Research Article |
Corresponding author: Anchalee Aowphol ( fsciacl@ku.ac.th ) Academic editor: Aaron Bauer
© 2019 Natee Ampai, Attapol Rujirawan, Perry L. Wood Jr, Bryan L. Stuart, Anchalee Aowphol.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Ampai N, Rujirawan A, Wood Jr PL, Stuart BL, Aowphol A (2019) Morphological and molecular analyses reveal two new insular species of Cnemaspis Strauch, 1887 (Squamata, Gekkonidae) from Satun Province, southern Thailand. ZooKeys 858: 127-161. https://doi.org/10.3897/zookeys.858.34297
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We describe two new insular gecko species of the genus Cnemaspis from Tarutao, Adang, and Rawi islands in Satun Province, southern Thailand. The new species are distinguished from their congeners in having a unique combination of morphological, scalation, and color pattern characters, and by genetic divergence in the mitochondrial NADH dehydrogenase subunit 2 (ND2) gene. Cnemaspis tarutaoensis sp. nov. was found to be a member of the C. kumpoli group, but is distinguished from all other species in that group by having 8–9 supralabials and 8 infralabials; 4–5 pore-bearing precloacal scales, pores rounded; 17–19 paravertebral tubercles randomly arranged; 27–29 subdigital lamellae under the fourth toe; subcaudal region yellowish, with smooth scales and a single enlarged median row; black gular markings in males and females; and 17.24–22.36% uncorrected pairwise sequence divergences. Cnemaspis adangrawi sp. nov. was found to be a member of the C. siamensis group, but is distinguished from all other species in that group by having 10 supralabials and 9 infralabials; 6–8 pore-bearing precloacal scales, pores rounded and arranged in a chevron shape; 23–25 randomly arranged, separated paravertebral tubercle rows; 26–28 subdigital lamellae under the fourth toe; subcaudal scales keeled, without enlarged median row; gular region, abdomen, limbs and subcaudal region yellowish in males only; gular marking absent in males and females; and 8.30–26.38 % uncorrected pairwise sequence divergences. Cnemaspis tarutaoensis sp. nov. occurs in karst formations on Tarutao Island, while Cnemaspis adangrawi sp. nov. is found near granitic, rocky streams on Adang and Rawi islands.
Island, rock geckos, species diversity, systematics
Southeast Asia is a global biodiversity hotspot with extraordinary levels of species endemism (
The rock gecko genus Cnemaspis Strauch, 1887 currently contains 57 recognized species distributed throughout Southeast Asia (
During recent fieldwork in 2017–2018 on Tarutao, Adang, and Rawi islands, Satun Province, southern Thailand, specimens of Cnemaspis were collected that differed from all other named species. Herein, we evaluate the morphological and molecular distinctiveness of these specimens.
Specimens of Cnemaspis were collected from Tarutao, Adang, and Rawi islands in Tarutao National Park, Mueang Satun District, Satun Province, Thailand (Fig.
Map illustrating the holotype locality (red star) and paratype localities (red circles) of Cnemaspis tarutaoensis sp. nov. at Tarutao Island, Satun Province, Thailand; the holotype locality (green star) and paratype localities (green circles) of Cnemaspis adangrawi sp. nov. at Adang and Rawi islands, Mueang Satun District, Satun Province, Thailand.
Only adult individuals were used in the morphological analysis, as determined by the presence of hemipenes or precloacal pores in males, and the presence of calcium glands or eggs in females. Measurements were taken by the first author on the left side of preserved specimens to the nearest 0.1 mm using digital calipers under a Nikon SMZ 445 dissecting microscope. Sixteen measurements were taken following
Meristic characters of scale counts and external observations of morphology were taken following
Comparative material was examined in the holdings of THNHM (Appendix
Genomic DNA was extracted from liver tissue of eight individuals of Cnemaspis (Table
Samples used in this study, including catalogue numbers, Genbank accession numbers and localities of voucher specimens. Voucher abbreviations are as follows: 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, Penang, Malaysia (USMHC), and Zoological Museum of Kasetsart University (ZMKU).
Species | Locality | Collection no. | Genbank accession no. | Reference |
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Cyrtodactylus intermedius | Cambodia, Kampot | FMNH 263228 | KT13107 |
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Hemidactylus garnotii | Myanmar, Mon State, Kyaihto Township, Kyait Hti Yo Wildlife Sanctuary. | CAS 222276 | EU68364 |
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Cnemaspis adangrawi sp. nov. | Thailand, Satun Province, Mueang Satun District, Adang Island | ZMKU R 00767 | MK862112 | This study |
Thailand, Satun Province, Mueang Satun District, Adang Island | THNHM 28207 | MK862113 | This study | |
Thailand, Satun Province, Mueang Satun District, Adang Island | ZMKU R 00770 | MK862114 | This study | |
Thailand, Satun Province, Mueang Satun District, Rawi Island | ZMKU R 00775 | MK862115 | This study | |
Thailand, Satun Province, Mueang Satun District, Rawi Island | ZMKU R 00776 | MK862116 | This study | |
Cnemaspis affinis | Malaysia, Penang, Pulau Pinang | LSUHC 6787 | KM024682 |
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Cnemaspis argus | Malaysia, Terengganu, Gunung Lawit | LSUHC 8304 | KM024687 |
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Malaysia, Terengganu, Gunung Lawit | LSUHC 10834 | KM024688 |
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Cnemaspis aurantiacopes | Vietnam, Kien Giang Province, Hon Dat Hill | LSUHC 8610 | KM024692 |
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Vietnam, Kien Giang Province, Hon Dat Hill | LSUHC 8611 | KM024693 |
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Cnemaspis biocellata | Malaysia, Perlis, Kuala Perlis | LSUHC 8817 | KM024707 |
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Malaysia, Perlis, Kuala Perlis | LSUHC 8817 | KM024708 |
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Malaysia, Perlis, Gua Kelam | LSUHC 8789 | KM024709 |
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Cnemaspis boulengerii | Vietnam, Ca Mau Province, Con Dao Archipelago | LSUHC9278 | KM024710 |
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Vietnam, Ca Mau Province, Con Dao Archipelago | LSUHC9279 | KM024711 |
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Cnemaspis caudanivea | Vietnam, Kien Giang Province, Hon Tre Island | LSUHC 8582 | KM024714 |
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Cnemaspis chanardi | Thailand, Nakhon Si Thammarat Province, Thum Thong Panra | LSUHC 9567 | KM024715 |
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Cnemaspis chanthaburiensis | Cambodia, Pursat Province, Phnom Dalai | LSUHC 9338 | KM024716 |
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Cnemaspis grismeri | Malaysia, Perak, Lenggong | LSUHC 9969 | KM024722 |
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Cnemaspis hangus | Malaysia, Pahang, Bukit Hangus | LSUHC 9358b | KM024728 |
