia, Perlis, Gua Wang Burma: LSUHC 09928
(female) and LSUHC 100075 (male).
Cyrtodactylus lekaguli: ailand, Trang Province, Na Yong District: ZMKU R 00918,
THNHM 017781, THNHM 017784, THNHM 017787, THNHM 017791 (5
males), and ZMKU R 00919, ZMKU R 00921, THNHM 017694, THNHM
017777, THNHM 017788, THNHM 017790 (6 females).
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