Research Article
Print
Research Article
A new rock gecko in the Cnemaspis siamensis group (Reptilia, Gekkonidae) from Kanchanaburi Province, western Thailand
expand article infoAttapol Rujirawan‡, Siriporn Yodthong§, Natee Ampai|, Korkhwan Termprayoon‡, Akrachai Aksornneam‡, Bryan L. Stuart, Anchalee Aowphol‡
‡ Kasetsart University, Bangkok, Thailand
§ Thaksin University, Phattalung, Thailand
| Srinakharinwirot University, Bangkok, Thailand
¶ North Carolina Museum of Natural Sciences, Raleigh, United States of America

Corresponding author: Anchalee Aowphol ( fsciacl@ku.ac.th )

Academic editor: Justin Bernstein
© 2022 Attapol Rujirawan, Siriporn Yodthong, Natee Ampai, Korkhwan Termprayoon, Akrachai Aksornneam, Bryan L. Stuart, Anchalee Aowphol.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Rujirawan A, Yodthong S, Ampai N, Termprayoon K, Aksornneam A, Stuart BL, Aowphol A (2022) A new rock gecko in the Cnemaspis siamensis group (Reptilia, Gekkonidae) from Kanchanaburi Province, western Thailand. Zoosystematics and Evolution 98(2): 345-363. https://doi.org/10.3897/zse.98.89591
ZooBank: urn:lsid:zoobank.org:pub:05AA43D4-8054-4884-B7E9-D5D572E6B3A6
Open Access

Abstract

We describe a new species of the gekkonid genus Cnemaspis from Erawan National Park in Kanchanaburi Province of western Thailand. Molecular phylogenetic analyses, based on the mitochondrial NADH dehydrogenase subunit 2 gene and flanking tRNAs, revealed that Cnemaspis auriventralis sp. nov. is nested within the C. siamensis group and is closely related to C. huaseesom, but has uncorrected pairwise genetic divergences of 12.12–27.92% from all other species in the C. siamensis group. The new species is also distinguished from other species in the C. siamensis group by having the combination of snout-vent length 36.7–38.6 mm in males (N = 3), 32.9–36.9 mm in females (N = 2); eight to ten supralabials; seven to nine infralabials; ventral scales smooth; six or seven precloacal pores in males; 16–17 paravertebral tubercles linearly arranged; tubercles on the lower flanks present; lateral caudal furrows present; no caudal tubercles in the lateral furrows; ventrolateral caudal tubercles present anteriorly; caudal tubercles not encircling tail; subcaudals smooth bearing a single median row of enlarged smooth scales; two postcloacal tubercles on each side; no shield-like subtibial scales; subtibial scales smooth; no enlarged submetatarsal scales; 23–27 subdigital lamellae on the fourth toe; sexually dimorphic for dorsal and ventral colour pattern; prescapular marking absent; gular marking absent; and yellow colouration in life of all ventral surfaces of head, body and tail in adult males. The new species is currently known only from upland karst habitat at its type locality.

Key Words

Cnemaspis auriventralis, Erawan National Park, karst formations, molecular phylogenetics, morphology

Introduction

The Southeast Asian Rock Gecko genus Cnemaspis Strauch, 1887 comprises a monophyletic clade of approximately 66 recognised species that are distributed from Laos, southern Vietnam westwards through southern Indochina, southwards through the Thai-Malay Peninsula, Sumatra, Java and eastwards to Borneo (Grismer 2010; Grismer et al. 2014, 2020; Wood et al. 2017; Riyato et al. 2019; Ampai et al. 2020; Quah et al. 2020; Nashriq et al. 2022; Uetz et al. 2022). Based on molecular and morphological data, Southeast Asian Cnemaspis are recovered in four major monophyletic clades that contain six species groups (Grismer et al. 2014). The Cnemaspis siamensis group is distributed across the Thai-Malay Peninsula and northwards to Kanchanaburi Province, western Thailand (Grismer et al. 2014, 2020; Ampai et al. 2019). This group currently contains 13 named species (Grismer et al. 2010, 2014, 2020; Wood et al. 2017; Ampai et al. 2019, 2020) including C. adangrawi Ampai, Rujirawan, Wood, Stuart & Aowphol, 2019, C. chanardi Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010, C. huaseesom Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010, C. kamolnorranathi Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010, C. lineatubercularis Ampai, Wood, Stuart & Aowphol, 2020, C. omari Grismer, Wood, Anuar, Riyanto, Ahmad, Muin, Sumontha, Grismer, Chan, Quah & Pauwels, 2014, C. phangngaensis Wood, Grismer, Aowphol, Aguilar, Cota, Grismer, Murdoch & Sites, 2017, C. punctatonuchalis Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010, C. roticanai Grismer & Chan, 2010, C. selenolagus Grismer, Yushchenko, Pawangkhanant, Nazarov, Naiduangchan, Suwannapoom & Poyarkov, 2020, C. siamensis (Smith, 1925), C. thachanaensis Wood, Grismer, Aowphol, Aguilar, Cota, Grismer, Murdoch & Sites, 2017 and C. vandeventeri Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010. Ecologically, many species of this genus are substrate or microhabitat specialists (e.g. granite, karst, vegetation or terrestrial) and restricted to activity periods (diurnal or nocturnal) and elevational zonation (lowlands or uplands) (Grismer et al. 2014; Wood et al. 2017; Ampai et al. 2020). In Kanchanaburi Province of western Thailand, C. huaseesom was discovered from Sai Yok National Park, Sai Yok District, based on specimens that were commonly found on hillsides in lowland areas having karst boulders (Grismer et al. 2010, 2014).

During recent herpetological surveys in Kanchanaburi Province of western Thailand, we collected five specimens of Cnemaspis from the karst formations in Erawan National Park. Molecular and morphological analyses revealed that the Erawan Cnemaspis were members of the C. siamensis group, but differed from all other recognised species. Herein, we describe it as a new species.

Materials and methods

Field collection of specimens

Field surveys were conducted at Erawan National Park, Tha Kradan Subdistrict, Si Sawat District, Kanchanaburi Province, Thailand, in November 2019 and November 2021 (Fig. 1). Direct observations were made during the day (09:00–17:00 h) and at night (19:00–22:00 h) and Cnemaspis specimens were collected by hand. Captured specimens were humanely euthanised using tricaine methanesulphonate (MS-222) within 24 hours of collection (Simmons 2015). Liver or muscle tissues were immediately removed from euthanised individuals, preserved in 95% ethyl alcohol and stored at -20 °C for molecular analysis. Euthanised specimens were fixed in 10% formalin and later transferred to 70% ethyl alcohol for permanent storage. Specimens and tissues were deposited in the herpetological collection of the Zoological Museum, Kasetsart University, Thailand (ZMKU). Geographic coordinates and elevations were recorded using a Garmin GPSMAP 64s with WGS84 datum. Ambient air temperature and relative humidity were collected with a Kestrel 4000 Weather Meter. Live animals and preserved specimens were photographed using a Nikon D700 or Z50 digital camera with an AF-S Micro Nikkor 60-mm f/2.8G ED lens and external flashes.

Figure 1. 

Map illustrating the type locality (yellow star) of Cnemaspis auriventralis sp. nov. at Erawan National Park, Si Sawat District, Kanchanaburi Province, Thailand and the Cnemaspis siamensis group samples used in the molecular analyses (asterisk) given in Table 2. Stars indicate type localities and circles represent additional localities.

Morphology

Morphological characters taken and their abbreviations were modified from recent studies of the genus Cnemaspis (Wood et al. 2017; Ampai et al. 2020; Grismer et al. 2020). Morphological measurements were taken with digital calipers to the nearest 0.1 mm. Scalation and other aspects of external morphology were examined using a Nikon SMZ745 stereomicroscope. Measurements were taken on the left side of the body, while scale counts were taken on both right and left sides (R/L) when possible. Measurements and meristic characters are shown in Table 1 and qualitative observations of external morphology evaluated are described below.

Table 1.
Download as
CSV
XLSX

Morphological characters and abbreviations used in this study.

