Research Article |
Corresponding author: Attapol Rujirawan ( fsciapr@ku.ac.th ) Academic editor: Thomas Ziegler
© 2022 L. Lee Grismer, Anchalee Aowphol, Siriporn Yodthong, Natee Ampai, Korkhwan Termprayoon, Akrachai Aksornneam, Attapol Rujirawan.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Grismer LL, Aowphol A, Yodthong S, Ampai N, Termprayoon K, Aksornneam A, Rujirawan A (2022) Integrative taxonomy delimits and diagnoses cryptic arboreal species of the Cyrtodactylus brevipalmatus group (Squamata, Gekkonidae) with descriptions of four new species from Thailand. ZooKeys 1129: 109-162. https://doi.org/10.3897/zookeys.1129.90535
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Species delimitation and species diagnosis must remain separate operations to avoid constructing taxonomies comprised of non-monophyletic species based on morphological similarity as opposed to phylogenetic propinquity. This is particularly true for highly specialized species such as the range-restricted upland taxa in the Cyrtodactylus brevipalmatus group of Indochina where strong selection pressure for an arboreal lifestyle has contributed to morphologically similar but distantly related species. This in turn, has resulted in a history of erroneous taxonomies that have actually obscured rather than revealed the diversity within this group. A Bayesian phylogeny of the C. brevipalmatus group recovered at least 15 putative species-level lineages, at least seven of which are undescribed, and of which four are described herein. A total evidence morphological data set comprised of 16 normalized morphometric, 15 meristic, and seven categorical characters scored across 51 individuals was subjected to a multiple factor analysis (MFA) and an analysis of variance (ANOVA) to diagnose the putative species. These new species descriptions contribute to focusing attention to the unrealized diversity in upland tropical ecosystems, which due to climate change, are some of the most impearled ecosystems on the planet. Thus, it is paramount that taxonomies do not conflate species identities and underrepresent true diversity.
geckos, genetics, morphology, Southeast Asia, taxonomy
A cornerstone of biodiversity conservation is a phylogenetic-based taxonomy where the names of the component species are consistent with the patterns and processes by which they evolved. Taxonomies constructed from paraphyletic or polyphyletic species misrepresent history, thus obscuring true diversity and potentially countermanding the effectiveness of conservation efforts. Before a newly discovered population can be given a new name, or the name of an existing species be successfully challenged, it should be properly delimited and diagnosed. Delimiting and diagnosing species are independent operations used together to construct taxonomies that reflect, and are consistent with evolutionary history. Unfortunately, these two operations are often conflated when analyses to diagnose species, which are most often rooted in morphological similarity, are equated with analyses to delimit species, which are rooted in phylogenetic propinquity. The unfortunate consequence of this is that taxonomies may be constructed using non-monophyletic species, thus obscuring rather than revealing the group’s actual diversity. This is especially true for taxonomies comprised of highly specialized cryptic species where, in the absence of a phylogeny, morphological convergence can be mistaken for common ancestry (see
Convergent morphology results from the specific resource requirements that necessitate a particular functional morphology (
Distribution of nominal species and unnamed populations and specimens of the Cyrtodactylus brevipalmatus group. Stars denote type localities. White circles are literature localities from which specimens were not examined and remain unidentified. Locality data for all material examined is in Table
The phylogenetic-based taxonomy of
The general lineage concept (GLC:
Species boundaries were cross-checked using a Bayesian Poisson Tree Process for species delimitation (bPTP;
Methods for DNA extraction, sequencing, and editing followed
Cyrtodactylus specimens examined in this study. Institutional abbreviations follow
Species | Location | Catalog no. | GenBank no. |
---|---|---|---|
C. brevipalmatus | Thailand, no data | LSUHC 1899 | not in tree |
C. brevipalmatus | Thailand, Nakhon Si Thammarat Province, Nopphitam District, Khao Nan National Park, Huay Lak Protected Unit | THNHM 10670 | not in tree |
C. brevipalmatus | Thailand, Nakhon Si Thammarat Province, Lan Saka District, Khao Luang National Park | THNHM 14112 | not in tree |
C. brevipalmatus | Thailand, Nakhon Si Thammarat Province, Khao Ram Mt. | AUP-00573 | OK626313 |
C. cf. brevipalmatus | Peninsular Malaysia, Kedah State, Pulau Langkawi, Gunung Raya | LSUHC 11788 | not in tree |
C. cf. brevipalmatus | Peninsular Malaysia, Kedah State, Pulau Langkawi, Gunung Raya | LSUHC 15076 | not in tree |
C. elok | Peninsular Malaysia, Pahang State, Fraser’s Hill, the Gap | ZRC 2.6091/LSUHC 6471 | JQ889180 |
C. elok | Peninsular Malaysia, Negeri Sembilan State | LSUHC 8238 | not in tree |
C. elok | Peninsular Malaysia, Pahang State, near Cameron Highlands | LSUHC 12180 | not in tree |
C. elok | Peninsular Malaysia, Pahang State, near Cameron Highlands | LSUHC 12181 | not in tree |
C. elok | Malaysian pet trade, no data | ZMMU R-16144 | not in tree |
Cyrtodactylus fluvicavus sp. nov. | Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00958 paratype | OP620036 |
Cyrtodactylus fluvicavus sp. nov. | Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00959 holotype | OP620037 |