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Cnemaspis harimau | Malaysia, Kedah, Gunung Jeri | LSUHC 9665 | KM024730 |
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Cnemaspis huaseesom | Thailand, Kanchanaburi Province, Sai Yok National Park | LSUHC 9455 | KM024733 |
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Thailand, Kanchanaburi Province, Sai Yok National Park | LSUHC 9457 | KM024734 |
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Thailand, Kanchanaburi Province, Sai Yok National Park | LSUHC 9458 | KM024735 |
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Cnemaspis karsticola | Malaysia, Kelantan, Gunung Reng | LSUHC 9054 | KM024736 |
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Malaysia, Kelantan, Gunung Reng | LSUHC 9055 | KM024737 |
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Cnemaspis kumpoli | Malaysia, Perlis, Perlis State Park | LSUHC 8847 | KM024745 |
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Malaysia, Perlis, Perlis State Park | LSUHC 8848 | KM024746 |
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Cnemaspis lineogularis | Thailand, Prachuap Khiri Khan Province, Kui Buri District, Wat Khao Daeng | BYU 62535 | KY091231 |
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Thailand, Prachuap Khiri Khan Province, Kui Buri District, Wat Khao Daeng | ZMKU R 00728 | KY091233 |
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Cnemaspis mahsuriae | Malaysia, Kedah, Pulau Langkawi, Gunung Raya | LSUHC 11829 | KT250634 |
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Cnemaspis mcguirei | Malaysia, Perak, Bukit Larut | LSUHC 8853 | KM024751 |
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Cnemaspis monachorum | Malaysia, Kedah, Langkawi Archipelago, Pulau Langkawi | LSUHC 9114 | KM024754 |
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Malaysia, Kedah, Langkawi Archipelago, Pulau Langkawi | LSUHC 10807 | KM024755 |
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Cnemaspis narathiwatensis | Malaysia, Perak, Belum-Temengor, Sungai Enam | USMHC 1347 | KM024762 |
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Malaysia, Perak, Belum-Temengor, Sungai Enam | USMHC 1348 | KM024763 |
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Cnemaspis neangthyi | Cambodia, Pursat Province, O’Lakmeas | LSUHC 8515 | KM024767 |
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Cambodia, Pursat Province, O’Lakmeas | LSUHC 8516 | KM024768 |
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Cnemaspis niyomwanae | Thailand, Trang Province, Thum Khao Ting | LSUHC 9568 | KM024773 |
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Thailand, Trang Province, Thum Khao Ting | LSUHC 9571 | KM024774 |
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Cnemaspis nuicamensis | Vietnam, An Giang Province, Nui Cam Hill | LSUHC 8646 | KM024775 |
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Vietnam, An Giang Province, Nui Cam Hill | LSUHC 8647 | KM024776 |
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Vietnam, An Giang Province, Nui Cam Hill | LSUHC 8648 | KM024777 |
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Cnemaspis omari | Thailand, Satun Province, Phuphaphet Cave | LSUHC 9565 | KM024780 |
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Malaysia, Perlis, Perlis State Park | LSUHC 9978 | KM024779 |
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Cnemaspis perhentianensis | Malaysia, Terengganu, Pulau Perhentian Besar | LSUHC 8699 | KM024820 |
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Cnemaspis phangngaensis | Thailand, Phangnga Province, Mueang Phangnga District, Khao Chang, Phung Chang Cave | BYU 62537 | KY091234 |
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Thailand, Phangnga Province, Mueang Phangnga District, Khao Chang, Phung Chang Cave | BYU 62538 | KY091235 |
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Cnemaspis punctatonuchalis | Thailand, Prachaup Khiri Khan Province, Thap Sakae | BYU 62539 | KY091236 |
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Thailand, Prachaup Khiri Khan Province, Thap Sakae | BYU 62540 | KY091237 |
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Cnemaspis roticanai | Malaysia, Kedah, Pulau Langkawi, Gunung Raya | LSUHC 9430 | KM024829 |
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Malaysia, Kedah, Pulau Langkawi, Gunung Raya | LSUHC 9431 | KM024830 |
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Malaysia, Kedah, Pulau Langkawi, Gunung Raya | LSUHC 9439 | KM024831 |
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Cnemaspis siamensis | Thailand, Chumpon Province, Pathio District | LSUHC 9474 | KM024838 |
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Thailand, Chumpon Province, Pathio District | LSUHC 9485 | KM024839 |
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Cnemaspis tarutaoensis sp. nov. | Thailand, Satun Province, Mueang Satun District, Tarutao Island | ZMKU R 00761 | MK862117 | This study |
Thailand, Satun Province, Mueang Satun District, Tarutao Island | ZMKUR 00763 | MK862118 | This study | |
Thailand, Satun Province, Mueang Satun District, Tarutao Island | ZMKU R 00764 | MK862119 | This study | |
Cnemaspis thachanaensis | Thailand, Surat Thani Province, Tha Chana District, Tham Khao Sonk Hill | BYU 62542 | KY091239 |
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Thailand, Surat Thani Province, Tha Chana District, Tham Khao Sonk Hill | BYU 62543 | KY091243 |
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Thailand, Surat Thani Province, Tha Chana District, Tham Khao Sonk Hill | BYU 62544 | KY091244 |
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Cnemaspis tucdupensis | Vietnam, An Giang Province, Tuc Dup Hill | LSUHC 8631 | KM024852 |
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Vietnam, An Giang Province, Tuc Dup Hill | LSUHC 8632 | KM024853 |
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Cnemaspis vandeventeri | Thailand, Ranong Province, Suk Saran District, Naka | BYU 62541 | KY091238 |
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Homologous sequences of 56 Cnemaspis and the outgroups Cyrtodactylus intermedius and Hemidactylus garnotii (following
Phylogenies were reconstructed with the maximum likelihood (ML) criterion using IQ-TREE v. 1.6.7 (
Phylogenies were also reconstructed with Bayesian Inference (BI) using MrBayes v. 3.2 on XSEDE on the Cyberinfrastructure for Phylogenetic Research (CIPRES;
The most likely tree in the ML analysis, and the 50% majority-rule consensus of the sampled trees from the BI analysis, were visualized using FigTree v. 1.4.3 (
The aligned dataset contained 1,296 characters of 64 individuals of Cnemaspis and two individuals of the outgroup species. The standard deviation of split frequencies among the two simultaneous BI runs was 0.001478. The ESS values were greater than or equal to 3,630 for all parameters. A single most likely tree resulted from the ML analysis.
The most likely ML tree and the 50% majority rule consensus tree from the BI analysis had similar topologies (Fig.