Abbreviations Characters
Measurement
SVL Snout-vent length, taken from tip of snout to the anterior margin of vent
TW Tail width at the base of the tail immediately posterior to the postcloacal swelling
TL Tail length, as distance from the vent to the tip of the tail, whether original, broken or regenerated
FL Forearm length, taken on the dorsal surface from the posterior margin of the elbow while flexed 90° to the inflection of the flexed wrist
TBL Tibia length, taken on the ventral surface from the posterior surface of the knee while flexed 90° to the base of the heel
HL Head length, as distance from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout
HW Head width at the angle of the jaws
HD Head depth, as the maximum height of head from the occiput to the throat
AG Axilla-groin length, taken from the posterior margin of the fore-limb at its insertion point on the body to the anterior margin of the hind-limb at its insertion point on the body
ED Eye diameter, as the maximum horizontal diameter of the eyeball
EE Eye-ear distance, measured from the anterior margin of the ear opening to the posterior edge of the eyeball
EL Ear length, taken from the greatest vertical distance of the ear opening
EN Eye-nostril distance, measured from the anterior most margin of the eyeball to the posterior margin of the external nares
ES Eye-snout distance, measured from the anterior margin of the eyeball to the tip of snout
IO Inner orbital distance, as the width of the frontal bone at the level of the anterior edges of the orbit
IN Internarial distance, measured between the medial margins of the nares across the rostrum
Scalation
SL Supralabial scales, counted from below the middle of the orbit to the rostral scale
IL Infralabial scales, counted from below the middle of the orbit to the mental scale
PVT The number of paravertebral tubercles between limb insertions, counted in a straight line immediately left of the vertebral column
4TL The number of subdigital lamellae beneath the fourth toe, counted from the base of the first phalanx to the claw
PP The total number of pore-bearing precloacal scales in males
PPS The number of postcloacal tubercles on each side of tail base

Additional character states evaluated were the general size (i.e. strong, moderate, weak) and arrangement (i.e. random or linear) of the dorsal body tubercles; the orientation and shape of precloacal pores; the number of precloacal scales lacking pores separating the left and right series of pore-bearing precloacal scales; the degree and arrangement of body and tail tuberculation; the relative size and morphology of the subcaudal scales, subtibial scales and submetatarsal scales beneath the first metatarsal. Sex and maturity were determined by the presence of secondary sexual characteristics, such as the presence of hemipenes or pore-bearing precloacal scales in males, the presence of calcium glands or eggs in females or sexually dimorphic colour patterns. Morphological data for comparisons were obtained from the original and expanded descriptions of other species in the C. siamensis group (Smith 1925; Grismer and Chan 2010; Grismer et al. 2010, 2014, 2020; Wood et al. 2017; Ampai et al. 2019, 2020).

DNA extraction and PCR amplification

We extracted genomic DNA from the liver tissue of five individuals of Cnemaspis from Erawan National Park, Kanchanaburi Province (Table 2) using the DNeasy Blood and Tissue Kit (Qiagen, Germany) according to the manufacturer’s protocol. A portion of the mitochondrial NADH dehydrogenase subunit 2 gene (ND2) and its flanking tRNAs was amplified via a double-stranded polymerase chain reaction (PCR), using the light strand primer L4437b (5’-AAGCAGTTGGGCCCATACC-3’; Macey et al. 1997) and heavy strand primer H5934 (5’ AGRGTGCCAATGTCTTTGTGRTT-3’; Macey et al. 1997). PCR reactions were executed in 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 55 °C for 35 s, followed by a cycle extension at 72 °C for 35 s, for 33–40 cycles with a final extension at 72 °C for 10 min. PCR products were purified using a QIAquick PCR Purification Kit (Qiagen, Germany). PCR products were sequenced in both forward and reverse directions using the same amplifying primers at Biobasic Asia Inc. (Singapore) on an ABI 3730XL automatic sequencer (Applied Biosystems, CA, USA). Sequences were visually checked and edited in Geneious R11 (Biomatters Ltd, Auckland, New Zealand). The protein-coding region of ND2 was translated to amino acids and checked to confirm the lack of premature stop codons. All new sequences were deposited in GenBank under accession numbers OP093974–OP093978 (Table 2).

Table 2.
Download as
CSV
XLSX

Samples used in the molecular analyses, including their locality, voucher number and GenBank accession number. Voucher abbreviations are the School of Agriculture and Natural Resources, University of Phayao (AUP), Monte L. Bean Life Science Museum at Brigham Young University (BYU), California Academy of Sciences (CAS), the Field Museum of Natural History, Chicago, Illinois, USA (FMNH), La Sierra University Herpetological Collection (LSUHC), Universiti Sains Malaysia Herpetological Collection at the Universiti Sains Malaysia Herpetological Collection at the Universiti Sains Malaysia, Penang, Malaysia (USMHC), Zoological Museum of Kasetsart University (ZMKU) and the Zoological Museum of Moscow University (ZMMU).