Cyrtodactylus fluvicavus sp. nov. | Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00960 paratype | OP620038 |
Cyrtodactylus fluvicavus sp. nov. | Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00961 paratype | OP620039 |
Cyrtodactylus fluvicavus sp. nov. | Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00962 paratype | OP620040 |
Cyrtodactylus fluvicavus sp. nov. | Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00963 paratype | OP620041 |
Cyrtodactylus fluvicavus sp. nov. | Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park | ZMKU R 00964 paratype | OP620042 |
C. interdigitalis | Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | THNHM 20226 paratype | not in tree |
C. interdigitalis | Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | THNHM 20227 paratype | not in tree |
C. interdigitalis | Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | THNHM 20228 paratype | not in tree |
C. interdigitalis | Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | THNHM 20229 paratype | not in tree |
C. interdigitalis | Thailand, Phetchabun Province, Nam Nao National Park, Tham Yai Nam Nao | YC000952 | ON055281 |
Cyrtodactylus kochangensis sp. nov. | Thailand, Ranong Province, Mueng Ranong District, Ko Phayam Subdistrict, Ko Chang | ZMKU R 00945 holotype | OP620023 |
C. cf. kochangensis sp. nov. | Thailand, Ranong Province, Khlong Nakha Wildlife Sanctuary | THNHM 01667 | not in tree |
C. ngati | Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, karst forest near Pa Thom Cave | HNUE-R00111 holotype | ON411220 |
C. ngati | Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, karst forest near Pa Thom Cave | HNUE-R00112 paratype | not in tree |
C. ngati | Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, karst forest near Pa Thom Cave | IEBR 4829 paratype | OK626318 |
C. ngati | Vietnam, Dien Bien Province, Dien Bien District, Pa Thom Commune, Pa Xa Lao Village, karst forest near Pa Thom Cave | VNUF R.2020.12 paratype | OK626319 |
C. ngati3 | Laos, Khammouane Province | VNUF R.2014.50 | ON411221 |
C. ngati3 | Laos, Khammouane Province, Phou Hin Poun National Biodiversity Conservation Area | FMNH 255454 | JQ889181 |
C. ngati3 | Laos, Khammouane Province, Phou Hin Poun National Biodiversity Conservation Area | FMNH 270492 | OK626315 |
C. ngati3 | Laos, Khammouane Province, Phou Hin Poun National Biodiversity Conservation Area | FMNH 270493 | not in tree |
C. ngati4 | Thailand, Loei Province, Nam San Noi River, Phu Luang Wildlife Sanctuary | FMNH 265806 | JX519471 |
C. cf. ngati1 | Laos, Xaignabouli Province, Ban Pha Liep, Houay Liep Stream | NCSM 79472 | OK626316 |
C. cf. ngati2 | Laos, Vientiane Province | ZMMU R-14917 | not in tree |
C. cf. ngati2 | Laos, Vientiane Province, tributary of Nam Pha River, Houay Wan Stream | NCSM 80100 | OK626317 |
Cyrtodactylus rivularis sp. nov. | Thailand, Prachuap Khiri Khan Province, Hua Hin District, Huai Sat Yai Subdistrict, Kaeng Krachan National Park, Pa La-U Waterfall | ZMKU R 00946 paratype | OP620024 |
Cyrtodactylus rivularis sp. nov. | Thailand, Prachuap Khiri Khan Province, Hua Hin District, Huai Sat Yai Subdistrict, Kaeng Krachan National Park, Pa La-U Waterfall | ZMKU R 00947 holotype | OP620025 |
C. rukhadeva | Thailand, Ratchaburi Province, Suan Phueng District, Khao Laem Mountain | ZMMU R-16851 holotype | OK626320 |
C. rukhadeva | Thailand, Ratchaburi Province, Suan Phueng District, Hoop Phai Tong | ZMMU R-16852 paratype | not in tree |
C. rukhadeva | Thailand, Ratchaburi Province, Suan Phueng District, Suan Phueng Subdistrict | ZMKU R 00948 | OP620026 |
C. cf. rukhadeva | Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 01807 | not in tree |
C. cf. rukhadeva | Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 03251 | not in tree |
C. cf. rukhadeva | Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 03252 | not in tree |
C. cf. rukhadeva | Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 03253 | not in tree |
C. cf. rukhadeva | Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 03254 | not in tree |
C. cf. rukhadeva | Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 24622 | not in tree |
C. cf. rukhadeva | Thailand, Phetchaburi Province, Kaeng Krachan National Park | THNHM 24838 | not in tree |
Cyrtodactylus uthaiensis sp. nov. | Thailand, Uthai Thani Province, Lan Sak District, Thung Na Ngam Subdistrict | ZMKU R 00949 holotype | OP620027 |
C. sp.9 | Thailand, Kanchanaburi Province, Thong Pha Phum District, Thong Pha Phum National Park | AUP-01715 | MT468909 |
C. sp.9 | Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00950 | OP620028 |
C. sp.9 | Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00951 | OP620029 |
C. sp.9 | Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00952 | OP620030 |
C. sp.9 | Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00953 | OP620031 |
C. sp.9 | Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00954 | OP620032 |
C. sp.9 | Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00955 | OP620033 |
C. sp.9 | Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00956 | OP620034 |
C. sp.9 | Thailand, Kanchanaburi Province, Thong Pha Phum District, Pilok Subdistrict, Thong Pha Phum National Park | ZMKU R 00957 | OP620035 |
C. sp.10 | Thailand, Tak Province, Tha Song Yang District, Mae Moei National Park, Chao Doi Waterfall | AUP-00680 | MT468902 |
C. sp.11 (C. cf. interdigitalis) | Thailand, Phitsanulok Province, Phu Hin Rong Kla National Park | ZMMU R-16492 | MW792061 |
C. sp.13 | Thailand, Tak Province, Umphang District, Thung Yai Naresuan Wildlife Sanctuary | THNHM 00104 | not in tree |
C. sp.13 | Thailand, Kanchanaburi Province, Thong Pha Phum District, Ban Saphan Lao | THNHM 27821 | not in tree |