The single best maximum likelihood tree of the mitochondrial NADH dehydrogenase subunit 2 (ND2) gene and flanking tRNAs from geckos of the genera Cnemaspis, Cyrtodactylus and Hemidactylus, shown in full view (above) and close-up view of relevant clades (below). Support values at nodes are bootstrap values from a maximum likelihood analysis of the same dataset followed by posterior probabilities of the Bayesian analysis.
Mean (minimum–maximum) uncorrected p-distances (%) within the Cnemaspis kumpoli group based on 1,296 bp of the mitochondrial ND2 gene and flanking tRNAs. Numbers in bold are within species divergence. n = number of individuals.
No. | Species | n | 1 | 2 | 3 | 4 | 5 |
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1 | C. tarutaoensis sp. n. | 3 | 0.16 (0.00–0.31) | ||||
2 | C. monachorum | 2 | 17.70 (17.24–18.17) | 0.54 (0.00–1.09) | |||
3 | C. biocellata | 3 | 20.34 (20.19–20.50) | 19.10 (19.79–19.41) | 3.57 (0.00–7.14) | ||
4 | C. kumpoli | 2 | 21.84 (21.74–21.89) | 22.28 (22.95–22.52) | 13.51 (13.51–13.51) | 0.16 (0.00–0.31) | |
5 | C. niyomwanae | 2 | 21.35 (20.34–22.36) | 21.20 (19.88–22.36) | 14.44 (13.20–15.68) | 12.89 (11.49–14.29) | 1.79 (0.00–3.57) |
Cnemaspis samples from Adang and Rawi islands represented a well-supported lineage (100 BS, 1.0 PP) within the siamensis group, and was recovered as being closely related to a clade containing C. chanardi, C. phangngaensis, C. omari, and C. roticanai (Fig.
Mean (minimum-maximum) uncorrected p-distances (%) within the Cnemaspsis siamensis group based on 1,296 bp of the mitochondrial ND2 gene and flanking tRNAs. Numbers in bold are within species divergence. n = number of individuals.
No. | Species | n | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 |
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1 | C. adangrawi sp. nov. | 5 | 2.81 | |||||||||
(0.00–4.68) | ||||||||||||
2 | C. chanardi | 1 | 11.40 | 0.00 | ||||||||
(10.85–11.91) | ||||||||||||
3 | C. omari | 2 | 9.36 | 11.81 | 2.13 | |||||||
(8.30–10.21) | (11.49–12.13) | (0.00–4.26) | ||||||||||
4 | C. phangngaensis | 2 | 10.19 | 11.38 | 11.17 | 0.11 | ||||||
(9.57–10.85) | (11.27–11.49) | (10.85–11.49) | (0.00–0.21) | |||||||||
5 | C. siamensis | 2 | 25.83 | 24.40 | 27.77 | 25.00 | 0.00 | |||||
(25.74–25.96) | (24.26–24.68) | (27.66–27.87) | (24.89–25.11) | |||||||||
6 | C. roticanai | 3 | 8.92 | 11.77 | 9.01 | 8.90 | 28.16 | 0.11 | ||||
(8.51–9.57) | (11.70–11.91) | (8.72–9.36) | (8.72–9.15) | (28.09–28.30) | (0.00–0.21) | |||||||
7 | C. vandeventeri | 1 | 24.26 | 24.04 | 26.60 | 25.21 | 12.34 | 26.88 | 0.00 | |||
(24.04–24.47) | (24.04–24.04) | (25.96–27.23) | (25.11–25.32) | (12.34–12.34) | (26.81–27.02) | |||||||
8 | C. thachanaensis | 3 | 25.50 | 24.40 | 28.30 | 26.13 | 13.35 | 27.66 | 14.47 | 0.53 | ||
(25.10–25.96) | (24.26–24.68) | (27.23–28.94) | (25.74–26.81) | (13.19–14.26) | (27.45–28.09) | (14.26–14.89) | (0.00–1.06) | |||||
9 | C. punctatonuchalis | 2 | 25.23 | 25.53 | 26.38 | 25.00 | 19.36 | 25.60 | 21.06 | 21.13 | 0.00 | |
(24.04–26.17) | (25.53–25.53) | (26.38–26.38) | (25.00–25.00) | (19.36–19.36) | (25.53–25.74) | (21.06–21.06) | (21.06–21.28) | |||||
10 | C. huaseesom | 3 | 26.00 | 26.17 | 28.19 | 23.72 | 19.36 | 27.52 | 20.64 | 20.99 | 16.95 | 0.43 |
(25.74–26.38) | (26.17–26.17) | (27.87–28.51) | (23.62–23.83) | (19.36–19.36) | (27.45–27.66) | (20.64–20.64) | (20.64–21.70) | (16.81–17.02) | (0.00–0.64) |
The Tarutao and Adang-Rawi samples of Cnemaspis differed from each other and all other congeners by being diagnosable in morphology and mitochondrial DNA. Owing to these independent lines of evidence, we hypothesize that the Tarutao and Adang-Rawi samples represent two new species to science, and are described below.
(Figs
(Figs
Paratypes of Cnemaspis tarutaoensis sp. nov. in preservative. A dorsal view B ventral view; from left to right, top panel (females): ZMKU R 00758, ZMKU R 00759, ZMKU R 00760, ZMKU R 00766, THNHM 28201, and THNHM 28202; bottom panel (males): ZMKU R 00761, ZMKU R 00762, ZMKU R 00765, ZMKU R 00764, THNHM 28203, and THNHM 28205.
THNHM 28204 (one juvenile), same data as holotype except collected 5 April 2018.
Cnemaspis tarutaoensis sp. nov. can be distinguished from all other Cnemaspis by having the following combination of characters: (1) adult males with maximum snout-vent length (SVL) 36.4 mm (mean 34.7 ± SD 1.5, n = 7) and females with maximum SVL 34.8 mm (mean 33.7 ± SD 0.6, n = 6); (2) 8–9 supralabials and 8 infralabials; (3) 4–5 pore-bearing precloacal scales, pores rounded; (4) 17–19 paravertebral tubercles, small in size, randomly arranged; (5) 27–29 subdigital lamellae under the 4th toe; (6) subcaudal region yellowish, scales smooth with a single enlarged median subcaudal row; (7) one postcloacal tubercles on each side; (8) no sexual dimorphism in dorsal and ventral patterns; and (9) black gular markings present in males and females. These differences are summarized for geographically close congeners in the kumpoli group (Table
Meristic character states and color patterns of species in the Cnemaspis kumpoli group. Measurements are taken in millimeters and measurement abbreviations are defined in the text. var = character variable; – = data unavailable, ant = anterior.
