Six new species of the Cyrtodactylus intermedius complex (Squamata: Gekkonidae) from the Cardamom Mountains and associated highlands of Southeast Asia

Abstract

An integrative taxonomic analysis using color pattern, morphology, and 1449 base pairs of the ND2 mitochondrial gene and its five flanking tRNAs demonstrated that eight species-level lineages occur within the Cyrtodactylus intermedius complex (Cyrtodactylus intermedius sensu stricto, C. phuquocensis and related populations) of the Cardamom mountains and associated highlands that have a sequence divergence ranging 3.4–8.9%. Additionally, each lineage is discretely diagnosable from one another based on morphology and color pattern and most occur in specific geographic regions (upland areas, karst formations or islands) that prevent or greatly restrict interpopulation gene flow. Six of these lineages were masquerading under the nomen C. intermedius and are described as the following: Cyrtodactylus auralensis sp. nov. endemic to Phnom Aural, the highest mountain in Cambodia; C. bokorensis sp. nov. endemic to the Bokor Plateau, Cambodia; C. cardamomensis sp. nov. from the main block of the Cardamom mountains; C. thylacodactylus sp. nov. endemic to Phnom Dalai the northernmost peak of the Cardamom mountains; C. laangensis sp. nov. endemic to the Phnom Laang karst formation, Cambodia; and C. septimontium sp. nov. from the Bảy Núi Hills of southwest Vietnam.

Full text

Accepted by M. Heinicke: 16 Nov. 2018; published: 8 Feb. 2019
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN
1175-5334
(online edition)
Copyright © 2019 Magnolia Press
Zootaxa 4554 (1): 001
–
062
https://www.mapress.com/j/zt/
Article
1
https://doi.org/10.11646/zootaxa.4554.1.1
http://zoobank.org/urn:lsid:zoobank.org:pub:4C844226-7C0D-4B16-B87C-9044B8BAA1D0
Six new species of the Cyrtodactylus intermedius complex (Squamata: Gekkonidae)
from the Cardamom Mountains and associated highlands of Southeast Asia
MATTHEW L. MURDOCH
1,*
, L. LEE GRISMER
2
, PERRY L. WOOD JR.
3
, THY NEANG
4
, NIKOLAY A.
POYARKOV
5
, NGO VAN TRI
6
, ROMAN A. NAZAROV
7
, ANCHALEE AOWPHOL
8
, OLIVIER S.G.
PAUWELS
9
, HUNG NGOC NGUYEN
10
& JESSE L. GRISMER
11
1
Department of Biology, Villanova University, 800 Lancaster Ave, Villanova, PA, USA
2
Herpetology Laboratory, Department of Biology, La Sierra University, 4500 Riverwalk Parkway, Riverside, California 92515 USA
3
Department of Ecology and Evolutionary Biology and Biodiversity Institute, University of Kansas, Dyche Hall, 1345 Jayhawk Blvd,
Lawrence, KS 66045-7561, USA
4
Wild Earth Allies, 77a, St. Betong, Bayap Village, Sk. Phnom Penh Thmei, Kh. Sen Sok, Phnom Penh, CAMBODIA
5
Department of Vertebrate Zoology, Biological Faculty, Lomonosov, Moscow State University, Leninskiye Gory, Moscow GSP-1,
119234, RUSSIA
Joint Russian-Vietnamese Tropical Research and Technological Center, 63 Nguyen Van Huyen Road, Nghia Do, Cau Giay, Hanoi,
VIETNAM
6
Department of Environmental Management and Technology, Institute of Tropical Biology, Vietnam Academy of Science and Technol-
ogy, Ho Chi Minh, Vietnam, 9/621 Hanoi Highway, Thu Duc Distrcit, Hochiminh City, VIETNAM
7
Zoological Museum, Moscow State University, B. Nikitskaya ul. 2, Moscow 125009, RUSSIA
8
Department of Zoology, Faculty of Science, Kasetsart University, Chatuchak, Bangkok, THAILAND
9
Département des Vertébrés Récents, Institut Royal des Sciences Naturelles de Belgique, 29 Rue Vautier, B-1000 Brussels, BELGIUM
10
Biodiversity Program, Taiwan International Graduate Program, Academia Sinica, Taipei, TAIWAN
Department of Zoology, Southern Institute of Ecology, Vietnam Academia of Science and Technology, Ho Chi Minh City, VIETNAM
Department of Life Science, National Taiwan Normal University, Taipei, TAIWAN
11
Department of Biological Sciences 331 Funchess Hall, Auburn University Auburn, Alabama 36849, USA
La Kretz Center for Californian Conservation Science, Institute of the Environment and Sustainability, La Kretz Hall, University of
California Los Angeles, Los Angeles, CA 90095, Suite 300, Box 951496. USA
*
Corresponding author. E-mail: mmurdoch@villanova.edu
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
Materials and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
Taxonomy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Cyrtodactylus intermedius Smith . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Cyrtodactylus auralensis sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Cyrtodactylus bokorensis sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
Cyrtodactylus cardamomensis sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32
Cyrtodactylus thylacodactylus sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .38
Cyrtodactylus laangensis sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .42
Cyrtodactylus septimontium sp. nov. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .46
Cyrtodactylus phuquocensis Ngo, Grismer, & Grismer 2010 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .54
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Key to the species of the Cyrtodactylus intermedius complex . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 58

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Abstract
An integrative taxonomic analysis using color pattern, morphology, and 1449 base pairs of the ND2 mitochondrial gene
and its five flanking tRNAs demonstrated that eight species-level lineages occur within the Cyrtodactylus intermedius
complex (Cyrtodactylus intermedius sensu stricto, C. phuquocensis and related populations) of the Cardamom mountains
and associated highlands that have a sequence divergence ranging 3.4–8.9%. Additionally, each lineage is discretely di-
agnosable from one another based on morphology and color pattern and most occur in specific geographic regions (upland
areas, karst formations or islands) that prevent or greatly restrict interpopulation gene flow. Six of these lineages were mas-
querading under the nomen C. intermedius and are described as the following: Cyrtodactylus auralensis sp. nov. endemic
to Phnom Aural, the highest mountain in Cambodia; C. bokorensis sp. nov. endemic to the Bokor Plateau, Cambodia; C.
cardamomensis sp. nov. from the main block of the Cardamom mountains; C. thylacodactylus sp. nov. endemic to Phnom
Dalai the northernmost peak of the Cardamom mountains; C. laangensis sp. nov. endemic to the Phnom Laang karst for-
mation, Cambodia; and C. septimontium sp. nov. from the Bảy Núi Hills of southwest Vietnam.
Key words: Cyrtodactylus, systematics, taxonomy, new species, Cardamom Mountains, Cambodia, Thailand, Vietnam
Introduction
Of the world’s gekkotan lineages, the genus Cyrtodactylus Gray is by far the most speciose with new species being
described at a prolific rate from across its range (e. g., Agarwal et al. 2016; Grismer et al. 2018a; Luu et al. 2016;
Nazarov et al. 2014; Oliver et al. 2016; Riyanto et al. 2015). Most of these descriptions resulted from field work in
previously unsurveyed areas throughout Southeast Asia and as such, the species having been described are known
only from a few specimens from their type localities. However, the description C. macrotuberculatus Grismer &
Norhayati (2008c) from an insular population in Malaysia previously considered to be C. pulchellus (Gray), started
a chain reaction demonstrating that some widely distributed species of Cyrtodactylus may represent species
complexes containing as many as 14 species (e.g. Grismer et al. 2012, 2014; Sumontha et al. 2012). As these taxa
are put under scrutiny the hidden diversity within Cyrtodactylus continues to be revealed (e.g. Grismer et al. 2012,
2014, 2016; Oliver et al. 2016; Welton et al. 2010). With the description of C. phuquocensis Ngo, Grismer &
Grismer (2010) it was first suggested that Cyrtodactylus intermedius Smith is a complex that consists of more than
one species. Grismer et al. (2015a) went on to propose that the variation in color pattern, morphology, and genetic
data in the remaining populations of C. intermedius sensu auctorum provided additional evidence that this species
was still a collection of multiple, yet undescribed lineages.
Historically years of civil conflict and security concerns stalled investigation into Cambodia’s biodiversity.
However, with the political reconciliation and integration of the late 1990’s there has been a resurgence in
herpetological field research in Cambodia with a special interest being taken in the Cardamom Mountains (Daltry
& Chheang 2000; Swan & Daltry 2002; Daltry & Traeholt 2003; Emmet & Olsson 2005; Stuart & Emmet 2006;
Grismer et al. 2007a; 2008a, b; Neang et al. 2010). As it is currently understood, Cyrtodactylus intermedius is a
forest-dwelling and scansorial species. Despite being commonly known as the Cardamom Bent-toed Gecko it has a
wide distribution, ranging from the southern boundary of the Khorat Plateau of northeastern Thailand through the
mountains of southeastern Thailand, eastward through the hills, plateaus and islands of southern Cambodia, and
into the Bảy Núi hills of the Mekong Delta in southern Vietnam (Fig. 1). However, morphologically distinct
populations have been noted from isolated upland areas, karst formations, and islands (Grismer et al. 2015a) where
gene flow between populations is currently highly unlikely. Although mitochondrial DNA phylogenies have
proven to be extremely valuable in delimiting species boundaries and identifying cryptic species, highly divergent
haplotypes can persist within populations due to incomplete lineage sorting and introgression, resulting in
phylogenies that are inconsistent with species histories (McGuire et al. 2007). Therefore, the addition of non-
recombinant data sources (morphology) to taxonomic analyses has proven to be highly informative in
disentangling the lineages of species complexes (e.g. Malhotra et al. 2011). Therefore, we employed an integrative
taxonomic approach (i.e. Padial et al. 2010) to test for, and to delimit species boundaries within the C. intermedius
complex (C. intermedius sensu lato and C. phuquocensis) in order to provide an evolutionary based taxonomy. The
results indicated that the geographic variation in color pattern and morphology between the populations of C.
intermedius was correlated with the existence of eight exclusive lineages bearing sequence divergences from one
another ranging from 3.7%–8.9% and each occurring within a specific geographic region. Six of these lineages are
described herein as new species.

Zootaxa 4554 (1) © 2019 Magnolia Press
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3
SIX NEW CY RTODACTYLUS FROM SOUTHEAST ASIA
FIGURE 1. Map illustrating the locations where members of the Cyrtodactylus intermedius complex have been recorded.
Thailand: 1= Sakaerat Biosphere Reserve; 2 = Sa Kaeo; 3 = Khao Soi Dao; 4 = Khao Sebab. Cambodia: 5 = Phnom Dalai; 6 =
Phnom Samkos; 7 = O’Som Village; 8 = Thmar Baing; 9 = Koh Rong Island; 10 = Phnom Aural; 11 = Kirirom National Park;
12 = Bokor Plateau; 13 = Phnom Laang. Vietnam: 14 = Phu Quoc Island; 15 = Nui Co To; 16 = Nui Cam.
Materials and methods
Species delimitation. The general lineage concept (GLC: De Queiroz 2007) used herein proposes that a species
constitutes a population of organisms independently evolving from other such populations owing to a lack of gene

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flow. By “independently”, it is meant that new mutations arising in one species cannot spread readily into another
species (Barraclough et al. 2003; De Queiroz 2007). Increasingly, integrative studies on the nature and origins of
species are using a wider range of empirical data to delimit species boundaries (Coyne & Orr 1998; Knowles &
Carstens 2007; Fontaneto et al. 2007; Feulner et al. 2007; Leach et al. 2009), rather than relying solely on
traditional taxonomic methods. Under the GLC herein, molecular phylogenies were used to infer species
boundaries and univariate (ANOVA) and multivariate (PCA) morphological analyses were used to describe those
boundaries. These boundaries were cross-checked using a Generalized Mixed Yule Coalescent (GMYC) approach
(Pons et al. 2006), thus providing an independent framework to complement the empirically based thresholds of
the morphological and molecular analyses. The GMYC approach is a method for delimiting species from single-
locus gene trees by detecting genetic clustering beyond the expected levels of a null hypothesis which infers that all
individuals of a population form a genetically interacting nexus. In clades where effective population sizes are
relatively low and divergence times among the populations are relatively high, the single-threshold version of the
model (such as that used herein) outperforms the multi-threshold version (Fujisawa & Barraclough 2013). The
GMYC relies on the prediction that independent evolution leads to the appearance of distinct genetic clusters,
separated by relatively longer internal branches (Barraclough et al. 2003; Acinas et al. 2004). Such groups
therefore diverge into discrete units of morphological and genetic variation that are recovered with surveys of
higher clades. The analysis was run on the web server at http://species.h-its.org/gmyc/.
Data and Analyses. Mitochondrial DNA.— The primary aim of this study was to use 1449 bp of the
mitochondrial gene NADH dehydrogenase subunit 2 (ND2) to evaluate the phylogenetic relationships of the disjunct
populations of Cyrtodactylus intermedius sensu lato that extend approximately 550 km from Sakaerat in eastern
Thailand to the Bảy Núi Mountains in southwestern Vietnam. Data were sampled as widely as possible and included
78 individuals from 15 localities from the extent of their known distribution (Table 1; Fig. 5). Based on the
phylogeny of Wood et al. (2012), C. hontreensis, C. oldhami, C. peguensis zebraicus, and C. tigroides were selected
as closely related outgroup species to root the tree. All sequences have been deposited in GenBank (Table 1).
TABLE 1. Taxon sampling for ingroup and outgroup, locality data, and GenBank accession numbers of the specimens
used in this study.
Voucher Taxon Locality GenBank no.
LSUHC 7346 C. auralensis sp. nov. Phnom Aural, Kampong Speu province, Cambodia KT013127
LSUHC 7396 C. auralensis sp. nov. Phnom Aural, Kampong Speu province, Cambodia KT013128
LSUHC 7398 C. auralensis sp. nov. Phnom Aural, Kampong Speu province, Cambodia KT013129
LSUHC 7399 C. auralensis sp. nov. Phnom Aural, Kampong Speu province, Cambodia KT013130
LSUHC 7400 C. auralensis sp. nov. Phnom Aural, Kampong Speu province, Cambodia KT013131
LSUHC 7401 C. auralensis sp. nov. Phnom Aural, Kampong Speu province, Cambodia KT013132
LSUHC 7409 C. auralensis sp. nov. Phnom Aural, Kampong Speu province, Cambodia KT013133
LSUHC 8541 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013146
LSUHC 8542 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013147
LSUHC 8543 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013148
LSUHC 8545 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013149
LSUHC 8546 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013150
LSUHC 8547 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013151
LSUHC 8548 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013152
LSUHC 8549 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013153
LSUHC 8550 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013154
LSUHC 8552 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013155
LSUHC 8553 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013156
LSUHC 8555 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013157
FMNH 263228 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013107
......continued on the next page

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SIX NEW CY RTODACTYLUS FROM SOUTHEAST ASIA
TABLE 1. (Continued)
Voucher Taxon Locality GenBank no.
FMNH 263229 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013108
FMNH 263230 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013109
FMNH 263232 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013110
FMNH 263233 C. bokorensis sp. nov. Bokor Plateau, Kampot province, Cambodia KT013111
FMNH 263345 C. cardamomensis sp. nov. Thmar Baing, Koh Kong province, Cambodia KT013115
FMNH 263346 C. cardamomensis sp. nov. Thmar Baing, Koh Kong province, Cambodia KT013116
LSUHC 10081 C. cardamomensis sp. nov. O'Som Village, Pursat province, Cambodia KT013118
LSUHC 10083 C. cardamomensis sp. nov. O'Som Village, Pursat province, Cambodia KT013119
LSUHC 10095 C. cardamomensis sp. nov. O'Som Village, Pursat province, Cambodia KT013120
LSUHC 10096 C. cardamomensis sp. nov. O'Som Village, Pursat province, Cambodia KT013121
LSUHC 10121 C. cardamomensis sp. nov. O'Som Village, Pursat province, Cambodia KT013122
LSUHC 7855 C. cardamomensis sp. nov. Phnom Samkos, Pursat province, Cambodia KT013134
LSUHC 7918 C. cardamomensis sp. nov. Phnom Samkos, Pursat province, Cambodia KT013135
LSUHC 7936 C. cardamomensis sp. nov. Phnom Samkos, Pursat province, Cambodia KT013136
LSUHC 7943 C. cardamomensis sp. nov. Phnom Samkos, Pursat province, Cambodia KT013137
LSUHC 9318 C. thylacodactylus sp. nov. Phnom Dalai, Pursat province, Cambodia KT013162
LSUHC 9319 C. thylacodactylus sp. nov. Phnom Dalai, Pursat province, Cambodia KT013163
LSUHC 9325 C. thylacodactylus sp. nov. Phnom Dalai, Pursat province, Cambodia KT013164
LSUHC 9326 C. thylacodactylus sp. nov. Phnom Dalai, Pursat province, Cambodia KT013165
LSUHC 8583 C. hontreensis Hon Tre Island, Kien Giang province, Vietnam JX440538
ZMKU R 0724/
AA02183
C. intermedius Thai clade Sakaerat, Nakhon Ratchasima, Thailand MH940239
ZMKU R 0725/
AA02184
C. intermedius Thai clade Sakaerat, Nakhon Ratchasima, Thailand MH940240
CUMZR 2005 07 30
139
C. intermedius Thai clade Sakaerat, Nakhon Ratchasima, Thailand GU550710
FMNH 265811 C. intermedius Thai clade Sa Kaeo, Thailand KT013117
FMNH 265812 C. intermedius Thai clade Sa Kaeo, Thailand JQ889182
LSUHC 9513 C. intermedius Thai clade Khao Soi Dao, Chanthaburi, Thailand JX519469
LSUHC 10535 C. intermedius complex
incertae sedis 1
Koh Rong Island, Sihanoukville province, Cambodia KT013123
LSUHC 10536 C. intermedius complex
incertae sedis 1
Koh Rong Island, Sihanoukville province, Cambodia KT013124
LSUHC 10562 C. intermedius complex
incertae sedis 1
Koh Rong Island, Sihanoukville province, Cambodia KT013125
LSUHC 10563 C. intermedius complex
incertae sedis 1
Koh Rong Island, Sihanoukville province, Cambodia KT013126
FMNH 263238 C. intermedius complex
incertae sedis 2
Phnom Sruoch, Kirirom National Park, Kampong
Speu, Cambodia
KT03112
FMNH 263239 C. intermedius complex
incertae sedis 2
Phnom Sruoch, Kirirom National Park, Kampong
Speu, Cambodia
KT03113
FMNH 263240 C. intermedius complex
incertae sedis 2
Phnom Sruoch, Kirirom National Park, Kampong
Speu, Cambodia
KT03114
LSUHC 8770 C. laangensis sp. nov. Phnom Laang, Kampot province, Cambodia KT013158
LSUHC 8771 C. laangensis sp. nov. Phnom Laang, Kampot province, Cambodia KT013159
......continued on the next page