C. sp.14 (C. cf. brevipalmatus) | Peninsular Malaysia, Kedah State, Pulau Langkawi, Gunung Raya | USMHC 2555 | OK626314 |
Morphological data included both meristic and morphometric characters. To reduce the degree of researcher bias, data were taken using the protocol of
Meristic characters evaluated were the number of supralabial scales (SL), counted from the largest scale at the corner of the mouth or posterior to the eye, to the rostral scale; infralabial scales (IL), counted from termination of enlarged scales at the corner of the mouth to the mental scale; number of paravertebral tubercles (PVT) between the limb insertions counted in a straight line immediately left of the vertebral column; number of longitudinal rows of body tubercles (LRT) counted transversely across the body midway between the limb insertions from one ventrolateral body fold to the other; number of longitudinal rows of ventral scales (VS) counted transversely across the abdomen midway between limb insertions from one ventrolateral fold to the other; number of transverse rows of ventral scales (VSM) counted along the midline of the body from the postmentals to just anterior to the cloacal opening, stopping where the scales become granular; number of expanded subdigital lamellae on the fourth toe proximal to the digital inflection (TL4E) counted from the base of the first phalanx where it contacts the body of the foot to the largest scale on the digital inflection–the large contiguous scales on the palmar and plantar surfaces were not counted; number of small, generally unmodified subdigital lamellae distal to the digital inflection on the fourth toe (TL4U) counted from the digital inflection to the claw including the claw sheath; total number of subdigital lamellae (TL4T) beneath the fourth toe (i.e. TL4E + TL4U = TL4T); number of expanded subdigital lamellae on the fourth finger proximal to the digital inflection (FL4E) counted the same way as with TL4E; number of small generally unmodified subdigital lamellae distal to the digital inflection on the fourth finger (FL4U) counted the same way as with TL4U; total number of subdigital lamellae (FL4T) beneath the fourth toe (i.e. FL4E + FL4U = FL4T); total number of enlarged femoral scales (FS) from each thigh combined as a single metric; number of enlarged precloacal scales (PCS);number of precloacal pores (PP) in males; the number of femoral pores (FP) in males from each thigh combined as a single metric; and the number of dark body bands (BB) between the dark band on the nape and the hind limb insertions on the body. A post-sacral or sacral band when present, was not counted. Categorical characters evaluated were the presence or absence of tubercles on the flanks (FKT); single enlarged, unmodified, medial subcaudal scales (SC2); enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginated (SC3); slightly enlarged medial subcaudals (SC1); large or small dorsolateral caudal tubercles (DCT) forming a narrow or wide ventrolateral caudal fringe (VLF1); ventrolateral caudal fringe scales generally homogenous or not (VLF2); and the cross-section of the tail round or square (TLcross).
An input file implemented in BEAUti (Bayesian Evolutionary Analysis Utility) v. 2.4.6 was run in BEAST (Bayesian Evolutionary Analysis Sampling Trees) v. 2.4.6 (
All statistical analyses were conducted using
Small sample sizes (N = 1 or 2) for some of the species/populations precluded them from statistical analyses. A Levene’s test for the normalized morphometric and meristic characters was conducted to test for equal variances across all groups. Characters with equal variances were analyzed with an analysis of variance (ANOVA) and TukeyHSD post hoc test for mean comparisons involving more than three groups. Those with unequal variances were subjected to Welch’s F-test and Games-Howell post hoc test to test for mean comparisons involving more than three groups.
Morphospatial clustering and positioning among the species/populations was analyzed using multiple factor analysis (MFA) on a concatenated data set comprised of 15 meristic characters, 16 normalized morphometric characters, and seven categorical characters (Suppl. material
A non-parametric permutation multivariate analysis of variance (PERMANOVA) from the vegan package 2.5–3 in R (
The BEAST phylogeny was projected onto the first two dimensions of the MFA plot using the phylomorphospace () command from the R package phytools (
Some of the populations examined had only genetic or morphological data. However, only phylogenetically delimited populations bearing morphological differences from other populations were described as new species. In some cases, populations represented by only morphological data but in close geographic proximity to named species from which they could not be separated morphologically, were considered conferre (cf.) to the named species pending further investigation.
The BEAST analysis recovered 8–10 new species and two major clades within the brevipalmatus group: a weakly supported clade (BPP = 0.82) from the southernmost portion of the Thai-Malay Peninsula south of the Isthmus of Kra comprised of Cyrtodactylus elok, C. brevipalmatus, and C. sp.14 and a well-supported clade (1.00) comprised of all other species north of the Isthmus of Kra (Figs
Uncorrected pairwise sequence divergences within the brevipalmatus group range from 2.84–22.84% (Table
Mean (minimum–maximum) percentages of uncorrected pairwise sequence divergence (p-distances) among the putative species of the Cyrtodactylus brevipalmatus group based on 1,386 base pairs of mitochondrial NADH dehydrogenase subunit 2 gene (ND2) and adjacent tRNAs. Intraspecific p-distance are in bold font. n/a = data not applicable.