Characters/Species | C. tarutaoensis sp. nov. | C. biocellata | C. kumpoli | C. monachorum | C. niyomwanae |
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Sample size | 13 | 25 | 13 | 12 | 5 |
Maximum SVL | 36.4 | 40.2 | 63.0 | 32.9 | 56.8 |
Supralabial scales | 8–9 | 6–10 | 7–9 | 7–8 | 8–11 |
Infralabial scales | 8 | 5–9 | 6–8 | 5–7 | 6–8 |
Ventral scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 |
No. of precloacal pores | 4–5 | 6–12 | 1–8 | 3 | 3 |
Precloacal pore continuous (1) or separated (0) | 0.1 | 1 | 0 | 1 | 0.1 |
No. of paravertebral tubercles | 17–19 | 21–27 | 28–35 | 11–20 | 26–31 |
Tubercles present (1) or absent (0) on lower flanks | 0 | 1 | 1 | 0 | 0 |
No. of 4th toe lamellae | 27–29 | 29–37 | 34–41 | 24–30 | 31–34 |
Lateral caudal furrows present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 |
Lateral caudal tubercle row present (1) or absent (0) | 0 | ant | 0 | ant | 0 |
Subcaudal scales keeled (1) or smooth (0) | 0 | 0 | 0 | 0 | 0 |
Enlarge submetatarsal scales on 1st toe (1) or not (0) | 0 | 0 | 0 | 0 | 0 |
Enlarge median subcaudal scales row (1) or not (0) | 1 | 1 | 1 | 1 | 1 |
No. of postcloacal tubercles in males | 1 | 1 | 2.3 | 1–2 | 1.2 |
Subcaudal region yellow present (1) or not (0) | 1 | var | 0 | 0 | 0 |
Ventral pattern sexually dimorphic present (1) or not (0) | 0 | 1 | 0 | 1 | – |
Dorsal color pattern sexually dimorphic (1) or not (0) | 0 | 1 | 1 | 0 | 1 |
Wide black and yellow bands on tail present (1) or not (0) | 1 | 0 | 0 | 0 | 0 |
Gular marking (1) or not (0) | 1 | 0 | 0 | 1 | 0 |
Adult male; SVL 36.3 mm; head moderate in size (HL/SVL 0.28), elongate, narrow (HW/SVL 0.15), flattened (HD/HL 0.33), distinct from neck; snout moderate (ES/HL 0.42), in lateral view slightly concave; postnasal region constricted medially; scales of rostrum, raised, smooth, larger than conical scales on occiput; faint supraorbital ridges; gular and throat scales raised, smooth and round; shallow frontorostral sulcus; canthus rostralis nearly absent, smoothly rounded; eye large (ED/HL 0.18); pupil round; ear opening oval, taller than wide; rostral slightly concave, dorsal 80% divided by longitudinal median groove; rostral bordered posteriorly by supranasal and laterally by first supralabial; 9, 9 (right, left) slightly raised supralabials decreasing in size posteriorly; 8, 8 (right, left) infralabials decreasing in size posteriorly; nostril elliptical, oriented posterodorsally, bordered posteriorly by small, granular postnasal scales; mental large, triangular, bordered posteriorly by three large postmentals.
Body slender, elongate (AG/SVL 0.39); small, raised and equal in sized, dorsal scales throughout body intermixed with several large, multicarinate tubercles randomly arranged; 19 paravertebral tubercles; tubercles absent on lower flanks; tubercles extend from occiput to base of tail; dorsal scales slightly raised and keeled; pectoral and abdominal scales smooth and round, flat to concave, slightly larger than dorsal scales and not larger posteriorly; ventral scales of brachia raised, smooth and juxtaposed; four pores-bearing precloacal scales arranged in a chevron, separated; precloacal pore rounded; precloacal depression absent; femoral pores absent.
Fore and hind limbs moderately long, slender; scales beneath forearm slightly raised, smooth and subimbricate; subtibial scales keeled; palmar scales smooth and juxtaposed; digits elongate, slender, inflected joint and bearing slightly recurved claws; subdigital lamellae unnotched; lamellae beneath first phalanges wide; lamellae beneath phalanx immediately following inflection granular; lamellae of distal phalanges wide; lamellae beneath inflection large; interdigital webbing absent; enlarge submetatarsal scales on 1st toe absent; fingers increase in length from first to fourth with fourth and fifth nearly equal in length; relative length of fingers IV>V>III>II>I; toes increase in length from first to fifth with fourth and fifth nearly equal in length; relative length of toes IV>V>III>II>I; total number of subdigital lamellae on 4th toe 28, 28 (right, left).
Caudal and subcaudal scales smooth, similar to dorsal scale size; lateral caudal furrow present; lateral caudal tubercle row absent; enlarge caudal tubercles at the base of tail not encircling tail; enlarged median subcaudal scales row present; tail length (TL) 34.3 mm with broken at tail tip; enlarged postcloacal tubercle 1, 1 (right, left) on lateral surface of hemipenial swellings at the base of tail.
(Figs
(Fig.
Most paratypes approximate the holotype in general aspects of color pattern (Fig.
Descriptive measurements in millimeters and characters of the type series of Cnemaspis tarutaoensis sp. nov. M = male; F = female; – = data unavailable or absent; b = broken; r = regenerated.