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Genomic DNA was isolated from liver or skeletal muscle samples stored in 95% ethanol using the Qiagen
DNeasy
TM
tissue kit (Valencia, CA, USA). ND2 was amplified using a double-stranded Polymerase Chain Reaction
(PCR) under the following conditions: 1.0 µl genomic DNA, 1.0 µl light strand primer 1.0 µl heavy strand primer,
1.0 µl dinucleotide pairs, 2.0 µl 5x buffer, MgCl 10x buffer, 0.1 µl Taq polymerase, and 7.56 µl ultra-pure H
2
O
(Table 2). PCR reactions were executed on an Eppendorf Mastercycler gradient thermocycler 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
10% agarose gel electrophoresis. Successful PCR products were vacuum purified using MANU 30 PCR plates
(Millipore) and purified products were resuspended in ultra-pure water. Purified PCR products were sequenced
using the ABI Big-Dye Terminator v3.1 Cycle Sequencing Kit in an ABI GeneAmp PCR 9700 thermal cycler.
Cycle sequencing reactions were purified with Sephadex G-50 Fine (GE Healthcare) and sequenced on an ABI
3730xl DNA Analyzer at the BYU DNA Sequencing center. Primers used for amplification and sequencing are
presented in Table 2. Sequences were analyzed from both the 3' and the 5' ends separately to confirm congruence
between the reads. Both the forward and the reverse sequences were uploaded and edited in Geneious
TM
version
v5.5.6 (Drummond et al. 2011) and were edited therein. The protein-coding region of the ND2 sequence was
aligned by eye. MacClade v4.08 (Maddison & Maddison 2005) was used to calculate the correct amino acid
reading frame and to confirm the lack of premature stop codons.
Phylogenetic analyses.— The phylogenetic analysis applied two model-based methods, Maximum Likelihood
(ML) and a time-calibrated Bayesian Evolutionary Analysis Sampling Trees (BEAST). The ML analysis was
performed using IQ-TREE (Nguyen et al. 2015) with 1000 bootstrap pseudoreplicates using the ultrafast bootstrap
TABLE 1. (Continued)
Voucher Taxon Locality GenBank no.
LSUHC 8772 C. laangensis sp. nov. Phnom Laang, Kampot province, Cambodia KT013160
LSUHC 8773 C. laangensis sp. nov. Phnom Laang, Kampot province, Cambodia KT013161
CUMZ
R2005.07.30.54
C. peguensis Khao Luang National Park, Thailand GU550727
LSUHC 8603 C. phuquocensis Phu Quoc Island, Kien Giang province, Vietnam KT013182
LSUHC 8688 C. phuquocensis Phu Quoc Island, Kien Giang province, Vietnam KT013183
LSUHC 9486 C. oldhami Chumpon province, Thailand MH940241
K79432C C. septimontium sp. nov. Nui Cam Mountain, An Giang province, Vietnam MH940226
ITBCZ 3026 C. septimontium sp. nov. Nui Cam Mountain, An Giang province, Vietnam MH940231
ITBCZ 3027 C. septimontium sp. nov. Nui Cam Mountain, An Giang province, Vietnam MH940225
ZMMU NAP 05272 C. septimontium sp. nov. Nui Cam Mountain, An Giang province, Vietnam MH940223
ZMMU NAP 05273 C. septimontium sp. nov. Nui Cam Mountain, An Giang province, Vietnam MH940227
ZMMU NAP 05274 C. septimontium sp. nov. Nui Cam Mountain, An Giang province, Vietnam MH940228
ZMMU NAP 05275 C. septimontium sp. nov. Nui Cam Mountain, An Giang province, Vietnam MH940229
ZMMU NAP 05276 C. septimontium sp. nov. Nui Cam Mountain, An Giang province, Vietnam MH940231
ZMMU NAP 05277 C. septimontium sp. nov. Nui Cam Mountain, An Giang province, Vietnam MH940224
K79432F C. septimontium sp. nov. Nui Co To Mountain, An Giang province, Vietnam MH940232
ZMMU NAP 05320 C. septimontium sp. nov. Nui Co To Mountain, An Giang province, Vietnam MH940234
ZMMU NAP 05321 C. septimontium sp. nov. Nui Co To Mountain, An Giang province, Vietnam MH940237
ZMMU NAP 05322 C. septimontium sp. nov. Nui Co To Mountain, An Giang province, Vietnam MH940233
ZMMU NAP 05323 C. septimontium sp. nov. Nui Co To Mountain, An Giang province, Vietnam MH940235
ZMMU NAP 05324 C. septimontium sp. nov. Nui Co To Mountain, An Giang province, Vietnam MH940236
ZMMU NAP 05325 C. septimontium sp. nov. Nui Co To Mountain, An Giang province, Vietnam MH940238
IRSNB 2380 C. tigroides Ban Tha, Sai-Yok district, Kanchanaburi province,
Thailand
JX440562

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approximation algorithm (Minh et al. 2013). The data set was partitioned by codon and the Bayesian Information
Criterion (BIC) implemented in IQ-TREE (Nguyen et al. 2015) indicated that K3Pu+G4 was the best-fit model of
evolution for position 1, HKY+G4 was the best-fit model for positions 2 and 3, and K3P+G4 was the appropriate
model for the tRNAs. Nodes having ultrafast ML bootstrap values (UBS) of 95 and above were considered well-
supported (Nguyen et al. 2015). Uncorrected pairwise sequence divergences were calculated in MEGA7.0 (Kumar
et al. 2016).
TABLE 2. Primers used for amplification and sequencing reactions for the ND2 gene and the flanking tRNA's
The time-calibrated Bayesian Inference analysis was implemented in the program Bayesian Evolutionary
Analysis Utility (BEAUti) version 2.4.7 and run on BEAST2 version 2.4.7 (Bouckaert et al. 2014). A previous run
of the data using a relaxed clock model indicated there was very little rate variation among the sites and so a strict
clock model was used for the final analysis employing unlinked site and linked tree models. A substitution model
of HKY+G4 was set to approximate the models of evolution used in the ML analysis with empirical frequencies
estimated so as to fix them to the proportions observed in the data. A coalescent exponential population prior was
employed because intraspecific relationships among many individuals were being assessed and it was not known a
priori which tips (individuals) would be grouped as species. Under the coalescent model, the default priors for
population growth (Laplace Distribution) and size (1/X) were left unchanged because these parameters were not
being estimated. The MCMC was run for 80 million generations and logged every 80000 generations. Fossil
calibrations from Wood et al. (2012) placed a mean date of 25.00 million years (95% HPD 18.42–32.03; (Wood.
unpuble. data)) on the node between the Cyrtodactylus intermedius and its sister species C. hontreensis and this
was used as a constraint prior to date the same node with a standard deviation of 4.2% and assuming a normal
distribution. Maximum clade credibility trees using mean heights at the nodes were generated using TreeAnnotator
v.1.8.0 (Rambaut & Drummond 2013) with a burnin of 1000 trees. The BEAST2 log files were visualized and
checked in Tracer 1.6.0 (Rambaut et al. 2014) to ensure effective sample size (ESS) values were above 200 for all
parameters. Nodes having Bayesian posterior probabilities (BPP) of 0.95 and above were considered well-
supported (Huelsenbeck et al. 2001; Wilcox et al. 2002). We note that it has been demonstrated that the third codon
position is susceptible to substitution saturation (Grismer et al. 2015b) and could contribute to overestimating node
ages. However, Grismer et al. (2015b) noted that although third codon position saturation was evident in their
study of the gekkonid genus Hemiphyllodactylus, it was not a significant factor in estimating node ages across
various codon and gene partition schemes and was consistent with similar node age estimates using nuclear genes
on the same taxa (Heinicke et al. 2011).
Morphological analyses.—Color notes were taken from living specimens and digital images of living
specimens of all possible age classes prior to preservation. Measurements were taken on the left side of the body
when possible to the nearest 0.1 mm by MLM using GlowGeek dial calipers under Leica ZOOM 2000 dissecting
microscope. Measurements taken 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, regenerated, or broken; tail width (TW), taken at the base of
the tail immediately posterior to the postcloacal swelling; forearm length (FL), taken on the ventral surface from
the posterior margin of the elbow while flexed 90 degrees 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 degrees 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 measured 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 orbit; eye to snout distance (ES), measured from anteriormost margin of
Primer name Primer reference Location Sequence
L4437b Macey & Schulte (1999) External 5’-AAGCAGTTGGGCCCATACC-3’
CyrtintF1 Siler et al. (2010) Internal 5’-TAGCCYTCTCYTCYATYGCCC-3’
CyrtintR1 Siler et al. (2010) Internal 5’-ATTGTKAGDGTRGCYAGGSTKGG-3’
H5934 Macey & Schulte (1999) External 5’- AGRGTGCCAATGTCTTTGTGRTT-3’

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the orbit to the tip of snout; eye to nostril distance (EN), measured from the anterior margin of the orbit to the
posterior margin of the external nares; inter orbital distance (IO), measured between the dorsal-most edges of the
orbit; and internarial distance (IN), measured between the nares across the rostrum.
FIGURE 2. Underside of right hand of (A) and right hind foot (C) of Cyrtodactylus thylacodactylus sp. nov. illustrating the
presence of interdigital pocketing in the webbing and underside of right hand (B) and right hind foot (D) of C. cardamomensis
sp. nov. illustrating the standard lack of interdigital webbing and pocketing.
Meristic characters evaluated were the numbers of supralabial scales (SL) counted from the largest scale
immediately below the middle of the eyeball to the rostral scale; infralabial scales (IL) counted from the mental to
the termination of enlarged scales just after the upturn of the mouth; the number of paravertebral tubercles (PVT)
between limb insertions counted in a straight line immediately left of the vertebral column; the number of
longitudinal rows of body tubercles (LRT) counted transversely across the center of the dorsum from one
ventrolateral fold to the other; the number of longitudinal rows of ventral scales (VS) counted transversely across
the center of the abdomen from one ventrolateral fold to the other; the number of expanded subdigital lamellae
proximal to the digital inflection on the fourth toe (TLE) counted from the base of the first phalanx where it
contacts the body of the foot to the largest scale on the digital inflection, large continuous scales on the palmar and
plantar surfaces were not counted; the number of small, generally unmodified subdigital lamellae distal to the
digital inflection on the fourth toe (TLU) counted from the digital inflection to the claw including the claw sheath
(Fig. 2); the total number of subdigital lamellae (TTL) beneath the fourth toe (i.e. TLE + TLU = TTL); and the total
number of enlarged femoral scales (FS) from each thigh combined as a single metric. In some species, only the
distalmost FS are greatly enlarged and the proximal scales are smaller whereas in others, the enlarged scales are
continuous with the enlarged precloacal scales and the separation of the two scales rows was determined to be at a

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point even with the lateral body margin (Fig. 3). The number of enlarged precloacal scales (PS); the number of
precloacal pores (PP) in males; the number of rows of post-precloacal scales (PPS) on the midline between the
enlarged precloacal scales and the vent; the number of postcloacal tubercles (PCT) on either side of the base of the
tail; and the number of dark body bands (BB) between the nuchal loop (dark band running from eye to eye) and the
hind limb insertions.
Non-meristic morphological characters evaluated were if enlarged femoral scales and precloacal scales were
separated by a diastema at the base of the femora or contiguous and if so if the proximal femoral scales were less
than half the size of those distal (Fig. 3); and the presence or absence of a pocket of skin webbing between the third
and fourth digit of the hind foot and two pockets of skin webbing on either side of the third digit of the forefoot
(Fig. 2).
FIGURE 3. Differences in, and arrangement of femoral and precloacal scale morphology. (A) Cyrtodactylus cardamomensis
sp. nov. illustrating contiguous femorals where those proximal are the same size as those distal, (B) C. auralensis sp. nov.
illustrating contiguous femorals where those proximal are less than half the size of those distal, (C) C. laangensis sp. nov.
illustrating variant where femorals not contiguous with precloacals, (D) C. laangensis sp. nov. illustrating variant with total
lack of enlarged femoral scales.
Color pattern characters evaluated were the posterior border of the nuchal loop being evenly rounded or
tapering to a chevron shaped point (Fig. 4); and the presence or absence of dark pigmented blotches on the top of
the head (Fig. 4).
Statistical analyses.—An analysis of variance (ANOVA) was performed to ascertain if statistically significant
mean differences among meristic characters (p<0.05) existed among the discrete populations delimited in the
phylogenetic analyses. ANOVAs having a p-value less than 0.05 indicating that statistical differences existed were
subjected to a Tukey HSD test to ascertain which population pairs differed significantly (p<0.05) from each other.
Principal Component Analysis (PCA) and Discriminant Analysis of Principal Components (DAPC) were used
to determine if populations from different localities occupied unique positions in morphospace and the degree to
which their variation in morphospace coincided with potential species boundaries predicted by the molecular
phylogenetic and univariate analyses. PCA, implemented by the prcomp command in R v 3.2.1 (R Core Team,
2015), searches for the best overall low-dimensional representation of significant morphological variation in the
data. Precloacal pore counts were excluded from the meristic PCA due to their presence only in males and tail
width was excluded due to variation between sexes. Hatchlings and juveniles were not included in the PCA and
only adults available to the principle investigator (MLM) were included to further standardize the data. Characters
used in the PCAs and DAPCs were discrete meristic data from the scale counts SL, IL, PVT, LRT, VS, TLU, TLE,

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TTL, FS, PS, PPS, PCT, and BB and the continuous mensural data from FL, TBL, AG, HL, HW, HD, ED, EE, ES,
EN, IO, and IN. Grismer et al. (2017a) demonstrated that concatenating meristic and mensural data produced the
most informative results in separating populations in morphospace. To remove allometric effects due to body size,
each character was adjusted using the following equation: Xadj=X-β(SVL-SVLmean), where Xadj=adjusted value;
X=measured value; β=unstandardized regression coefficient for each OTU; SVL=measured snout-vent-length;
SVLmean=overall average SVL of all OTU’s (Lleonart et al. 2000; Thorpe, 1975, 1983; Turan, 1999). All PCA
data were natural log-transformed prior to analysis and scaled to their standard deviation 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 transform meristic and mensural data into comparable units for analysis. To
characterize clustering and separation in morphospace, a DAPC was performed to search for linear combinations of
morphological variables having the greatest between-group variance and the smallest within-group variance
(Jombart et al., 2010). DAPC relies on log transformed data from the PCA as a prior step to ensure that variables
analyzed are not correlated and number fewer than the sample size. Principal components with eigenvalues of 1.0
or greater were retained for the DAPC according to the criterion of Kaiser (1960). All statistical analyses were
performed using the platform R v 3.2.1 (R Core Team, 2015).
FIGURE 4. Presence of pigmented blotches on top of head and rounded posterior border of nuchal loop (A) Cyrtodactylus
auralensis sp. nov. and (B) C. bokorensis sp. nov. illustrating lack of pigmented blotches on top of head and a pointed chevron
shaped posterior border of the nuchal loop.
Results
The ML and BEAST analyses recovered trees bearing identical topologies (Fig. 5). All nodes in the BEAST tree
were strongly supported (BPP ≥ 0.95; Fig. 5) and all but five of the relatively deeper nodes in the ML were strongly
supported (UBS ≥ 95; Fig. 5). The phylogeny reveals a high degree of phylogeographic substructuring within
Cyrtodactylus intermedius sensu lato across its range that is closely associated with topographic features (Figs.
1,5). The phylogeny reveals C. intermedius is composed of two major groups: a western group containing the Thai,
Phnom Dalai, Phnom Samkos, O’Som, Thmar Baing, Phnom Aural, and Koh Rong Island populations and an
eastern group containing the Bokor Plateau, Kirirom, Phnom Laang, Phu Quoc Island, and Bảy Núi hills
populations. These clades are separated on opposite sides of a wide, lowland drainage bisecting the central
Cardamoms of Cambodia (Figs. 1, 5). In Thailand, the three populations sampled form a clade but occur in

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different mountain ranges across 190 km (Figs. 1, 5) and bear uncorrected pairwise sequence divergences of 2.0–
8.0% (Table 3). Tissue samples from the type locality of Khao Sebab, Chanthaburi Province were unobtainable but
due to this population’s morphological equivalency to the adjacent population from Khao Soi Dao we infer that it
falls within the distinct Thai clade. Despite evidence of additional undescribed species within the Thai clade, the
taxonomy of this group will remain unchanged until further analysis (Aowphol et al. in prep.). In Cambodia and
Vietnam, populations from the isolated mountains of Phnom Aural; Phnom Dalai; Kirirom; the Phnom Laang karst
tower; the Bokor Plateau; and the Bảy Núi Mountains (Nui Co To and Nui Cam) as well as the islands of Koh Rong
and Phu Quoc form separate clades. Phnom Aural, Phnom Dalai, Koh Rong Island, Cardamom Mountains and
Thai populations form a western monophyletic group and bear uncorrected pairwise sequence divergences among
them of 3.5%–9.3% (Table 4) across 440 km (Figs. 1,5). Phnom Laang, Kirirom, Bokor Plateau, Phu Quoc Island
and the Bảy Núi Mountain populations of Nui Co To and Nui Cam form the eastern clade and bear uncorrected
pairwise sequence divergences among them of 2.9%–8.9% (Table 4) across 130 km (Figs. 1, 5).
FIGURE 5. Tree: maximum likelihood topology illustrating the relationships of the Cyrtodactylus intermedius complex. Black
circles represent nodes supported by BPP and UBS values of 95% and 95, respectively. Gray circles denote nodes supported
only by BPP 95% or above. Map: distribution of the species of the Cyrtodactylus intermedius complex from the Cardamom
Mountains and uplands of Thailand, Cambodia, and Vietnam. Localities of new species represented by circles, localities of
known species represented by triangles, type locality of C. intermedius represented by a star, and locations of specimens
included in genetic analyses but are considered incertae sedis represented by squares.

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TABLE 3. Percent uncorrected pairwise sequence divergence for the different Thai populations of Cyrtodactylus
intermedius sensu stricto calculated from 1449 base pairs of the mitochondrial gene ND2.
TABLE 4. Percent uncorrected pairwise sequence divergence for species of the Cyrtodactylus intermedius complex
calculated from 1449 base pairs of the mitochondrial gene ND2.
The PCA of the concatenated data set reveals the most distant morphospatial separation occurs in the Phnom
Dalai and Phnom Laang populations with general separation of the Phnom Aural population and a cluster
consisting of the Bảy Núi populations and C. intermedius and a second cluster consisting of the Bokor and
Cardamom populations and C. phuquocensis (Fig. 6). PC1 accounted for 29% of the variation in the data set and
loaded most heavily for tibia length, head length, eye diameter, and eye to snout distance (TBL, HL, ED and ES;
Table 5). PC2 accounted for 13% of the variation in the data set and loaded most heavily for infralabial scales,
femoral scales, head width, and ear to eye distance (IL, FS, HW, and EE; Table 5). The DAPC showed even greater
separation among the Phnom Laang, Phnom Dalai and Bokor populations with general clustering of the remaining
populations (Fig. 7). The univariate and multivariate morphological analyses support the molecular analyses by
indicating that the new species within the group are well-separated from each other in morphospace and bear a
number of statistically significant mean differences in varying combinations of meristic and color pattern
characters (Table 6). Additionally, notable consistent differences in scale morphology and aspects of coloration and
pattern also provide reliable diagnostic character differences among the species (Table 7).
The preponderance of data suggests that the pronounced phylogeographic structure within Cyrtodactylus
intermedius sensu lato that bear deep genetic divergences, statistically different character state means, and
generally wide morphospatial separation among the sampled populations is indicative of a species complex and as
such we consider each population to be specifically distinct and describe them below.
The GMYC species delimitation independently recovered the same species inferred from the above analyses
but considered the C. intermedius Thai clade to be composed of three species and the Phnom Aural population,
central Cardamom clade, and Bokor Plateau population to be composed of two species each (Fig. 8). Sukumaran &
Knowles (2017) demonstrated that species delimitation methods generally overestimate species diversity by
recovering clades not species and that additional criteria such as morphology should be used in conjunction with
these analyses. Fujisawa & Barraclough (2013) also note that the GMYC approach should be used in conjunction
with additional independent data. We agree with these recommendations and with the exception of C. intermedius
Thai clade, which requires further research, believe the GMYC recovered noteworthy intrapopulational genetic
structure among individuals collected just meters apart (in the case of Phnom Aural) but due to their morphological
similarity we do not consider them separate species. Therefore, we base our species delimitation on a more
integrative approach.
1. Sakaerat 2. Sa Kaeo 3. Chanthaburi
1. Sakaerat *** *** ***
2. Sa Kaeo 8 *** ***
3. Chanthaburi 8 7.4 ***
12345678910
1. C. auralensis sp. nov. *** *** *** *** *** *** *** *** *** ***
2. C. bokorensis sp. nov. 7.6 *** *** *** *** *** *** *** *** ***
3. C. cardamomensis sp. nov. 7.7 7.9 *** *** *** *** *** *** *** ***
4. C. thylacodactylus sp. nov. 8 8.4 3.7 *** *** *** *** *** *** ***
5. C. intermedius Thai clade 7.1 7 5.4 6.2 *** *** *** *** *** ***
6. C. laangensis sp. nov. 7.6 3.9 7.8 8.1 6.9 *** *** *** *** ***
7. C. phuquocensis 8.7 7.7 7.6 8.9 7.5 6.9 *** *** *** ***
8. C. septimontium sp. nov. 8.5 5.9 6.8 8.2 7.1 5.4 7.4 *** *** ***
9. C. intermedius complex
incertae sedis 1
9 9.1 3.5 5 6.9 9.3 9 8.1 *** ***
10. C. intermedius complex
incertae sedis 2
8.56.37.68.97.26.67.42.98.8***

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SIX NEW CY RTODACTYLUS FROM SOUTHEAST ASIA
FIGURE 6. Principal component analysis (PCA) of the species of the Cyrtodactylus intermedius complex showing ordination
along the first two principal components.
FIGURE 7. Discriminant analysis of principal components (DAPC) of the species of the Cyrtodactylus intermedius complex
emphasizing between-group variance and minimizing within-group variance based on retention of the first eight components.
Circles represent 95% confidence intervals.