C. brevipalmatus | C. cf. ngati1 | C. cf. ngati2 | C. elok | Cyrtodactylus fluvicavus sp. nov. | C. interdigitalis | Cyrtodactylus kochangensis sp. nov. | C. ngati, C. ngati3 and C. ngati4 | Cyrtodactylus rivularis sp. nov. | C. rukhadeva | C. sp.9 (Thong Pha Phum) | C. sp.10 | C. sp.11 | C. sp.14 | Cyrtodactylus uthaiensis sp. nov. | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
C. brevipalmatus N = 1 | n/a | ||||||||||||||
C. cf. ngati1 N = 1 | 21.03 | n/a | |||||||||||||
C. cf. ngati2 N = 1 | 21.68 | 4.39 | n/a | ||||||||||||
C. elok N = 1 | 20.77 | 22.58 | 21.42 | n/a | |||||||||||
Cyrtodactylus fluvicavus sp. nov. N = 7 | 18.86 | 10.64 | 11.02 | 20.15 | 0.10 | ||||||||||
(18.84–18.97) | (10.58–10.84) | (10.97–11.23) | (20.13–20.26) | (0.00–0.26) | |||||||||||
C. interdigitalis N = 1 | 20.77 | 6.97 | 9.16 | 22.84 | 12.02 | n/a | |||||||||
(12.00–12.13) | |||||||||||||||
Cyrtodactylus kochangensis sp. nov. N = 1 | 19.35 | 14.58 | 14.71 | 20.90 | 12.31 | 15.23 | n/a | ||||||||
(12.26–12.31) | |||||||||||||||
C. ngati, C. ngati3 and C. ngati4 N = 7 | 20.70 | 3.30 | 3.71 | 21.11 | 11.34 | 8.13 | 14.58 | 0.84 | |||||||
(20.65–20.90) | (2.84–4.00) | (3.35–4.26) | (20.90–21.42) | (11.10–11.87) | (7.74–8.65) | (14.45–14.84) | (0.00–1.55) | ||||||||
Cyrtodactylus rivularis sp. nov. N = 2 | 20.00 | 15.87 | 15.03 | 21.61 | 12.57 | 15.48 | 12.26 | 15.03 | 0.52 | ||||||
(19.74–20.26) | (15.61–16.13) | (14.84–15.23) | (21.42–21.81) | (12.26–13.03) | (15.23–15.74) | (12.00–12.52) | (14.71–15.48) | ||||||||
C. rukhadeva N = 2 | 20.65 | 15.42 | 15.48 | 21.61 | 12.25 | 16.00 | 13.10 | 15.23 | 4.65 | 1.55 | |||||
(20.13–21.16) | (14.84–16.00) | (14.84–16.13) | (21.16–22.06) | (11.61–13.03) | (15.35–16.65) | (12.52–13.68) | (14.19–16.23) | (3.61–5.68) | |||||||
C. sp.9 (Thong Pha Phum) N = 9 | 20.34 | 7.93 | 9.51 | 22.02 | 9.75 | 8.96 | 13.22 | 8.81 | 13.12 | 13.25 | 0.22 | ||||
(20.13–20.65) | (7.74–8.00) | (9.42–9.55) | (21.81–22.32) | (9.55–9.94) | (8.77–9.03) | (13.03–13.29) | (8.13–9.68) | (12.77–13.42) | (12.52–13.94) | (0.00–0.52) | |||||
C. sp.10 N = 1 | 19.87 | 9.29 | 10.84 | 21.94 | 10.12 | 10.19 | 13.68 | 10.21 | 13.94 | 14.32 | 8.06 | n/a | |||
(10.06–10.32) | (10.06–10.45) | (13.68–14.19) | (13.68–14.97) | (7.87–8.13) | |||||||||||
C. sp.11 N = 1 | 20.39 | 7.23 | 8.90 | 22.19 | 11.12 | 3.87 | 14.58 | 8.28 | 15.35 1) | 15.61 | 8.96 | 10.45 | n/a | ||
(11.10–11.23) | (8.00–8.65) | (15.10–15.6 | (14.97–16.26) | (8.77–9.03) | |||||||||||
C. sp.14 N = 1 | 6.45 | 20.90 | 20.65 | 20.00 | 18.34 | 20.13 | 19.10 | 20.52 | 19.74 | 20.00 | 19.60 | 18.84 | 19.61 | n/a | |
(18.32–18.45) | (20.26–20.65) | (19.48–20.00) | (19.48–20.52) | (19.48–19.87) | |||||||||||
Cyrtodactylus uthaiensis sp. nov. N = 1 | 19.74 | 5.81 | 8.13 | 21.16 | 10.12 | 7.1 | 13.94 | 6.97 | 13.94 | 13.94 | 7.80 | 8.39 | 6.58 | 19.48 | n/a |
(10.06–10.32) | (6.58–7.61) | (13.68–14.19) | (13.29–14.58) | (7.61–7.87) |
The bPTP species delimitation analysis recovered 16 putative species within the brevipalmatus group with varying degrees of support (Table
Species | NMI |
---|---|
C. brevipalmatus | 1 |
C. cf. ngati1 | 1 |
C. cf. ngati2 | 0.967 |
C. elok | 1 |
Cyrtodactylus fluvicavus sp. nov. | 0.957 |
C. interdigitalis | 0.997 |
Cyrtodactylus kochangensis sp. nov. | 1 |
C. ngati | 0.841 |
Cyrtodactylus rivularis sp. nov. | 0.575 |
C. rukhadeva ZMKU R 00948 | 0.947 |
C. rukhadeva ZMMU R-16851 | 0.947 |
Cyrtodactylus uthaiensis sp. nov | 1 |
C. sp.9 (Thong Pha Phum) | 0.565 |
C. sp.10 | 1 |
C. sp.11 | 0.997 |
C. sp.14 | 1 |
The ANOVA and TukeyHSD post hoc analyses of the adjusted morphometric and meristic characters were consistent with the phylogenetic and pairwise distance data in recovering a number of statistically significant differences between the Cyrtodactylus fluvicavus sp. nov. and C. interdigitalis as well as others (Table
A MFA of the species-level lineages based on the BEAST phylogeny (Fig.
The phylomorphospace analysis illustrates that the morphological similarities among the species/populations in the MFA are discordant with their phylogenetic placement in the tree (Fig.
Given the phylogenetic delimitation of the Si Sawat, Prachuap Khiri Khan, Uthai Thani, and Ko Chang populations (Figs
Cyrtodactylus sp. Yodthong, Rujirawan, Stuart, Grismer, Aksornneam, Termprayoon, Ampai & Aowphol, 2022: 161.
Adult male ZMKU R 00959 from Tham Than Lot Noi-Tham Than Lot Yai Nature Trail, Chaloem Rattanakosin National Park, Khao Chot Subdistrict, Si Sawat District, Kanchanaburi Province, Thailand (14.66930°N, 99.29060°E, 526 m a.s.l.), collected by Korkhwan Termprayoon, Akrachai Aksornneam, Natee Ampai, and Siriporn Yodthong on 20 April 2019.
Adult males ZMKU R 00958 and ZMKU R 00960 and adult females ZMKU R 00961–64 bear the same collection site as the holotype.
