Zoosyst. Evol. 98 (2) 2022, 345-363 | DOI 10.3897/zse.98.89591 Ate BERLIN A new rock gecko in the Cnemaspis siamensis group (Reptilia, Gekkonidae) from Kanchanaburi Province, western Thailand Attapol Rujirawan', Siriporn Yodthong*, Natee Ampai°, Korkhwan Termprayoont, Akrachai Aksornneam’, Bryan L. Stuart*, Anchalee Aowphol! Department of Zoology, Faculty of Science, Kasetsart University, Bangkok 10900, Thailand Department of Biology, Faculty of Science, Thaksin University, Pa Phayom, Phattalung 93210, Thailand Department of Biology, Faculty of Science, Srinakharinwirot University, Bangkok 10110, Thailand North Carolina Museum of Natural Sciences, 11 West Jones Street, Raleigh, North Carolina 27601, USA FB W MY fF https://zoobank. org/05AA43 D4-8054-4884-B7E9-D5D572E6B3A6 Corresponding author: Anchalee Aowphol (fsciacl@ku.ac.th) Academic editor: Justin Bernstein # Received 28 June 2022 # Accepted 15 August 2022 Published 14 September 2022 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 (NV = 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 et al. 2022; Uetz et al. 2022). Based on molecular and morphological data, Southeast Asian Cnemaspis are re- The Southeast Asian Rock Gecko genus Cnemaspis Strauch, 1887 comprises a monophyletic clade of approx- imately 66 recognised species that are distributed from Laos, southern Vietnam westwards through southern In- dochina, 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 covered in four major monophyletic clades that contain SIX Species groups (Grismer et al. 2014). The Cnemas- pis siamensis group 1s 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 spe- cies (Grismer et al. 2010, 2014, 2020; Wood et al. 2017; Ampai et al. 2019, 2020) including C. adangrawi Ampai, Copyright Rujirawan, A. et al. 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. 346 Rujirawan, A. et al.: A new rock gecko in the Cnemaspis siamensis group from western Thailand Rujirawan, Wood, Stuart & Aowphol, 2019, C. chanar- di Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010, C. huaseesom Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010, C. kamolnor- ranathi Grismer, Sumontha, Cota, Grismer, Wood, Pau- wels & Kunya, 2010, C. lineatubercularis Ampai, Wood, Stuart & Aowphol, 2020, C. omari Grismer, Wood, An- uar, Rryanto, Ahmad, Muin, Sumontha, Grismer, Chan, Quah & Pauwels, 2014, C. phangngaensis Wood, Gris- mer, Aowphol, Aguilar, Cota, Grismer, Murdoch & Sites, 2017, C. punctatonuchalis Grismer, Sumontha, Cota, Grismer, Wood, Pauwels & Kunya, 2010, C. rot- icanai 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 zo- nation (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 speci- mens 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 speci- mens of Cnemaspis from the karst formations in Erawan National Park. Molecular and morphological analyses re- vealed 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, Kanchanab- uri Province, Thailand, in November 2019 and Novem- ber 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. Cap- tured specimens were humanely euthanised using tr- icaine 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 de- posited in the herpetological collection of the Zoologi- cal Museum, Kasetsart University, Thailand (ZMKU). Geographic coordinates and elevations were recorded zse.pensoft.net using a Garmin GPSMAP 64s with WGS84 datum. Ambient air temperature and relative humidity were col- lected 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. 