Museum number | ZMKU | ZMKU | ZMKU | ZMKU | ZMKU | THNHM | THNHM | ZMKU | ZMKU | THNHM | ZMKU | ZMKU | THNHM |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|
R 00763 | R 00761 | R 00762 | R 00764 | R 00765 | 28203 | 28205 | R 00766 | R 00758 | 28201 | R 00759 | R 00760 | 28202 | |
Type series | Holotype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype |
Sex | M | M | M | M | M | M | M | F | F | F | F | F | F |
SVL | 36.3 | 33.3 | 35.2 | 32.6 | 35.2 | 33.8 | 36.4 | 33.3 | 34.8 | 33.4 | 33.5 | 33.8 | 33.6 |
TL | 34.3b | 8.7b | 42.3r | b | 40.8r | b | 17.3b | 47.7 | 45.3 | 50.3 | 52.4 | 44.1r | 13.6b |
TW | 3.5 | 3.4 | 3.5 | 3.2 | 3.5 | 3.2 | 3.6 | 3.5 | 3.8 | 3.5 | 3.4 | 3.4 | 3.3 |
FL | 5.3 | 5.1 | 5.2 | 5.1 | 5.3 | 5.1 | 5.3 | 5.2 | 5.2 | 5.0 | 5.1 | 5.1 | 5.1 |
TBL | 6.3 | 6.2 | 6.3 | 6.0 | 6.3 | 6.2 | 6.3 | 6.0 | 6.3 | 6.0 | 6.1 | 6.2 | 6.1 |
AG | 14.3 | 14.1 | 14.3 | 14.1 | 14.3 | 14.1 | 14.3 | 14.3 | 14.2 | 14.0 | 14.1 | 14.0 | 14.0 |
HL | 10.0 | 9.8 | 10.1 | 9.7 | 10.0 | 9.6 | 10.1 | 9.5 | 9.9 | 9.7 | 9.7 | 9.7 | 9.9 |
HW | 5.6 | 5.4 | 5.5 | 5.4 | 5.6 | 5.3 | 5.6 | 5.5 | 5.6 | 5.2 | 5.3 | 5.4 | 5.3 |
HD | 3.3 | 3.1 | 3.2 | 3.2 | 3.3 | 3.2 | 3.3 | 3.2 | 3.2 | 3.0 | 3.1 | 3.2 | 3.0 |
ED | 1.8 | 1.9 | 1.9 | 1.8 | 1.8 | 1.8 | 1.9 | 1.8 | 1.8 | 1.8 | 1.9 | 1.9 | 1.9 |
EE | 2.9 | 2.9 | 3.0 | 2.9 | 3.0 | 2.9 | 3.0 | 2.8 | 2.8 | 2.7 | 2.8 | 2.8 | 2.8 |
ES | 4.2 | 4.1 | 4.1 | 4.0 | 4.2 | 4.0 | 4.2 | 4.1 | 4.2 | 4.0 | 4.1 | 4.0 | 4.0 |
EN | 3.5 | 3.2 | 3.4 | 3.2 | 3.5 | 3.2 | 3.6 | 3.4 | 3.4 | 3.2 | 3.2 | 3.3 | 3.2 |
IO | 2.2 | 2.1 | 2.2 | 2.0 | 2.1 | 2.0 | 2.2 | 1.9 | 2.2 | 2.1 | 2.1 | 2.1 | 2.0 |
EL | 0.6 | 0.7 | 0.7 | 0.6 | 0.7 | 0.6 | 0.7 | 0.6 | 0.7 | 0.7 | 0.7 | 0.6 | 0.6 |
IN | 0.8 | 0.7 | 0.8 | 0.7 | 0.8 | 0.8 | 0.8 | 0.7 | 0.7 | 0.7 | 0.8 | 0.8 | 0.7 |
Supralabials | 9 | 9 | 8 | 9 | 9 | 9 | 9 | 9 | 8 | 8 | 8 | 8 | 9 |
Infralabials | 8 | 8 | 8 | 8 | 8 | 8 | 8 | 8 | 8 | 8 | 8 | 8 | 8 |
No. of precloacal pores | 4 | 4 | 4 | 4 | 4 | 4 | 5 | – | – | – | – | – | – |
Precloacal pore continuous (1) or separated (0) | 0 | 0 | 0 | 0 | 0 | 0 | 1 | – | – | – | – | – | – |
No. of paravertebral tubercles | 19 | 19 | 18 | 18 | 19 | 18 | 19 | 17 | 18 | 19 | 18 | 19 | 18 |
No. of 4th toe lamellae | 28 | 29 | 29 | 29 | 29 | 29 | 28 | 29 | 27 | 27 | 27 | 27 | 28 |
Gular marking (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Cnemaspis tarutaoensis sp. nov. is known only from the type locality on Tarutao Island, approximately 40 km off the coast of Thailand. All specimens were found in karst forest near mangroves and karst outcrops near a stream (Fig.
Paratypes found during the day (ZMKU R 00759 and 00760, ZMKU R 00762 and 00763, ZMKU R 00765–00766, and THNHM 28202–28204) were in shaded areas, cracks, and crevices of rock boulders. When disturbed, some individuals would retreat into cracks and crevices, or hide in shaded areas of the rock boulder. Paratypes found at night (ZMKU R 00758, ZMKU R 00761, ZMKU R 00764, THNHM 28201 and THNHM 28205) were in deep crevices, within cracks on the shaded (by day) surfaces of boulders, or perched on vegetation near karst. Three gravid females (ZMKU R 00758, ZMKU R 00760, and THNHM 28202) contained two eggs during November 2017. THNHM 28204 (juvenile) was observed on vegetation near a rock boulder on 5 April 2018. At night, Cyrtodactylus cf. astrum was found in syntopy on rock boulders and karst formations with C. tarutaoensis sp. nov.
The specific epithet refers to the type locality of the new species.
Cnemaspis tarutaoensis sp. nov. can be distinguished from all other members of the kumpoli group (C. biocellata, C. kumpoli, C. monachorum, and C. niyomwanae) by having a maximum SVL of 36.4 mm (vs 32.9 mm in C. monachorum, 40.2 mm in C. biocellata, 63.0 mm in C. kumpoli, and 56.8 mm in C. niyomwanae).
Cnemaspis tarutaoensis sp. nov. is further distinguished from C. monachorum by having eight infralabial scales (vs 5–7 in C. monachorum). The new species is further distinguished from C. biocellata, C. monachorum and C. niyomwanae by having 4–5 precloacal pores (vs 6–12 in C. biocellata and three in C. monachorum and C. niyomwanae). The new species is further distinguished from C. biocellata, C. kumpoli, and C. niyomwanae by having 17–19 paravertebral tubercles (vs 21–27 in C. biocellata, 28–35 in C. kumpoli and 26–31 in C. niyomwanae). The new species is further distinguished from C. biocellata and C. kumpoli by lacking tubercles on lower flanks (vs present in C. biocellata and C. kumpoli). The new species is further distinguished from C. biocellata, C. kumpoli and C. niyomwanae by having 26–29 lamellae under the 4th toe (vs 29–37 in C. biocellata, 34–41 in C. kumpoli, and 31–34 in C. niyomwanae).
Cnemaspis tarutaoensis sp. nov. is further distinguished from C. kumpoli, C. monachorum and C. niyomwanae by having yellow coloration in the subcaudal region and wide black and yellow bands on tail (vs lacking in C. kumpoli, C. monachorum, and C. niyomwanae). The new species is further distinguished from C. biocellata, C. kumpoli, and C. niyomwanae by lacking a sexually dimorphic dorsal color pattern (vs present in C. biocellata, C. kumpoli, and C. niyomwanae). The new species is further distinguished from C. monachorum and C. biocellata by lacking lateral caudal tubercle row (vs present in C. monachorum and C. biocellata). The new species is distinguished from C. biocellata, C. kumpoli, and C. niyomwanae by having gular marking (vs lacking in C. biocellata, C. kumpoli, and C. niyomwanae).
(Figs
(Figs
ZMKU R 00772 and THNHM 28212–28215 (five juveniles), same data as holotype except collected 6 April 2018.