Cyrtodactylus fluvicavus sp. nov. can be separated from all other species of the brevipalmatus group by the combination of having 11–13 supralabials, 9 or 10 infralabials, 26–30 paravertebral tubercles, 14–18 rows of longitudinally arranged tubercles, 30–39 transverse rows of ventrals, 154–175 longitudinal rows of ventrals, 9–11 expanded subdigital lamellae on the fourth toe, 10–13 unexpanded subdigital lamellae on the fourth toe, 19–22 total subdigital lamellae on the fourth toe; 7–9 expanded subdigital lamellae on the fourth finger, 9–11 unexpanded subdigital lamellae on the fourth finger, 17–19 total subdigital lamellae on the fourth finger; 9–12 total enlarged femoral scales, 8–11 total femoral pores in males; 14 or 15 precloacal pores in males; 14 or 15 enlarged precloacals; enlarged femorals and enlarged precloacals not continuous; proximal femorals less than one-half the size of the distal femorals; small tubercles on forelimbs and flanks; small dorsolateral caudal tubercles and narrow ventrolateral caudal fringe; ventrolateral caudal fringe composed scales of different size; tail circular in cross-section; slightly enlarged unpaired medial subcaudals not posteromedially furrowed; maximum SVL 78.2 mm; three dark transverse body bands (Tables
(Fig.
Holotype of Cyrtodactylus fluvicavus sp. nov. ZMKU R 00959 (field no. AA 07001) from Thailand, Kanchanaburi Province, Si Sawat District, Khao Chot Subdistrict, Chaloem Rattanakosin National Park A dorsal view B ventral view C dorsal view of head D ventral view of femoral and precloacal regions E dorsal view of tail F ventral view of tail.
Body relatively short (AG/SVL 0.46) with well-defined ventrolateral folds; dorsal scales small, granular interspersed with larger, conical, semi-regularly arranged, weakly keeled tubercles; tubercles extend from occipital region onto base of tail and slightly beyond as paravertebral rows; smaller tubercles extend anteriorly onto nape and occiput, diminishing in size anteriorly; approximately 17 longitudinal rows of tubercles at midbody; approximately 30 paravertebral tubercles; small tubercles on flanks; 34 longitudinal rows of flat, imbricate, ventral scales much larger than dorsal scales; 155 transverse rows of ventral scales; 15 large, pore-bearing, precloacal scales; no deep precloacal groove or depression; and two rows of post-precloacal scales on midline.
Forelimbs moderate in stature, relatively short (ForL/SVL 0.14); granular scales of forearm slightly larger than those on body, interspersed with large tubercles; palmar scales rounded, slightly raised; digits well-developed, relatively short, inflected at basal interphalangeal joints; digits narrower distal to inflections; subdigital lamellae wide, transversely expanded proximal to joint inflections, narrower transverse lamellae distal to joint inflections; claws well-developed, claw base sheathed by a dorsal and ventral scale; 8R/8L expanded and 10R/10L unexpanded lamellae beneath the fourth finger; hind limbs larger and thicker than forelimbs, moderate in length (TibL/SVL 0.16), covered dorsally by granular scales interspersed with moderately sized, conical tubercles dorsally and posteriorly and anteriorly by flat, slightly larger, subimbricate scales; ventral scales of thigh flat, subimbricate, larger than dorsals; subtibial scales flat, imbricate; one row of 5R/6L enlarged pore-bearing femoral scales not continuous with enlarged pore-bearing precloacal scales, terminating distally at knee; proximal femoral scales smaller than distal femorals, the former forming an abrupt union with much smaller, rounded, ventral scales of posteroventral margin of thigh; plantar scales flat; digits relatively long, well-developed, inflected at basal interphalangeal joints; 9R/9L wide, transversely expanded subdigital lamellae on fourth toe proximal to joint inflection extending onto sole, and 11R/11L unexpanded lamellae beneath the fourth toe; and claws well-developed, claw base sheathed by a dorsal and ventral scale.
Tail original, 97.6 mm long (TL/SVL 1.34), 5.2 mm in width at base, tapering to a point; sub-circular or nearly round in cross-section; dorsal scales flat, square bearing tubercles forming paravertebral rows and small tubercles forming a dorsolateral longitudinal row; slightly larger, posteriorly directed, semi-spinose tubercles forming narrow but distinct ventrolateral caudal fringe; larger scales of ventrolateral fringe occur at regular intervals; medial subcaudals slightly enlarged but not paired, distinctly enlarged single medial subcaudals absent; subcaudals, larger than dorsal caudals; base of tail bearing hemipenial swellings; 3R/2L conical postcloacal tubercles at base of hemipenial swellings; and postcloacal scales flat, imbricate.
(Fig.
(Fig.
Cyrtodactylus fluvicavus sp. nov. is currently known from the type locality at Tham Than Lot Noi-Tham Than Lot Yai Nature Trail in Chaloem Rattanakosin National Park, Si Sawat District, Kanchanaburi Province, western Thailand (Fig.
The specific epithet fluvicavus comes from the Latin fluvius, meaning stream, river, or flow and the Latin cavus, meaning hollow or hole and refers to a landmark cave in the Chaloem Rattanakosin National Park which has a stream that flows through it.
Cyrtodactylus fluvicavus sp. nov. is the sister species to a clade composed of ten lineages in a phylogenetic sequence of C. sp.9, C. sp.10, Cyrtodactylus uthaiensis sp. nov., C. sp.11, C. interdigitalis, C. cf. ngati1, C. cf. ngati2, C. ngati3, and the sister lineages C. ngati4 and C. ngati (Fig.
All individuals were found in karst forests bearing mixed deciduous and dry evergreen trees amidst rocky streams and a nearby waterfall (Fig.
Summary statistics of the normalized morphometric data for the putative species of the Cyrtodactylus brevipalmatus group. SD = ± standard deviation. Character abbreviations are listed in the Materials and methods.