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TABLE 5. Summary statistics and principal component analysis scores for the Cyrtodactylus intermedius complex. Abbreviations are listed in the Materials and methods.
PC1 PC2 PC3 PC4 PC5 PC6 PC7 PC8 PC9 PC10 PC11
Std. dev. 2.708248751
1.772460844
1.608170552
1.37751533 1.200117232
1.071606082
1.020218107
0.964176828
0.936531076
0.8579313 0.839591646
Prop. of Var.
0.29338 0.12566 0.10345 0.0759 0.05761 0.04593 0.04163 0.03719 0.03508 0.02944 0.0282
Cum. Prop. 0.29338 0.41905 0.5225 0.5984 0.65601 0.70194 0.74358 0.78076 0.81585 0.84529 0.87349
eigen 7.3346113 3.141617443
2.586212524
1.897548485
1.44028137 1.148339595
1.040844986
0.929636956
0.877090456
0.736046115
0.704914131
SL -0.249892007
-0.070051752
0.009139269
0.155815282
-0.098621498
0.268846951
-0.150742355
0.021387215
-0.254418769
0.38392346 -0.243080333
IL -0.080435719
-0.364161975
0.094722263
-0.003409587
-0.364448858
0.299903527
0.040939479
-0.170056841
0.054810766
0.177441659
0.041665308
PVT 0.064487165
0.024935928
-0.385495168
-0.267469994
0.012083812
-0.032097154
0.003533113
-0.433327218
0.24016661 0.288724745
0.332813977
LRT 0.142364863
0.180959303
-0.202420548
0.069393992
-0.372850438
-0.085912527
-0.255218066
-0.195260561
0.398124405
-0.153299926
-0.231738228
VS -0.061169472
0.239807221
-0.208181778
0.379259091
-0.131075404
-0.189174016
0.195242183
0.12443262 -0.174796361
-0.034834468
0.305630232
TLE -0.244936583
-0.095508463
-0.211153849
-0.073912327
-0.339073609
-0.299343716
0.180843681
-0.045332812
-0.048281539
-0.039127721
-0.259125402
TLU -0.201089178
0.110418058
-0.288452339
-0.104436138
0.282062732
0.305303378
0.103187197
-0.150195505
-0.237273197
-0.174148871
-0.253853204
TTL -0.270709647
0.01748156 -0.310135657
-0.103787705
-0.013756196
0.019742649
0.178027957
-0.171729569
-0.165737733
-0.127609469
-0.339090731
FS -0.015543723
0.495912564
0.002919347
0.094843969
0.089332639
0.263566234
0.090860304
0.059440602
-0.09133612 -0.053352433
0.106294739
PS 0.115230214
-0.017113601
-0.186459474
-0.413088224
-0.291825553
0.15270163 -0.047735281
0.342587662
-0.114737714
0.222901778
0.167278813
PPS 0.035586394
-0.033377961
-0.310496284
0.293233876
-0.161831494
0.316117636
-0.396422414
0.296034008
0.112266813
-0.014371016
-0.120706147
PCT 0.071735376
-0.15899535 -0.339576559
0.328977619
0.069134121
-0.131907868
0.083721047
-0.175414216
-0.237046326
0.309255687
0.249607933
BB 0.090655224
-0.015266864
-0.142882876
0.006784957
-0.16752484 0.32180889 0.675929425
0.283355676
0.319112829
-0.118184457
0.042845118
FL -0.159302026
-0.262652603
-0.230689885
0.232393657
0.223688934
-0.059720259
-0.047054273
0.284809606
0.21263504 -0.01403368 -0.002184817
TBL -0.338122144
-0.063922919
-0.092514141
0.099885119
0.057647328
-0.062394636
-0.113486854
-0.044310948
0.165193587
-0.00257225 0.034857166
AG 0.198552027
-0.16484075 -0.204293924
-0.022679528
0.347506283
0.344777143
-0.181890873
-0.119267638
0.113212596
-0.211154619
0.13412633
HL -0.305836199
0.173524713
-0.037821941
0.094509663
-0.060601916
-0.024642406
-0.047004235
0.056157169
0.107119836
0.080481474
0.243115158
HW 0.112861155
0.316801959
-0.25725635 -0.134044705
-0.073702408
-0.037178041
-0.130218506
-0.098824205
-0.059366483
-0.154883029
0.022629565
HD -0.262670613
0.043941264
-0.007480447
-0.293880105
0.01885399 -0.153944329
-0.155582879
0.201409209
0.197373443
-0.100282454
0.04505675
ED -0.316174428
-0.068665686
-0.03531402 -0.10343076 0.018088753
-0.115667621
-0.034613374
0.100494919
-0.047053493
-0.216304799
0.216277072
EE -0.138686487
0.409262089
0.083562685
-0.158534758
-0.008260572
0.151266942
-0.077932614
0.059116368
-0.009993057
0.399674246
-0.029957116
ES -0.333905931
0.078019152
0.027992498
-0.008549493
0.108176655
0.03790824 -0.104537497
0.113963428
0.209845273
0.035746728
0.159977773
EN -0.161724469
-0.063887768
0.152126927
0.05284934 -0.377516134
0.2501879 -0.152516347
-0.21517264 -0.27689763 -0.448225237
0.375966681
IO 0.254301255
0.175560127
-0.043598209
0.22298964 -0.124638937
-0.158652704
-0.111857371
0.113537353
-0.096631476
-0.045270468
-0.128427769
BN -0.158549611
0.176873432
0.248799031
0.297142705
-0.021896587
0.138625672
0.142087128
-0.344931461
0.368324182
0.080500365
-0.069067113

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TABLE 6. Matirx of statistically significant mean differences of characters between species pairs of the C. intermedius complex. Abbreviations are listed in the Materials and
Methods.
1 2 3 4 5 6 7
1. C. auralensis sp. nov *** *** *** *** *** *** ***
2. C. bokorensis sp. nov. SL, LRT, TLU *** *** *** *** *** ***
3. C. cardamomensis sp. nov. SL, VS, TTL, PS,
PCT
VS, TLE, FS, PS,
PCT
*** *** *** *** ***
4. C. thylacodactylus sp. nov SL, IL, VS, TLE,
TLU, TTL, FS
SL, VS, TLE, TLU,
TTL, FS
SL, TLU, TTL, FS,
PS, PCT
*** *** *** ***
5. C. laangensis sp. nov. LRT, TLU, FS IL, VS, FS IL, TLE, FS, PS,
PCT
SL, IL, TLE, TTL,
FS
*** *** ***
6. C. intermedius *** FS TLE, TLU, TTL TLE, TTL FS *** ***
7. C. phuquocensis TLE, TTL, PS TLE, PS PS FS, PS TLE, FS TLE, TTL, PS ***
8. C. septimontium sp. nov. PVT, TLU, TTL,
PCT
SL, PVT, VS, PCT SL, PVT, FS, PS,
PCT
SL, PVT, TLE,
TTL, FS, PCT
PVT, FS PVT, TTL, FS, PS,
PCT
PVT, TLE, P CT

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TABLE 7. Meristic descriptive character summary statistics, morphological characters, and color pattern characters of the species of the Cyrtodactylus intermedius complex.
C. auralensis
sp. nov.
C. bokorensis
sp. nov.
C. cardamomensis
sp. nov.
C. thylacodactylus
sp. nov.
C. laangensis
sp. nov.
C. septimontium
sp. nov.
C. phuquocensis C. intermedius
supralabials (SL)
Mean 8.6 8 7.8 7 8.5 8.8 8 8
SD 0.5 0 0.63 0 0.84 0.54 0 0
Range 8 or 9 8 7–9 7 7–9 8–10 8 8
N 11 14 11 6 6 19 3 3
infralabials (IL)
Mean 9.9 9.3 9.5 8.8 10.7 9.8 9.3 9.3
SD 0.7 0.61 0.52 0.75 0.52 0.76 0.58 0.58
Range 9–11 8–10 9 or 10 8–10 10 or 11 9–11 9 or 10 9 or 10
N 11 14 11 6 6 19 3 3
paravertebral tubercles (PVT)
Mean 30 31.7 31.1 30 30.8 26.9 30.7 31.7
SD 1.1 0.99 1.45 0.63 1.47 2.16 0.58 1.15
Range 29–32 30–33 29–34 29–31 29–32 23–30 30 or 31 31–33
N 11 14 11 6 19 3 3
longitudinal rows of tubercles (LRT) 6
Mean 17.5 19.2 18.3 18.8 17.5 18.2 18.3 19.3
SD 0.52 0.89 1.35 0.98 0.55 1.08 0.58 0.58
Range 17 or 18 18–20 17–21 18–20 17–18 16–20 18 or 19 19 or 20
N 11 14 11 6 6 19 3 3
ventral scales (VS)
Mean 41.9 43.2 38.8 38.7 39.3 40.6 40.7 42.7
SD 1.38 0.89 2.04 2.66 1.21 2.09 1.15 1.15
Range 37–44 42–48 36–43 36–42 37–40 37–46 40–42 42–44
N 11 14 11 6 6 19 3 3
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SIX NEW CY RTODACTYLUS FROM SOUTHEAST ASIA
TABLE 7. (Continued)
C. auralensis
sp. nov.
C. bokorensis
sp. nov.
C. cardamomensis
sp. nov.
C. thylacodactylus
sp. nov.
C. laangensis
sp. nov.
C. septimontium
sp. nov.
C. phuquocensis C. intermedius
expanded subdigital lamellae on 4th toe (TLE)
Mean 6.5 6.6 5.8 5.2 7 6.5 5.3 7.3
SD 0.52 0.51 0.40 0.41 0 0.51 0.58 0.58
Range 6 or 7 6 or 7 5 or 6 5 or 6 7 6 or 7 5 or 6 7 or 8
N 11 14 11 6 6 19 3 3
unmodified subdigital lamellae on 4th toe
(TLU)
Mean 13.5 12.5 12.7 11.2 12.2 11.9 12.7 13
SD 0.82 0.76 0.47 0.75 0.75 0.74 0.58 1
Range 12–15 11–14 12 or 13 10–11 11–13 11–13 12 or 13 12–14
N 11 14 11 6 6 19 3 3
total subdigital lamellae on 4th toe (TTL)
Mean 20 19.1 18.5 16.3 19.2 18.4 18 20.3
SD 0.89 0.73 0.82 1.03 0.75 0.83 1 1.15
Range 19–21 18–20 17–19 15–18 18–20 17–20 17–19 19–21
N 11 14 11 6 6 19 3 3
enlarged femoral scales (FS)
Mean 26.3 27.9 24.2 19.7 14 27.7 27.7 23.7
SD 1.9 1.38 1.72 2.25 1.1 2.58 0.58 0.58
Range 23–28 26–30 23–28 17–22 0–16 24–33 27 or 28 23 or 24
N 11 14 11 6 6 19 3 3
precloacal scales (PS)
Mean 8.3 8.4 9.5 8.3 8 7.7 6.7 9
SD 0.65 0.74 0.69 0.82 0.63 0.58 0.58 1
Range 7–9 7–10 8–10 7–9 7–9 6–8 6 or 7 8–10
N 11 14 11 6 6 19 3 3
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TABLE 7. (Continued)
C. auralensis
sp. nov.
C. bokorensis
sp. nov.
C. cardamomensis
sp. nov.
C. thylacodactylus
sp. nov.
C. laangensis
sp. nov.
C. septimontium
sp. nov.
C. phuquocensis C. intermedius
post-precloacal scales rows (PPS)
Mean 2.9 2.9 2.8 3 2.8 2.8 2.7 3
SD 0.3 0.27 0.40 0 0.41 0.37 0.58 0
Range 2–3 2 or 3 2 or 3 3 2 or 3 2 or 3 2 or 3 3
N 11 14 11 6 6 19 3 3
postcloacal tubercles (PCT)
Mean 3 2.9 2.2 3 2.8 2 2.7 2.7
SD 0 0.36 0.40 0 0.75 0 0.58 0.58
Range 2 or 3 2 or 3 2 or 3 1 - 3 2 - 4 2 2 or 3 2 or 3
N 11 14 11 6 6 19 3 3
body bands (BB)
Mean 4.2 4.1 4.3 4.2 4 4 4 4
SD 0.4 0.27 0.47 0.41 0 0 0 0
Range 4 or 5 4 or 5 4 or 5 4 or 5 4 4 4 4
N 11 14 11 6 6 19 3 3
Morphology
femoral and precloacal scales continuous yes variable yes yes no yes yes yes
proximal femoral < 1/2 size of distal
femorals
yes yes no no yes yes yes yes
pocketing between digits of hindfeet no no no yes no no no no
pocketing between digits of forefeet no no no yes no no no no
Color Pattern
dark pigmented blotches on top of head
present
yes variable no no variable no no no
posterior border of the nuchal loop rounded
or pointed
rounded pointed rounded rounded rounded rounded rounded rounded
Maximum SVL (mm) 84.3 93 84.1 74.6 82.2 90.4 84.2 78.7
Sample Size 11 14 11 6 6 19 3 3

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SIX NEW CY RTODACTYLUS FROM SOUTHEAST ASIA
FIGURE 8. GMYC species delimitation. Lineages delimited as species by the GMYC are illustrated in red. Species recognized
in this analysis are those blocked off in different colors and named.
CUMZ R refers to the reptile collection at Chulalongkorn University Museum of Natural History, Bangkok,
Thailand; FMNH refers to the Field Museum of Natural History, Chicago, Illinois, USA; ITBCZ refers to the
Institute of Tropical Biology Collection of Zoology in Ho Chi Minh City, Vietnam and K refers to Ngo Van Tri
field numbers; LSUHC refers to La Sierra University Herpetological Collection and LSUDPC refers to the La
Sierra University Digital Photo Collection, La Sierra University, Riverside, California, USA; MCZ R refers to
Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts, USA; T refers to Neang Thy
field numbers; UMMZ refers to University of Michigan Museum of Zoology, Ann Arbor, Michigan, USA; ZMKU
refers to the Zoological Museum, Kasetsart University, Bangkok, Thailand and AA refers to corresponding
Anchalee Aowphol field numbers; ZMMU R refers to the reptile collection at Zoological Museum of Moscow
University, Moscow, Russia and NAP refers to the corresponding Nikolay A. Poyarkov field numbers.

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Taxonomy
Cyrtodactylus intermedius Smith
Intermediate Bow-fingered Gecko
Fig. 9, Table 8
Gymnodactylus intermedius Smith 1917: 221
Cyrtodactylus intermedius Taylor 1963: 734
Gymnodactylus (Cyrtodactylus) intermedius Wermuth 1965: 54
Cyrtodactylus intermedius Cox et al. 1998: 86
Cyrtodactylus (Cyrtodactylus) intermedius Rösler 2000: 66
Cyrtodactylus intermedius Nguyen et al. 2009
Syntypes. MCZ R-39040 and UMMZ 78687; collected by M.A. Smith, March 1916 from “Khao Sebab, SE Siam”
Additional specimen. FMNH 215981; collected by D.L. Damman, September of 1979 from Khao Soi Daow
Wildlife Sanctuary
Diagnosis. Adult males reaching 78.7 mm SVL; eight supralabials, 9 or 10 infralabials; 31–33 paravertebral
tubercles; 19 or 20 longitudinal rows of dorsal tubercles; 42–44 rows of ventral scales; seven or eight expanded
subdigital lamellae proximal to the digital inflection, 12–14 unmodified, distal, subdigital lamellae; 19–21 total
subdigital lamellae on fourth toe; enlarged femoral and precloacal scales continuous; 23 or 24 enlarged femoral
scales; proximal femoral scales one-half the size of distal femorals; 8–10 enlarged precloacal scales with pores on
each in males; three rows of enlarged post-precloacal scales; two or three postcloacal tubercles; no pocketing
between digits of hind or forefeet; dark pigmented blotches absent on top of head; posterior border of nuchal loop
rounded; and four dark body bands (Table 8). These characters are scored across all species of the Cyrtodactylus
intermedius complex in Table 7.
FIGURE 9. Cyrtodactylus intermedius from Khao Kitchakoot, Chanthaburi, Thailand. (photo by P. L. Wood Jr.)