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. Additional character states evaluated were the general size (1.e. strong, moderate, weak) and arrangement (i.e. random or linear) of the dorsal body tubercles; the orienta- tion and shape of precloacal pores; the number of precloa- cal scales lacking pores separating the left and right series of pore-bearing precloacal scales; the degree and arrange- ment 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 mitochon- drial 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 Mas- tercycler gradient thermocycler under the following con- ditions: initial denaturation at 95 °C for 2 min, followed Zoosyst. Evol. 98 (2) 2022, 345-363 15°N 13°N 11°N 9°N 7°N 0 50 100km —— g: 98°E 100°E Figure 1. Map illustrating the type locality (yellow star) of Cnemaspis auriventralis sp. nov. at Erawan National Park, Si Sawat 347 oh, so a Cambodia ‘ae % te ; se ‘ 4 3 ai Cnemaspis species © auriventralis sp. nov. @ adangrawi @ chanardi @ huaseesom @ kamolnorranathi @ /ineotubercularis © omari @ phangngaensis @ punctatonuchalis C) roticanai © selenolagus © siamensis © thachanaensis @ vandeventeri =e = Type locality ©) = Additional locality * = Molecular data 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. 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 visual- ly 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 con- firm the lack of premature stop codons. All new sequenc- es were deposited in GenBank under accession numbers OP093974—OP093978 (Table 2). Phylogenetic analyses Additional homologous sequences of 68 individuals be- longing to the C. affinis group, C. boulengerii group, the C. argus group, the C. chanthaburiensis group, the zse.pensoft.net 348 Rujirawan, A. et al.: A new rock gecko in the Cnemaspis siamensis group from western Thailand Table 1. 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 10 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 C. kumpoli group, the C. siamensis group and outgroups were downloaded from GenBank. Cyrtodactylus boko- rensis Murdoch, Grismer, Wood, Neang, Poyarkov, Tr1, 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 sequenc- es were aligned using the default options in the MUS- CLE (Edgar 2004) plug-in in Geneious R11 (Biomatters Ltd, Auckland, New Zealand). The aligned dataset was partitioned into four partitions consisting of 1‘-3™ 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 deter- mined using the Bayesian information criterion (BIC) im- plemented in ModelFinder (Kalyaanamoorthy et al. 2017). The best-fit evolutionary models were TPM2u+F+I+G4 for tRNAs and TVM+F+1I+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) us- ing default priors and models of evolution that were select- ed by the BIC and used in the ML analysis. Two independ- ent 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 zse.pensoft.net 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 anal- yses were visualised using FigTree v. 1.4.4 (http://tree. bio. ed.ac.uk/software/figtree/). Nodes having ultrafast boot- strap support values (UFB) > 95 and Bayesian posterior probabilities (BPP) > 0.95 were considered highly support- ed (Huelsenbeck and Ronquist 2001; Wilcox et al. 2002; Minh et al. 2013). Uncorrected pairwise sequence diver- gences (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 Bl analysis. The best tree in the ML analysis had a Maximum Likelihood value (InL) 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 Zoosyst. Evol. 98 (2) 2022, 345-363 349 Table 2. 