Species | SVL | AG | HumL | ForL | FemL | TibL | HL | HW | HD | ED | EE | ES | EN | IO | EL | IN |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
C. brevipalmatus (N = 5) | ||||||||||||||||
Mean | 1.82 | 1.49 | 0.96 | 0.96 | 1.06 | 1.02 | 1.27 | 1.10 | 0.87 | 0.67 | 0.75 | 0.86 | 0.73 | 0.68 | 0.07 | 0.32 |
SD | 0.020 | 0.027 | 0.031 | 0.023 | 0.024 | 0.036 | 0.012 | 0.006 | 0.017 | 0.041 | 0.025 | 0.008 | 0.019 | 0.043 | 0.058 | 0.041 |
Lower | 1.80 | 1.46 | 0.92 | 0.92 | 1.02 | 0.97 | 1.25 | 1.10 | 0.86 | 0.63 | 0.71 | 0.85 | 0.70 | 0.62 | -0.02 | 0.25 |
Upper | 1.85 | 1.52 | 0.99 | 0.98 | 1.09 | 1.07 | 1.29 | 1.11 | 0.89 | 0.73 | 0.77 | 0.87 | 0.75 | 0.73 | 0.13 | 0.36 |
C. elok (N = 4) | ||||||||||||||||
Mean | 1.91 | 1.59 | 0.80 | 1.05 | 1.14 | 1.13 | 1.34 | 1.20 | 1.00 | 0.71 | 0.83 | 0.95 | 0.79 | 0.70 | 0.19 | 0.43 |
SD | 0.016 | 0.012 | 0.341 | 0.026 | 0.022 | 0.024 | 0.001 | 0.012 | 0.008 | 0.017 | 0.020 | 0.008 | 0.010 | 0.070 | 0.062 | 0.033 |
Lower | 1.89 | 1.57 | 0.34 | 1.02 | 1.11 | 1.10 | 1.34 | 1.18 | 0.98 | 0.68 | 0.81 | 0.94 | 0.78 | 0.61 | 0.15 | 0.40 |
Upper | 1.93 | 1.60 | 1.09 | 1.08 | 1.16 | 1.15 | 1.34 | 1.21 | 1.00 | 0.72 | 0.85 | 0.95 | 0.80 | 0.76 | 0.28 | 0.48 |
Cyrtodactylus fluvicavus sp. nov. (N = 7) | ||||||||||||||||
Mean | 1.86 | 1.52 | 0.95 | 1.01 | 1.11 | 1.03 | 1.31 | 1.14 | 0.91 | 0.69 | 0.78 | 0.92 | 0.79 | 0.73 | 0.20 | 0.38 |
SD | 0.027 | 0.009 | 0.013 | 0.017 | 0.018 | 0.012 | 0.007 | 0.011 | 0.011 | 0.007 | 0.016 | 0.008 | 0.011 | 0.006 | 0.04 | 0.02 |
Lower | 1.82 | 1.51 | 0.94 | 0.98 | 1.10 | 1.01 | 1.30 | 1.13 | 0.90 | 0.68 | 0.76 | 0.91 | 0.78 | 0.72 | 0.14 | 0.35 |
Upper | 1.89 | 1.54 | 0.98 | 1.04 | 1.15 | 1.05 | 1.32 | 1.16 | 0.93 | 0.70 | 0.81 | 0.94 | 0.81 | 0.74 | 0.24 | 0.40 |
C. interdigitalis (N = 4) | ||||||||||||||||
Mean | 1.86 | 1.50 | 0.98 | 1.01 | 1.10 | 1.08 | 1.30 | 1.12 | 0.80 | 0.72 | 0.76 | 0.90 | 0.77 | 0.68 | 0.13 | 0.33 |
SD | 0.060 | 0.016 | 0.032 | 0.015 | 0.020 | 0.007 | 0.012 | 0.006 | 0.187 | 0.026 | 0.026 | 0.024 | 0.034 | 0.0307 | 0.033 | 0.035 |
Lower | 1.78 | 1.48 | 0.94 | 0.99 | 1.07 | 1.07 | 1.28 | 1.12 | 0.56 | 0.69 | 0.73 | 0.88 | 0.74 | 0.66 | 0.10 | 0.29 |
Upper | 1.91 | 1.52 | 1.01 | 1.02 | 1.12 | 1.09 | 1.31 | 1.13 | 0.96 | 0.75 | 0.79 | 0.94 | 0.81 | 0.73 | 0.17 | 0.38 |
Cyrtodactylus kochangensis sp. nov. (N = 1) | ||||||||||||||||
Value | 1.78 | 1.51 | 1.00 | 0.95 | 1.05 | 1.02 | 1.30 | 1.14 | 0.91 | 0.69 | 0.76 | 0.91 | 0.78 | 0.68 | 0.05 | 0.36 |
C. cf. kochangensis sp. nov. (N = 1) | ||||||||||||||||
Value | 1.85 | 1.50 | 1.02 | 0.94 | 1.08 | 1.08 | 1.27 | 1.09 | 0.90 | 0.72 | 0.70 | 0.88 | 0.75 | 0.60 | 0.12 | 0.36 |
C. ngati (N = 3) | ||||||||||||||||
Mean | 1.83 | 1.47 | 0.91 | 0.99 | 1.06 | 1.05 | 1.31 | 1.08 | 0.85 | 0.57 | 0.76 | 0.86 | 0.81 | 0.74 | -0.12 | 0.43 |
SD | 0.009 | 0.002 | 0.004 | 0.007 | 0.000 | 0.006 | 0.001 | 0.003 | 0.010 | 0.035 | 0.016 | 0.015 | 0.004 | 0.009 | 0.019 | 0.008 |
Lower | 1.82 | 1.47 | 0.91 | 0.98 | 1.06 | 1.04 | 1.31 | 1.08 | 0.84 | 0.55 | 0.74 | 0.85 | 0.80 | 0.73 | -0.13 | 0.42 |
Upper | 1.84 | 1.47 | 0.91 | 1.00 | 1.06 | 1.05 | 1.31 | 1.09 | 0.86 | 0.61 | 0.77 | 0.88 | 0.81 | 0.75 | -0.09 | 0.43 |
C. ngati3 (N = 3) | ||||||||||||||||
Mean | 1.88 | 1.58 | 0.94 | 1.00 | 1.12 | 1.07 | 1.32 | 1.12 | 0.95 | 0.69 | 0.81 | 0.93 | 0.80 | 0.77 | 0.08 | 0.41 |
SD | 0.039 | 0.001 | 0.002 | 0.02 | 0.001 | 0.017 | 0.007 | 0.002 | 0.014 | 0.003 | 0.004 | 0.007 | 0.006 | 0.022 | 0.016 | 0.01 |
Lower | 1.85 | 1.58 | 0.93 | 0.98 | 1.12 | 1.05 | 1.31 | 1.12 | 0.94 | 0.69 | 0.80 | 0.92 | 0.79 | 0.74 | 0.07 | 0.40 |