Species Locality Voucher GenBank accession no. Reference
Outgroup
Cyrtodactylus bokorensis Cambodia, Kampot FMNH 263228 KT013107 Grismer et al. (2015b)
Hemidactylus garnotii Myanmar, Mon State, Kyaihto Township, CAS 222276 EU68364 Bauer et al. (2008)
Ingroup
Cnemaspis adangrawi Thailand, Satun Province, Mueang Satun District, Adang Island ZMKU R 00767 MK862112 Ampai et al. (2019)
Cnemaspis adangrawi Thailand, Satun Province, Mueang Satun District, Adang Island THNHM 28207 MK862113 Ampai et al. (2019)
Cnemaspis adangrawi Thailand, Satun Province, Mueang Satun District, Adang Island ZMKU R 00770 MK862114 Ampai et al. (2019)
Cnemaspis affinis Malaysia, Penang, Pulau Pinang LSUHC 6787 KM024682 Grismer et al. (2014)
Cnemaspis argus Malaysia, Terengganu, Gunung Lawit LSUHC 8304 KM024687 Grismer et al. (2014)
Cnemaspis argus Malaysia, Terengganu, Gunung Lawit LSUHC 10834 KM024688 Grismer et al. (2014)
Cnemaspis aurantiacopes Vietnam, Kien Giang Province, Hon Dat Hill LSUHC 8610 KM024692 Grismer et al. (2014)
Cnemaspis aurantiacopes Vietnam, Kien Giang Province, Hon Dat Hill LSUHC 8611 KM024693 Grismer et al. (2014)
Cnemaspis auriventralis sp. nov. Thailand, Kanchanaburi Province, Si Sawat District, Tha Kradan Subdisctrict, Erawan National Park ZMKU R 00999 OP093974 This study
Cnemaspis auriventralis sp. nov. Thailand, Kanchanaburi Province, Si Sawat District, Tha Kradan Subdisctrict, Erawan National Park ZMKU R 01000 OP093975 This study
Cnemaspis auriventralis sp. nov. Thailand, Kanchanaburi Province, Si Sawat District, Tha Kradan Subdisctrict, Erawan National Park ZMKU R 01001 OP093976 This study
Cnemaspis auriventralis sp. nov. Thailand, Kanchanaburi Province, Si Sawat District, Tha Kradan Subdisctrict, Erawan National Park ZMKU R 01002 OP093977 This study
Cnemaspis auriventralis sp. nov. Thailand, Kanchanaburi Province, Si Sawat District, Tha Kradan Subdisctrict, Erawan National Park ZMKU R 01003 OP093978 This study
Cnemaspis biocellata Malaysia, Perlis, Kuala Perlis LSUHC 8817 KM024707 Grismer et al. (2014)
Cnemaspis biocellata Malaysia, Perlis, Kuala Perlis LSUHC 8817 KM024708 Grismer et al. (2014)
Cnemaspis boulengerii Vietnam, Ca Mau Province, Con Dao Archipelago LSUHC9278 KM024710 Grismer et al. (2014)
Cnemaspis boulengerii Vietnam, Ca Mau Province, Con Dao Archipelago LSUHC9279 KM024711 Grismer et al. (2014)
Cnemaspis caudanivea Vietnam, Kien Giang Province, Hon Tre Island LSUHC 8582 KM024714 Grismer et al. (2014)
Cnemaspis chanardi Thailand, Nakhon Si Thammarat Province, Tham Thong Panra LSUHC 9567 KM024715 Grismer et al. (2014)
Cnemaspis chanthaburiensis Cambodia, Pursat Province, Phnom Dalai LSUHC 9338 KM024716 Grismer et al. (2014)
Cnemaspis grismeri Malaysia, Perak, Lenggong LSUHC 9969 KM024722 Grismer et al. (2014)
Cnemaspis hangus Malaysia, Pahang, Bukit Hangus LSUHC 9358 KM024728 Grismer et al. (2014)
Cnemaspis harimau Malaysia, Kedah, Gunung Jeri LSUHC 9665 KM024730 Grismer et al. (2014)
Cnemaspis huaseesom Thailand, Kanchanaburi Province, Sai Yok National Park LSUHC 9455 KM024733 Grismer et al. (2014)
Cnemaspis huaseesom Thailand, Kanchanaburi Province, Sai Yok National Park LSUHC 9457 KM024734 Grismer et al. (2014)
Cnemaspis huaseesom Thailand, Kanchanaburi Province, Sai Yok National Park LSUHC 9458 KM024735 Grismer et al. (2014)
Cnemaspis karsticola Malaysia, Kelantan, Gunung Reng LSUHC 9054 KM024736 Grismer et al. (2014)
Cnemaspis karsticola Malaysia, Kelantan, Gunung Reng LSUHC 9055 KM024737 Grismer et al. (2014)
Cnemaspis kumpoli Malaysia, Perlis, Perlis State Park LSUHC 8847 KM024745 Grismer et al. (2014)
Cnemaspis kumpoli Malaysia, Perlis, Perlis State Park LSUHC 8848 KM024746 Grismer et al. (2014)
Cnemaspis lineatubercularis Thailand, Nakhon Si Thammarat Province, Lan Saka District, Wang Mai Pak Waterfall ZMKU R 00825 MT112890 Ampai et al. (2020)
Cnemaspis lineatubercularis Thailand, Nakhon Si Thammarat Province, Lan Saka District, Wang Mai Pak Waterfall ZMKU R 00828 MT112891 Ampai et al. (2020)
Cnemaspis lineatubercularis Thailand, Nakhon Si Thammarat Province, Lan Saka District, Wang Mai Pak Waterfall ZMKU R 00829 MT112892 Ampai et al. (2020)
Cnemaspis lineogularis Thailand, Prachuap Khiri Khan Province, Kui Buri District, Wat Khao Daeng BYU 62535 KY091231 Wood et al. (2017)
Cnemaspis lineogularis Thailand, Prachuap Khiri Khan Province, Kui Buri District, Wat Khao Daeng ZMKU R 00728 KY091233 Wood et al. (2017)
Cnemaspis mahsuriae Malaysia, Kedah, Pulau Langkawi,Gunung Raya LSUHC 11829 KT250634 Grismer et al. (2015a)
Cnemaspis mcguirei Malaysia, Perak, Bukit Larut LSUHC 8853 KM024751 Grismer et al. (2014)
Cnemaspis monachorum Malaysia, Kedah, Langkawi Archipelago, Pulau Langkawi LSUHC 9114 KM024754 Grismer et al. (2014)
Cnemaspis monachorum Malaysia, Kedah, Langkawi Archipelago, Pulau Langkawi LSUHC 10807 KM024755 Grismer et al. (2014)
Cnemaspis narathiwatensis Malaysia, Perak, Belum-Temengor, Sungai Enam USMHC 1347 KM024762 Grismer et al. (2014)
Cnemaspis narathiwatensis Malaysia, Perak, Belum-Temengor, Sungai Enam USMHC 1348 KM024763 Grismer et al. (2014)
Cnemaspis neangthyi Cambodia, Pursat Province, O’Lakmeas LSUHC 8515 KM024767 Grismer et al. (2014)
Cnemaspis neangthyi Cambodia, Pursat Province, O’Lakmeas LSUHC 8516 KM024768 Grismer et al. (2014)
Cnemaspis niyomwanae Thailand, Trang Province, Tham Khao Ting LSUHC 9568 KM024773 Grismer et al. (2014)
Cnemaspis niyomwanae Thailand, Trang Province, Tham Khao Ting LSUHC 9571 KM024774 Grismer et al. (2014)
Cnemaspis nuicamensis Vietnam, An Giang Province, Nui Cam Hill LSUHC 8646 KM024775 Grismer et al. (2014)
Cnemaspis nuicamensis Vietnam, An Giang Province, Nui Cam Hill LSUHC 8647 KM024776 Grismer et al. (2014)
Cnemaspis omari Thailand, Satun Province, Phuphaphet Cave LSUHC 9565 KM024780 Grismer et al. (2014)
Cnemaspis omari Malaysia, Perlis, Perlis State Park LSUHC 9978 KM024779 Grismer et al. (2014)
Cnemaspis perhentianensis Malaysia, Terengganu, Pulau Perhentian Besar LSUHC 8699 KM024820 Grismer et al. (2014)
Cnemaspis phangngaensis Thailand, Phangnga Province, Mueang Phangnga District, Khao Chang, Phung Chang Cave BYU 62537 KY091234 Wood et al. (2017)
Cnemaspis phangngaensis Thailand, Phangnga Province, Mueang Phangnga District, Khao Chang, Phung Chang Cave BYU 62538 KY091235 Wood et al. (2017)
Cnemaspis punctatonuchalis Thailand, Prachaup Khiri Khan Province, Thap Sakae BYU 62539 KY091236 Wood et al. (2017)
Cnemaspis punctatonuchalis Thailand, Prachaup Khiri Khan Province, Thap Sakae BYU 62540 KY091237 Wood et al. (2017)
Cnemaspis roticanai Malaysia, Kedah, Pulau Langkawi, Gunung Raya LSUHC 9430 KM024829 Grismer et al. (2014)
Cnemaspis roticanai Malaysia, Kedah, Pulau Langkawi, Gunung Raya LSUHC 9431 KM024830 Grismer et al. (2014)
Cnemaspis selenolagus Thailand, Ratchaburi Province, Suan Phueng District ZMMU
R-16391		 MW051887 Grismer et al. (2020)
Cnemaspis selenolagus Thailand, Ratchaburi Province, Suan Phueng District AUP-00767 MW051888 Grismer et al. (2020)
Cnemaspis siamensis Thailand, Chumpon Province, Pathio District LSUHC 9474 KM024838 Grismer et al. (2014)
Cnemaspis siamensis Thailand, Chumpon Province, Pathio District LSUHC 9485 KM024839 Grismer et al. (2014)
Cnemaspis tarutaoensis Thailand, Satun Province, Mueang Satun District, Tarutao Island ZMKU R 00761 MK862117 Ampai et al. (2019)
Cnemaspis tarutaoensis Thailand, Satun Province, Mueang Satun District, Tarutao Island ZMKUR 00763 MK862118 Ampai et al. (2019)
Cnemaspis tarutaoensis Thailand, Satun Province, Mueang Satun District, Tarutao Island ZMKU R 00764 MK862119 Ampai et al. (2019)
Cnemaspis thachanaensis Thailand, Surat Thani Province, Tha Chana District, Tham Khao Sonk Hill BYU 62542 KY091239 Wood et al. (2017)
Cnemaspis thachanaensis Thailand, Surat Thani Province, Tha Chana District, Tham Khao Sonk Hill BYU 62543 KY091243 Wood et al. (2017)
Cnemaspis thachanaensis Thailand, Surat Thani Province, Tha Chana District, Tham Khao Sonk Hill BYU 62544 KY091244 Wood et al. (2017)
Cnemaspis tubaensis West Malaysia, Kedah, Langkawi Archipelago, Tuba Island USMHC 2527 MT028175 Quah et al. (2020)
Cnemaspis tubaensis West Malaysia, Kedah, Langkawi Archipelago, Tuba Island USMHC 2528 MT028176 Quah et al. (2020)
Cnemaspis tucdupensis Vietnam, An Giang Province, Tuc Dup Hill LSUHC 8631 KM024852 Grismer et al. (2014)
Cnemaspis tucdupensis Vietnam, An Giang Province, Tuc Dup Hill LSUHC 8632 KM024853 Grismer et al. (2014)
Cnemaspis vandeventeri Thailand, Ranong Province, Suk Saran District, Naka BYU 62541 KY091238 Wood et al. (2017)

Phylogenetic analyses

Additional homologous sequences of 68 individuals belonging to the C. affinis group, C. boulengerii group, the C. argus group, the C. chanthaburiensis group, the C. kumpoli group, the C. siamensis group and outgroups were downloaded from GenBank. Cyrtodactylus bokorensis Murdoch, Grismer, Wood, Neang, Poyarkov, Tri, Nazarov, Aowphol, Pauwels, Nguyen & Grismer, 2019 and Hemidactylus garnotii Duméril & Bibron, 1836 were selected as outgroups to root the tree following Ampai et al. (2020) and Quah et al. (2020). The five newly-generated and downloaded Cnemaspis sequences were aligned using the default options in the MUSCLE (Edgar 2004) plug-in in Geneious R11 (Biomatters Ltd, Auckland, New Zealand). The aligned dataset was partitioned into four partitions consisting of 1st–3rd ND2 codon positions and tRNAs.

Maximum Likelihood (ML) and Bayesian Inference (BI) were used to estimate phylogenetic relationships. Best-fit models of evolution for each partition was determined using the Bayesian information criterion (BIC) implemented in ModelFinder (Kalyaanamoorthy et al. 2017). The best-fit evolutionary models were TPM2u+F+I+G4 for tRNAs and TVM+F+I+G4, TIM3+F+G4 and GTR+F+G4 for ND2 codon positions 1, 2 and 3, respectively. The ML analysis was performed using the IQ-TREE webserver 1.6.12 (Trifinopoulos et al. 2016) with 1,000 bootstrap pseudo-replicates using the ultrafast bootstrap analysis (Minh et al. 2013; Hoang et al. 2018). The BI analysis was implemented in MrBayes v.3.2 (Ronquist et al. 2012) on the CIPRES Science Gateway V.3.3 (Miller et al. 2010) using default priors and models of evolution that were selected by the BIC and used in the ML analysis. Two independent runs, each with three heated and one cold chain, were performed using Markov Chain Monte Carlo (MCMC). The MCMC chains were run for 10,000,000 generations and trees sampled every 1,000 generations with the first 25% of each run discarded as burn-in. Stationarity was evaluated by ensuring effective sample sizes (ESS) were above 200 for all parameters in Tracer v. 1.7 (Rambaut et al. 2018). The phylogenetic trees from the ML and BI analyses were visualised using FigTree v. 1.4.4 (http://tree.bio.ed.ac.uk/software/figtree/). Nodes having ultrafast bootstrap support values (UFB) ≥ 95 and Bayesian posterior probabilities (BPP) ≥ 0.95 were considered highly supported (Huelsenbeck and Ronquist 2001; Wilcox et al. 2002; Minh et al. 2013). Uncorrected pairwise sequence divergences (p-distances) were calculated in MEGA 11 (Tamura et al. 2021) using the pairwise deletion option to remove gaps and missing data from the alignment prior to analysis.