Cnemaspis adangrawi sp. nov. can be distinguished from all other Cnemaspis by having the following combination of characters: (1) adult males with maximum SVL length 44.9 mm (mean 41.8 ± SD 2.5, n = 11) and females with maximum SVL 43.8 mm (mean 42.5 ± SD 1.5, n = 4); (2) 10 supralabials and 9 infralabials; (3) 6–8 pore-bearing precloacal scales with rounded pores arranged in chevron shape and separated; (4) 23–25 paravertebral tubercles randomly arranged; (5) 26–28 subdigital lamellae under the 4th toe; (6) subcaudal scales keeled and lacking enlarge median row; (7) one postcloacal tubercle each side; (8) gular region, abdomen, limbs and subcaudal region yellowish in males only; (9) mid-gular marking absent in males and females. These differences are summarized for geographically close congeners in the siamensis group (Table
Meristic character state and color pattern of species in the Cnemapsis siamensis group. Measurements are taken in millimeters and measurement abbreviations are defined in the text. – = data unavailable, w = weak.
Characters/Species | C. adangrawi sp. nov. | C. chanardi | C. huaseesom | C. omari | C. phangngaensis | C. punctatonuchalis | C. roticanai | C. siamensis | C. thachanaensis | C. vandeventeri |
---|---|---|---|---|---|---|---|---|---|---|
Sample size | 15 | 25 | 5 | 8 | 2 | 5 | 8 | 12 | 6 | 3 |
Maximum SVL | 44.9 | 40.9 | 43.5 | 41.3 | 42.0 | 49.6 | 47.0 | 39.7 | 39.0 | 44.7 |
Supralabial scales | 10 | 8–10 | 7–10 | 8–9 | 10 | 8 | 8–9 | 8–9 | 10–11 | 8,9 |
Infralabial scales | 9 | 8 | 6–9 | 7–8 | 10 | 7–8 | 7–8 | 6–8 | 9–11 | 7–9 |
Ventral scales keeled (1) or smooth (0) | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 1 |
No. of precloacal pores | 6–8 | 6–8 | 5–8 | 3–6 | 4 | 0 | 3–6 | 0 | 0 | 4 |
Precloacal pore continuous (1) or separated (0) | 0 | 0 | 1 | 0 | 1 | – | 0 | – | – | 0 |
Precloacal pores elongate (1) or round (0) | 0 | 0 | 0 | 0 | 0 | – | 0 | – | – | 0 |
No. of paravertebral tubercles | 23–25 | 22–25 | 18–24 | 22–29 | 22 | 24–27 | 25–27 | 19–25 | 15–19 | 25–29 |
Paravertebral tubercles linearly arranged (1) or more random (0) | 0 | 0 | w,0 | w,0 | 1 | w | 0 | 0 | 1 | 0 |
Tubercles present (1) or absent (0) on lower flanks | 0 | 1 | 1 | w,1 | 0 | 1 | 1 | 1 | 1 | 1 |
No. of 4th toe lamellae | 26–28 | 26–29 | 21–31 | 25–28 | 29 | 29–31 | 26–29 | 24–26 | 24 | 24–28 |
Lateral caudal furrows present (1) or absent (0) | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 0 |
Subcaudal keeled (1) or smooth (0) | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 1 | 1 | 1 |
Enlarge median subcaudal scales row (1) or not (0) | 0 | 1 | 0 | 0 | 0 | 1 | w | 1 | 0 | 1 |
No. of postcloacal tubercles in males | 1 | 1 | 1,2 | 1 | 2 | 1–3 | 1–2 | 1–2 | 0 | 1–3 |
Subtibial scales keeled (1) or smooth (0) | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | 1 |
Subcaudal region yellow present (1) or not (0) | 1 | 1 | 1 | 1 | 1 | 0 | 1 | 0 | 0 | 0 |
Ventral pattern sexually dimorphic present (1) or not (0) | 1 | 1 | 1 | – | 0 | 1 | 1 | 1 | 1 | 1 |
Dorsal color pattern sexually dimorphic (1) or not (0) | 0 | 0 | 1 | 0 | 0 | 1 | 1 | 0 | 0 | 0 |
Lineate gular marking (1) or not (0) | 0 | 0 | 0 | 0 | 0 | – | 0 | 1 | 1 | 0 |
Adult male; SVL 44.6 mm; head moderately sized (HL/SVL 0.26), narrow (HW/SVL 0.15), flattened (HD/HL 0.38), and head distinct from neck; snout moderate (ES/HL 0.47), snout slightly concave in lateral view; postnasal region constricted medially; scales of rostrum smooth, larger than conical scales on occiput; weak supraorbital ridges; lineate gular marking absent; gular and throat scales raised, keeled and round; shallow frontorostral sulcus; canthus rostralis nearly absent, smoothly rounded; eye large (ED/HL 0.20); pupil round; ear opening oval, taller than wide; rostral slightly concave; rostral bordered posteriorly by supranasals; 10, 10 (right, left) supralabials decreasing in size posteriorly; 9, 9 (right, left) infralabials decreasing in size posteriorly; nostril elliptical, oriented posterodorsally, bordered by small postnasal scales; mental large, triangular, concave bordered posteriorly by three large postmentals.
Body slender, elongate (AG/SVL 0.42); small, keeled, dorsal scales equal in size throughout body intermixed with several large, keeled, multicarinate tubercles randomly arranged; 24 paravertebral tubercles; tubercles absent on lower flanks; tubercles extend from occiput to base of tail; dorsal scales raised and keeled; pectoral and abdominal scales keeled, round, flat to concave, slightly larger than dorsal and not larger posteriorly; ventral scales of brachia smooth, raised and juxtaposed; eight separated pore-bearing precloacal scales with rounded pores; precloacal depression absent; femoral pores absent.
Fore and hind limbs moderately long, slender; scales beneath forearm slightly raised, smooth and subimbricate; subtibial scales keeled; palmar scales smooth and juxtaposed; digits elongate, slender, inflected joint and bearing slightly recurved claws; subdigital lamellae unnotched; lamellae beneath first phalanges wide; lamellae beneath phalanx immediately following inflection granular; lamellae of distal phalanges wide; lamellae beneath inflection large; interdigital webbing absent; enlarged submetatarsal scales on 1st toe absent; fingers increase in length from first to fourth with fourth and fifth nearly equal in length; relative length of fingers IV>V>III>II>I; toes increase in length from first to fifth with fourth and fifth nearly equal in length; relative length of toes IV>V>III>II>I; total subdigital lamellae on 4th toe 28, 28 (right, left).
Caudal and subcaudal scales keeled, similar to dorsal scale size; lateral caudal furrow present; enlarge caudal tubercles arranged in segmented whorls, not encircling tail; enlarge median subcaudal scales row absent; caudal tubercles present on lateral furrow; tail length (TL) 58.3 mm with regenerated tail; enlarge, flat, postcloacal tubercle 1, 1 (right, left) on lateral surface of hemipenial swellings at the base of tail.