Upper | 1.92 | 1.58 | 0.94 | 1.02 | 1.12 | 1.09 | 1.33 | 1.12 | 0.96 | 0.69 | 0.81 | 0.94 | 0.80 | 0.78 | 0.10 | 0.42 |
C. ngati4 (N = 1) | ||||||||||||||||
Value | 1.87 | 1.50 | 0.87 | 1.02 | 1.13 | 1.07 | 1.33 | 1.11 | 0.91 | 0.72 | 0.79 | 0.94 | 0.83 | 0.75 | 0.51 | 0.43 |
C. cf. ngati1 (N = 1) | ||||||||||||||||
Value | 1.89 | 1.59 | 0.96 | 1.03 | 1.13 | 1.12 | 1.34 | 1.12 | 0.95 | 0.85 | 0.76 | 0.96 | 0.82 | 0.73 | 0.28 | 0.49 |
C. cf. ngati2 (N = 2) | ||||||||||||||||
Mean | 1.92 | 1.59 | 1.01 | 1.02 | 1.17 | 1.1 | 1.34 | 1.15 | 0.95 | 0.72 | 0.80 | 0.94 | 0.80 | 0.65 | 0.03 | 0.37 |
SD | 0.035 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 |
Lower | 1.89 | 1.59 | 1.01 | 1.02 | 1.17 | 1.10 | 1.34 | 1.15 | 0.95 | 0.72 | 0.80 | 0.94 | 0.80 | 0.65 | 0.03 | 0.37 |
Upper | 1.94 | 1.59 | 1.01 | 1.02 | 1.17 | 1.10 | 1.34 | 1.15 | 0.95 | 0.72 | 0.80 | 0.94 | 0.80 | 0.65 | 0.03 | 0.37 |
Cyrtodactylus rivularis sp. nov. (N = 2) | ||||||||||||||||
Mean | 1.85 | 1.53 | 0.89 | 0.98 | 1.04 | 1.03 | 1.30 | 1.16 | 0.91 | 0.76 | 0.80 | 0.91 | 0.77 | 0.75 | 0.05 | 0.34 |
SD | 0.025 | 0.01 | 0.002 | 0.007 | 0.007 | 0.005 | 0.005 | 0.001 | 0.013 | 0.007 | 0.024 | 0.005 | 0.004 | 0.025 | 0.019 | 0.016 |
Lower | 1.83 | 1.53 | 0.89 | 0.97 | 1.03 | 1.03 | 1.29 | 1.16 | 0.91 | 0.75 | 0.79 | 0.91 | 0.77 | 0.74 | 0.04 | 0.33 |
Upper | 1.87 | 1.54 | 0.90 | 0.98 | 1.04 | 1.04 | 1.30 | 1.16 | 0.92 | 0.76 | 0.82 | 0.91 | 0.78 | 0.77 | 0.06 | 0.35 |
C. rukhadeva and C. cf. rukhadeva (N = 10) | ||||||||||||||||
Mean | 1.85 | 1.49 | 1.01 | 0.95 | 1.05 | 1.02 | 1.30 | 1.14 | 0.92 | 0.71 | 0.75 | 0.91 | 0.77 | 0.67 | 0.09 | 0.35 |
SD | 0.026 | 0.028 | 0.055 | 0.029 | 0.027 | 0.023 | 0.009 | 0.018 | 0.025 | 0.036 | 0.034 | 0.014 | 0.015 | 0.087 | 0.069 | 0.022 |
Lower | 1.79 | 1.45 | 0.91 | 0.90 | 0.99 | 0.97 | 1.29 | 1.10 | 0.87 | 0.62 | 0.70 | 0.89 | 0.74 | 0.46 | 0.00 | 0.32 |
Upper | 1.88 | 1.54 | 1.08 | 1.00 | 1.09 | 1.05 | 1.32 | 1.16 | 0.95 | 0.75 | 0.80 | 0.92 | 0.79 | 0.73 | 0.23 | 0.38 |
Cyrtodactylus uthaiensis sp. nov. (N = 1) | ||||||||||||||||
Value | 1.76 | 1.80 | 0.95 | 0.99 | 1.09 | 1.02 | 1.28 | 1.10 | 0.76 | 0.72 | 0.74 | 0.88 | 0.75 | 0.68 | 0.19 | 0.32 |
C. sp.9 (Thong Pha Phum) (N = 8) | ||||||||||||||||
Mean | 1.86 | 1.53 | 0.91 | 0.97 | 1.08 | 1.02 | 1.30 | 1.16 | 0.89 | 0.71 | 0.77 | 0.90 | 0.77 | 0.74 | 0.09 | 0.34 |
SD | 0.024 | 0.016 | 0.034 | 0.021 | 0.032 | 0.019 | 0.014 | 0.017 | 0.008 | 0.015 | 0.004 | 0.012 | 0.007 | 0.012 | 0.061 | 0.023 |
Lower | 1.81 | 1.50 | 0.86 | 0.93 | 1.02 | 0.99 | 1.27 | 1.14 | 0.88 | 0.69 | 0.76 | 0.89 | 0.76 | 0.72 | 0.00 | 0.30 |
Upper | 1.88 | 1.55 | 0.95 | 1.00 | 1.12 | 1.05 | 1.32 | 1.19 | 0.90 | 0.74 | 0.78 | 0.93 | 0.78 | 0.76 | 0.18 | 0.38 |
C. sp.11 (N = 1) | ||||||||||||||||
Value | 1.83 | 1.53 | 1.01 | 0.99 | 1.13 | 1.09 | 1.30 | 1.14 | 0.75 | 0.70 | 0.78 | 0.92 | 0.78 | 0.61 | 0.08 | 0.35 |
C. sp.13 (N = 2) | ||||||||||||||||
Mean | 1.83 | 1.45 | 0.94 | 0.94 | 1.07 | 1.01 | 1.27 | 1.11 | 0.91 | 0.67 | 0.75 | 0.88 | 0.76 | 0.74 | 0.15 | 0.34 |
SD | 0.040 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 | 0.000 |
Lower | 1.80 | 1.45 | 0.94 | 0.94 | 1.07 | 1.01 | 1.27 | 1.11 | 0.91 | 0.67 | 0.75 | 0.88 | 0.76 | 0.74 | 0.15 | 0.34 |
Upper | 1.86 | 1.45 | 0.94 | 0.94 | 1.07 | 1.01 | 1.27 | 1.11 | 0.91 | 0.67 | 0.75 | 0.88 | 0.76 | 0.74 | 0.15 | 0.34 |
Adult female ZMKU R 00947 from Pa La-U Waterfall, Kaeng Krachan National Park, Huai Sat Yai Subdistrict, Hua Hin District, Prachuap Khiri Khan Province, Thailand (12.53685°N, 99.45972°E, 368 m a.s.l.), collected by Attapol Rujirawan, Siriporn Yodthong, Korkhwan Termprayoon, Natee Ampai, and Piyawan Puanprapai on 15 September 2017.