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 Mu- seum 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 Collec- tion 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 Reference accession no. Outgroup Cyrtodactylus bokorensis Cambodia, Kampot FMNH 263228 KTO13107 ~— 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 KMO24687 — 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 Thailand, Kanchanaburi Province, Si Sawat District, Tha Kradan ZMKU R 00999 = OP093974 This study sp. nov. Subdisctrict, Erawan National Park Cnemaspis auriventralis Thailand, Kanchanaburi Province, Si Sawat District, Tha Kradan ZMKURO1000 OP093975 This study sp. nov. Subdisctrict, Erawan National Park Cnemaspis auriventralis Thailand, Kanchanaburi Province, Si Sawat District, Tha Kradan ZMKURO1001 OP093976 This study Sp. nov. Subdisctrict, Erawan National Park Cnemaspis auriventralis Thailand, Kanchanaburi Province, Si Sawat District, Tha Kradan ZMKUR0O1002 OP093977 This study sp. nov. Subdisctrict, Erawan National Park Cnemaspis auriventralis Thailand, Kanchanaburi Province, Si Sawat District, Tha Kradan ZMKUR0O1003 OP093978 This study Sp. nov. Subdisctrict, Erawan National Park 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 KMO24710 ~~ 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 Cambodia, Pursat Province, Phnom Dalai LSUHC 9338 KM024716 — Grismer et al. (2014) chanthaburlensis Cnemaspis grismeri Malaysia, Perak, Lenggong LSUHC 9969 KMO24722 —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 KMO24736 — 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 Thailand, Nakhon Si Thammarat Province, Lan Saka District, Wang Mai ZMKURO0825 MT112890 Ampai et al. (2020) lineatubercularis Pak Waterfall Cnemaspis Thailand, Nakhon Si Thammarat Province, Lan Saka District, Wang Mai ZMKUR0OO0828 £MT112891 Ampai et al. (2020) lineatubercularis Pak Waterfall Cnemaspis Thailand, Nakhon Si Thammarat Province, Lan Saka District, Wang Mai ZMKUR0O0829 MT112892 Ampai et al. (2020) lineatubercularis Pak Waterfall Cnemaspis lineogularis Thailand, Prachuap Khiri Khan Eee Kui Buri District, Wat Khao BYU 62535 KY091231 Wood et al. (2017) aeng Cnemaspis lineogularis Thailand, Prachuap Khiri Khan aes Kui Buri District, Wat Khao = ZMKUROO728 KY091233 Wood et al. (2017) aeng Cnemaspis mahsuriae Malaysia, Kedah, Pulau Langkawi,Gunung Raya LSUHC 11829 KT250634 ~~ Grismer et al. (2015a) Cnemaspis mcguirei Malaysia, Perak, Bukit Larut LSUHC 8853 KMO24751 ~~ Grismer et al. (2014) Cnemaspis monachorum Malaysia, Kedah, Langkawi Archipelago, Pulau Langkawi LSUHC 9114 KMO24754 ~~ Grismer et al. (2014) Cnemaspis monachorum Malaysia, Kedah, Langkawi Archipelago, Pulau Langkawi LSUHC 10807 KM024755 ~~ Grismer et al. (2014) Cnemaspis Malaysia, Perak, Belum-Temengor, Sungai Enam USMHC 1347 KMO24762 ~— Grismer et al. (2014) narathiwatensis Cnemaspis Malaysia, Perak, Belum-Temengor, Sungai Enam USMHC 1348 KMO24763 — Grismer et al. (2014) narathiwatensis 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) zse.pensoft.net 350 Species Cnemaspis niyomwanae Cnemaspis nuicamensis Cnemaspis nuicamensis Cnemaspis omari Cnemaspis omari Cnemaspis perhentianensis Cnemaspis phangngaensis Cnemaspis phangngaensis Cnemaspis punctatonuchalis Cnemaspis punctatonuchalis Cnemaspis roticanai Cnemaspis roticanai Cnemaspis selenolagus Cnemaspis selenolagus Cnemaspis siamensis Cnemaspis siamensis Cnemaspis tarutaoensis Cnemaspis tarutaoensis Cnemaspis tarutaoensis Cnemaspis thachanaensis Cnemaspis thachanaensis Cnemaspis thachanaensis Cnemaspis tubaensis Cnemaspis tubaensis Cnemaspis tucdupensis Cnemaspis tucdupensis Cnemaspis vandeventeri Rujirawan, A. et al.: A new rock gecko in the Cnemaspis siamensis group from western Thailand Locality Voucher GenBank Reference accession no. Thailand, Trang Province, Tham Khao Ting LSUHC 9571 KMO24774 — Grismer et al. (2014) Vietnam, An Giang Province, Nui Cam Hill LSUHC 8646 KM024775 — Grismer et al. (2014) Vietnam, An Giang Province, Nui Cam Hill LSUHC 8647 KM024776 ~~ Grismer et al. (2014) Thailand, Satun Province, Phuphaphet Cave LSUHC 9565 KM024780 ~~ Grismer et al. (2014) Malaysia, Perlis, Perlis State Park LSUHC 9978 KM024779 — Grismer et al. (2014) Malaysia, Terengganu, Pulau Perhentian Besar LSUHC 8699 KM024820 _ Grismer et