Results

The final alignment of ND2 and flanking tRNAs contained 1,327 characters of 71 individuals of Cnemaspis and two individuals of outgroup species (Table 2). The average standard deviation of split frequencies was 0.000732 and the ESS of all parameters were ≥ 5,153 for all parameters in the BI analysis. The best tree in the ML analysis had a Maximum Likelihood value (lnL) of -22,848.995. The ML and BI analyses recovered trees with topologies similar to each other and to those recovered by Ampai et al. (2020) (Fig. 2). The five samples from Erawan National Park formed a strongly supported monophyletic lineage (≥ 95 UFB, ≥ 0.95 BPP) within the C. siamensis group. The Erawan National Park population was strongly supported (≥ 95 UFB, ≥ 0.95 BPP) to be the sister taxon of C. huaseesom from Sai Yok National Park, Sai Yok District, Kanchanaburi Province. Uncorrected pairwise genetic divergences (p-distances) within the Erawan National Park population were 0.00–0.65%. The Erawan National Park population had uncorrected p-distances of 12.12–12.55% from C. huaseesom and 15.15–27.92% from the other species in the siamensis group. The p-distances amongst species in the C. siamensis group ranged from 8.23–29.00% (Table 3).

Table 3.
Download as
CSV
XLSX

Mean (minimum-maximum) percentages of uncorrected pairwise sequence divergences (p-distances) of Cnemaspis species in the C. siamensis group compared to C. auriventralis sp. nov., based on 1,327 aligned characters of the mitochondrial NADH dehydrogenase subunit 2 gene and flanking tRNAs. Intraspecific p-distances are in bold font.

Species N C. auriventralis sp. nov. C. adangrawi C. chanardi C. huaseesom C. lineatubercularis C. omari C. phangngaensis C. punctatonuchalis C. roticanai C. selenolagus C. siamensis C. thachanaensis C. vandeventeri
C. auriventralis sp. nov. 5 0.30 (0.00–0.65)
C. adangrawi 3 25.57 (25.11–25.97) 3.03 (0.00–4.55)
C. chanardi 1 26.23 (25.97–26.41) 11.40 (11.02–12.12) –
C. huaseesom 3 12.29 (12.12–12.55) 26.26 (26.19–26.41) 26.62 (26.62–26.62) 0.43 (0.00–0.65)
C. lineatubercularis 3 25.19 (24.89–25.54) 17.97 (17.32–19.05) 17.32 (17.10–17.53) 27.95 (27.71–28.35) 0.29 (0.22–0.43)
C. omari 2 27.68 (27.27–27.92 9.63 (8.23–10.39) 11.90 (11.69–12.12) 28.57 (28.14–29.00) 19.05 (18.61–19.48) 4.11 (4.11–4.11)
C. phangngaensis 2 23.94 (23.81–24.03) 10.57 (9.74–11.04) 11.58 (11.47–11.69) 24.13 (24.03–24.24) 17.93 (17.75–18.18) 11.26 (11.04–11.47) 0.22 (0.22–0.22)
C. punctatonuchalis 2 15.54 (15.37–15.80) 24.68 (24.03–25.97) 25.54 (25.54–25.54) 16.81 (16.67–16.88) 25.97 (25.76–26.19) 26.30 (26.19–26.41) 25.00 (24.89–25.11) 0.00 (0.00–0.00)
C. roticanai 2 26.39 (26.19–26.62) 9.05 (8.66–9.74) 12.01 (11.90–12.12) 28.03 (27.92–28.14) 16.13 (15.80–16.45) 9.09 (8.66–9.52) 9.09 (8.87–9.31) 25.65 (25.54–25.76) 0.22 (0.22–0.22)
C. selenolagus 2 15.54 (15.15–16.02) 26.01 (25.54–26.62) 26.73 (26.41–27.06) 18.40 (18.18–18.61) 26.73 (26.19–27.27) 27.06 (26.62–27.49) 24.68 (24.24–25.11) 15.91 (15.58–16.23) 25.97 (25.54–26.41) 0.65 (0.65–0.65)
C. siamensis 2 18.92 (18.61–19.26) 25.61 (25.54–25.76) 25.32 (25.32–25.32) 19.05 (19.05–19.05) 27.56 (27.49–27.71) 27.49 (27.49–27.49) 24.78 (24.68–24.89) 18.61 (18.61–18.61) 28.03 (27.92–28.14) 20.67 (20.56–20.78) 0.00 (0.00–0.00)
C. thachanaensis 3 19.71 (19.26–20.56) 25.18 (24.89–25.54) 23.95 (23.81–24.24) 20.49 (20.13–21.21) 27.13 (26.84–27.71) 27.81 (27.49–28.35) 25.72 (25.32–26.41) 20.20 (20.13–20.35) 27.31 (27.06–27.71) 23.48 (22.94–24.24)) 13.35 (12.99–14.07) 0.72 (0.00–1.08)
C. vandeventeri 1 19.13 (18.61–19.48) 24.10 (24.03–24.24) 23.81 (23.81–23.81) 19.84 (19.70–19.91) 25.40 (25.32–25.54) 26.30 (25.76–26.84) 25.00 (24.89–25.11) 20.35 (20.35–20.35) 26.73 (26.62–26.84) 22.19 (22.08–22.29) 12.55 (12.55–12.55) 14.29 (14.07–14.72) –
Figure 2. 

The best tree resulting from Maximum Likelihood analysis of 1,327 aligned characters of the mitochondrial NADH dehydrogenase subunit 2 gene and flanking tRNAs of Cnemaspis species. Nodal support is indicated by Ultrafast bootstrap (UFB) values and Bayesian posterior probabilities (BPP) from a separate Bayesian Inference analysis, respectively. GenBank accession numbers and locality data for sequenced samples are provided in Table 2.

Taxonomic hypotheses

Cnemaspis samples from Erawan National Park, Tha Kradan Subdistrict, Si Sawat District, Kanchanaburi Province differed from congeners in mtDNA analyses and diagnostic morphological characters (see “Comparisons”). Based on these corroborating lines of evidence, we hypothesise that the Erawan National Park population represents a previously unnamed species, which is described below.

Taxonomy

Cnemaspis auriventralis sp. nov.

https://zoobank.org/899EA1FF-67AC-453A-87A0-31E55C4A242D
Figs 3, 4, 5, 6, 7, 8

Cnemaspis huaseesom Yodthong, Rujirawan, Stuart, Grismer, Aksornneam, Termprayoon, Ampai & Aowphol, 2022: 160.

Holotype

(Figs 3–6). ZMKU R 01001, adult male from Thailand, Kanchanaburi Province, Si Sawat District, Tha Kradan Subdistrict, Erawan National Park, Tham Phra That Protection Unit (14.39730N, 99.0818E; 747 m elevation), collected 18 November 2021 by Attapol Rujirawan, Siriporn Yodthong, Natee Ampai and Akrachai Aksornneam.

Figure 3. 

Adult male holotype of Cnemaspis auriventralis sp. nov. (ZMKU R 01001) in life.

Figure 4. 

Adult male holotype of Cnemaspis auriventralis sp. nov. (ZMKU R 01001) immediately after euthanasia. A. Dorsal and ventral views; B. Subcaudal region and C. Precloacal region with precloacal pores (outlined in black).

Figure 5. 

Adult male holotype of Cnemaspis auriventralis sp. nov. (ZMKU R 01001) immediately after euthanasia. A. Lateral view of head; B. Dorsal and ventral views of head (supranasal and internasal scales in dorsal view and mental, postmental and first infralabial scales in ventral view; outlined in black); C. Palmar view of the right manus; D. Plantar view of the right pes.

Figure 6. 

Dorsal and ventral views of adult male holotype of Cnemaspis auriventralis sp. nov. (ZMKU R 01001) in preservative.

Paratypes

(Figs 7, 8). Two adult males and two adult females. ZMKU R 01002 (adult female), same collection data as the holotype. ZMKU R 01003 (adult female), same collection data as the holotype, except collected 20 November 2021. ZMKU R 00999–01000 (two adult males) same collection data as the holotype, except collected 26 November 2019 by Attapol Rujirawan, Siriporn Yodthong, Korkhwan Termprayoon and Akrachai Aksornneam.

Figure 7. 

Adult female paratype of Cnemaspis auriventralis sp. nov. (ZMKU R 01002). A. Dorsolateral view in life; B. Dorsal and ventral views immediately after euthanasia; C. Dorsal and ventral views in preservative.

Figure 8. 

Cnemaspis auriventralis sp. nov. paratypes in life. A. Adult male ZMKU R 00999; B. Adult male ZMKU R 01000; C. Adult female ZMKU R 01003.

Diagnosis

Cnemaspis auriventralis sp. nov. can be distinguished from all other species in the C. siamensis group by having the following combination of morphological and colour pattern characters: SVL 36.7–38.6 mm in adult males (N = 3), 32.9–36.9 mm in adult females (N = 2); eight to ten supralabials; seven to nine infralabials; ventral scales smooth; six or seven precloacal pores in males; 16–17 paravertebral tubercles linearly arranged; tubercles on the lower flanks present; lateral caudal furrows present; no caudal tubercles in the lateral furrows; ventrolateral caudal tubercles present anteriorly; caudal tubercles not encircling tail; subcaudals smooth bearing a single median row of enlarged smooth scales; two postcloacal tubercles on each side; no shield-like subtibial scales; subtibial scales smooth; no enlarged submetatarsal scales; 23–27 subdigital lamellae on the fourth toe; sexually dimorphic in dorsal and ventral colour pattern; prescapular marking absent; gular marking absent; and yellow colouration in life on all ventral surfaces of head, body and tail in adult males.