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SIX NEW CY RTODACTYLUS FROM SOUTHEAST ASIA
TABLE 8. Meristic, mensural, and color pattern data of Cyrtodactylus intermedius from Khao Soi Dao and the type
locality of Khao Sebab. Abbreviations are listed in the Materials and Methods. R = right, L = left, / = data unobtainable,
r = regenerated, b = broken.
Remarks. No tissues were available from the type locality of Khao Sebab, Chanthaburi, Thailand however a
specimen, FMNH 215981, and genetic material, LSUHC 9513, from Khao Soi Dao Wildlife Sanctuary in the
adjacent mountain 30km north of the type locality were available. Upon morphological comparison of the two
syntypes, UMMZ 78687 and MCZ R-39040, with FMNH 215981 from Khao Soi Dao the two populations show no
UMMZ 78687 MCZ R-39040 FMNH 215981
locality Khao Sebab Khao Sebab Khao Soi Dao
sex M m m
supralabials 8 8 8
infralabials 10 9 9
paravertebral tubercles 33 31 31
longitudinal rows of tubercles 20 19 19
ventral scales 44 42 42
expanded subdigital lamellae on 4th toe 8 7 7
unmodified subdigital lamellae on 4th toe 13 12 14
total subdigital lamellae on 4th toe 21 19 21
enlarged femoral scales (R/L) R12L12 R11L12 R12L12
precloacal scales 10 9 8
precloacal pores 10 9 8
post-precloacal scales rows 3 3 3
postcloacal tubercles 3 3 2
body bands 4 4 4
femoral and precloacal scales continuous Yes yes yes
proximal femoral scales < 1/2 size of distal femorals Yes yes yes
pocketing between digits of hindfeet No no no
pocketing between digits of forefeet No no no
dark pigmented blotches on top of head present No no no
posterior border of nuchal loop rounded or pointed Rounded rounded rounded
SVL 73.4 78.7 76.1
TL 78 64r 8.5b
TW 5.9 6.5 6.7
FL 11.4 11.9 11.5
TBL 14 14.4 13.9
AG 34 35 33.4
HL 21.4 22.5 21.8
HW 13.7 15 14.6
HD 8.2 9 8.1
ED 5 5.5 5.1
EE 6.1 6.4 6.3
ES 8.5 8.8 8.3
EN 6.4 7 6.3
IO 3.2 3.5 3.4
IN 2.2 2.2 2.3

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morphological differences and are thus recognized here as conspecific. Thus, it is highly probable that if genetic
analyses of the Khao Sebab population were conducted it would be closely related to or conspecific with the Khao
Soi Dao population which falls within the Thai clade. The Thai clade is a member of the western group and the
sister clade to a lineage containing [C. thylacodactylus sp. nov. [C. intermedius complex incertae sedis 1 and C.
cardamomensis sp. nov.]] from which it is separated by 5.4%–6.9% sequence divergence (Table 4).
Cyrtodactylus auralensis sp. nov.
Phnom Aural Bent-toed Gecko
Figs. 10 & 11, Table 9.
Cyrtodactylus intermedius Swan, & Daltry 2002: 23–40; Daltry & Traeholt 2003: 89–90; Grismer, Chav, Neang, Wood,
Grismer, Youmans, Ponce, Daltry, Kaiser 2007: 223.
Holotype. Adult male LSUHC 7399 collected on 6 August 2005 by Neang Thy, L. Lee Grismer, Perry L. Wood,
Jr., Jesse L. Grismer, Thou Chav, Timothy M. Youmans, and Armando Ponce from Camp 1, Phnom Aural Wildlife
Sanctuary, Kampong Speu Province, Cambodia (1201’N, 10408’E; 549 m in elevation).
Paratypes. Adult male LSUHC 7396, adult females LSUHC 7397–98 and 7410 all bear the same collection
data as the holotype. Adult female LSUHC 7346 was collected at the same location as the holotype on 5 August
2005 by the same collectors.
Diagnosis. Adult males reaching 83.5 mm SVL, adult females reaching 84.3 mm SVL; eight or nine
supralabials, 9–11 infralabials; 29–32 paravertebral tubercles; 17 or 18 longitudinal rows of dorsal tubercles; 37–44
rows of ventral scales; six or seven expanded subdigital lamellae proximal to the digital inflection, 12–15
unmodified, distal, subdigital lamellae; 19–21 total subdigital lamellae on fourth toe; enlarged femoral and
precloacal scales continuous; 23–28 enlarged femoral scales; proximal femoral scales less than half the size of
distal proximal scales; 7–9 enlarged precloacal scales with pores on each in males; two or three rows of enlarged
post-precloacal scales; three postcloacal tubercles; no pocketing present between digits of hind or forefeet; dark
pigmented blotches present on top of head; posterior border of nuchal loop rounded, not chevron-shaped; four or
five dark body bands (Table 9). These characters are scored across all species of the Cyrtodactylus intermedius
complex in Table 7.
Description of holotype. Adult male SVL 75.9 mm; head moderate in length (HL/SVL 0.29) and width (HW/
HL 0.66), somewhat flattened (HD/HL 0.36), distinct from neck, and triangular in dorsal profile; lores concave
anteriorly, weakly inflated posteriorly, prefrontal region deeply concave, canthus rostralis rounded; snout elongate
(ES/HL 0.38), rounded in dorsal profile; eye large (ED/HL 0.24); ear opening elliptical, obliquely oriented,
moderate in size (EL/HL 0.10); eye to ear distance greater than diameter of eye; rostral rectangular, partially
divided dorsally by inverted Y-shaped furrow, bordered posteriorly by large left and right supranasals and three
smaller internasals, bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally
by large supranasal, posteriorly by three moderately sized postnasals, bordered ventrally by first supralabial;
8(R,L) rectangular supralabials extending to below midpoint of eye, second supralabial slightly larger than first;
10(R,L) infralabials tapering smoothly to below and slightly past the termination of enlarged supralabials; scales of
rostrum and lores flat to slightly raised, larger than granular scales on top of head and occiput; scales of occiput
intermixed with distinct, enlarged tubercles; dorsal superciliaries not elongate or keeled; 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 slightly enlarged, elongate sublabials extending
posteriorly to seventh infralabial; gular and throat scales small, granular, grading posteriorly into slightly larger,
flatter, smooth, imbricate, pectoral and ventral scales.
Body relatively short (AG/SVL 0.46) with poorly defined ventrolateral folds; dorsal scales small, granular
interspersed with relatively large, conical, semi-regularly arranged, weakly keeled tubercles; tubercles extend from
occiput onto original portion of tail; similarly sized and spaced tubercles continue onto nape of the neck and
occiput but diminish in size and distinction on top of head; approximately 17 longitudinal rows of tubercles at
midbody between ventrolateral, body folds; 31 paravertebral tubercles; 42 flat, imbricate, ventral scales between
ventrolateral body folds, ventral scales much larger than dorsal scales; eight large, pore-bearing, precloacal scales;
no deep precloacal groove or depression; and three rows of post-precloacal scales.

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FIGURE 10. Type series and additional specimens examined of Cyrtodactylus auralensis sp. nov. Upper: Type series from left
to right: holotype LSUHC 7399, paratypes LSUHC 7346, 7396-98, 7410. Lower: additional specimens examined.
Forelimbs moderate in stature, relatively short (FL/SVL 0.16); granular scales of forearm larger than those on
body, interspersed with large, conical tubercles; palmar scales rounded, slightly raised; interdigital pocketing
absent between digits of the forefeet; digits well-developed, inflected at basal, interphalangeal joints; digits slightly
more narrow distal to inflections; subdigital lamellae transversely expanded proximal to joint inflections, more
granular distal to inflection; claws well-developed, claw base sheathed by a dorsal and ventral scale; hind limbs
more robust than forelimbs, moderate in length (TBL/SVL 0.19), covered dorsally by granular scales interspersed
with large, conical tubercles and anteriorly by flat, slightly larger scales; ventral scales of thigh flat, imbricate,
larger than dorsals; ventral tibial scales flat, imbricate; one row of 13(R,L) enlarged femoral scales in contact with
enlarged precloacal scales, terminate just before the inflection of the knee; femoral pores absent; proximal femoral
scales one fourth the size of those distally and form abrupt union with smaller, rounded, ventral scales of
posteroventral margin of thigh; subtibial scales flat, imbricate; plantar scales flat; interdigital pocketing absent
between digits of the hind feet; digits relatively long, well-developed, inflected at basal, interphalangeal joints;
7(R,L) transversely expanded subdigital lamellae on fourth toe proximal to joint inflection extends onto the sole;

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14(R,L) unmodified lamellae distal to inflection; 21(R,L) total subdigital lamellae; and claws well-developed,
sheathed by a dorsal and ventral scale at base.
Tail 73.9 mm in length, first 6.8 mm original, last 67.1 mm regenerated, 6.4 mm in width at base, tapering to a
point; dorsal scales of original portion of tail flat, square; regenerated portion of tail covered with small, smooth
rectangular scales dorsally; median row of transversely expanded subcaudal scales, significantly larger than dorsal
caudal scales; caudal tubercles present on original portion; base of tail bearing hemipenal swellings; three
postcloacal tubercles on either side of base of hemipenal swellings; and postcloacal scales flat, imbricate.
FIGURE 11. Cyrtodactylus auralensis sp. nov. Upper left: LSUDPC 1180; adult female. Upper right: forested microhabitat
from the slopes of Phnom Aural at camp II. Lower right: forested microhabitat from the slopes of Phnom Aural at camp I.
Bottom right: LSUDPC 1178 hatchling. All photos by L. Grismer.
Coloration in alcohol. Dorsal ground color of head, body, limbs, and tail brown; dark-brown blotches on top
of head; dark-brown nuchal loop with rounded posterior border extends from posterior margin of one eye to
posterior margin of other eye; nuchal loop edged with thin, light, lines; five similarly colored dorsal bands lacking
lightened centers occur between limb insertions, first band terminates at shoulders, second, third and fourth bands
terminate dorsal to ventrolateral fold, fifth band terminates at anterior margin of hind limb insertions; light edging
on body bands generally restricted to the tubercles giving edging a spotted appearance; body band/interspace ratio
1.25; one additional dark-brown band posterior to hind limbs; regenerated tail a uniform brown color (Fig. 10).
Vari a t i o n. The paratypes closely approach the holotype in preserved coloration (Fig. 10). All paratypes have
four dorsal bands instead of five. LSUHC 7397 has a darker overall color pattern. LSUHC 7396–97 and LSUHC
7410 have light tubercles bordering the dark blotches on the top of their heads. LSUHC 7397 third dorsal band
bifurcates just dorsal to the ventrolateral fold. Coloration in life shows light edging of dark dorsal bands and
tubercles yellow (Fig. 11). Tubercles bordering dark dorsal bands and on dorsal margin of thigh yellow. Coloration
of hatchlings much lighter than that of adults (Fig. 11). Meristic differences in the type series and additional
specimens examined are presented in Table 9.

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TABLE 9. Meristic, mensural, and color pattern data from the type series and additional specimens of Cyrtodactylus auralensis sp. nov. Abbreviations are listed in the Materials and Methods. R = right,
L = left, / = data unobtainable, r = regenerated, b = broken.
LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC
7346 7396 7397 7398 7399 7410 7365 7400 7401 7409 7459
Paratype Paratype Paratype Paratype Holotype Paratype
locality Phnom
Aural
Phnom
Aural
Phnom
Aural
Phnom
Aural
Phnom
Aural
Phnom
Aural
Phnom
Aural
Phnom
Aural
Phnom
Aural
Phnom
Aural
Phnom
Aural
sex f m f f m F / f m f juv /
supralabials 9 9 9 8 8 9 9 8 9 8 9
infralabials 11 10 10 10 10 9 11 9 10 9 10
paravertebral tubercles 32 30 29 30 31 31 29 29 29 31 29
longitudinal rows of tubercles 17 18 18 18 17 18 17 17 17 18 17
ventral scales 41 42 44 40 42 42 42 41 37 44 40
expanded subdigital lamellae on 4th toe 6 7 7 6 7 6 7 6 7 6 6
unmodified subdigital lamellae on 4th toe 13 12 14 13 14 13 14 15 13 14 14
total subdigital lamellae on 4th toe 19 19 21 19 21 19 21 21 20 20 20
enlarged femoral scales (R/L) R13L13 R12L11 R14L13 R13L12 R13L13 R14L14 14R14L R14L14 R11L12 R14L14 R13L14
precloacal scales 9 9 9 8 8 9 8 8 7 8 8
precloacal pores / 9 / / 8 / / / 7 / /
post-precloacal scales rows 3 3 3 3 3 3 3 3 2 3 3
postcloacal tubercles 3 3 3 3 3 3 3 3 3 3 3
body bands 4 4 4 4 5 4 4 4 5 4 4
femoral and precloacal scales continuous yes yes yes yes yes Yes yes yes yes yes no
proximal femoral scales < 1/2 size of distal
femorals
yes yes yes yes yes Yes yes yes yes yes yes
pocketing between digits of hindfeet no no no no no No no no no no no
pocketing between digits of forefeet no no no no no No no no no no no
dark pigmented blotches on top of head yes yes yes yes yes Yes yes yes yes yes yes
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TABLE 9. (Continued)
LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC
7346 7396 7397 7398 7399 7410 7365 7400 7401 7409 7459
Paratype Paratype Paratype Paratype Holotype Paratype
posterior border of the nuchal loop rounded
or pointed
rounded rounded rounded rounded rounded rounded rounded rounded rounded rounded rounded
SVL 80.9 83.5 83.1 84.3 75.9 77.9 33.2 59.6 64.9 54.2 47.6
TL 10.1b 75.1r 6.9b 70.5r 73.9r 96
TW 6.1 7.6 6.6 6.7 6.4 6.4
FL 12.1 13.1 13.6 13.7 12 12.6
TBL 14.3 15.4 15.7 15.4 14.1 15
AG 37.6 39.4 39.1 40.7 35.2 36.5
HL 22.8 24.2 23.6 24.9 22 22
HW 15.3 16 15.7 16.2 14.5 14.3
HD 8.3 8.6 8.2 8.6 7.9 8.2
ED 5 5.4 5.2 5.3 5.2 5.2
EE 6.5 6.5 6.6 6.9 5.9 6.1
ES 9 9.6 9.3 10.2 8.4 8.5
EN 7.6 7.6 7.1 8 6.6 6.6
IO 3.3 3.6 3.3 3.7 3.1 3.3
IN 2.6 2.5 2.9 2.6 2.4 2.4

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Additional specimens examined. Four additional specimens from the type locality were examined (Table 9).
In addition, a juvenile female LSUHC 7459 was collected on 8 August 2005 at Camp 2, Phnom Aural, Kampong
Speu Province, Cambodia (1201’N, 10409’E; 1121 m in elevation) by the same collectors.
Distribution. Cyrtodactylus auralensis sp. nov. is known only from Phnom Aural Wildlife Sanctuary and
associated Mountainous areas nearby, Kampong Speu Province, Cambodia.
Etymology. The specific epithet, auralensis, is a noun in apposition in reference to the type locality of Phnom
Aural mountain, the highest mountain in Cambodia, to which it presumably endemic.
Natural history. Grismer et al. (2007a) reported that all specimens were collected at night in microhabitats
ranging from rocks along the edges of streams, tree trunks, up to 2 m above the ground in vegetation, to leaf litter
(Fig. 11). However, lizards were more common in areas where rocks were present. This is in accord with the
observations made by Stuart and Emmett (2006) with populations C. cardamomensis sp. nov. (see description
below) from the central Cardamoms. Grismer et al. (2007a) noted that C. auralensis sp. nov. was far more
abundant in hill evergreen forests at elevations between 500–600 m than at elevations above 1100 m in the cooler,
damper cloud forests.
Comparisons. Cyrtodactylus auralensis sp. nov. is a member of the western group and the sister species to a
lineage containing the remaining species of that group C. intermedius Thai clade [C. thylacodactylus sp. nov. [C.
intermedius complex incertae sedis 1 [C. cardamomensis sp. nov.]]] from which it is separated by 7.1%–9.0%
sequence divergence (Table 4). The PCA shows Cyrtodactylus auralensis sp. nov. is separated in morphospace
along PC1 and PC2 from all other members of the C. intermedius complex with the exception of C. cardamomensis
sp. nov. with which there is slight overlap (Fig. 6). The DAPC analysis shows the 95% confidence intervals of
Cyrtodactylus auralensis sp. nov. as distinct from C. thylacodactylus sp. nov., C. laangensis sp. nov. and C.
bokorensis sp. nov., overlapping slightly with C. intermedius and C. phuquocensis and grouping closely with C.
septimontium sp. nov. and C. cardamomensis sp. nov. (Fig. 7). Cyrtodactylus auralensis sp. nov. is well-
differentiated from all other members of the C. intermedius complex, with the exception of C. intermedius, by
having varying combinations of statistically different mean values of supralabial and infralabial scales;
paravertebral tubercles; longitudinal rows of tubercles; ventral scales; unmodified, expanded, and total number of
subdigital lamellae; enlarged femoral scales, precloacal scales, and postcloacal tubercles (Table 6). It differs further
from all other members of the complex in having the consistent presence of distinct pigmented blotches on top of
the head, although some individuals of C. bokorensis sp. nov., and C. laangensis sp. nov. have blotches but they
are not consistently present. Rounded posterior border of the nuchal loop separates C. auralensis sp. nov. from C.
bokorensis sp. nov. Continuous contact between precloacal and femoral scales differentiates it from C. laangensis
sp. nov. and some individuals of C. bokorensis sp. nov. in which contact is variable. Proximal femoral scales being
less than one-half the size of the distal femorals differentiates it from C. cardamomensis sp. nov. and C.
thylacodactylus sp. nov. Lack of interdigital pocketing separates it from C. thylacodactylus sp. nov. (Table 7).
Cyrtodactylus bokorensis sp. nov.
Bokor Plateau Bent-toed Gecko
Figs. 12 & 13, Table 10.
Gymnodactylus intermedius Smith 1935:44.
Cyrtodactylus intermedius Stuart & Emmett 2006:17.
Holotype. Adult male LSUHC 8554 collected on 10 July 2007 by L. Lee Grismer and Neang Thy from the Bokor
National Park Field Station, Kampot Province, Cambodia (10 37.611'N 104 01.541'E; 1033 m in elevation).
Paratypes. Adult males LSUHC 8541–42, adult females LSUHC 8548–50 and 8555 bear the same collection
data as the holotype.
Diagnosis. Adult males reaching 81.9 mm SVL, adult females reaching 93 mm SVL; eight supralabials, 8–10
infralabials; 30–33 paravertebral tubercles; 18–20 longitudinal rows of dorsal tubercles; 42–48 rows of ventral
scales; six or seven expanded subdigital lamellae proximal to the digital inflection, 11–14 unmodified, distal,
subdigital lamellae; 18–20 total subdigital lamellae on fourth toe; enlarged femoral and precloacal scales vary from
being separated by a diastema to continuous; 26–30 enlarged femoral scales; proximal femoral scales less than half
the size of distal proximal scales; 7–10 enlarged precloacal scales with pores on each in males; two or three rows of

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enlarged post-precloacal scales; two or three postcloacal tubercles; no pocketing present between digits of hind or
forefeet; dark pigmented blotches on top of head vary from present to absent; posterior border of nuchal loop
pointed, chevron-shaped; and four or five dark body bands (Table 10). These characters are scored across all
species of the Cyrtodactylus intermedius complex in Table 7.
Description of holotype. Adult male SVL 81.9 mm; head moderate in length (HL/SVL 0.28) and width (HW/
HL 0.65), somewhat flattened (HD/HL 0.36), distinct from neck, and triangular in dorsal profile; lores concave
anteriorly, weakly inflated posteriorly, prefrontal region moderately concave, canthus rostralis rounded; snout
elongate (ES/HL 0.39), rounded in dorsal profile; eye large (ED/HL 0.23); ear opening elliptical, obliquely
oriented, moderate in size (EL/HL 0.08); eye to ear distance greater than diameter of eye; rostral rectangular,
partially divided dorsally by a linear furrow, bordered posteriorly by large left and right supranasals and single
moderately sized internasal, bordered laterally by first supralabials; external nares bordered anteriorly by rostral,
dorsally by large supranasal, posteriorly by three moderately sized postnasals, bordered ventrally by first
supralabial; 8(R,L) rectangular supralabials extending to below midpoint of eye, second supralabial slightly larger
than first; 9(R,L) infralabials tapering smoothly to below and slightly past the termination of enlarged supralabials;
scales of rostrum and lores flat to slightly raised, larger than granular scales on top of head and occiput; scales of
occiput intermixed with distinct, enlarged tubercles; dorsal superciliaries not elongate or keeled; 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 slightly enlarged, elongate sublabials extending
posteriorly to seventh infralabial; gular and throat scales small, granular, grading posteriorly into slightly larger,
flatter, smooth, imbricate, pectoral and ventral scales.
FIGURE 12. Type series and additional specimens examined of Cyrtodactylus bokorensis sp. nov. Upper: Type series from left
to right: holotype LSUHC 8554, paratypes LSUHC 8541-42, 8548-50, 8555. Lower: additional specimens examined.