al. (2014) Thailand, Phangnga Province, Mueang Phangnga District, Knao Chang, BYU 62537 KY091234 Wood et al. (2017) Phung Chang Cave Thailand, Phangnga Province, Mueang Phangnga District, Khao Chang, BYU 62538 KY091235 Wood et al. (2017) Phung Chang Cave Thailand, Prachaup Khiri Khan Province, Thap Sakae BYU 62539 KY091236 Wood et al. (2017) Thailand, Prachaup Khiri Khan Province, Thap Sakae BYU 62540 KY091237 Wood et al. (2017) Malaysia, Kedah, Pulau Langkawi, Gunung Raya LSUHC 9430 KM024829 — Grismer et al. (2014) Malaysia, Kedah, Pulau Langkawi, Gunung Raya LSUHC 9431 KM024830 — Grismer et al. (2014) Thailand, Ratchaburi Province, Suan Phueng District BH asa MW051887 — Grismer et al. (2020) Thailand, Ratchaburi Province, Suan Phueng District AUP-00767 MW051888 — Grismer et al. (2020) Thailand, Chumpon Province, Pathio District LSUHC 9474 KM024838 — Grismer et al. (2014) Thailand, Chumpon Province, Pathio District LSUHC 9485 KM024839 — Grismer et al. (2014) Thailand, Satun Province, Mueang Satun District, Tarutao Island ZMKUROO761 MK862117 Ampai et al. (2019) Thailand, Satun Province, Mueang Satun District, Tarutao Island ZMKUR 00763 = =MK862118 Ampai et al. (2019) Thailand, Satun Province, Mueang Satun District, Tarutao Island ZMKU R 00764 MK862119 Ampai et al. (2019) Thailand, Surat Thani Province, aan District, Tham Khao Sonk BYU 62542 KY091239 Wood et al. (2017) Thailand, Surat Thani Province, viasvalg District, Tham Khao Sonk BYU 62543 KY091243 Wood et al. (2017) Thailand, Surat Thani Province, heen District, Tham Khao Sonk BYU 62544 KY091244 Wood et al. (2017) West Malaysia, Kedah, Langkawi Archipelago, Tuba Island USMHC 2527 MT028175 Quah et al. (2020) West Malaysia, Kedah, Langkawi Archipelago, Tuba Island USMHC 2528 MT028176 Quah et al. (2020) Vietnam, An Giang Province, Tuc Dup Hill LSUHC 8631 KM024852 ~~ Grismer et al. (2014) Vietnam, An Giang Province, Tuc Dup Hill LSUHC 8632 KM024853 — Grismer et al. (2014) Thailand, Ranong Province, Suk Saran District, Naka BYU 62541 KY091238 Wood et al. (2017) 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). Taxonomic hypotheses Cnemaspis samples from Erawan National Park, Tha Kra- dan Subdistrict, Si Sawat District, Kanchanaburi Province differed from congeners in mtDNA analyses and diagnos- tic 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/899EA 1 FF-67AC-453A-87A0-31E55C4A242D Figs 3-8 Cnemaspis huaseesom Yodthong, Rujirawan, Stuart, Grismer, Aksorn- neam, Termprayoon, Ampai & Aowphol, 2022: 160. zse.pensoft.net 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, Siri- porn Yodthong, Natee Ampai and Akrachai Aksornneam. 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 No- vember 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. Diagnosis. Cnemaspis auriventralis sp. nov. can be distinguished from all other species in the C. siamensis group by having the following combination of morpho- logical and colour pattern characters: SVL 36.7—38.6 mm in adult males (NV = 3), 32.9-36.9 mm in adult females (N = 2); eight to ten supralabials; seven to nine infralabi- als; ventral scales smooth; six or seven precloacal pores in males; 16—17 paravertebral tubercles linearly arranged; tubercles on the lower flanks present; lateral caudal fur- rows 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 post- Zoosyst. Evol. 98 (2) 2022, 345-363 Sheu EU268364 Hemidactylus gamotii KT013107 Cyrtodactylus bokorensis KM024710 Cnemapis boulengeri KM024711 Cnemapis boulengeri ee | chanthaburiensis group ZMKU R 00999 Cnemaspis < ZMKU R 01000 Cnemaspis a ZMKU R 01003 Cnemaspis ai ZMKU R 01001 Cnemaspis a ZMKU R 01002 Cnemaspis 66/0.91 KM024734 Cnemaspis huaseesom & f KM024735 Cnemaspis huaseesom 79/1 KM024733 Cnemaspis huaseesom aS MW051887 Cnemaspis selenolagus MW051888 