Description of holotype

Adult male; SVL 38.0 mm; head oblong in dorsal profile, moderate in size (HL/SVL 0.28), somewhat narrow (HW/SVL 0.19), flat (HD/HL 0.40), distinct from neck; snout moderate (ES/HL 0.42), snout slightly concave in lateral profile; postnasal region concave medially; scales of rostrum round, weakly keeled, raised, larger than similarly-shaped scales on occiput; weak supraorbital ridges; weak frontorostral sulcus; canthus rostralis smoothly rounded; eye large (ED/HL 0.24); extra-brillar fringe scales small in general, but slightly larger anteriorly; pupil round; ear opening oval, taller than wide; rostral concave dorsally, dorsal 80% divided by longitudinal groove; rostral bordered posteriorly by supranasals, one small azygous internasal and nostrils; bordered laterally by first supralabials; 8R/9L (right/left) raised supralabials of similar size, but smallest posteriorly; 8R/8L infralabials, decreasing gradually in size posteriorly; nostrils small, elliptical, orientated dorsolaterally; bordered posteriorly by single, flat, enlarged postnasal scales; mental large, triangular, flat, extending to level of second infralabials, bordered posteriorly by three postmentals, medial postmental smaller than laterals; gular scales smooth, flat, round or oval, juxtaposed; throat scales smooth, raised, round, juxtaposed to subimbricate.

Body slender, elongate (AG/SVL 0.42); small, raised, weakly keeled, dorsal scales generally equal in size throughout body, intermixed with numerous, large, multi-keeled, linearly arranged tubercles; enlarged, multi-keeled, conical tubercles on flanks; tubercles extend from the occiput to base of the tail and continue on tail in whorls; body tubercles slightly smaller anteriorly; 17 paravertebral tubercles; pectoral and abdominal scales smooth, flat, imbricate; abdominal scales larger than pectoral and dorsal scales; seven contiguous, pore-bearing, precloacal scales; precloacal pores round to elongate.

Fore-limbs moderately long, slender; dorsal scales raised, weakly keeled, juxtaposed; ventral scales of brachia smooth, raised, juxtaposed; scales beneath forearm smooth, slightly raised, subimbricate; digits long with an inflected joint; claws recurved; subdigital lamellae unnotched; subdigital lamellae wide throughout length of digits, bearing a larger scale at digital inflections; interdigital webbing absent; fingers increase in length from first to fifth, with fourth and fifth nearly equal in length; relative length of fingers I < II < III < V ≤ IV; total subdigital lamellae on fingers I–V: 12–18–21–25–22 (right), 12–18–22–broken-broken (left). Hind-limbs slightly longer and thicker than fore-limbs; dorsal scales keeled, raised, juxtaposed; ventral scales of thigh and subtibial scales smooth, flat, imbricate; plantar scales smooth, raised, subimbricate; enlarged submetatarsal scales beneath first toes absent; digits elongate with an inflected joint; claws recurved; subdigital lamellae unnotched; lamellae wide throughout length of digits; enlarged scales at digital inflections; interdigital webbing absent; toes increase in length from first to fourth and fifth nearly equal in length; relative length of toes I < II < III < V ≤ IV; total subdigital lamellae on toes I–V: 11–18–22–23–22 (right), 11–17–21–24–21 (left).

Tail original (broken at tip), long, slender, TL = 45.9 mm (TL/SVL 1.21); dorsal, caudal scales arranged in segmented whorls; caudal scales keeled, raised, juxtaposed; mid-dorsal and lateral, caudal furrows present; subcaudals smooth; median row of enlarged subcaudal scales present; paravertebral, dorsolateral and lateral rows of large, keeled, caudal tubercles extend length of tail; ventrolateral rows of tubercles present only anteriorly; caudal tubercle rows do not encircle tail; tubercles absent from lateral caudal furrow; enlarged postcloacal tubercles 2R/2L on lateral surface of hemipenial swellings at base of tail.

Colouration in life

(Figs 3–5). Dorsal ground colour of head, nape and fore-limbs grey; dorsal ground colour of trunk and hind-limbs yellowish-grey; dorsal ground colour of tail yellow; top of head bearing small, diffuse, faint, dark and light markings; dark postorbital stripes faint extending to occiput; pair of dark, diffuse, blotches on nape; large, light, irregularly-shaped, vertebral blotches extend from nape to base of tail, continuing on to tail as light yellow caudal bands; small, light, irregularly-shaped blotches in shoulder regions and flanks; limbs mottled with small, diffuse light and dark blotches; digits light grey bearing thin, dark bands; all ventral surfaces of head, body and tail yellow; ventral surfaces of limbs light grey with yellow speckling.

Colouration in preservative

(Fig. 6). Dorsal and lateral surfaces of head, body, limbs and tail darker grey than in life, with some fading of markings. Ventral surfaces of head, body, limbs and tail creamy-white, with minute dark speckling on gular region, limbs and tail regions.

Variation

Cnemaspis auriventralis sp. nov. shows significant sexual dimorphism in colour pattern. All female paratypes lack yellow colouration on the tail and ventral surfaces. Female paratype ZMKU R 01003 was darker coloured in life than other members of the type series. Two male paratypes ZMKU R 00999–01000 were lighter coloured than the holotype in life. Dark markings on the dorsum of all paratypes are more prominent than the holotype. The female paratypes (ZMKU R 01002–01003) lack precloacal pores and have postcloacal tubercles that are relatively smaller than those in males. One male paratype (ZMKU R 01000) has six (2R/4L) pore-bearing precloacal scales separated by a single scale lacking pore. Variation in morphometric and meristic data amongst specimens in the type series are presented in Table 4.

Table 4.
Download as
CSV
XLSX

Descriptive measurements (millimetres) and meristics (right/left) of the type series of Cnemaspis auriventralis sp. nov. Key: M = adult male; F = adult female; Cont. = continuous; Sep. = separated; NA = data unavailable or not applicable. Morphological abbreviations are defined in Table 1.

Museum number ZMKU R 01001 ZMKU R 00999 ZMKU R 01000 ZMKU R 01002 ZMKU R 01003 All males All females
Type holotype paratype paratype paratype paratype N = 3 N = 2
Sex M M M F F Mean ± SD (min-max) Mean ± SD (min-max)
SVL 38.0 38.6 36.7 32.9 36.9 37.8 ± 0.97 (36.7–38.6) 34.9 ± 2.83 (32.9–36.9)
TW 3.3 3.1 3.4 3.3 3.0 3.3 ± 0.15 (3.1–3.4) 3.2 ± 0.21 (3.0–3.3)
TL 45.9 45.6 48.1 36.3 50.0 46.5 ± 0.21 (45.6–48.1)a 50.0b
Tail condition broken original original broken original NA NA
FL 6.4 6.4 6.2 5.8 6.3 6.3 ± 0.12 (6.2–6.4) 6.1 ± 0.35 (5.8–6.3)
TBL 8.3 8.2 8.0 7.3 8.7 8.2 ± 0.15 (8.0–8.3) 8.0 ± 0.99 (7.3–8.7)
HL 10.5 10.9 10.4 10.2 11.2 10.6 ± 0.26 (10.4–10.9) 10.7 ± 0.71 (10.2–11.2)
HW 7.2 7.3 7.2 6.3 7.3 7.2 ± 0.06 (7.2–7.3) 6.8 ± 0.71 (6.3–7.3)
HD 4.2 4.1 3.9 4.0 4.1 4.1 ± 0.15 (3.9–4.2) 4.1 ± 0.07 (4.0–4.1)
AG 16.1 16.0 16.2 15.7 16.4 16.1 ± 0.10 (16.0–16.2) 16.1 ± 0.49 (15.7–16.4)
ED 2.5 2.5 2.2 2.0 2.2 2.4 ± 0.17 (2.2–2.5) 2.1 ± 0.14 (2.0–2.2)
EE 2.8 3.0 2.8 2.7 3.1 2.9 ± 0.12 (2.8–3.0) 2.9 ± 0.28 (2.7–3.1)
EL 1.3 0.9 1.1 1.2 1.1 1.1 ± 0.20 (0.9–1.3) 1.2 ± 0.07 (1.1–1.2)
EN 3.7 3.7 3.6 3.3 3.4 3.7 ± 0.06 (3.6–3.7) 3.4 ± 0.07 (3.3–3.4)
ES 4.4 4.8 4.6 4.6 4.5 4.6 ± 0.20 (4.4–4.8) 4.6 ± 0.07 (4.5–4.6)
IO 2.3 2.4 2.5 2.2 2.2 2.4 ± 0.10 (2.3–2.5) 2.2 ± 0.00 (2.2–2.2)
IN 1.1 1.1 1.0 0.9 1.0 1.1 ± 0.06 (1.0–1.1) 1.0 ± 0.07 (0.9–1.0)
HL/SVL 0.28 0.28 0.28 0.31 0.30 0.28 ± 0.00 (0.28–0.28) 0.31 ± 0.00 (0.30–0.31)
HW/SVL 0.19 0.19 0.20 0.19 0.20 0.19 ± 0.00 (0.19–0.20) 0.19 ± 0.00 (0.19–0.20)
HD/HL 0.40 0.38 0.38 0.39 0.37 0.38 ± 0.01 (0.38–0.40) 0.38 ± 0.02 (0.37–0.39)
ES/HL 0.42 0.44 0.44 0.45 0.40 0.43 ± 0.01 (0.42–0.44) 0.43 ± 0.03 (0.40–0.45)
ED/HL 0.24 0.23 0.21 0.20 0.20 0.23 ± 0.01 (0.21–0.24) 0.20 ± 0.00 (0.20–0.20)
AG/SVL 0.42 0.41 0.44 0.48 0.44 0.43 ± 0.01 (0.41–0.44) 0.46 ± 0.02 (0.44–0.48)
TL/SVL 1.21 1.18 1.31 1.10 1.36 1.23 ± 0.07 (1.18–1.31) 1.23 ± 0.18 (1.10–1.36)
Scalation Min-max
SL 8R/9L 9R/9L 9R/9L 10R/9L 8R/8L 8–10
IL 8R/8L 9R/7L 8R/8L 9R/8L 7R/7L 7–9
PVT 17 17 16 16 16 16–17
4TL 23R/24L 25R/26L 26R/27L 26R/27L 27R/27L 23–27
PP 7 7 6 (2R/4L) absent absent 6–7
Pore arrangement Cont. Cont. Sep. NA NA NA
PPS 2R/2L 2R/2L 2R/2L 2R/2L 2R/2L 2