(Figs
(Fig.
Most paratypes approximate the holotype in general aspects of morphology (Figs
Descriptive measurements in millimeters and characters of the type series of Cnemaspis adangrawi sp. nov. M = male; F = female; – = data unavailable or absent; b = broken; r = regenerated.
Museum number | ZMKU | ZMKU | THNHM | THNHM | THNHM | ZMKU | THNHM | ZMKU | ZMKU | THNHM | ZMKU | ZMKU | ZMKU | ZMKU | THNHM |
R 00767 | R 00769 | 28206 | 28207 | 28208 | R 00770 | 28209 | R 00773 | R 00775 | 28210 | R 00776 | R 00768 | R 00771 | R 0074 | 28211 | |
Type series | Holotype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype | Paratype |
Sex | M | M | M | M | M | M | M | M | M | M | M | F | F | F | F |
SVL | 44.6 | 44.9 | 37.9 | 39.7 | 43.7 | 42.2 | 41.5 | 42.8 | 42.3 | 37.6 | 43.1 | 40.6 | 43.8 | 43.7 | 41.9 |
TL | 58.3r | 55.7 | 47.7 | 53.1 | 39.7r | 56.5 | 49.8 | 56.2 | 51.0 | 45.6r | 42.3r | 50.5 | 50.3 | 42.6 | 40.5b |
TW | 4.3 | 4.3 | 3.9 | 3.8 | 4.3 | 4.1 | 4.0 | 4.1 | 4.0 | 3.8 | 4.2 | 4.0 | 4.1 | 4.3 | 4.1 |
FL | 6.6 | 6.6 | 5.9 | 6.0 | 6.6 | 6.4 | 6.3 | 6.4 | 6.4 | 5.9 | 6.5 | 6.2 | 6.5 | 6.5 | 6.4 |
TBL | 8.5 | 8.5 | 7.7 | 8.1 | 8.5 | 8.4 | 8.3 | 8.3 | 8.4 | 7.6 | 8.4 | 8.1 | 8.4 | 8.4 | 8.3 |
AG | 18.8 | 18.7 | 15.2 | 17.2 | 18.5 | 18.3 | 18.3 | 18.4 | 18.3 | 15.2 | 18.5 | 18.3 | 18.6 | 18.6 | 18.4 |
HL | 11.6 | 11.6 | 10.9 | 10.9 | 11.6 | 11.2 | 11.0 | 11.2 | 11.2 | 10.6 | 11.4 | 11.0 | 11.1 | 11.3 | 11.1 |
HW | 7.0 | 7.0 | 6.4 | 6.5 | 7.0 | 6.8 | 6.7 | 6.8 | 6.8 | 6.4 | 6.9 | 6.6 | 6.7 | 6.9 | 6.8 |
HD | 4.5 | 4.2 | 3.9 | 3.9 | 4.5 | 4.3 | 4.0 | 4.5 | 4.5 | 2.8 | 4.5 | 4.2 | 4.4 | 4.5 | 4.4 |
ED | 2.4 | 2.4 | 2.1 | 1.9 | 2.4 | 2.2 | 2.1 | 2.3 | 2.3 | 2.0 | 2.4 | 2.2 | 2.4 | 2.4 | 2.2 |
EE | 3.7 | 3.8 | 3.2 | 3.3 | 3.6 | 3.8 | 3.6 | 3.8 | 3.7 | 3.3 | 3.7 | 3.5 | 3.8 | 3.8 | 3.7 |
ES | 5.5 | 5.4 | 4.5 | 4.6 | 5.2 | 5.2 | 5.0 | 5.1 | 5.1 | 4.4 | 5.1 | 5.0 | 5.2 | 5.2 | 5.0 |
EN | 4.4 | 4.3 | 3.3 | 3.4 | 4.1 | 4.0 | 4.0 | 4.2 | 4.1 | 3.2 | 4.2 | 4.2 | 4.1 | 4.2 | 4.1 |
IO | 2.9 | 2.9 | 2.6 | 2.6 | 2.9 | 2.8 | 2.7 | 2.8 | 2.8 | 2.7 | 2.8 | 2.7 | 2.9 | 2.8 | 2.7 |
EL | 0.9 | 0.9 | 0.8 | 0.8 | 0.9 | 0.9 | 0.9 | 0.8 | 0.9 | 0.8 | 0.9 | 0.8 | 0.9 | 0.9 | 0.8 |
IN | 0.9 | 0.9 | 0.9 | 0.9 | 0.9 | 0.9 | 0.9 | 0.9 | 0.8 | 0.8 | 0.9 | 0.8 | 0.8 | 0.9 | 0.9 |
Supralabials | 10 | 10 | 10 | 10 | 10 | 10 | 10 | 10 | 10 | 10 | 10 | 10 | 10 | 10 | 10 |
Infralabials | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 | 9 |
No. of precloacal pores | 8 | 6 | 6 | 6 | 6 | 6 | 6 | 8 | 6 | 8 | 6 | – | – | – | – |
Precloacal pore continuous (1) or separated (0) | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | – | – | – | – |
No. of paravertebral tubercles | 24 | 25 | 25 | 24 | 25 | 24 | 25 | 23 | 24 | 23 | 25 | 25 | 25 | 23 | 23 |
No. of 4th toe lamellae | 28 | 28 | 27 | 28 | 28 | 27 | 28 | 27 | 27 | 27 | 28 | 26 | 28 | 26 | 28 |
Cnemaspis adangrawi sp. nov. is known only from Adang and Rawi islands, 60 and 61 km off the coast of Thailand, respectively (Fig.
Paratypes found during the day (ZMKU R 00767–00768, ZMKU R 00770–00772, ZMKU R 00775–00776, THNHM 28206–28209, and THNHM 28211–28215) were in crevices of boulders, shaded areas with holes in the soil at the base of a rock wall near a stream, and on boulder outcrops near streams. When disturbed, some individuals would retreat into rock crevices or into holes in the soil at the base of a rock wall. Paratypes found at night (ZMKU R 00769, ZMKU R 00773–00774 and THNHM 28210) were in shaded areas (by day), deep at the base of boulders, or perched on vegetation near a rocky stream. Two gravid females (ZMKU R 00771 and THNHM 28211) contained one or two eggs during November 2017. Some juveniles (not collected) were found in holes in the soil and perched on vegetation near a stream at Rawi Island on 8 April 2018. At night, Cyrtodactylus macrotuberculatus was found in syntopy on the rock wall and vegetation near a stream at Jonsalad Waterfall, Adang Island, with Cnemaspis adangrawi sp. nov.
The specific epithet refers to Adang and Rawi islands where the new species is found, and is a noun in apposition.