Cnemaspis selenolagus KY091236 Cnemaspis punctatonuchalis KY091237 Cnemaspis punctatonuchalis KY091238 Cnemaspis vandeventeri KM024838 Cnemaspis siamensis KM024839 Cnemaspis siamensis KY091243 Cnemaspis thachanaensis f KY091239 Cnemaspis thachanaensis KY091244 Cnemaspis thachanaensis MT112892 Cnemaspis lineatubercularis P MT112890 Cnemaspis lineatubercularis ' M7T112891 Cnemaspis lineatubercularis KM024829 Cnemaspis roticanai KM024830 Cnemaspis roticanai MK862114 Cnemaspis adangrawi MK862112 Cnemaspis adangrawi MK862113 Cnemaspis adangrawi siamensis group Nodal support = UFB/BPP @ urs>95andBPP>0.95 — leon 58/ — 34/— ae’ 86/0.87 65/0.99 0.2 subsititutions/site KM024779 Cnemaspis omari KM024780 Cnemaspis omari KM024715 Cnemaspis chanardi KY091234 Cnemaspis phangngaensis KY091235 Cnemaspis phangngaensis ——__| affinis group <—_| argus group ———— | :sOkumpoli group Figure 2. The best tree resulting from Maximum Likelihood analysis of 1,327 aligned characters of the mitochondrial NADH dehy- drogenase 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. cloacal tubercles on each side; no shield-like subtibial scales; subtibial scales smooth; no enlarged submetatar- sal 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 C(HD/HL 0.40), distinct from neck; snout moderate (ES/HL 0.42), snout slightly concave in lateral profile; postnasal re- gion concave medially; scales of rostrum round, weakly keeled, raised, larger than similarly-shaped scales on oc- ciput; weak supraorbital ridges; weak frontorostral sul- cus; 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% di- vided 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 posterior- ly; 8R/8L infralabials, decreasing gradually in size poste- riorly; nostrils small, elliptical, orientated dorsolaterally; bordered posteriorly by single, flat, enlarged postnasal scales; mental large, triangular, flat, extending to lev- el of second infralabials, bordered posteriorly by three postmentals, medial postmental smaller than laterals; gu- lar 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, mul- ti-keeled, linearly arranged tubercles; enlarged, mul- ti-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 pec- toral 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 zse.pensoft.net 352 Rujirawan, A. et al.: A new rock gecko in the Cnemaspis siamensis group from western Thailand Table 3. 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 i! 2 = = el £ & Nd © 2 3 3 Es eT) 2 ® o oe 8 2 25 2 Sa 3 S Ss s 3s” © . = ® G 8 i S = oO C. auriventralis 5 0.30 sp. nov. (0.00- 0.65) C. adangrawi a 25.67 3.03 (25.11- (0.00- 25.97) 4.55) C. chanardi le 26.75 11.40 - (25.97- (11.02- 26.41) 12.12) C. huaseesom B= "L229 26.26 26.62 0.43 (12.12— (26.19- (26.62- (0.00- 12:55). -26:41)° 26:62) 0.65) ( SF. 25195 AA97 1332 27.95 0.29 lineatubercularis (24.89- (17.32- (17.10- (27.71- (0.22- 25.54) 19.05) 17753) 28.35) 0.43) C. omari 2 27.68 9.63 11.90 283.57 19.05 (27.27- (8.23- (11.69- (28.14- (18.61- 2E QZ «= M1039) 2 S212) 29.00) 19.48) C. phangngaensis 2 23.94 10:57 11.58 24.13 17.93 (23.81- (9.74- (11.47- (24.03- (17.75- 24.03) 11.04) 11.69) 24.24) 18.18) G: 2 15.54 24.68 25-54. 16.81 25.97 punctatonuchalis (15.37— (24.03- (25.54- (16.67- (25.76- 15.80) 25.97) 25.54) 16.88) 26.19) C. roticanai 2 26.39 9.05 12.01 28.03 16.13 (26.19- (8.66- (11.90- (27.92- (15.80- 26.62) 9.74) 1212) 28.14) 16.45) C. selenolagus 2 1554 26.01 26.73 18.40 26.73 (15.15- (25.54- (26.41- (18.18- (26.19- 16.02) 26.62) 27.06) 18.61) 27.27) C. siamensis 2 18.92 25.61 25.32 19.05 27.56 (18.61- (25.54- (25.32- (19.05- (27.49- 19.26) 25.76) 25.32) 19.05) 274M) C. thachanaensis 3 19.71 25.18 23.95 20.49 27.13 (19.26- (24.89- (23.81- (20.13- (26.84- 20.56) 25.54) 24.24) 21.21) 27.71) C. vandeventeri 1 19.13 