a Data of original tails from two males. b Data of original tail from one female.

Distribution

This species is known only from the type locality. The type series was collected from a karst formation that is part of the Tenasserim Mountain Range in Erawan National Park, Tha Kradan Subdistrict, Si Sawat District, Kanchanaburi Province, Thailand (Figs 1, 9).

Figure 9. 

Habitat of Cnemaspis auriventralis sp. nov. at the type locality at Erawan National Park, Si Sawat District, Kanchanaburi Province, Thailand. A. Microhabitat in karst boulder crevice (white arrow); B. Microhabitat on undersides of karst boulder (white arrow).

Natural history

The type locality of Cnemaspis auriventralis sp. nov. is a karst forest at 747 m elevation. The type series was collected in November 2019 at 14:00–14:30 h with temperature 25.4 °C and relative humidity 62.1% and in November 2021 at 16:00–20:00 h with temperature 25.3–26.8 °C and relative humidity 79.9–88.0%. All specimens were found on karst boulders along the nature trail to the Tham (= cave) Phra That. Most were observed clinging upside down to the undersides of large boulders (around 1–3 m2) or in narrow crevices of the boulders. Four individuals (ZMKU R 00999–01002) were found during the daytime (14:00–16:30 h) and one individual (ZMKU R 01003) was found at night (20:30 h). Most observed individuals were found during the daytime and they were active and rapidly escaped from disturbances. The one individual found at night was relatively inactive (slow-moving). At night, the new species was found to co-occur with five other species of gekkonid lizards, Cyrtodactylus monilatus Yodthong, Rujirawan, Stuart, Grismer, Aksornneam, Termprayoon, Ampai & Aowphol, 2022, Cyrtodactylus tigroides Bauer, Sumontha & Pauwels, 2003, Gehyra mutilata (Wiegmann, 1834), Dixonius hangseesom Bauer, Sumontha, Grossmann, Pauwels & Vogel, 2004 and Dixonius siamensis (Boulenger, 1899).

Etymology

The species epithet auriventralis is derived from aurum (L.) for gold and ventralis (L.) for venter in reference to the new species having distinct yellow colouration on all ventral surfaces of the head, body and tail in adult males. We propose “Erawan Rock Gecko” for the common English name and “จิ้งจกนิ้วยาวเอราวัณ” (Jing Jok Niew Yao Erawan) for the common Thai name of the new species.

Comparisons

Cnemaspis auriventralis sp. nov. is distinguishable from all other members of the C. siamensis group by a combination of morphological and colour pattern characteristics (see Table 5 for additional comparisons).

Table 5.
Download as
CSV
XLSX

Diagnostic morphological and colour pattern characteristics distinguishing Cnemaspis auriventralis sp. nov. from other species of the C. siamensis group. Grey highlight indicates differences with the new species. Key: Cont. = continuous; Sep. = separated; NA = data unavailable or not applicable.

Characters C. auriventralis sp. nov. C. adangrawi C. chanardi C. huaseesom C. kamolnorranathi C. lineatubercularis C. omari C. phangngaensis C. punctatonuchalis C. roticanai C. selenolagus C. siamensis C. thachanaensis C. vandeventeri
Morphology
Max SVL 38.6 44.9 40.1 43.5 37.8 41.8 41.3 42.0 49.6 47.0 36.2 39.7 39.0 44.7
Supralabial 8–10 10 7–10 7–10 8–9 8–9 8–9 10 8 8–9 10–11 8–9 10–11 8–9
Infralabial 7–9 9 6–8 6–9 7–8 8–9 7–8 10 7–8 7–8 10 6–8 9–11 7–9
Ventral scales smooth keeled keeled smooth weakly keeled or smooth keeled keeled keeled smooth keeled smooth keeled keeled keeled
No. of precoacal pores 6–7 6–8 6–8 5–8 7 4–7 4 4 0 3–6 6–7 0 0 4
Precloacal pores arrangement Cont. or Sep. Sep. Sep. Cont. Cont. Sep. Sep. Cont. NA Sep. Cont. NA NA Sep.
No. of paravertebral tubercles 16–17 23–25 20–30 18–24 19–24 19–21 22–29 22 24–27 25–27 16–18 19–25 15–19 25–29
Tubercles arranged linearly randomly randomly weakly linear or randomly semi-linearly linearly semi-linearly or randomly linearly semi-linearly randomly randomly randomly linearly randomly
Tubercles on lower flanks present absent present present present present generally present absent present present absent present generally present absent
Lateral caudal furrows present present present present present present present present present present absent present present present
Caudal tubercles in lateral furrow absent present absent present present absent absent absent absent absent NA absent absent absent
Ventrolateral caudal tubercles anteriorly present present absent absent absent present absent present present absent absent absent present absent
Enlarged median subcaudal scale row present absent present absent weak absent absent absent present present absent present absent present
Subcaudals smooth keeled keeled smooth keeled keeled keeled or smooth keeled smooth keeled smooth keeled keeled keeled
Single median row of subcaudals smooth keeled smooth smooth keeled keeled smooth keeled smooth keeled NA keeled keeled weakly keeled
Caudal tubercles encircle tail no no no no no no yes no no no yes no no no
No. of postcloacal tubercles in males 2 1 1 1 or 2 1 or 2 1 1 2 1–3 1 or 2 2 1 or 2 0 1–3
Subtibial scales smooth keeled keeled smooth keeled or smooth keeled keeled keeled smooth keeled smooth keeled keeled keeled
Shield-like subtibial scales absent absent absent absent absent absent absent absent absent absent absent absent absent absent
Enlarged submetatarsal scales on 1st toe absent absent absent absent absent absent absent absent absent absent present absent present absent
No. of 4th toe lamellae 23–27 26–28 25–30 21–31 24–28 27–29 25–28 29 29–31 26–29 22 24–26 23–25 24–28
Colouration and pattern
Dorsal colour pattern sexually dimorphic yes no no yes no no no yes yes yes NA no yes no
Ventral pattern sexually dimorphic yes yes yes yes NA yes yes yes yes yes NA yes yes yes
Anterior 1/2 of body orange-yellow, posterior 1/2 grey no no no no no no no no no no yes no no no
Ocelli on brachium and side of neck no no no no no no no no yes no yes no no no
Light or yellowish, prescapular crescent no yes yes no variable yes yes yes no yes yes no no yes
Fore-limbs yellow in males no no no yes no no no no no no yes no no no
Original tail yellow in males yes no no yes no no no no no no no no no no
Regenerated tail yellow no no no no no no no no NA yes no no no no
Lineate gular markings no no no no no no no no no no no yes yes no
All ventral surfaces of head, body and tail yellow in males yes no no no no no no no no yes no no no no

Cnemaspis auriventralis sp. nov. differs from C. adangrawi by having a smaller maximum SVL of 38.6 mm (vs. 44.9 mm); ventral scales smooth (vs. keeled); 16–17 paravertebral tubercles (vs. 23–25); paravertebral tubercles linearly arranged (vs. randomly); tubercles on lower flanks present (vs. absent); caudal tubercles in lateral furrow absent (vs. present); enlarged median subcaudal scales row present (vs. absent); subcaudal scales smooth (vs. keeled); single median row of subcaudals smooth (vs. keeled); two postcloacal tubercles on each side in males (vs. one); subtibial scales smooth (vs. keeled); sexual dimorphism of dorsal colour pattern present (vs. absent); light or yellowish prescapular crescent absent (vs. present); yellow colouration on original tail in males present (vs. absent); and yellow colouration on all ventral surfaces of head, body and tail in males (vs. yellowish colouration only on gular region, abdominal region and caudal region).