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TABLE 10. Meristic, mensural, and color pattern data from the type series and additional specimens of Cyrtodactylus bokorensis sp. nov. Abbreviations are listed in the Materials
and Methods. R = right, L = left, / = data unobtainable, r = regenerated, b = broken.
LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC
8541 8542 8548 8549 8550 8554 8555 8538 8543 8544 8545 8547 8551 8553
Paratype Paratype Paratype Paratype Paratype Holotype Paratype
locality Bokor Bokor Bokor Bokor Bokor Bokor Bokor Bokor Bokor Bokor Bokor Bokor Bokor Bokor
sex m m f f f m f m f f f f m m
supralabials 8 8 8 8 8 8 8 8 8 8 8 8 8 8
infralabials 9 9 9 9 8 9 10 10 10 9 9 9 10 10
paravertebral tubercles 31 31 32 33 32 33 31 30 33 31 32 33 31 31
longitudinal rows of tubercles 20 20 19 19 18 20 20 18 19 20 18 20 20 18
ventral scales 43 42 46 43 42 42 46 43 48 42 42 42 42 42
expanded subdigital lamellae on
4th toe
7 7 7 6 6 6 6 7 7 7 6 7 7 6
unmodified subdigital lamellae on
4th toe
12 13 13 14 13 12 12 12 12 12 13 13 11 13
total subdigital lamellae on 4th
toe
19 20 20 20 19 18 18 19 19 19 19 20 18 19
enlarged femoral scales (R/L) R16L14 R13L14 R13L15 R14L14 R13L13 R14L13 R13L13 R16L13 R13L15 R13L14 R14L14 R15L15 R15L15 R14L13
precloacal scales 8 7 9 8 8 9 8 8 10 8 9 8 9 8
precloacal pores 8 7 / / / 9 / 8 / / / / 9 8
post-precloacal scales rows 3 3 3 3 3 3 3 3 3 3 3 3 3 2
postcloacal tubercles 2 3 3 3 3 3 3 3 3 3 2 3 3 3
body bands 4 4 5 4 4 4 4 4 4 4 4 4 4 4
femoral and precloacal scales
continuous
yes yes yes yes yes yes no yes yes yes yes yes yes yes
proximal femoral scales < 1/2 size
of distal femorals
yes yes yes yes yes yes yes yes yes yes yes yes yes yes
pocketing between digits of
hindfeet
no no no no no no no no no no no no no no
pocketing between digits of
forefeet
no no no no no no no no no no no no no no
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TABLE 10. (Continued)
LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC
8541 8542 8548 8549 8550 8554 8555 8538 8543 8544 8545 8547 8551 8553
Paratype Paratype Paratype Paratype Paratype Holotype Paratype
dark pigmented blotches on top of
head
no no no no no yes no no no yes no no yes no
posterior border of the nuchal
loop rounded or pointed
pointed pointed pointed pointed pointed pointed pointed pointed pointed pointed pointed pointed pointed pointed
SVL 80.1 77.9 93 88 81.2 81.9 77.5 64.7 73 64.7 63 54 74.7 70.5
TL 60r 33.5b 120 99r 47.9r 119 105
TW 7.3 6.6 6.8 6.5 6.7 8.1 6.1
FL 11.9 12.3 13.2 13.3 13 12.4 11.9
TBL 14.3 14.8 16.6 15.8 15 14.5 14.8
AG 33.9 37.3 43.6 43.3 38.8 38.5 37.5
HL 22.9 22.1 24.9 24.4 22.4 22.8 22.3
HW 15.5 15.4 16.4 16.1 15.4 14.8 14.8
HD 8.2 8.1 9.1 8.8 8.6 8.1 7.9
ED 5.2 5.5 5.7 5.6 5 5.3 4.8
EE 6.6 5.8 6.3 7 6.5 6.6 5.9
ES 8.4 8.3 9.4 9.1 9.1 8.8 8.6
EN 6.5 6.7 7.8 7.2 2.9 6.8 6.6
IO 3.3 3.8 4.3 4.2 3.6 4.4 3.7
IN 3.1 2.4 3.1 3 2.4 2.8 3

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Body relatively short (AG/SVL 0.47) with poorly defined ventrolateral folds; dorsal scales small, granular
interspersed with relatively large, conical, semi-regularly arranged, weakly keeled tubercles; tubercles extend from
occiput to caudal constriction and onto tail where they occur in transverse rows separated by six small flat scales;
caudal tubercles largest dorsally, weak laterally, and absent ventrally; similarly sized and spaced tubercles continue
onto occiput and nape of the neck but diminish in size and distinction on top of the head; approximately 20
longitudinal rows of tubercles at midbody between ventrolateral, body folds; 33 paravertebral tubercles; 42 flat,
imbricate, ventral scales between ventrolateral body folds, ventral scales much larger than dorsal scales; nine large,
pore-bearing, precloacal scales; no deep precloacal groove or depression; and three rows of post-precloacal scales.
FIGURE 13. Cyrtodactylus bokorensis sp. nov. Upper left: adult male (photo by Neang Thy). Upper right: wind-swept grassy
microhabitat from the top of the Bokor Plateau (photo by L. Grismer). Lower left: Granite boulder strewn stream bed
microhabitat from the Bokor Plateau (photo by L. Grismer). Lower right: adult male (photo by Neang Thy).
Forelimbs moderate in stature, relatively short (FL/SVL 0.15); granular scales of forearm larger than those on
body, interspersed with large, conical tubercles; palmar scales rounded, slightly raised; interdigital pocketing
absent on the forefeet; digits well-developed, inflected at basal, interphalangeal joints; digits slightly more narrow
distal to inflections; subdigital lamellae transversely expanded proximal to joint inflections, more granular distal to
inflection; claws well-developed, claw base sheathed by a dorsal and ventral scale; hind limbs more robust than
forelimbs, moderate in length (TBL/SVL 0.18), covered dorsally by granular scales interspersed with large, conical
tubercles and anteriorly by flat, slightly larger scales; ventral scales of thigh flat, imbricate, larger than dorsals;
subtibial scales flat, imbricate; one row of 14(R), 13(L) enlarged femoral scales in contact with enlarged precloacal
scales, terminate just medial to inflection of knee; femoral pores absent; proximal femoral scales one-fourth the
size of those distal, form abrupt union with smaller, rounded, ventral scales of posteroventral margin of thigh;
plantar scales flat; interdigital pocketing absent on hind feet; digits relatively long, well-developed, inflected at
basal, interphalangeal joints; 6(R,L) transversely expanded subdigital lamellae on fourth toe proximal to joint
inflection that extend onto the sole, 12(R,L) unmodified lamellae distal to inflection; and claws well-developed,
sheathed by a dorsal and ventral scale at base.

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Tail original, 119 mm in length, 8.1 mm in width at base, tapering to a point; dorsal scales small, square, in
transverse rows; tail segmented with five transverse scale rows per segment; posterior margin of segments
bordered by four large, keeled tubercles dorsally extending to anterior two-thirds of tail; subcaudal region bearing
large, transverse scales for first half of tail, second half bearing smaller, irregularly shaped scales; base of tail
bearing hemipenal swellings; three postcloacal tubercles on either side of hemipenal swellings; and postcloacal
scales smooth, flat, large, imbricate.
Coloration in alcohol. Dorsal ground color of head, body, limbs and tail dark-brown; dark-brown blotches on
top of head ringed with white tubercles; dark-brown nuchal loop with pointed chevron-shaped posterior border
extending from posterior margin of one eye to posterior margin of other eye; nuchal loop edged with thin, white,
lines; four similarly colored dorsal bands with slightly lightened centers occur between limb insertions; first band
terminates at shoulders; second and third bands bear notches on their anterior borders, terminate dorsal to
ventrolateral fold; fourth band terminates at anterior margin of hind limb insertions; light edging on body bands
generally restricted to tubercles giving edging a spotted appearance; body band/interspace ratio 1.00; additional
dark-brown band posterior to hind limbs; 13 dark, caudal bands; and 12 light caudal bands infused with dark
pigment (Fig 11).
Vari a t i o n. The paratypes closely approach the holotype in coloration (Fig 11). LSUHC 8548 has five bands
instead of four. All paratypes have a pointed posterior border of the nuchal loop but vary between sharply and
bluntly pointed. All paratypes lack pigmented blotches on the top of their heads. Living specimens have light
edging along the dark dorsal bands and the tubercles are white. Tubercles border the dark dorsal bands and the
dorsal margin of the high is white. Dark blotches occur on the dorsal margin of the thigh (Fig 12). Meristic
differences among the type specimens and additional specimens examined are presented in Table 10.
Additional specimens examined. Seven additional specimens from the type locality were examined (Table
10).
Distribution. Cyrtodactylus bokorensis sp. nov. is known only from Bokor National park, Kampot province,
Cambodia.
Natural history. The type locality is located at 1033 m in elevation near the center of the Bokor Plateau in a
wide, open area among low hills. Summit habitat of the plateau is composed of scattered sandstone rocks and
boulders among highly disturbed wind-blown scrub vegetation. Also abundant at 350 m elevation in wet evergreen
forest associated with rocky areas on the southern side of Bokor NP and therefore probably common throughout
the park (Fig. 12). All specimens were collected at night during light intermittent rain showers in the vicinity of the
Bokor National Park Field Station. Lizards were found moving across the ground in the open, on the sides of
boulders, and on rock walls associated with the field station. Stuart & Emmet (2006) noted that some specimens
were collected on vertical walls of a reservoir and one was collected on a piece of sandstone in grassland.
Etymology. The specific epithet, bokorensis sp. nov., is an adjective in reference to type locality of the Bokor
Plateau to which it is presumably endemic.
Comparisons. Cyrtodactylus bokorensis sp. nov. is a member of the eastern group and the sister species of
Cyrtodactylus laangensis sp. nov. from Phnom Laang from which it is separated by 3.9% sequence divergence
(Table 4). The PCA analysis indicates Cyrtodactylus bokorensis sp. nov. is distinct in morphospace from all other
species with the exceptions of C. septimontium sp. nov. and C. cardamomensis sp. nov. with which there is overlap
(Fig 6). The concatenated DAPC analysis Cyrtodactylus bokorensis sp. nov. shows complete separation from all
other species (Fig. 7). Cyrtodactylus bokorensis sp. nov. is well-differentiated from all other members of the C.
intermedius complex by having varying combinations of statistically different mean values of supralabial and
infralabial scales, paravertebral tubercles, longitudinal rows of tubercles, ventral scales, unmodified, expanded, and
total number of subdigital lamellae, enlarged femoral scales, precloacal scales, and postcloacal tubercles (Table 6).
It differs further from all other members of the complex in the posterior border of the nuchal loop being chevron
shaped and relatively pointed as opposed to rounded. It is differentiated from C. thylacodactylus sp. nov. in lacking
interdigital pocketing (Table 7).
Cyrtodactylus cardamomensis sp. nov.
Cardamom Mountains Bent-toed Gecko
Figs. 14 & 15, Table 11.

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Cyrtodactylus intermedius Daltry and Traeholt 2003: 89–90; Emmet and Olsson 2005: 34–35; Stuart and Emmet 2006: 17;
Grismer, Thy, Chav, Wood, Oaks, Holden, Grismer, Szutz, and Youmans 2008: 165.
Holotype. Adult male LSUHC 7947 collected on 10 August 2006 by L. Lee Grismer, Neang Thy, Thou Chav,
Perry L. Wood Jr., Jamie R. Oaks, Jeremy Holden, Jesse L. Grismer, Thomas R. Szutz, Timothy M. Youmans from
Camp 3, Phnom Samkos Wildlife Sanctuary, Pursat Province, Cambodia (1211’52’’N, 10303’10’’E; 336 m in
elevation).
Paratypes. Adult female LSUHC 7936 and female juvenile LSUHC 7943 bear the same collection data. Adult
female LSUHC 7918 was collected at camp 2 (1212’N, 10304’E, 331 m in elevation) by the same collectors. Adult
female LSUHC 10096 was collected near O’Som village Pursat Province, Cambodia (1204’N 10309’E) on 23
August 2011 by the same collectors plus Evan S. H. Quah. Adult male FMNH 263344 was collected in the Thmar
Baing district, Koh Kong Province, Cambodia (1140’12’’N 10342’46’’E) 820 m in elevation on 28 February, 2004
by Bryan L. Stuart. Adult male FMNH 263345 was collected in the Thmar Baing district, Koh Kong, Cambodia
along the Russei Chrum river (1157’24’’N 10318’43’’E) 420 m in elevation 31 December, 2003 by Bryan L. Stuart.
Diagnosis. Adult males reaching 80.6 mm SVL, adult females reaching 84.1 mm SVL; 7–9 supralabials, 8–10
infralabials; 29–34 paravertebral tubercles; 17–21 longitudinal rows of dorsal tubercles; 36–43 rows of ventral
scales; five or six expanded subdigital lamellae proximal to the digital inflection, 12 or 13 unmodified, distal,
subdigital lamellae; 17–19 total subdigital lamellae on fourth toe; enlarged femoral and precloacal continuous; 23–
28 enlarged femoral scales; proximal femoral scales ranging from greater than one-half the size to the same size as
distal femoral scales; nine or 10 enlarged precloacal scales with pores in each in males; two or three rows of
enlarged post-precloacal scales; two or three postcloacal tubercles; no interdigital pocketing present; dark
pigmented blotches absent from top of head; posterior border of nuchal loop rounded; and four or five dark body
bands (Table 11). These characters are scored across all species of the Cyrtodactylus intermedius complex in Table
7.
Description of holotype. Adult male SVL 80.6 mm; head moderate in length (HL/SVL 0.28) and width (HW/
HL 0.70), somewhat flattened (HD/HL 0.37), distinct from neck, and triangular in dorsal profile; lores concave
anteriorly, weakly inflated posteriorly, prefrontal region deeply concave, canthus rostralis rounded; snout elongate
(ES/HL 0.38), rounded in dorsal profile; eye large (ED/HL 0.24); ear opening elliptical, obliquely oriented,
moderate in size (EL/HL 0.07); eye to ear distance greater than diameter of eye; rostral rectangular, partially
divided dorsally by L-shaped furrow, bordered posteriorly by large left and right supranasals and one large azygous
internasal, bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by large
supranasal, posteriorly by two moderately sized postnasals, bordered ventrally by first supralabial; 8(R,L)
rectangular supralabials extending to below midpoint of eye, second supralabial slightly larger than first; 10(R,L)
infralabials tapering smoothly to just below and slightly past the termination of enlarged supralabials; scales of
rostrum and lores flat to slightly raised, larger than granular scales on top of head and occiput; scales of occiput
intermixed with distinct, enlarged tubercles; dorsal superciliaries not elongate or keeled; 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 slightly enlarged, elongate sublabials extending
posteriorly to seventh infralabial; gular and throat scales small, granular, grading posteriorly into slightly larger,
flatter, smooth, imbricate, pectoral and ventral scales.
Body relatively short (AG/SVL 0.47) with poorly defined ventrolateral folds; dorsal scales small, granular
interspersed with relatively large, conical, semi-regularly arranged, weakly keeled tubercles; tubercles extend from
occiput onto base of tail but end at regenerated tail; similarly sized and spaced tubercles continue onto nape and
occiput but diminish in size and distinction on top of head; approximately 17 longitudinal rows of tubercles at
midbody between ventrolateral, body folds; 32 paravertebral tubercles; 37 flat, imbricate, ventral scales between
ventrolateral body folds, ventral scales much larger than dorsal scales; nine large, pore-bearing, precloacal scales;
no deep precloacal groove or depression; and two rows of post-precloacal scales.

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TABLE 11. Meristic, mensural, and color pattern data from the type series and additional specimens of Cyrtodactylus cardamomensis sp. nov Abbreviations are listed in the Materials and Methods. R =
right, L = left, / = data unobtainable, r = regenerated, b = broken.
LSUHC LSUHC LSUHC LSUHC LSUHC FMNH FMNH LSUHC LSUHC LSUHC FMNH
7918 7936 7943 7947 10096 263344 263345 7855 10081 10082 263346
Paratype Paratype Paratype Holotype Paratype Paratype Paratype
locality Phnom
Samkos
Phnom
Samkos
Phnom
Samkos
Phnom
Samkos
O'Som Thmar
Baing
Thmar
Baing
Phnom
Samkos
O'Som O'Som Thmar
Baing
sex f f f m f m m hatchling m f f
supralabials 7 8 7 8 8 8 9 8 7 8 8
infralabials 9 9 9 10 10 9 10 10 9 10 9
paravertebral tubercles 34 30 30 32 32 30 32 32 31 29 31
longitudinal rows of tubercles 18 17 17 17 21 19 18 17 20 18 18
ventral scales 39 43 40 37 36 39 39 38 36 40 40
expanded subdigital lamellae on 4th toe 6 5 6 6 6 6 6 5 6 6 6
unmodified subdigital lamellae on 4th toe 12 13 13 13 13 13 13 12 12 13 13
total subdigital lamellae on 4th toe 18 17 19 19 19 19 19 17 18 19 19
enlarged femoral scales (R/L) R12L12 R14L12 R12L11 R13L13 R12L11 R12L11 R12L11 R11L12 R12L12 R14L14 R12L11
precloacal scales 9 10 9 9 10 10 10 9 10 10 8
precloacal pores / / / 9 / 10 10 / / / /
post-precloacal scales rows 3 3 2 2 3 3 3 3 3 3 3
postcloacal tubercles 2 2 2 2 2 2 2 2 3 2 3
body bands 4 4 4 4 5 4 4 4 5 4 5
femoral and precloacal scales continuous yes yes yes yes yes yes yes yes yes yes yes
proximal femoral scales < 1/2 size of
distal femorals
no no no no no no no no no no no
pocketing between digits of hindfeet no no no no no no no no no no no
pocketing between digits of forefeet no no no no no no no no no no no
dark pigmented blotches on top of head
present
no no no no no no no no no no no
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TABLE 11. (Continued)
LSUHC LSUHC LSUHC LSUHC LSUHC FMNH FMNH LSUHC LSUHC LSUHC FMNH
7918 7936 7943 7947 10096 263344 263345 7855 10081 10082 263346
Paratype Paratype Paratype Holotype Paratype Paratype Paratype
posterior border of the nuchal loop
rounded or pointed
rounded rounded rounded rounded rounded rounded rounded rounded rounded rounded rounded
SVL 82.8 79 55.8 80.6 84.1 74 72
TL 107.8 76.5r 78 103r 61.5r 4.5b 80r
TW 6.8 5.8 4.6 6.4 6.1 6.4 5.5
FL 12.5 12.2 8.2 11.5 12.9 11.4 11
TBL 14.7 13.8 9.4 13.6 14.9 13.4 13.2
AG 40.2 38 24.2 38.2 42.7 35.3 34.6
HL 23.3 22.2 15.7 22.2 23 20.1 20.1
HW 15.9 14.6 10.5 15.5 15.6 14.1 14
HD 9.2 8.5 6 8.3 9.6 7.7 7.6
ED 5.6 5.5 3.4 5.3 5.4 5 4.7
EE 6.9 6 4.2 7.1 6.7 5.7 5.4
ES 9.3 8.7 6 8.5 9 8.1 8
EN 7.2 6.4 2.1 6.6 6.8 6.5 6.3
IO 3.4 3.4 2.5 3.3 3.4 3.1 2.9
IN 2.7 2.3 1.9 2.7 2.6 2.2 2.1

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FIGURE 14. Type series and additional specimens examined of Cyrtodactylus cardamomensis sp. nov. Upper: Type series
from left to right: holotype LSUHC 7947, paratypes LSUHC 7918, 7936, 7943,10096, FMNH 263344. Lower Right: paratype
FMNH 263345. Lower Left: additional specimens examined.
FIGURE 15. Cyrtodactylus cardamomensis sp. nov. Upper left: adult male from Phnom Samkos, Pursat province, Cambodia
(photo by Neang Thy). Upper right adult female from O’Som village, Pursat province, Cambodia LSUDPC 6072 (photo by L.
Grismer). Lower left: adult male from O’Som village, Pursat province, Cambodia LSUDPC 6071 (photo by L. Grismer).
Lower right: forested microhabitat from the slopes of Phnom Samkos (photo by L. Grismer).