24.10 23.81 19.84 25.40 (18.61- (24.03- (23.81- (19.70- = (25.32- 19.48) 24.24) 23.81) 19.91) 25.54) 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 through- out length of digits, bearing a larger scale at digital in- flections; 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 3 oD = a ig S © o [s) = > £ S 2 S < £ © ® S 2 s ° & S ce : a) = ® w = o = = w = § a =] 1S) A (o) = f 5 Qa (Ss) oO (6) 2. G 4.11 (4.11- 4.11) 11.26 0.22 (11.04- (0.22- 11.47) 0.22) 26.30 25.00 0.00 (26.19- (24.89- (0.00- 26.41) 25.11) 0.00) 9.09 9.09 25.65 0.22 (8.66- (8.87- (25.54— (0.22- 9.52) 9.31) 25.76) 0.22) 27.06 24.68 15.91 25.97 0.65 (26.62— (24.24— (15.58- (25.54- (0.65- 27.49) 25.11) 16.23) 26.41) 0.65) 27.49 24.78 18.61 28.03 20.67 0.00 (27.49- (24.68- (18.61- (27.92- (20.56- (0.00- 27.49) 24.89) 18.61) 28.14) 20.78) 0.00) 2F 31: 2572 20.20 ae Re al 23.48 13335 0.72 (27.49- (25.32- (20.13- (27.06- (22.94- (12.99- (0.00- 28.35) 26.41) 20.35) 27.71) 24.24)) 14.07) 1.08) 26.30 25.00 20.35 26.73 22.19 12.55 14.29 - 25.76- (24.89- (20.35- (26.62- (22.08- (12.55- (14.07- 26.84) 25.11) 20.35) 26.84) 22.29) 12.55) 14.72) digital 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 < I] < II < V Data of original tail from one female. = English name and “avaninenaasviam” (Jing Jok Niew Yao Erawan) for the common Thai name of the new species. Comparisons. Cnemaspis auriventralis sp. nov. 1s dis- tinguishable from all other members of the C. siamensis group by a combination of morphological and colour pat- tern characteristics (see Table 5 for additional comparisons). Cnemaspis auriventralis sp. nov. differs from C. adan- grawi by having a smaller maximum SVL of 38.6 mm (vs. 44.9 mm); ventral scales smooth (vs. keeled); 16-17 par- avertebral tubercles (vs. 23-25); paravertebral tubercles linearly arranged (vs. randomly); tubercles on lower flanks present (vs. absent); caudal tubercles in lateral furrow ab- sent (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); sub- tibial scales smooth (vs. keeled); sexual dimorphism of dorsal colour pattern present (vs. absent); light or yellow- ish prescapular crescent absent (vs. present); yellow col- ouration 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); subcau- dal 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 cres- cent 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 relat- ed 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 zse.pensoft.net 360 Rujirawan, A. et al.: A new rock gecko in the Cnemaspis siamensis group from western Thailand 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 gu- lar region, throat, pectoral region, underside of fore-limbs and subcaudal region). Cnemaspis auriventralis sp. nov. differs from C. ka- molnorranathi 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 di- morphism 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 coloura- tion on ventral surfaces). Cnemaspis auriventralis sp. nov. differs from C. linea- tubercularis by having ventral scales smooth (vs. keeled); 16-17 paravertebral tubercles (vs. 19-21); enlarged me- dian subcaudal scales row present (vs. absent); subcaudal scales smooth (vs. keeled); single median row of subcau- dals smooth (vs. keeled); two postcloacal tubercles on each side in males (vs. one); subtibial scales smooth (vs. keeled); sexual dimorphism of dorsal colour pattern pres- ent (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. om- ari 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 an- teriorly present (vs. absent); enlarged median subcaudal scales row present (vs. absent); caudal tubercles not en- circling 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 cres- cent absent (vs. present); and yellow colouration on all ventral surfaces of head, body and tail in males (vs. yel- low 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); sub- caudal 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 sur- faces of head, body and tail in males (vs. yellow coloura- zse.pensoft.net tion only on anterior gular region, abdomen and subcau- dal 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. pres- ent); yellow colouration on original tail in males present (vs. absent); and yellow colouration on all ventral surfac- es of body and tail in males (vs. orange colouration on throat and subcaudal region). Cnemaspis auriventralis sp. nov. differs from C. roti- canai by having a smaller maximum SVL of 38.6 mm (vs. 47.0 mm); ventral scales smooth (vs. keeled); 16—17 par- avertebral tubercles (vs. 25—27); paravertebral tubercles linearly arranged (vs. randomly); ventrolateral caudal tu- bercles 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. pres- ent); 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. sele- nolagus 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 anteri- orly 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. pres- ent); 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. siamen- sis by having ventral scales smooth (vs. keeled); 6—7 pre- cloacal pores in males (vs. 0); 16-17 paravertebral tuber- cles (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); sub- tibial scales smooth (vs. keeled); sexual dimorphism of dor- sal 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 colour- ation 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 me- dian subcaudal scales row present (vs. absent); subcaudal scales smooth (vs. keeled); single median row of subcaudals Zoosyst. Evol. 98 (2) 2022, 345-363 361 Table 5. 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. = sepa- rated; NA = data unavailable or not applicable. Characters Morphology Max SVL Supralabial Infralabial Ventral scales No. of precoacal pores Precloacal pores arrangement No. of paravertebral tubercles Tubercles arranged Tubercles on lower flanks Lateral caudal furrows Caudal tubercles in lateral furrow Ventrolateral caudal tubercles anteriorly Enlarged median subcaudal scale row Subcaudals Single median row of subcaudals Caudal tubercles encircle tail No. of postcloacal tubercles in males Subtibial scales ShieldHike subtibial scales Enlarged submetatarsal scales on 1‘ toe No. of 4" toe lamellae C. auriventralis sp. nov. 38.6 8-10 7-9 smooth 6-7 Cont. or Sep. 16-17 linearly present present absent present present smooth smooth no smooth absent absent 23-27 Colouration and pattern Dorsal colour pattern sexually dimorphic Ventral pattern sexually dimorphic Anterior 1/2 of body orange- yellow, posterior 1/2 grey Ocelli on brachium and side of neck Light or yellowish, prescapular crescent yes yes no no no C. adangrawi 44.9 10 9 keeled 6-8 Sep. Zo=eD C. chanardi 40.1 7-10 6-8 keeled 6-8 Sep. 20-30 C. huaseesom ALSy AS) 7-10 6-9 smooth 5-8 Cont. 18-24 randomly randomly weakly absent present present present absent keeled keeled no keeled absent absent 26-28 no yes no no yes present present absent absent present keeled smooth no keeled absent absent 25-30 no yes no no yes linear or randomly present present present absent absent smooth smooth no lor2 smooth absent absent 21-31 yes yes no no no C. kamolnorranathi 37.8 8-9 7-8 weakly keeled or smooth 7 Cont. 19-24 sem- linearly present present present absent weak keeled keeled no lor2 keeled or smooth absent absent 24-28 no NA no no variable C. lineatubercularis 41.8 8-9 8-9 keeled 4-7 Sep. 19-21 linearly present present absent present absent keeled keeled no keeled absent absent 27-29 no yes no no yes C. omari 41.3 8-9 7-8 keeled Ae) sem- linearly or randomly generally present present absent absent absent keeled or smooth smooth yes 1 keeled