Cnemaspis auriventralis sp. nov. differs from C. chanardi by having ventral scales smooth (vs. keeled); 16–17 paravertebral tubercles (vs. 20–30); paravertebral tubercles linearly arranged (vs. randomly); ventrolateral caudal tubercles anteriorly present (vs. absent); subcaudal scales smooth (vs. keeled); two postcloacal tubercles on each side in males (vs. one); subtibial scales smooth (vs. keeled); sexual dimorphism of dorsal colour pattern present (vs. absent); light or yellowish prescapular crescent absent (vs. present); yellow colouration on original tail in males present (vs. absent); and yellow colouration on all ventral surfaces of head, body and tail in males (vs. yellow colouration only on gular region, belly, underside of hind-limbs and subcaudal region).

Cnemaspis auriventralis sp. nov. is most closely related in mitochondrial DNA to C. huaseesom (Fig. 2), but differs in morphology from C. huaseesom by having a smaller maximum SVL of 38.6 mm (vs. 43.5 mm); 16–17 paravertebral tubercles (vs. 18–24); caudal tubercles in lateral furrow absent (vs. present); ventrolateral caudal tubercles anteriorly present (vs. absent); enlarged median subcaudal scales row present (vs. absent); yellow dorsal colouration on fore-limbs in males absent (vs. present); and yellow colouration on all ventral surfaces of head, body and tail in males (vs. yellow colouration only on gular region, throat, pectoral region, underside of fore-limbs and subcaudal region).

Cnemaspis auriventralis sp. nov. differs from C. kamolnorranathi by having 16–17 paravertebral tubercles (vs. 19–24); caudal tubercles in lateral furrow absent (vs. present); ventrolateral caudal tubercles anteriorly present (vs. absent); subcaudal scales smooth (vs. keeled); single median row of subcaudals smooth (vs. keeled); sexual dimorphism of dorsal colour pattern present (vs. absent); yellow colouration on original tail in males present (vs. absent); and yellow colouration on all ventral surfaces of head, body and tail in males (vs. lacking yellow colouration on ventral surfaces).

Cnemaspis auriventralis sp. nov. differs from C. lineatubercularis by having ventral scales smooth (vs. keeled); 16–17 paravertebral tubercles (vs. 19–21); enlarged median subcaudal scales row present (vs. absent); subcaudal scales smooth (vs. keeled); single median row of subcaudals smooth (vs. keeled); two postcloacal tubercles on each side in males (vs. one); subtibial scales smooth (vs. keeled); sexual dimorphism of dorsal colour pattern present (vs. absent); light or yellowish prescapular crescent absent (vs. present); yellow colouration on original tail in males present (vs. absent); and yellow colouration on all ventral surfaces of head, body and tail in males (vs. yellowish colouration only on anterior gular, abdominal and subcaudal regions).

Cnemaspis auriventralis sp. nov. differs from C. omari by having ventral scales smooth (vs. keeled); 6–7 precloacal pores in males (vs. 4); 16–17 paravertebral tubercles (vs. 22–29); ventrolateral caudal tubercles anteriorly present (vs. absent); enlarged median subcaudal scales row present (vs. absent); caudal tubercles not encircling the tail (vs. encircling); two postcloacal tubercles on each side in males (vs. one); subtibial scales smooth (vs. keeled); sexual dimorphism of dorsal colour pattern present (vs. absent); light or yellowish prescapular crescent absent (vs. present); and yellow colouration on all ventral surfaces of head, body and tail in males (vs. yellow colouration only on gular region, belly, underside of hind-limbs, and subcaudal region).

Cnemaspis auriventralis sp. nov. differs from C. phangngaensis by having 7–9 infralabials (vs. 10); ventral scales smooth (vs. keeled); 6–7 precloacal pores in males (vs. 4); 16–17 paravertebral tubercles (vs. 22); tubercles on lower flanks present (vs. absent); enlarged median subcaudal scales row present (vs. absent); subcaudal scales smooth (vs. keeled); single median row of subcaudals smooth (vs. keeled); subtibial scales smooth (vs. keeled); 23–27 subdigital lamellae on the fourth toe (vs. 29); light or yellowish prescapular crescent absent (vs. present); and yellow colouration on all ventral surfaces of head, body and tail in males (vs. yellow colouration only on anterior gular region, abdomen and subcaudal region).

Cnemaspis auriventralis sp. nov. differs from C. punctatonuchalis by having a smaller maximum SVL of 38.6 mm (vs. 49.6 mm); 6–7 precloacal pores in males (vs. 0); 16–17 paravertebral tubercles (vs. 24–27); 23–27 subdigital lamellae on the fourth toe (vs. 29–31); ocelli on brachium and side of neck in males absent (vs. present); yellow colouration on original tail in males present (vs. absent); and yellow colouration on all ventral surfaces of body and tail in males (vs. orange colouration on throat and subcaudal region).

Cnemaspis auriventralis sp. nov. differs from C. roticanai by having a smaller maximum SVL of 38.6 mm (vs. 47.0 mm); ventral scales smooth (vs. keeled); 16–17 paravertebral tubercles (vs. 25–27); paravertebral tubercles linearly arranged (vs. randomly); ventrolateral caudal tubercles anteriorly present (vs. absent); subcaudal scales smooth (vs. keeled); single median row of subcaudals smooth (vs. keeled); subtibial scales smooth (vs. keeled); light or yellowish prescapular crescent absent (vs. present); yellow colouration on original tail in males present (vs. absent); and yellow colouration on regenerated tail absent (vs. present).

Cnemaspis auriventralis sp. nov. differs from C. selenolagus by having 7–9 infralabials (vs. 10); paravertebral tubercles linearly arranged (vs. randomly); tubercles on lower flanks present (vs. absent); lateral caudal furrow present (vs. absent); ventrolateral caudal tubercles anteriorly present (vs. absent); enlarged median subcaudal scales row present (vs. absent); caudal tubercles not encircling the tail (vs. encircling); enlarged submetatarsal scales on the first toe absent (vs. present); 23–27 subdigital lamellae on the fourth toe (vs. 22); orange-yellow colouration on anterior 1/2 of body in males absent (vs. present); ocelli on brachium and side of neck in males absent (vs. present); light or yellowish prescapular crescent absent (vs. present); yellow dorsal colouration on fore-limbs in male absent (vs. present); yellow colouration on original tail in males present (vs. absent); and having yellow colouration on all ventral surfaces of head, body and tail in males (vs. yellow colouration only on anterior part of body).

Cnemaspis auriventralis sp. nov. differs from C. siamensis by having ventral scales smooth (vs. keeled); 6–7 precloacal pores in males (vs. 0); 16–17 paravertebral tubercles (vs. 19–25); paravertebral tubercles linearly arranged (vs. randomly); ventrolateral caudal tubercles anteriorly present (vs. absent); subcaudal scales smooth (vs. keeled); single median row of subcaudals smooth (vs. keeled); subtibial scales smooth (vs. keeled); sexual dimorphism of dorsal colour pattern present (vs. absent); lineate gular marking absent (vs. present); and yellow colouration on all ventral surfaces of head, body and tail in males (vs. yellow colouration only on gular region, throat and pectoral region).

Cnemaspis auriventralis sp. nov. differs from C. thachanaensis by having ventral scales smooth (vs. keeled); 6–7 precloacal pores in males (vs. 0); enlarged median subcaudal scales row present (vs. absent); subcaudal scales smooth (vs. keeled); single median row of subcaudals smooth (vs. keeled); two poscloacal tubercles on each side in males (vs. 0); subtibial scales smooth (vs. keeled); lineate gular marking absent (vs. present); and yellow colouration on all ventral surfaces of head, body and tail in males (vs. yellowish-orange colouration only on gular region).

Cnemaspis auriventralis sp. nov. differs from C. vandeventeri by having a smaller maximum SVL of 38.6 (vs. 44.7 mm); ventral scales smooth (vs. keeled); 6–7 precloacal pores in males (vs. 4); 16–17 paravertebral tubercles (vs. 25–29); paravertebral tubercles linearly arranged (vs. randomly); tubercles on lower flanks present (vs. absent); ventrolateral caudal tubercles anteriorly present (vs. absent); subcaudal scales smooth (vs. keeled); single median row of subcaudals smooth (vs. weakly keeled); subtibial scales smooth (vs. keeled); sexual dimorphism of dorsal colour pattern present (vs. absent); light or yellowish prescapular crescent absent (vs. present); and yellow colouration on all ventral surfaces of head, body and tail in males (vs. orange colouration on gular region, throat, pectoral region, underside of limbs, belly and subcaudal region).