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Forelimbs moderate in stature, relatively short (FL/SVL 0.14); granular scales of forearm larger than those on
body, interspersed with large, conical tubercles; palmar scales rounded, slightly raised; interdigital pocketing
absent; digits well-developed, inflected at basal, interphalangeal joints; digits slightly more narrow distal to
inflections; subdigital lamellae transversely expanded proximal to joint inflections, more granular distal to
inflection; claws well-developed, claw base sheathed by a dorsal and ventral scale; hind limbs more robust than
forelimbs, moderate in length (TBL/SVL 0.17), covered dorsally by granular scales interspersed with large, conical
tubercles and anteriorly by flat, slightly larger scales; ventral scales of thigh flat, imbricate, larger than dorsals;
subtibial scales flat, imbricate; one row of 13(R,L) enlarged femoral scales in contact with enlarged precloacal
scales, terminate just before the inflection of the knee; femoral pores absent; proximal femoral scales same size as
distal femorals, form abrupt union with smaller, rounded, ventral scales of posteroventral margin of thigh; plantar
scales flat; interdigital pocketing absent; digits relatively long, well-developed, inflected at basal, interphalangeal
joints; 6(R,L) transversely expanded subdigital lamellae on fourth toe proximal to joint inflection that extends onto
the sole; 13(R,L) unmodified lamellae distal to inflection; 18 total number of subdigital lamellae, and claws well-
developed, sheathed by a dorsal and ventral scale at base.
Tail 103 mm in length, first 34 mm original, last 69 mm regenerated, 6.4 mm in width at base, tapering to a
point; dorsal scales of original portion of tail, flat, square; regenerated portion of tail covered with small, smooth
rectangular scales dorsally; median row of transversely expanded subcaudal scales, significantly larger than dorsal
caudal scales; slightly keeled caudal tubercles present on original portion; base of tail bearing hemipenal swellings;
two postcloacal tubercles on either side of base of hemipenal swellings; and postcloacal scales flat, imbricate.
Coloration in alcohol. Dorsal ground color of head, body, limbs, and tail brown; dark-brown nuchal loop with
rounded posterior border extends from posterior margin of one eye to posterior margin of other eye; nuchal loop
edged with thin, light, lines; four similarly colored dorsal bands with slightly lightened centers occur between limb
insertions; second band slightly hourglass shaped; first band terminates at shoulders; second and third bands
terminate dorsal to ventrolateral fold; fourth band terminates at anterior margin of hind limb insertions; a single
similarly colored blotch present to the right of dorsal midpoint; tubercles bordering body bands bright white in
color giving a speckled appearance; body band/interspace ratio 1.2; three dark and two light caudal bands present
before giving way to uniform brown regenerated tail (Fig 14).
Vari a t i o n. The paratypes closely approach the holotype in coloration (Fig. 14). LSUHC 7918 and 7943 have
rounded distal body band borders terminating with a clear lightly colored border before the ventrolateral fold.
LSUHC 10096 has five body bands as opposed to four. In life, there is light edging along the dark dorsal bands and
the tubercles are pale yellow to a darker butterscotch color. Tubercles on dorsal margin of thigh slightly lighter than
base color (Fig 15). Meristic differences among the type specimens and additional specimens examined are
presented in Table 11.
Additional specimens examined. Three additional specimens examined (LSUHC 10081–82) were collected
near O’Som village Pursat Province, Cambodia (1204’N 10309’E) on 23 August, 2011. LSUHC 7855 was
collected 7 August 2006 at camp 1, Phnom Samkos, Pursat Province, Cambodia (1208'N 10308E; 331 m in
elevation). FMNH 263346 was collected at Tatai Leu, Thmar Baing district, Koh Kong (11 48’59’’N 103 31’51’’E)
430 m on 27 January 2004 (Table 11).
Distribution. Cyrtodactylus cardamomensis sp. nov. is presently known from three locations ranging across
the Cardamom Mountains from Phnom Samkos Wildlife Sanctuary, and the forests around O’Som and Thmar
Baing. The population from Koh Rong may prove to be C. cardamomensis sp. nov. and if this is the case the
species should be present in the hills south of the known populations.
Etymology. The specific epithet, cardamomensis, is an adjective in reference to the mountain range where the
type locality, Phnom Samkos, is located. While the Cardamom mountains refer broadly to all highlands located in
southwestern Cambodia we have chosen this group to be named as such as it occurs broadly through a large and
mostly contiguous segment of the mountain range rather than being restricted to specific isolated geological
features such as Phnom Aural, Phnom Dalai or the Bokor Plateau.
Natural history. The type locality is a former Khmer Rouge hideout situated along an unnamed river at the
base of Phnom Samkos at 1290 m elevation in a transitional zone between dry dipterocarp forest and hill evergreen
forest. No sign of extensive logging was evident at the time of collection and the understory was relatively free of
vegetation. Specimens were collected at night and found on the ground among small rocks and on vegetation no
higher than one meter above the ground. All material collected from secondary lowland forest at O’Som was also

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found at night in low vegetation. Stuart & Emmet (2006) note that specimens collected from Thmar Baing were
found in hill evergreen forest as well as disturbed lowland dry evergreen forest.
Comparisons. Cyrtodactylus cardamomensis sp. nov. is a member of the western group and the sister to C.
intermedius complex incertae sedis 1 from Koh Rong Island, Cambodia from which it is separated by 3.5%
sequence divergence (Table 4). The PCA analysis shows Cyrtodactylus cardamomensis sp. nov. is separated in
morphospace from all species with the exceptions of C. phuquocensis, C. auralensis sp. nov., and C. bokorensis sp.
nov. with which there is overlap. (Fig 6). The DAPC analysis shows C. cardamomensis sp. nov. as distinct from all
other species within the complex with the exceptions of C. intermedius, C. septimontium sp. nov., and C.
auralensis sp. nov. with which there is some overlap (Fig. 7). Cyrtodactylus cardamomensis sp. nov. is well-
differentiated from all species of the C. intermedius complex by having combinations of statistically different mean
values of supralabial and infralabial scales, paravertebral tubercles, longitudinal rows of tubercles, ventral scales,
unmodified, expanded, and total number of subdigital lamellae, enlarged femoral scales, precloacal scales, and
postcloacal tubercles (Table 6). It differs from C. auralensis sp. nov. by lacking any dark pigmented blotches on the
top of the head. It differs from C. bokorensis sp. nov. in having a nuchal loop with a rounded posterior border. It
differs from C. laangensis sp. nov. in having contact between femoral and precloacal scales. It differs from all
other species with the exception of C. thylacodactylus sp. nov. in having its proximal and distal femoral scales
being of approximately the same size and shape as opposed to having proximal femoral scales less than one-half
the size of the distal scales. It is further differentiated from C. thylacodactylus sp. nov. in lacking interdigital
pocketing (Table 7).
Remarks. The populations from the Cardamom mountains and Koh Rong Island have a sequence divergence
of 3.5% (Table 3) despite a physical distance of 100km, 15km of which is open ocean. Conversely, C.
cardamomensis sp. nov. and C. thylacodactylus sp. nov. have a 3.7% sequence divergence but are only separated
by 20km with significant morphological differentiation. No specimens of the population from Koh Rong Island
were available for morphological analysis so differences with C. cardamomensis sp. nov. could not be assessed.
Additionally, no specimens have been collected in the 100km gap, so it is unknown if there are intermediate
populations allowing for gene flow. Thus, for the purpose of this research the population from Koh Rong Island
will be referred to as incertae sedis 1.
Cyrtodactylus thylacodactylus sp. nov.
Phnom Dalai, Pocket-fingered, Bent-toed Gecko
Figs. 16 & 17, Table 12.
Cyrtodactylus intermedius Daltry and Chheang 2000: 107; Neang, Grismer, Chan, Grismer, Wood, and Youmans 2010: 137.
Holotype. Adult male LSUHC 9319 collected on 6 July 2009 by Neang Thy, L. Lee Grismer, Chan Kin Onn, Jesse
L. Grismer, Perry L. Wood Jr., and Timothy M. Youmans from Camp 1, Phnom Dalai, Pursat Province, Cambodia
(1225’23”N, 10304’07”E, 556 m in elevation).
Paratypes. Juvenile male LSUHC 9318 bears the same collection data as the holotype; adult females LSUHC
9326, 9336 and juvenile female LSUHC 9325 were collected on 7 July 2009, by the same collectors at camp 2,
Phnom Dalai, Pursat Province, Cambodia (1226’13”N, 10303’21”E, 968 m in elevation). Juvenile female 9349
was collected from camp 2 by the same collectors on 8 July 2009.
Diagnosis. Adult males reaching 71.5 mm SVL, adult females reaching 74.6 mm SVL; seven supralabials, 8–
10 infralabials; 29–31 paravertebral tubercles; 18–20 longitudinal rows of dorsal tubercles; 36–42 rows of ventral
scales; five or six expanded subdigital lamellae proximal to the digital inflection, 10–11 unmodified, distal,
subdigital lamellae; 15–18 total subdigital lamellae on fourth toe; enlarged femoral and precloacal continuous; 17–
22 enlarged femoral scales; proximal femoral scales the same size as distal femorals; 7–9 enlarged precloacal
scales with pores on each in males; three rows of enlarged post-precloacal scales; 1–3 postcloacal tubercles; deep
interdigital pocketing on either side of digit III on forefoot; deep interdigital pocketing present between third and
fourth digits of hind feet; dark pigmented blotches absent from top of head; posterior border of nuchal loop
rounded; and four or five dark body bands (Table 12). These characters are scored across all species of the
Cyrtodactylus intermedius complex in Table 7.

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TABLE 12. Meristic, mensural, and color pattern data from the type series of Cyrtodactylus thylacodactylus sp. nov.
Abbreviations are listed in the Materials and Methods. R = right, L = left, / = data unobtainable, r = regenerated, b =
broken.
LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC
9319 9326 9336 9318 9325 9349
Holotype Paratype Paratype Paratype Paratype Paratype
locality Phnom
Dalai
Phnom
Dalai
Phnom
Dalai
Phnom
Dalai
Phnom
Dalai
Phnom
Dalai
sex m f f m f f
supralabials 777777
infralabials 9 10 9 8 9 8
paravertebral tubercles 31 30 30 30 30 29
longitudinal rows of tubercles 20 18 20 18 19 18
ventral scales 36 41 42 36 37 40
expanded subdigital lamellae on 4th toe 5 5 5 5 6 5
unmodified subdigital lamellae on 4th toe 11 10 11 12 12 11
total subdigital lamellae on 4th toe 16 15 16 17 18 16
enlarged femoral scales (R/L) R8L10 R11L11 R11L10 R11L11 R8L10 R9L8
precloacal scales 9 9 9 7 8 8
precloacal pores 9 / / 7 / /
post-precloacal scales rows 3 3 3 3 3 3
postcloacal tubercles 3 3 3 3 3 3
body bands 4 5 4 4 4 4
femoral and precloacal scales continuous yes yes yes yes yes yes
proximal femoral scales < 1/2 size of
distal femorals
no no no no no no
pocketing between digits of hindfeet yes yes yes yes yes yes
pocketing between digits of forefeet yes yes yes yes yes yes
dark pigmented blotches on top of head
present
no no no no no no
posterior border of nuchal loop rounded
or pointed
rounded rounded rounded rounded rounded rounded
SVL 71.5 73.4 74.6 47.6 44.5 52.4
TL 74r 75r 100
TW 5.2 5.3 5.7
FL 10.2 11 10.7
TBL 12.9 12.2 12.4
AG 33.3 33.3 36
HL 20 20.4 20.1
HW 13.3 13.3 14.2
HD 7.9 7.6 7.7
ED 4.3 4.6 4.2
EE 5.5 5.8 5.5
ES 8 7.9 7.6
EN 6.2 6.1 5.9
IO 3.1 3.2 3.8
IN 2.6 2.6 2.4

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Description of holotype. Adult male SVL 71.5 mm; head moderate in length (HL/SVL 0.28) and width (HW/
HL 0.67), somewhat flattened (HD/HL 0.40), distinct from neck, and triangular in dorsal profile; lores concave
anteriorly, weakly inflated posteriorly, prefrontal region deeply concave, canthus rostralis rounded; snout elongate
(ES/HL 0.40), rounded in dorsal profile; eye large (ED/HL 0.22); ear opening elliptical, obliquely orientated,
moderate in size (EL/HL 0.12); eye to ear distance greater than diameter of eye; rostral rectangular, partially
divided dorsally by linear furrow, bordered posteriorly by large left and right supranasals and single smaller
internasal, bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by large
supranasal, posteriorly by two moderately sized postnasals, bordered ventrally by first supralabial; 7(R,L)
rectangular supralabials extending to below midpoint of eye, second supralabial slightly larger than first; 9(R,L)
infralabials tapering smoothly to below and slightly past termination of enlarged supralabials; scales of rostrum and
lores flat to slightly raised, larger than granular scales on top of head and occiput, scales of occiput intermixed with
distinct, enlarged tubercles; dorsal superciliaries not elongate or keeled; 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, elongate sublabials extending posteriorly to seventh
infralabial; gular and throat scales small, granular, grading posteriorly into slightly larger, flatter, smooth,
imbricate, pectoral and ventral scales.
FIGURE 16. Type series of Cyrtodactylus thylacodactylus sp. nov. from left to right: holotype LSUHC 9319, paratypes
LSUHC 9318, 9325-26, 9336, 9349.
Body relatively short (AG/SVL 0.47) with poorly defined ventrolateral folds; dorsal scales small, granular
interspersed with relatively large, conical, semi-regularly arranged, weakly keeled tubercles; tubercles extend from
occiput onto base of tail but terminate at regenerated tail; similar sized and spaced tubercles continue onto nape of
neck and occiput but diminish in size and distinction on top of head; approximately 20 longitudinal rows of
tubercles at midbody between ventrolateral, body folds; 31 paravertebral tubercles; 36 flat, imbricate, ventral scales
between ventrolateral body folds, ventral scales much larger than dorsal scales; nine large, pore-bearing, precloacal
scales; and no deep precloacal groove or depression.
Forelimbs moderate in stature, relatively short (FL/SVL 0.14); granular scales of forearm larger than those on
body, interspersed with moderately sized, conical tubercles; palmar scales rounded, slightly raised; significant
inpocketing of interdigit-webbing present on either side of digit III of the fore foot; digits well-developed, inflected
at basal, interphalangeal joints; digits slightly more narrow distal to inflections; subdigital lamellae transversely
expanded proximal to joint inflections, more granular distal to inflection; claws well-developed, claw base
sheathed by a dorsal and ventral scale; hind limbs more robust than forelimbs, moderate in length (TBL/SVL 0.18),
covered dorsally in granular scales interspersed with large, conical tubercles and covered anteriorly by flat, slightly
larger scales; ventral scales of thigh flat, imbricate, larger than dorsals; subtibial scales flat, imbricate; one row of
eight right and 10 left enlarged femoral scales in contact with enlarged precloacal scales, terminating just proximal
to knee; femoral pores absent; proximal femoral scales same size as distal femoral scales, form abrupt union with
smaller, rounded, ventral scales of posteroventral margin of thigh; plantar scales flat; significant inpocketing of
interdigit-webbing present between digits III and IV of the hind foot; digits relatively long well-developed,

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inflected at basal, interphalangeal joints; 5(R,L) transversely expanded subdigital lamellae on fourth toe proximal
to joint inflection extending onto the sole, 11(R,L) unmodified lamellae distal to inflection; and claws well-
developed, sheathed by a dorsal and ventral scale at base.
Tail 74 mm in length, first 11.8 mm original, last 62.6 mm regenerated, 5.2 mm in width at base, tapering to a
point; dorsal scales of original portion of tail flat, square; regenerated portion of tail covered with small, smooth
rectangular scales dorsally; median row of transversely expanded subcaudal scales, significantly larger than dorsal
caudal scales; caudal tubercles present on original portion; base of tail bearing hemipenal swellings; three
postcloacal tubercles on either side of base of hemipenal swellings; and postcloacal scales flat, imbricate.
Coloration in alcohol. Dorsal ground color of head, body, limbs, and tail brown; dark-brown nuchal loop with
rounded posterior border extends from posterior margin of one eye to posterior margin of other eye; nuchal loop
edged with thin, light, lines; four similarly colored dorsal bands with slightly lightened centers occur between limb
insertions, first band terminates at shoulders; second and third bands terminate dorsal to ventrolateral fold; fourth
band terminates at anterior margin of hind limb insertions; light edging on body bands with brighter white on
tubercles giving edging a slightly spotted appearance; body band/interspace ratio 1.3; additional dark-brown band
posterior to hind limbs; regenerated tail a uniform brown color (Fig 16).
FIGURE 17. Cyrtodactylus thylacodactylus sp. nov. Top: adult male from Phnom Dalai, Pursat province, Cambodia LSUDPC
5079. Bottom: forested boulder strewn stream bed microhabitat from the slopes of Phnom Dalai. All photos by L. Grismer.

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Va ri at io n . The paratypes closely approach the holotype in preserved coloration (Fig 16). LSUHC 9336 has
very light bordering on the body bands with distinct lightening at the center of each body band. LSUHC 9318 has
only three complete body bands with two small spots where the fourth would be between the limb insertions.
LSUHC has five body bands with the third and the fourth bands coming into contact in the vertebral region. In life
specimens have light-colored edging of the dark dorsal bands and the tubercles cream colored. Tubercles bordering
dark dorsal bands and on dorsal margin of thigh cream colored (Fig 17). Meristic differences among the type
specimens and additional specimens examined are presented in Table 12.
Distribution. Cyrtodactylus thylacodactylus sp. nov. is known only from Phnom Dalai in Samkos Wildlife
Sanctuary, Pursat Province, Cambodia.
Etymology. The specific epithet, thylacodactylus, is a Latinized adjective in masculine gender, derived from
Greek thylakos meaning “pouch”, and daktylos meaning “digit” in reference to the unique morphology of the new
species having deep interdigital pocketing between some of the digits.
Natural history. Neang et al. (2010) report that the type series was collected between 556–968 m in elevation
in evergreen hill forest in the vicinity of rocky streams. All specimens were collected at night at the base of trees
and on leaves and branches 1–1.5 m above the ground. Lizards were most commonly seen at the higher elevations
of Camp 11.
Comparisons. Cyrtodactylus thylacodactylus sp. nov. is a member of the western group and the sister species
to the clade comprised of C. intermedius complex incertae sedis 1 and C. cardamomensis sp. nov. from which it is
separated by 3.7%–5.0% sequence divergence (Table 4). The PCA and DAPC plots Cyrtodactylus thylacodactylus
sp. nov. as widely separated in morphospace from all other species within the complex (Figs. 6, 7). Cyrtodactylus
thylacodactylus sp. nov. is well-differentiated from all other members of the C. intermedius complex, by having
varying combinations of statistically different mean values of supralabial and infralabial scales, paravertebral
tubercles, longitudinal rows of tubercles, ventral scales, unmodified, expanded, and total number of subdigital
lamellae, enlarged femoral scales, precloacal scales, and postcloacal tubercles (Table 6). It differs from C.
auralensis sp. nov. in lacking dark pigmented blotches on the top of the head. It is separated from C. bokorensis sp.
nov. in having a rounded posterior border of the nuchal loop as opposed to it being pointed. Continuous contact
between the precloacal and femoral scales differentiates it from C. laangensis sp. nov. and some individuals of C.
bokorensis sp. nov. in which contact is variable. It is separated from all other members of the C. intermedius
complex, with the exception of C. cardamomensis sp. nov. in having proximal femoral scales approximately the
same size as the distal scales as opposed to being less than half the size of the distal scales. It is further
distinguished from all other members of the C. intermedius complex in having deep interdigital pocketing between
some of the digits of the front and rear feet (Table 7).
Remarks. Cyrtodactylus thylacodactylus sp. nov. occupies the most isolated position in morphospace within
the C. intermedius complex (Figs. 6, 7). The majority of variation in the data along PC1 and PC2 accounting for
body form (i.e., tibia length, head length, head width, eye diameter, eye to snout distance, and ear to eye distance)
whereas only one meristic character (number of femoral scales) accounted for notable variation along PC2 (Table
5). Overall, C. thylacodactylus sp. nov. is significantly smaller (Table 7) and more robust in body stature, indicated
by shorter hind limb and head length ratios to Snout Vent Length, than that of any other species in the complex.
Robustness was measured via an ANOVA of the ratios of Head Length (HL) and Tibia Length (TBL) to Snout Vent
Length (SVL) yielding a significant mean difference, adjusted P values <0.05.
Cyrtodactylus laangensis sp. nov.
Phnom Laang Bent-toed Gecko
Figs. 18 & 19, Table 13.
Holotype. Adult male LSUHC 8773 collected on 15 December 2007 by Neang Thy, L. Lee Grismer, and Jeremy
Holden from Phnom Laang, Kampot Province, Cambodia (10 42’13’’N, 104 20’57’’ E).
Paratypes. Adult males LSUHC 8771–72, adult females LSUHC 8770, 8774, and T 4653 bear the same
collection data as the holotype.