Cnemaspis adangrawi sp. nov. can be distinguished from other members of the siamensis group (C. chanardi, C. huaseesom, C. omari, C. phangngaensis, C. punctatonuchalis, C. roticanai, C. siamensis, C. thachanaensis, and C. vandeventeri) by having a smaller maximum SVL of 44.9 mm (vs 47.0 mm in C. roticanai, 49.6 mm in C. punctatonuchalis) and by having a larger maximum SVL 44.9 mm (vs 40.9 mm in C. chanardi, 43.5 mm in C. huaseesom, 41.3 mm in C. omari, 42.0 mm in C. phangngaensis, 39.7 mm in C. siamensis, 39.0 mm in C. thachanaensis, and 44.7 mm in C. vandeventeri).
Cnemaspis adangrawi sp. nov. is distinguished from C. omari, C. punctatonuchalis, C. roticanai, C. siamensis, and C. vandeventeri by having 10 supralabial scales (vs eight in C. punctatonuchalis and 8–9 in C. omari, C. roticanai, C. siamensis, and C. vandeventeri). This species is distinguished from C. chanardi, C. omari, C. phangngaensis, C. punctatonuchalis, C. roticanai, and C. siamensis by having 9 infralabial scales (vs 8 in C. chanardi, 7–8 in C. omari, C. punctatonuchalis, C. roticanai, 10 in C. phangngaensis, and 6–8 in C. siamensis). This species is distinguished from C. huaseesom and C. punctatonuchalis by having keeled ventral and subcaudal scales (vs smooth ventral and subcaudal scales in C. huaseesom and C. punctatonuchalis).
Cnemaspis adangrawi sp. nov. is distinguished from C. phangngaensis and C. vandeventeri by having 6–8 precloacal pores (vs 4 in C. phangngaensis and C. vandeventeri). This species is distinguished from C. punctatonuchalis, C. siamensis, and C. thachanaensis by presence of precloacal pores (vs precloacal pores absent in C. punctatonuchalis, C. siamensis, and C. thachanaensis). This species is distinguished from C. huaseesom and C. phangngaensis by having a separated row of precloacal pores (vs continuous in C. huaseesom and C. phangngaensis).
Cnemaspis adangrawi sp. nov. is distinguished from C. phangngaensis and C. thachanaensis by having 23–25 paravertebral tubercles (vs 22 in C. phangngaensis and 15–19 in C. thachanaensis). This species is distinguished from C. huaseesom, C. omari, C. punctatonuchalis, C. roticanai, C. siamensis, C. thachanaensis, and C. vandeventeri by lacking tubercles on lower flanks (vs present in C. huaseesom, C. omari, C. punctatonuchalis, C. roticanai, C. siamensis, C. thachanaensis, and C. vandeventeri). This species is distinguished from C. phangngaensis, C. punctatonuchalis, and C. thachanaensis by having 26–28 lamellae under 4th toe (vs 29 in C. phangngaensis, 29–31 in C. punctatonuchalis, and 24 in C. thachanaensis). This species is distinguished from C. chanardi and C. vandeventeri by having lateral caudal furrows (vs lacking in C. chanardi and C. vandeventeri).
Cnemaspis adangrawi sp. nov. can be further distinguished from C. chanardi, C. punctatonuchalis, C. siamensis, and C. vandeventeri by lacking enlarged median subcaudal scales (vs present in C. chanardi, C. punctatonuchalis, C. siamensis, and C. vandeventeri). This species is distinguished from C. huaseesom by having keeled subtibial scales (vs smooth subtibial scales in C. huaseesom). This species is distinguished from C. siamensis and C. thachanaensis by lacking lineate gular marking (vs present in C. siamensis and C. thachanaensis).
Studies on the taxonomy and systematics of Cnemaspis in Southeast Asia have increased in the past two decades (
Previously, the reported geographic distribution of Cnemaspis in Thailand was mostly restricted to the mainland (
The complex geological history in Thailand created a large number of limestone and granitic formations in southern Thailand (
This work was supported by grants from the Thailand Research Fund (DBG6080010) and the Center of Excellence on Biodiversity (BDC), Office of Higher Education Commission (BDC-PG4-160022) to AA, and a grant from Faculty of Science, Kasetsart University (50th Anniversary of Faculty of Science) to NA. This research was approved by the Institutional Animal Care and Use Committee of Faculty of Science, Kasetsart University (project number ACKU60-SCI-004). Department of National Parks, Wildlife and Plant Conservation, Thailand, granted permission to conduct the research. Kanchanapan Kamhang and Phanphong Kongkaew (Tarutao National Park) facilitated the fieldwork. Wachara Sanguansombat and Sunchai Makchai (Thailand Natural History Museum) made specimens in their care available for study. Korkhwan Termprayoon, Piyawan Puanprapai, and Siriporn Yodthong provided assistance with fieldwork. Evan S. H. Quah, and Vinh Q. Luu improved the manuscript. This paper is contribution number 892 of the Auburn University Museum of Natural History.
List of comparative specimens examined.
Cnemaspis chanardi: Thailand, Trang Province, Nayong District, Ban Chong: THNHM 06983 (male holotype); Krabi Province, Klong Thom District: THNHM 012439–012440 (males); Mueang Krabi District: THNHM 012436 and 012437 (males), THNHM 012438 (female); Nakhon Si Thammarat Province, Tha Sala District: THNHM 020992 (male); Lansaka district: THNHM 014111 (immature male); Noppitam district: THNHM 013838 (male), THNHM 010705 (male); Surat Thani Province, Ang Thong Island, Mueang Surat Thani District: THNHM 016074 (female).
Cnemaspis huaseesom: Thailand, Kanchanaburi Province, Sai Yok District, Sai Yok National Park: THNHM 15909 (male holotype).
Cnemaspis niyomwanae: Thailand, Trang Province, Palean District, Thum Khao Ting: THNHM 15909 (female holotype).
Cnemaspis punctatonuchalis: Thailand, Prachuap Khiri Khan Province, Thap Sakae District, Huay Yang National Park: THNHM 02001 (male holotype)
Cnemaspis siamensis: Thailand, Nakhon Si Thammarat Province, Lansaka District: THNHM 013828 (male); Tha Sala District: THNHM 018265 (male); Phetchabun Province, Nam Nao District: THNHM 01336 (female), THNHM 01337 (male); Phetchaburi Province, Cha-am District: THNHM 01448 (male), THNHM 01449 (immature male); Chumpon Province, Mueang Chumpon District: THNHM 0372 (male); Phato District: THNHM 01086 (male); Surat Thani Province, Vibhawadee District: THNHM 01084 (female); Ang Thong Island, Mueang Surat Thani District: THNHM 015624 (female).
Cnemaspis vandeventeri: Thailand, Ranong Province, Kapur District, Klong Naka: THNHM 08261 (male holotype), THNHM 08260 (female).