absent absent 25-28 no yes no no yes C. phangngaensis 42.0 10 10 keeled Cont. aes linearly absent present absent present absent keeled keeled no keeled absent absent ee) yes yes no no yes C. punctatonuchalis 49.6 8 7-8 smooth 24-27 sem linearly present present absent present present smooth smooth no 1-3 smooth absent absent 29-31 yes yes no yes no C. roticanai 47.0 8-9 7-8 keeled 3-6 Sep. 25-27 randomly present present absent absent present keeled keeled no lor2 keeled absent absent 26-29 yes yes no no yes C. selenolagus 36.2 10-11 10 smooth 6-7 Cont. 16-18 randomly randomly absent absent NA absent absent smooth NA yes smooth absent present 22 NA NA yes yes yes C. siamensis 39,7 8-9 6-8 keeled NA foo23 present present absent absent present keeled keeled no lor2 keeled absent absent 24-26 no yes no no no 2 nse 2 Sg gs 3 S I a oss 3 oO 39.0 44.7 10-11 8-9 9-11 7-9 keeled keeled 0 4 NA Sep. 15-19 25-29 linearly randomly generally absent present present present absent absent present absent absent present keeled keeled keeled weakly keeled no no 0 1-3 keeled keeled absent absent present absent 23-25 24-28 yes no yes yes no no no no no yes zse.pensoft.net 362 Characters = 2 o P=] he = = ey E = $ £ 3 c 5 a £ © ee ee 2 o g 2 & > = a ao} S © [s) ~~ zo © = S o GS S ? S £ = cS) cS) Forelimbs yellow in no no no yes no no males Original tail yellow yes no no yes no no in males Regenerated tail no no no no no no yellow Lineate gular no no no no no no markings All ventral surfaces yes no no no no no of head, body and tail yellow in males 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. van- deventeri by having a smaller maximum SVL of 38.6 (vs. 44.7 mm); ventral scales smooth (vs. keeled); 6-7 preclo- acal 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. ab- sent); 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 pres- capular crescent absent (vs. present); and yellow coloura- tion 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 au- riventralis sp. nov. belongs to the C. siamensis group and is closely related to C. huaseesom from Sai Yok Nation- al Park, Kanchanaburi Province, approximately 25 km to the west. Ecologically, the new species and its close relative C. huaseesom occur in similar habitats and sub- strates (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 spe- cies 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 (Gris- mer et al. 2014; Uetz et al. 2022). Cnemaspis auriventra- lis sp. nov. 1s the seventh new Thai Cnemaspis species de- scribed in the last five years (Ampai et al. 2020; Grismer et al. 2020; Uetz et al. 2022). The number of known Thai zse.pensoft.net Rujirawan, A. et al.: A new rock gecko in the Cnemaspis siamensis group from western Thailand ” 2 = 2 = 2 — — S A 2 ® <; o S © i) a o = S ob = 5 7" Fi s g E © £ S 2 £ © 3 S 2 $ 3 D S Ss zg a] = o 7) 3 = e) ry = o = SY a =| (S) i (S) ~ . a. {S) ro (S) 2 G no no no no yes no no no no no no no no no no no no no NA yes no no no no no no no no no yes yes no no no no yes no no no no 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 Universi- ty Research and Development Institute (KURDD, 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 Thai- land Science Research and Innovation through the Kaset- sart 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 Fac- ulty of Science, Kasetsart University (ACKU61-SCI-008; ACKU64-SCI-005). We would like to thank the Department of National Parks, Wildlife and Plant Conservation, Thai- land for issuing research permits and, in particular, we thank Peerawat Sirothphiphat, Superintendent of Erawan National Park, for facilitating the fieldwork. Supryanit Maiphae pro- vided 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 Zoosyst. 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