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TABLE 13. Meristic, mensural, and color pattern data from the type series of Cyrtodactylus laangensis sp. nov.
Abbreviations are listed in the Materials and Methods. R = right, L = left, / = data unobtainable, r = regenerated, b =
broken.
LSUHC LSUHC LSUHC LSUHC LSUHC T
8770 8771 8772 8773 8774 4653
Paratype Paratype Paratype Holotype Paratype Paratype
locality Phnom
Laang
Phnom
Laang
Phnom
Laang
Phnom
Laang
Phnom
Laang
Phnom
Laang
sex f mmmf f
supralabials 879999
infralabials 11 11 11 10 11 10
paravertebral tubercles 29 32 32 32 29 31
longitudinal rows of tubercles 18 18 17 17 17 18
ventral scales 39 40 37 40 40 40
expanded subdigital lamellae on 4th toe 7 77777
unmodified subdigital lamellae on 4th toe 12 11 13 13 12 12
total subdigital lamellae on 4th toe 19 18 20 20 19 19
enlarged femoral scales (R/L) R8L5 R6L8 R8L6 R0L0 R8L8 R6L7
precloacal scales 7 98888
precloacal pores / 988/ /
post-precloacal scales rows 3 23333
postcloacal tubercles 3 23423
body bands 444444
femoral and precloacal scales continuous no no no no no no
proximal femoral scales < 1/2 size of distal
femorals
yes yes yes / yes yes
pocketing between digits of hindfeet no no no no no no
pocketing between digits of forefeet no no no no no no
dark pigmented blotches on top of head no no no no no yes
posterior border of the nuchal loop rounded
or pointed
rounded rounded rounded rounded rounded rounded
SVL 77.6 76.2 74.4 76.9 82.2 76.5
TL 115.5 91.6r 88.8 116 / 106.7
TW 5.6 5.8 6.1 6.4 6.7 5.5
FL 12.3 12.7 12.4 12.7 12.7 12.5
TBL 15.3 15 15.4 15.4 15 15.2
AG 35 33.9 34.6 36.2 40.3 34.2
HL 21.9 21.2 21 21.9 22.8 21.6
HW 14 14.5 14.6 14.5 14.3 14
HD 8.6 8.3 8 8.8 8.2 8.2
ED 5.4 5.2 5.2 5.5 5.7 5.3
EE 5.6 5.4 5.7 6 5.6 5.8
ES 8.8 8.4 8.2 8.6 8.7 8.7
EN 7 6.3 6.1 6.9 6.6 7.1
IO 3 3 3.1 3.3 3.2 3.1
IN 2.6 2.4 2.6 2.3 2.4 3

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Diagnosis. Adult males reaching 76.9 mm SVL, adult females reaching 82.2 mm SVL; 7–9 supralabials, 10 or
11 infralabials; 29–32 paravertebral tubercles; 17 or 18 longitudinal rows of dorsal tubercles; 37–40 rows of ventral
scales; seven expanded subdigital lamellae proximal to the digital inflection, 11–13 unmodified, distal, subdigital
lamellae; 18–20 total subdigital lamellae on fourth toe; enlarged femoral scales either absent or separated from
precloacal scales by a diastema of smaller scales; when present 13–16 enlarged femoral scales with proximal
enlarged femoral scales less than half the size of distal femoral scales; 7–9 enlarged precloacal scales with pores on
each in males; two or three rows of enlarged post-precloacal scales; 2–4 postcloacal tubercles; no interdigital
pocketing; dark pigmented blotches on top of head present or absent; posterior border of nuchal loop rounded; and
four dark body bands (Table 13). These characters are scored across all species of the Cyrtodactylus intermedius
complex in Table 7.
Description of holotype. Adult male SVL 76.9 mm; head moderate in length (HL/SVL 0.28) and width (HW/
HL 0.66), somewhat flattened (HD/HL 0.40), distinct from neck, and triangular in dorsal profile; lores concave
anteriorly, weakly inflated posteriorly, prefrontal region moderately concave, canthus rostralis rounded; snout
elongate (ES/HL 0.39), rounded in dorsal profile; eye large (ED/HL 0.25); ear opening elliptical, obliquely
oriented, moderate in size (EL/HL 0.08); eye to ear distance greater than diameter of eye; rostral rectangular,
partially divided dorsally by linear furrow, bordered posteriorly by moderately sized left and right supranasals and
two internasals, bordered laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by
large supranasal, posteriorly by four moderately sized postnasals, bordered ventrally by first supralabial; 9(R,L)
rectangular supralabials extending to below midpoint of eye, second supralabial slightly larger than first; 10(R,L)
infralabials tapering smoothly to below and slightly past the termination of enlarged supralabials; scales of rostrum
and lores flat to slightly raised, larger than granular scales on top of head and occiput; scales of occiput intermixed
with distinct, enlarged tubercles; dorsal superciliaries not elongate or keeled; 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 slightly enlarged, elongate sublabials extending posteriorly to
seventh infralabial; gular and throat scales small, granular, grading posteriorly into slightly larger, flatter, smooth,
imbricate, pectoral and ventral scales.
Body relatively short (AG/SVL 0.47) with poorly defined ventrolateral folds; dorsal scales small, granular
interspersed with relatively large, conical, semi-regularly arranged, weakly keeled tubercles; tubercles extend from
occiput to caudal constriction and onto tail where they occur in transverse rows separated by seven or eight small,
flat scales; caudal tubercles largest dorsally, weak laterally, and absent ventrally; approximately 17 longitudinal
rows of tubercles at midbody between ventrolateral body folds; 32 paravertebral tubercles; 40 flat, imbricate,
ventral scales between ventrolateral body folds much larger than dorsal scales; eight large, pore-bearing, precloacal
scales; and no deep precloacal groove or depression.
Forelimbs moderate in stature, relatively short (FL/SVL 0.17); granular scales of forearm larger than those on
body, interspersed with moderately sized, conical tubercles; palmar scales rounded, slightly raised; interdigital
pocketing absent; digits well-developed, inflected at basal, interphalangeal joints; digits slightly more narrow distal
to inflections; subdigital lamellae transversely expanded proximal to joint inflections, more granular distal to
inflection; claws well-developed, claw base sheathed by a dorsal and ventral scale; hind limbs more robust than
forelimbs, moderate in length (TBL/SVL 0.20), covered dorsally by granular scales interspersed with large, conical
tubercles and anteriorly by flat, slightly larger scales; ventral scales of thigh flat, rounded, larger than dorsals,
decrease smoothly in size towards the posteroventral margin of thigh; subtibial scales flat, imbricate; enlarged
femoral scales absent; plantar scales flat; interdigital pocketing absent; digits relatively long, well-developed,
inflected at basal, interphalangeal joints; 7(R,L) transversely expanded subdigital lamellae on fourth toe proximal
to joint inflection that extend onto the sole, 13(R,L) unmodified lamellae distal to inflection; 20 total subdigital
lamellae; and claws well-developed, sheathed by a dorsal and ventral scale at base.
Tail original, 116 mm in length, 6.4 mm in width at base, tapering to a point; dorsal scales small, square, in
transverse rows; tail segmented with 6 or 7 transverse scale rows per segment; posterior margin of segments
bordered by four large, flat tubercles dorsally extending anterior one-third of tail; subcaudal region bearing large,
transverse scales for anterior one-half of tail, second half bearing smaller, irregularly shaped scales; base of tail
bearing hemipenal swellings; four postcloacal tubercles on either side of hemipenal swellings; and postcloacal
scales smooth, flat, imbricate.
Coloration in alcohol. Dorsal ground color of head, body, limbs, and tail light-brown; brown nuchal loop with

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rounded posterior border extends from posterior margin of one eye to posterior margin of other eye; nuchal loop
edged with thin, light, lines; four similarly colored dorsal bands with slightly lightened centers occur between limb
insertions; first band terminates at shoulders; second and third bands terminate dorsal to ventrolateral fold; fourth
band terminates at anterior margin of hind limb insertions; faint, light, edging on body bands; body band/interspace
ratio 0.90; one brown postsacral band posterior to hind limbs; original tail begins with dark bands with light
interspaces but fades to a uniform light-brown color (Fig 18).
Va ri at io n . The paratypes closely approach the holotype in preserved coloration (Fig. 18). Color in life shows
dark pigmented blotches on top of head ringed by light tubercles and lightened centers in the dark body bands and
dark blotches within the light intersperses as well as light and dark reticulations on limbs. Caudal interspaces begin
light-tan and change to a much brighter white near the tip (Fig 19).
Distribution. Cyrtodactylus laangensis sp. nov. is known only from Phnom Laang, an isolated karst formation
located in Kampot province, Cambodia.
Natural history. The type locality, Phnom Laang, is a small, isolated karst hill rising precipitously from the
flat lowlands of the eastern Mekong Delta region. The base of the hill is surrounded by highly disturbed scrub
vegetation but the hill itself bears typical ultramafic vegetation on its cliff-faces. The west face of Phnom Laang
opens up into a large open cave-like chamber that extends to near the center of the formation. The floor of the
chamber is covered with large, eroded karst boulders and rocks and its walls bear several large cracks and alcoves,
all of which provide appropriate microhabitat for karst-dwelling Cyrtodactylus (e.g. Grismer et al. 2016, 2017b,
2018a, b). At the base of the formation on the west side is a subterranean pond with clear water. The type series was
collected at night and all specimens were found on and amongst the karst microhabitat. Some were found crawling
on both large boulders and small rocks and others were found as high as 5 m above the ground on the chamber
walls.
FIGURE 18. Type s er ies of Cyrtodactylus laangensis sp. nov. from left to right: holotype LSUHC 8773, paratypes LSUHC
8770-72, 8774, T 4653.
Etymology. The specific epithet, laangensis, is an adjective in reference to type locality of the karst formation
Phnom Laang to which it is presumably endemic.
Comparisons. Cyrtodactylus laangensis sp. nov. is a member of the eastern group and the sister species to
Cyrtodactylus bokorensis sp. nov. from the Bokor Plateau, Kampot Province, Cambodia from which it is separated
by 3.9% sequence divergence (Table 4). The PCA and DAPC analyses indicate that Cyrtodactylus laangensis sp.
nov. is very isolated in morphospace from all other species within the complex (Fig. 6, 7). Cyrtodactylus
laangensis sp. nov. is well-differentiated from all other members of the C. intermedius complex by having varying
combinations of statistically different mean values of supralabial and infralabial scales, paravertebral tubercles,
ventral scales, expanded, and total number of subdigital lamellae, enlarged femoral scales, precloacal scales, and
postcloacal tubercles (Table 6). It differs from C. bokorensis by having a rounded posterior border of the nuchal

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loop as opposed to a pointed border. It is differentiated from all other members of the C. intermedius complex in
having a diastema between the enlarged femoral scales and the precloacal scales, with the exception of C.
bokorensis for which contact is variable. It is further differentiated from C. cardamomensis and C. thylacodactylus
in that the proximal most scales of its short femoral scale row are less than half the size of the distal scales. It is
further differentiated from C. thylacodactylus in lacking interdigital pocketing (Table 7).
FIGURE 19. Cyrtodactylus laangensis sp. nov. adult male T 4653 from Phnom Laang, Kampot province, Cambodia.
LSUDPC 4016 (photo by L. Grismer).
Parallel evolution of karst-association in Cyrtodactylus. Karst habitats are proving to be an extremely
important habitat for a growing number of endemic and microendemic species of amphibians and reptiles (Grismer
et al. 2016, 2017b, 2018a, b). Of all herpetofauna associated with these habitats, Cyrtodactylus has the greatest
diversity and shows a number of body form adaptations for this lifestyle. Cyrtodactylus laangensis sp. nov. is
another example of a site-specific karst-adapted endemic species. Cyrtodactylus laangensis sp. nov has traits noted
in other karst-associated Cyrtodactylus such as a decrease in enlarged femoral scale size and count as well elongate
hind limbs, this was measured via an ANOVA of the ratio of Tibia Length (TBL) to Snout Vent Length (SVL)
which yielded a significant mean difference, adjusted P values < 0.05, for all other members of the C. intermedius
complex with the exception of C. septimontium sp. nov. Grismer & Grismer (2017b) demonstrated that these
differences are functionally adaptive for locomoting on rocky surfaces in various planes of orientation. Despite
being separated by only 25km from its sister species, C. bokorensis sp. nov., the two species have an uncorrected
sequence divergence of 3.9% and the morphological and morphometric differences between the two species are
larger than between species having greater sequence divergence (Table 4, Fig. 5).
Cyrtodactylus septimontium sp. nov.
Bảy Núi (Seven Mountains) Bent-toed Gecko
Figs. 20 & 21, Table 14.
Cyrtodactylus intermedius Ngo and Grismer 2006: 328; Bobrov & Semenov 2008: 236; Nguyen, Ho, & Nguyen 2009: 768;
Nguyen et al. 2013; Nguyen et al. 2014.

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Holotype. Adult male LSUHC 12484 collected on 5 December 2005 by Ngo Van Tri at Nui Cam Mountain, An
Giang province, Vietnam.
Paratypes. Adult male LSUHC 8244 and adult female 8243 bear the same collection data as holotype. Adult
female LSUHC 12485 was collected at the same location. Adult male LSUHC 12488 and adult females LSUHC
12486–87 were collected on the adjacent hill Ta Soc Mountain, An Giang province, Vietnam. Adult males NAP–
05274, 05277, and adult females NAP–05272–73, 05275–76, all registered as ZMMU R–14773, were collected at
Nui Cam mountain, An Giang Province, Vietnam, on 26 August 2015 by N.A. Poyarkov and Nguyen Ngoc Hung
(N 10.51169; E 104.97193; 436 meters in elevation). Adult males NAP–05320–21, 05323, 05325, and adult
females NAP–05322, 05324, all registered as ZMMU R–14775, were collected at Nui Co To mountain, An Giang
Province, Vietnam, on 25 August 2015 by N.A. Poyarkov and Nguyen Ngoc Hung (N 10.38607; E 104.99169; 388
meters in elevation).
Diagnosis. Adult males reaching 85.5 mm SVL, adult females reaching 90.4 mm SVL; 8–10 supralabials, 9–
11 infralabials; 23–30 paravertebral tubercles; 16–20 longitudinal rows of dorsal tubercles; 37–46 rows of ventral
scales; six or seven expanded subdigital lamellae proximal to the digital inflection, 11–13 unmodified, distal,
subdigital lamellae; 17–20 total subdigital lamellae on fourth toe; enlarged femoral and precloacal scales
continuous; 24–33 enlarged femoral scales; proximal femoral scales less than one-half the size of distal proximal
scales; 6–8 enlarged precloacal scales with pores on each in males; two or three rows of enlarged post-precloacal
scales; two postcloacal tubercles; interdigital pocketing absent; dark pigmented blotches absent on top of head;
posterior border of nuchal loop rounded; four dark body bands (Table 14). These characters are scored across all
species of the Cyrtodactylus intermedius complex in Table 7.
Description of holotype. Adult male SVL 80.3 mm; head moderate in length (HL/SVL 0.29) and width (HW/
HL 0.69), somewhat flattened (HD/HL 0.39), distinct from neck, and triangular in dorsal profile; lores concave
anteriorly, weakly inflated posteriorly, prefrontal region deeply concave, canthus rostralis rounded; snout elongate
(ES/HL 0.39), rounded in dorsal profile; eye large (ED/HL 0.24); ear opening elliptical, obliquely oriented,
moderate in size (EL/HL 0.07); eye to ear distance greater than diameter of eye; rostral rectangular, partially
divided dorsally by an linear furrow, bordered posteriorly by large left and right supranasals and one large azygous
internasal, laterally by first supralabials; external nares bordered anteriorly by rostral, dorsally by large supranasal,
posteriorly by three moderately sized postnasals, ventrally by first supralabial; eight right, nine left rectangular
supralabials extending to below midpoint of eye, second supralabial slightly larger than first; eight right, nine left
infralabials tapering smoothly to below and slightly past the termination of enlarged supralabials; scales of rostrum
and lores flat to slightly raised, larger than granular scales on top of head and occiput; scales of occiput intermixed
with distinct, enlarged tubercles; dorsal superciliaries not elongate or keeled; 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 slightly enlarged, elongate sublabials extending posteriorly to sixth
infralabial; gular and throat scales small, granular, grading posteriorly into slightly larger, flatter, smooth,
imbricate, pectoral and ventral scales.
Body relatively short (AG/SVL 0.46) with poorly defined ventrolateral folds; dorsal scales small, granular
interspersed with relatively large, conical, semi-regularly arranged, weakly keeled tubercles; tubercles extend from
occiput to caudal constriction and onto tail where they occur in transverse rows separated by six or seven small, flat
scales; similarly sized and spaced tubercles continue onto nape and occiput but diminish in size and distinction on
top of head; approximately 19 longitudinal rows of tubercles at midbody between ventrolateral, body folds; 30
paravertebral tubercles; 42 flat, imbricate, ventral scales between ventrolateral body folds much larger than dorsal
scales; eight large, pore-bearing, precloacal scales; and no deep precloacal groove or depression.
Forelimbs moderate in stature, relatively short (FL/SVL 0.16); granular scales of forearm larger than those on
body, interspersed with large, conical tubercles; palmar scales rounded, slightly raised; interdigital pocketing
absent; digits well-developed, inflected at basal, interphalangeal joints; digits slightly more narrow distal to
inflections; subdigital lamellae transversely expanded proximal to joint inflections, more granular distal to
inflection; claws well-developed, claw base sheathed by a dorsal and ventral scale; hind limbs more robust than
forelimbs, moderate in length (TBL/SVL 0.19), covered dorsally by granular scales interspersed with large, conical
tubercles and anteriorly by flat, slightly larger scales; ventral scales of thigh flat, imbricate, larger than dorsals;
subtibial scales flat, imbricate; one row of 13(R) 14(L) enlarged femoral scales in contact with enlarged precloacal
scales, terminating just proximal to the inflection of the knee; femoral pores absent; proximal femoral scales one-
fourth the size of the distal scales, forming abrupt union with smaller, rounded, ventral scales of posteroventral margin