Discussion

Our phylogenetic analyses indicated that Cnemaspis auriventralis sp. nov. belongs to the C. siamensis group and is closely related to C. huaseesom from Sai Yok National Park, Kanchanaburi Province, approximately 25 km to the west. Ecologically, the new species and its close relative C. huaseesom occur in similar habitats and substrates (karst associated areas). However, the new species was found at 747 m elevation, whereas C. huaseesom was found in lowland areas (Grismer et al. 2010, 2014). Therefore, the geographic boundaries of these two species could be separated by elevation zonation (upland [> 600 m] and lowland [< 600 m] species; Grismer et al. 2014). The description of C. auriventralis sp. nov. brings the total number of Thai Cnemaspis to 21 species (Grismer et al. 2014; Uetz et al. 2022). Cnemaspis auriventralis sp. nov. is the seventh new Thai Cnemaspis species described in the last five years (Ampai et al. 2020; Grismer et al. 2020; Uetz et al. 2022). The number of known Thai Cnemaspis species continues to increase, likely as a result of new field research in poorly known areas and the use of integrative taxonomic approaches to delimit species in this genus (Wood et al. 2017; Ampai et al. 2019, 2020; Grismer et al. 2020). Additional field surveys and further taxonomic investigations using multiple lines of evidence in western Thailand are needed to determine the extent of the geographic range of the new species and to improve documentation of the herpetofaunal diversity in Thailand.

Acknowledgements

This work was financially supported by Kasetsart University Research and Development Institute (KURDI), Kasetsart University (grant no. 17.64) and the Office of the Permanent Secretary, Ministry of Higher Education, Science, Research and Innovation (grant no. RGNS 64-038). SY, KT, BLS and AA were supported by the Office of the Ministry of Higher Education, Science, Research and Innovation; and the Thailand Science Research and Innovation through the Kasetsart University Reinventing University Program 2021. NA was supported by a Srinakharinwirot University Research Grant (no. 596/2564). This research was approved by the Institutional Animal Care and Use Committee of the Faculty of Science, Kasetsart University (ACKU61-SCI-008; ACKU64-SCI-005). We would like to thank the Department of National Parks, Wildlife and Plant Conservation, Thailand for issuing research permits and, in particular, we thank Peerawat Sirothphiphat, Superintendent of Erawan National Park, for facilitating the fieldwork. Supiyanit Maiphae provided helpful suggestions for this research and Evan Quah and an anonymous reviewer improved the manuscript.

References

  • Ampai N, Rujirawan A, Wood Jr PL, Stuart BL, Aowphol A (2019) Morphological and molecular analyses reveal two new species of Cnemaspis Strauch, 1887 (Squamata, Gekkonidae) from Satun Province, southern Thailand. ZooKeys 858: 127–161. https://doi.org/10.3897/zookeys.858.34297
  • Ampai N, Wood Jr PL, Stuart BL, Aowphol A (2020) Integrative taxonomy of the rock-dwelling gecko Cnemaspis siamensis complex (Squamata, Gekkonidae) reveals a new species from Nakhon Si Thammarat Province, southern Thailand. ZooKeys 932: 129–159. https://doi.org/10.3897/zookeys.932.50602
  • Bauer AM, Giri VB, Greenbaum E, Jackman T, Dharne MS, Shouche SY (2008) On the systematics of the gekkonid genus Teratolepis Günther, 1869: Another one bites the dust. Hamadryad 32(2): 90–104.
  • Edgar RC (2004) MUSCLE: Multiple sequence alignment with high accuracy and high throughput. Nucleic Acids Research 32(5): 1792–1797. https://doi.org/10.1093/nar/gkh340
  • Grismer LL, Sumontha M, Cota M, Grismer JL, Wood Jr PL, Pauwels OS, Kunya K (2010) A revision and redescription of the rock gecko Cnemaspis siamensis (Taylor 1925) (Squamata: Gekkonidae) from Peninsular Thailand with descriptions of seven new species. Zootaxa 2576(1): 1–55. https://doi.org/10.11646/zootaxa.2576.1.1
  • Grismer LL, Wood Jr PL, Shahrul A, Awal R, Norhayati A, Muin M, Sumontha M, Grismer JL, Chan KO, Quah ES, Pauwels OSL (2014) Systematics and natural history of Southeast Asian Rock Geckos (genus Cnemaspis Strauch, 1887) with descriptions of eight new species from Malaysia, Thailand, and Indonesia. Zootaxa 3880(1): 1–147. https://doi.org/10.11646/zootaxa.3880.1.1
  • Grismer LL, Wood Jr PL, Quah ES, Anuar S, Ngadi E, Norhayati A (2015a) A new insular species of Rock Gecko (Cnemaspis Boulenger) from Pulau Langkawi, Kedah, Peninsular Malaysia. Zootaxa 3985(2): 203–218. https://doi.org/10.11646/zootaxa.3985.2.2
  • Grismer LL, Wood Jr PL, Tri N, Murdoch ML (2015b) The systematics and independent evolution of cave ecomorphology in distantly related clades of Bent-toed Geckos (Genus Cyrtodactylus Gray, 1827) from the Mekong Delta and islands in the Gulf of Thailand. Zootaxa 3980(1): 106–126. https://doi.org/10.11646/zootaxa.3980.1.6
  • Grismer LL, Yushchenko PV, Pawangkhanant P, Nazarov RA, Naiduangchan M, Suwannapoom C, Poyarkov NA (2020) A new species of Cnemaspis Strauch (Squamata: Gekkonidae) of the C. siamensis group from Tenasserim Mountains, Thailand. Zootaxa 4852(5): 547–564. https://doi.org/10.11646/zootaxa.4852.5.3
  • Hoang DT, Chernomor O, von Haeseler A, Minh BQ, Vinh LS (2018) UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35(2): 518–522. https://doi.org/10.1093/molbev/msx281
  • Kalyaanamoorthy S, Minh BQ, Wong TKF, von Haeseler A, Jermiin LS (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14(6): 587–589. https://doi.org/10.1038/nmeth.4285
  • Macey JR, Larson A, Ananjeva NB, Papenfuss TJ (1997) Evolutionary shifts in three major structural features of the mitochondrial genome among iguanian lizards. Journal of Molecular Evolution 44(6): 660–674. https://doi.org/10.1007/PL00006190
  • Miller MA, Pfeiffer W, Schwartz T (2010) Creating the CIPRES Science Gateway for inference of large phylogenetic trees. In: 2010 Gateway Computing Environments Workshop (GCE), 1–8. https://doi.org/10.1109/GCE.2010.5676129
  • Minh Q, Nguyen MAT, von Haeseler A (2013) Ultrafast approximation for phylogenetic bootstrap. Molecular Biology and Evolution 30(5): 1188–1195. https://doi.org/10.1093/molbev/mst024
  • Quah ESH, Wood Jr PL, Anuar S, Muin MA, Grismer LL (2020) A new species of Cnemaspis Strauch 1887 (Squamata: Gekkonidae) from the Langkawi Archipelago, Kedah, Peninsular Malaysia with an updated checklist of the herpetofauna of Tuba Island. Zootaxa 4767(1): 138–160. https://doi.org/10.11646/zootaxa.4767.1.6
  • Rambaut A, Drummond AJ, Xie D, Baele G, Suchard MA (2018) Posterior summarization in Bayesian phylogenetics using Tracer 1.7. Systematic Biology 67(5): 901–904. https://doi.org/10.1093/sysbio/syy032
  • Riyato A, Munir M, Martamenggala AI, Fitriana YS, Hamidy A (2019) Hiding in plain sight on Gunung Muria: a new species and first record of rock gecko (Cnemaspis Strauch, 1887; Squamata, Gekkonidae) from Java, Indonesia. Zootaxa 4608(1): 155–173. https://doi.org/10.11646/zootaxa.4608.1.9
  • Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542. https://doi.org/10.1093/sysbio/sys029
  • Simmons JE (2015) Herpetological Collecting and Collections Management, 3rd Edn. Society for the Study of Amphibians and Reptiles Herpetological Circular No. 42. Salt Lake City, UT, 191 pp.
  • Smith MA (1925) IV. – Contributions to the herpetology of Borneo. The Sarawak Museum Journal 3(8): 15–34.
  • Trifinopoulos J, Nguyen LT, von Haeseler A, Minh BQ (2016) W-IQ-TREE: A fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research 44(W1): W232–W235. https://doi.org/10.1093/nar/gkw256
  • Wilcox TP, Zwickl DJ, Heath TA, Hillis DM (2002) Phylogenetic relationships of the dwarf boas and a comparison of Bayesian and bootstrap measures of phylogenetic support. Molecular Phylogenetics and Evolution 25(2): 361–371. https://doi.org/10.1016/S1055-7903(02)00244-0
  • Wood Jr PL, Grismer LL, Aowphol A, Aguilar CA, Cota M, Grismer MS, Murdoch ML, Sites Jr JW (2017) Three new karst-dwelling Cnemaspis Strauch, 1887 (Squamata; Gekkonidae) from Peninsular Thailand and the phylogenetic placement of C. punctatonuchalis and C. vandeventeri. PeerJ 5: e2884. https://doi.org/10.7717/peerj.2884
  • Yodthong S, Rujirawan A, Stuart BL, Grismer LL, Aksornneam A, Termprayoon K, Ampai N, Aowphol A (2022) A new species in the Cyrtodactylus oldhami group (Squamata, Gekkonidae) from Kanchanaburi Province, western Thailand. ZooKeys 1103: 139–169. https://doi.org/10.3897/zookeys.1103.84672
login to comment