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TABLE 14. Meristic, mensural, and color pattern data from the type series of Cyrtodactylus septimontium sp. nov. Abbreviations are listed in the Materials and Methods. R =
right, L = left, / = data unobtainable, r = regenerated, b = broken.
LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC ZMMU
R-14773
ZMMU
R-14773
ZMMU
R-14773
8243 8244 12484 12485 12486 12487 12488 NAP-05277 NAP-05276 NAP-05275
Paratype Paratype Holotype Paratype Paratype Paratype Paratype Paratype Paratype Paratype
locality Nui Cam Nui Cam Nui Cam Nui Cam Nui
Co To
Nui
Co To
Nui
Co To
Nui Cam Nui Cam Nui Cam
sex f m m f f m f m f f
supralabials 9 9 8 9 9 9 8 8 8 9
infralabials 10 11 9 10 9 9 9 9 9 11
paravertebral tubercles 27 30 30 30 30 28 28 23 26 27
longitudinal rows of tubercles 18 19 19 19 18 17 16 18 19 18
ventral scales 40 42 42 40 40 42 40 40 41 38
expanded subdigital lamellae on 4th toe 6 6 6 6 7 7 6 6 6 7
unmodified subdigital lamellae on 4th toe 11 12 12 13 12 13 13 12 12 11
total subdigital lamellae on 4th toe 17 18 18 19 19 20 19 18 18 18
enlarged femoral scales (R/L) R13L14 R14L14 R13L14 R12L12 R13L12 R14L14 R14L14 R12L12 R14L15 R12L13
precloacal scales 8 8 8 8 8 7 7 7 8 8
precloacal pores 8r 8 8 8r 8r 7 7r 7 8 8r
post-precloacal scales rows 3 3 3 3 2 2 3 3 3 3
postcloacal tubercles 2 2 2 2 2 2 2 2 2 2
Body bands 4 4 4 4 4 4 4 4 4 4
femoral and precloacal scales continuous yes yes yes yes yes yes yes yes yes yes
proximal femoral scales < 1/2 size of distal
femorals
yes yes yes yes yes yes yes yes yes yes
pocketing between digits of hindfeet no no no no no no no no no no
pocketing between digits of forefeet no no no no no no no no no no
dark pigmented blotches on top of head present no no no no no no no no no no
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TABLE 14. (Continued)
LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC LSUHC ZMMU
R-14773
ZMMU
R-14773
ZMMU
R-14773
8243 8244 12484 12485 12486 12487 12488 NAP-05277 NAP-05276 NAP-05275
Paratype Paratype Holotype Paratype Paratype Paratype Paratype Paratype Paratype Paratype
posterior border of the nuchal loop rounded or
pointed
rounded rounded rounded rounded rounded rounded rounded rounded rounded rounded
SVL 68.4 83.2 80.3 84.8 81.7 80 83.3 79.8 90.4 79.9
TL 5.2b 74.7r 114 99r 85 119 77.4r
TW 6.3 6.9 6.2 6.5 6.2 6 6.3
FL 10.5 12.3 12.5 13.2 12.6 12 13
TBL 13.5 15.3 14.9 15.4 15.7 14.6 15.9
AG 30.8 36 36.9 37 35 34 36.7
HL 20.3 23.6 23 23.8 24.7 22.1 24.2
HW 13.1 15.4 15.9 15.3 15.5 14.2 15.1
HD 7.8 8.6 8.9 9.1 8.6 8.8 8.9
ED 4.9 5.3 5.5 5.1 5.2 5.6 5.8
EE 6.3 6.9 6.3 6.7 6.9 6.1 6.7
ES 8.4 9.2 8.9 9.3 9.5 9 9.6
EN 6.4 7.1 6.8
7.2 7.5 7.2 7.8
IO 3.3 3.4 3.7 3.6 3.5 3.5 3.6
IN 2.9 3 2.8 2.6 2.6 2.9 2.7
……continued on the next page

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TABLE 14. (Continued).
ZMMU
R-14773
ZMMU
R-14773
ZMMU
R-14773
ZMMU
R-14775
ZMMU
R-14775
ZMMU
R-14775
ZMMU
R-14775
ZMMU
R-14775
ZMMU
R-14775
NAP-
05274
NAP-
05273
NAP-
05272
NAP-
05321
NAP-
05320
NAP-
05322
NAP-
05324
NAP-
05323
NAP-
05325
Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype
locality Nui Cam Nui Cam Nui Cam Nui Co To Nui Co To Nui Co To Nui Co To Nui Co To Nui Co To
sex m f f m m f f m m
supralabials 9 9 9 9 9 8 10 9 9
infralabials 10 10 10 10 9 10 11 10 11
paravertebral tubercles 26 23 25 28 26 25 26 26 27
longitudinal rows of tubercles 20 18 16 19 18 18 18 18 20
ventral scales 42 42 46 42 42 38 40 37 38
expanded subdigital lamellae on 4th toe 6 6 7 7 7 6 7 7 7
unmodified subdigital lamellae on 4th toe 13 11 12 12 12 11 11 11 12
total subdigital lamellae on 4th toe 19 17 19 19 19 17 18 18 19
enlarged femoral scales (R/L) R12L13 R15L14 R14L14 R14L12 R16L16 R15L16 R17L16 R14L13 R16L14
precloacal scales 8 8 8 8 8 6 7 8 8
precloacal pores 8r 8 8 8 8 6 7 8 8
post-precloacal scales rows 3 3 3 2 3 3 3 3 3
postcloacal tubercles 2 2 2 2 2 2 2 2 2
Body bands 4 4 4 4 4 4 4 4 4
femoral and precloacal scales continuous yes yes yes yes yes yes yes yes yes
proximal femoral scales < 1/2 size of distal
femorals
yes yes yes yes yes yes yes yes yes
pocketing between digits of hindfeet no no no no no no no no no
pocketing between digits of forefeet no no no no no no no no no
dark pigmented blotches on top of head present no no no no no no no no no
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TABLE 14. (Continued).
ZMMU
R-14773
ZMMU
R-14773
ZMMU
R-14773
ZMMU
R-14775
ZMMU
R-14775
ZMMU
R-14775
ZMMU
R-14775
ZMMU
R-14775
ZMMU
R-14775
NAP-
05274
NAP-
05273
NAP-
05272
NAP-
05321
NAP-
05320
NAP-
05322
NAP-
05324
NAP-
05323
NAP-
05325
Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype Paratype
posterior border of the nuchal loop rounded or
pointed
rounded rounded rounded rounded rounded rounded rounded rounded rounded
SVL 67 65.3 69.3 85.5 84.4 89.8 86.8 82.5 59.5
TL
TW
FL
TBL
AG
HL
HW
HD
ED
EE
ES
EN
IO
IN

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FIGURE 20. Type series of Cyrtodactylus septimontium sp. nov. from left to right: Top: holotype LSUHC 12484, paratypes
LSUHC 8243-44, 12485-86, 12488, 12487. Middle: NAP 05276-77, 05275, 05274, 05273, 05272. Bottom: paratypes NAP
05321, 05320, 05322, 05324, 05323, 05323.

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FIGURE 21. Cyrtodactylus septimontium sp. nov. Top: adult male (Holotype LSUHC 12484) from Nui Cam Mountain, An
Giang province, Vietnam (photo by Ngo Van Tri). Middle: adult female from Nui Cam Mountain, An Giang Province, Vietnam
(photo by Nikolay A. Poyarkov). Bottom: Granite boulder forest microhabitat from the slopes of Nui Cam Mountain (photo by
L. Grismer).
of thigh; plantar scales round, slightly raised; interdigital pocketing absent; digits relatively long, well-developed,
inflected at basal, interphalangeal joints; 6(R,L) transversely expanded subdigital lamellae on fourth toe proximal to
joint inflection that extend onto the sole, 12(R,L) unmodified lamellae distal to inflection; 18 total subdigital lamellae;
and claws well-developed, sheathed by a dorsal and ventral scale at base.
Tail original, 114 mm in length, 6.2 mm in width at base, tapering to a point; dorsal scales of tail flat, square, in
transverse rows; tail segmented with six or seven transverse scale rows per segment; posterior margin of segments
bordered by three or four moderately sized, flat tubercles dorsally extending to anterior one-third of tail; subcaudal
region bearing large, transverse scales for first one-half of tail, second half bearing smaller, irregularly shaped
scales; base of tail bearing hemipenal swellings; two postcloacal tubercles on either side of base of hemipenal
swellings; and postcloacal scales smooth, flat, large, imbricate.
Coloration in alcohol. Dorsal ground color of head, body, limbs, and tail light-brown; brown nuchal loop with
rounded posterior border extends from posterior margin of one eye to posterior margin of other eye, edged with
thin, light, lines; four similarly colored dorsal body bands with slightly lightened centers occur between limb
insertions; first band terminates at shoulders; second and third bands terminate dorsal to ventrolateral fold; fourth
band terminates at anterior margin of hind limb insertions; light edging on body bands generally restricted to the

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tubercles producing a spotted appearance; body band/interspace ratio 1.3; additional dark-brown postsacral band;
and 11 dark and 12 light caudal bands (Fig 20).
Va ri at io n . The paratypes closely approach the holotype in coloration (Fig 20). In life, the base a tan color.
Edging of dark dorsal bands and tubercles white. The interiors of the body bands are nearly as light as base body
coloration. The light caudal bands also contain dark centers (Fig 21).
Distribution. Cyrtodactylus septimontium sp. nov. is known from the Bảy Núi Hill range in western An Giang
Province, Vietnam. Bảy Núi translates to “Seven Mountains” in Vietnamese in reference to the hills’ seven
prominent peaks. Specimens were collected from two of these hills, Nui Cam and Nui Co To, but it is presumed
that the species is found throughout the range.
Etymology. The specific epithet, septimontium, is a noun in apposition in reference to a Latinization of the
Vietnamese name for the hill range Bảy Núi meaning the “Seven Mountains”.
Natural history. The type locality of the new species, Nui Cam Mountain is part of the Bảy Núi Mountains –
a small range of seven larger mountains and several smaller hills of volcanic origin surrounded by the vast
agricultural lowlands of the Mekong Delta flood plain. Nui Cam is the highest of these mountains reaching 710 m
in elevation. Bảy Núi mountains are covered by old growth secondary semideciduous forest and highly disturbed
forest with numerous granitic outcroppings, large rocks with caverns and caves between them.
At the type locality specimens of Cyrtodactylus septimontium sp. nov. were collected at night (from 19:00
h to 24:00 h) mostly on large granite rocks, occasionally specimens were recorded on tree trunks up to 2.5 m above
the ground, or on the forest floor hiding in the leaf litter. Most specimens were recorded on vertical surfaces of
large granite rocks or in caverns between them. On Nui Cam Mountain Cyrtodactylus septimontium sp. nov. was
most abundant at elevations from ca. 300 to 490 m a.s.l. Sympatric species of lizards recorded with the new species
include Cnemaspis nuicamensis Grismer & Ngo, Gehyra mutilata (Wiegmann), Hemiphyllodactylus sp., Calotes
versicolor (Daudin), Lipinia vittigera (Boulenger) and Scincella cf. rufocaudata (Darevsky & Nguyen). On Nui Co
To Mountai n Cyrtodactylus septimontium sp. nov. was also recorded mostly on large granite rocks at elevations
350 – 450 m a.s.l., where it was found in sympatry with Cyrtodactylus grismeri Ngo and Cnemaspis tucdupensis
Grismer & Ngo.
Comparisons. Cyrtodactylus septimontium sp. nov. is a member of the eastern group sister species to C.
intermedius complex incertae sedis 2 from Kirirom National Park from which it is separated by 2.9% sequence
divergence (Table 4). The PCA analysis indicates C. septimontium sp. nov. is well-separated from all other species
in morphospace except for overlap with C. bokorensis sp. nov. and C. intermedius (Figs. 6). DAPC analysis shows
C. septimontium as distinct from C. bokorensis sp. nov., C. laangensis sp. nov., and C. thylacodactylus sp. nov. but
having general overlap with the remaining members of the complex. C. septimontium sp. nov. is well-differentiated
from all other members of the C. intermedius complex by having varying combinations of statistically different
mean values of supralabial and infralabial scales, paravertebral tubercles, ventral scales, unmodified, expanded,
and total number of subdigital lamellae, enlarged femoral scales, pre-cloacal scales, and post-cloacal tubercles
(Table 6). It differs from C. auralensis sp. nov. by lacking dark blotches on the top of the head. It differs from C.
bokorensis sp. nov. in having a nuchal loop with a rounded posterior border. It differs from C. laangensis in having
contact between femoral and precloacal scales. It differs from C. cardamomensis and C. thylacodactylus sp. nov. in
having proximal femoral scales that are less than half the size of the distal scales. It is further differentiated from C.
thylacodactylus sp. nov. in lacking interdigital pocketing (Table 7).
Remarks. Analysis of five specimens from Kirirom National Park, Cambodia, FMNH 263238-42, shows no
morphological differences and they may be conspecific with the populations form the Bảy Núi Hills. The two
populations have a sequence divergence of 2.9% (Table 4) despite a physical distance of 130km. However, for the
scope of this research the specimens from Kirirom will not be placed within septimontium sp. nov. due to the great
distance and lack of known intermediate populations where gene flow may exist.
Cyrtodactylus phuquocensis Ngo, Grismer, & Grismer 2010
Phu Quoc Island Bent-toed Gecko
Fig. 22, Table 15.
Cyrtodactylus phuquocensis Ngo, Grismer, & Grismer 2010

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FIGURE 22. Cyrtodactylus phuquocensis. Upper Left: juvenile eating a moth (photo by L. Grismer). Upper right: adult male
(photo by L. Grismer). Middle right: adult female (photo by Nikolay A. Poyarkov. Bottom: Granite boulder forest microhabitat
from Phu Quoc Island (photo by L. Grismer).

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TABLE 15. Meristic, mensural, and color pattern data from the type series of Cyrtodactylus phuquocensis.
Abbreviations are listed in the Materials and Methods. R = right, L = left, / = data unobtainable, r = regenerated, b =
broken.
LSUHC LSUHC LSUHC
11403 11404 11405
locality Phu Quoc Phu Quoc Phu Quoc
sex m f m
supralabials 8 8 8
infralabials 9 10 9
paravertebral tubercles 31 30 31
longitudinal rows of tubercles 18 19 18
ventral Scales 40 40 42
expanded subdigital lamellae on 4th toe 5 6 5
unmodified subdigital lamellae on 4th toe 12 13 13
total subdigital lamellae on 4th toe 17 19 18
enlarged femoral scales (R/L) R14L14 R14L13 R14L14
precloacal scales 7 7 6
precloacal pores 7 / 6
post-precloacal scales rows 2 3 3
postcloacal tubercles 3 2 3
Body bands 4 4 4
femoral and precloacal scales continuous yes yes yes
proximal femoral < 1/2 size of distal femorals yes yes y es
pocketing between digits of hindfeet no no no
pocketing between digits of forefeet no no no
dark pigmented blotches on top of head present no no no
posterior border of nuchal loop rounded or pointed rounded rounded rounded
SVL 81.6 84.2 59.9
TL 85r 94 80
TW 7.1 6.4 5.1
FL 12.1 13.2 9.4
TBL 14.8 15.2 11.2
AG 38.1 42.6 28.8
HL 22.7 23.6 16.9
HW 15.3 15.7 10.9
HD 8.5 8.5 6.4
ED 5.4 5.3 3.8
EE 6.3 6.5 4.4
ES 8.8 9.6 6.5
EN 6.8 7.5 5.3
IO 3.1 3.7 2.5
IN 2.8 3 2.2

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Holotype. LSUHC 5975; “Phu Quoc National Park, Phu Quoc Island, Kien Giang Province Biosphere Reserve,
Vie tna m ”.
Diagnosis. Adult males reaching 81.6 mm SVL, adult females reaching 85.8 mm SVL; eight supralabials, 9 or
10 infralabials; 30 or 31 paravertebral tubercles; 18 or 19 longitudinal rows of dorsal tubercles; 40–42 rows of
ventral scales; five or six expanded subdigital lamellae proximal to the digital inflection, 12 or 13 unmodified,
distal, subdigital lamellae; 17–19 total subdigital lamellae on fourth toe; enlarged femoral and precloacal scales
continuous; 27 or 28 enlarged femoral scales; proximal femoral scales less than half the size as distal femorals; six
or seven enlarged precloacal scales with pores on each in males; two or three rows of enlarged post-precloacal
scales; two or three postcloacal tubercles; interdigital pocketing absent; dark pigmented blotches absent from top of
head; posterior border of nuchal loop rounded; and four dark body bands (Table 15). These characters are scored
across all species of the Cyrtodactylus intermedius complex in Table 7.
Remarks. Cyrtodactylus phuquocensis is endemic to Phu Quoc Island, Kien Giang Province, Vietnam (Ngo et al.
2010) and is the sister species to the remaining taxa in the eastern group from which it bears a sequence difference of
5.4–7.7% (Table 4).
Discussion
The discovery of six new species of Cyrtodactylus previously considered to be under the same nomen cannot really
be taken as a surprise and adds to a growing body of data demonstrating that most widely distributed species
showing a great deal of intraspecific variation are actually species complexes. In Cyrtodactylus, this trend is most
notable in the C. pulchellus group that was originally recognized as one species but is now at least 15 species
(Grismer et al. 2012, 2014, 2016; Sumontha et al. 2012). As in the pulchellus group, morphological specializations
associated with scansorial behavior in karst ecosystems occur in the intermedius group with the slender and gracile
C. laangensis sp. nov. Additionally, and equally specialized in morphology—but in the opposite direction as C.
laangensis sp. nov. by bearing a significantly smaller but more robust body stature—is C. thylacodactylus sp. nov.
Additional field observations will be required to note how these adaptations bear on its life history. Nonetheless,
partitioning species complexes within Cyrtodactylus continue to be associated with distinctive ecomorphologies
and life histories. Not only does this further illustrate the trend that species richness within the genus Cyrtodactylus
is currently underestimated (Grismer et al. 2012, 2014, 2016; Oliver et al. 2016; Welton et al. 2010) but it also
punctuates the importance of the upland areas of Southeast Asia to biodiversity (see Grismer et al. 2016, 2017a and
references therein). Historic civil tensions and strife prevented biodiversity research in the Cardamom Mountains
of Cambodia. However, with the political reconciliation and integration of the late 1990’s, field work was
permitted to recommence. In the past two decades, numerous surveys have been made in the Cardamoms and
adjacent upland areas of Cambodia resulting in the discovery of no less than 12 new endemic species of
amphibians and reptiles (Ohler et al. 2002; Daltry & Wüster 2002; Grismer et al. 2007b, 2010; Wood et al 2010;
Neang et al. 2011; Mahony 2011; Neang et al. 2012; Geissler et al. 2014; Neang et al. 2014). The growing number
of species from these mountains highlights the need for continued field research and conservation in these unique
areas. There are still a number of isolated granitic and karstic areas within the western lowlands of the Mekong
Delta spanning the border between Vietnam and Cambodia and we have been shown photographs of an
undescribed karst-dwelling species from this area.
Key to the species of the Cyrtodactylus intermedius complex
The following key can be used as a guide to the identification of the species of the Cyrtodactylus intermedius
complex. However, it should be used in conjunction with the descriptions, diagnoses, and distribution section in the
text as some of the characters are sexually dimorphic.
1 Presence of interdigital pocketing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . thylacodactylus
- Absence of interdigital pocketing . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2 Posterior border of the nuchal loop pointed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . bokorensis
- Posterior border of the nuchal loop rounded . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3 Enlarged femoral scales not contiguous with precloacal scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . laangensis

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- Enlarged femoral scales contiguous with precloacal scales . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4 Dark pigmented blotches on top of head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .auralensis
- Dark pigmented blotches absent from top of head . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5 Proximal femoral scales same size as the distal femorals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . cardamomensis
- Proximal femoral scales less than one-half the size of the distal femorals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
6 Found West of the Cardamom Mountains. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . intermedius
- Found East of the Cardamom Mountains . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
7 Found on the mainland of South East Asia . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . septimontium
- Found on Phu Quoc Island . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . phuquocensis
Acknowledgements
For our work in Cambodia the authors wish to thank Fauna and Flora International and the Department of Nature
Conservation and Protection of the Ministry of the Environment, Cambodia, for facilitating our field work. We
would also like to thank His Excellency Say Samal the Senior Minister of Environment, His Excellency Mok
Mareth, the Senior Minister of Environment at the time of the surveys, His Excellency Chay Samith, Delegate of
the Royal Government of Cambodia in charge as General Director of Department of Administration for Nature
Conservation and Protection, and Mr Sy Ramony, Director of the Department of Wildlife Sanctuaries. Our work in
Thailand was supported by Thailand Research Fund (DBG6080010) to AA. We thank the Department of National
Parks, Wildlife and Plant Conservation, Thailand and Sakaerat Environmental Research Station for granting
research permission to AA. We also acknowledge Dr. Taksin Artchawakom the former director of Sakaerat
Environmental Research Station for permission to conduct research at SERS. For our work in Vietnam we would
like to thank Dr. Hoang Nghia Son, Director of Institute of Tropical Biology and Dr. Vu Ngoc Long for funding and
the permission letter, and Mr. Shaun Martin, Director of Educational Program for Nature, WWF in US funded a
small grant. NAP and HNN thank Andrei N. Kuznetsov (Hanoi) and Leonid P. Korzoun (Moscow) and Lin Si Min
(Taipei) for support of their work in Vietnam, also it was partially supported by the Russian Science Foundation