Adult female ZMKU R 00946 bearing the same data as the holotype.
Cyrtodactylus rivularis sp. nov. can be separated from all other species of the brevipalmatus group by the combination of having 12 or 13 supralabials, 9–11 infralabials, 33 or 34 paravertebral tubercles, 18–20 rows of longitudinally arranged tubercles, 34–37 transverse rows of ventrals, 160–166 longitudinal rows of ventrals, nine expanded subdigital lamellae on the fourth toe, 12 or 13 unexpanded subdigital lamellae on the fourth toe, 21 or 22 total subdigital lamellae on the fourth toe; eight expanded subdigital lamellae on the fourth finger, 10–12 unexpanded subdigital lamellae on the fourth finger, 18–20 total subdigital lamellae on the fourth finger; 14–16 total enlarged femoral scales; 15 enlarged precloacals; enlarged femorals and enlarged precloacals not continuous, and lacking pores; proximal femorals less than one-half the size of the distal femorals; small tubercles on forelimbs and flanks; large dorsolateral caudal tubercles and wide ventrolateral caudal fringe; ventrolateral caudal fringe composed generally homogeneous scales; tail square in cross-section; single enlarged unpaired medial subcaudals not posteromedially furrowed; maximum SVL 73.9 mm; three or four dark transverse body bands (Tables
(Fig.
Holotype of Cyrtodactylus rivularis sp. nov. ZMKU R 00947 (field no. AA 04887) from Thailand, Prachuap Khiri Khan Province, Hua Hin District, Huai Sat Yai Subdistrict, Kaeng Krachan National Park, Pa La-U Waterfall A dorsal view B ventral view C dorsal view of head D ventral view of femoral and precloacal regions E dorsal view of tail F ventral view of tail.
Body relatively short (AG/SVL 0.47) with well-defined ventrolateral folds; dorsal scales small, granular, interspersed with larger conical, semi-regularly arranged, weakly keeled tubercles; tubercles extend from occipital region onto base of tail and slightly beyond as paravertebral rows; tubercles of nape and occiput small; approximately 20 longitudinal rows of tubercles at midbody; approximately 34 paravertebral tubercles; tubercles on flanks nearly same size as those on dorsum; 34 longitudinal rows of flat, imbricate, ventral scales much larger than dorsal scales; 160 transverse rows of ventral scales; no pore-bearing, precloacal scales; 15 enlarged precloacal scales; no deep precloacal groove or depression; and three rows of post-precloacal scales on midline.
Forelimbs moderate in stature, relatively short (ForL/SVL 0.13); granular scales of forearm slightly larger than those on body, interspersed with tubercles; palmar scales rounded, slightly raised; digits well-developed, relatively short, inflected at basal interphalangeal joints; digits narrower distal to inflections; subdigital lamellae wide, transversely expanded proximal to joint inflections, narrower transverse lamellae distal to joint inflections; claws well-developed, claw base sheathed by a dorsal and ventral scale; 8R/8L expanded and 11R/11L unexpanded lamellae beneath the fourth finger; hind limbs larger and thicker than forelimbs, moderate in length (TibL/SVL 0.15), covered dorsally by granular scales interspersed with moderately sized, conical tubercles dorsally and posteriorly and anteriorly by flat, slightly larger, subimbricate scales; ventral scales of thigh flat, subimbricate, larger than dorsals; subtibial scales flat, imbricate; no pore-bearing femoral scales; 8R/8L enlarged femoral scales; enlarged femoral scales not contiguous with enlarged precloacal scales, terminating distally at knee; proximal femoral scales smaller than distal femoral scales, the former forming an abrupt union with much smaller, rounded, ventral scales of posteroventral margin of thigh; plantar scales flat; digits relatively long, well-developed, inflected at basal interphalangeal joints; 9R/9L wide, transversely expanded subdigital lamellae on fourth toe proximal to joint inflection that extend onto sole, and 13R/13L unexpanded lamellae beneath the fourth toe; and claws well-developed, sheathed by a dorsal and ventral scale at base.
Posterior one-half of tail regenerated, tail long 91.5 mm (TL/SVL 1.24), 4.8 mm in width at base, tapering to a point; nearly square in cross-section; dorsal scales flat, square bearing large tubercles forming a discontinuous dorsolateral longitudinal row; slightly larger, posteriorly directed, semi-spinose tubercles forming large distinct ventrolateral caudal fringe; scales of ventrolateral fringe generally homogeneous; single medial subcaudals enlarged but not paired; subcaudal scales, larger than dorsal caudal scales; base of tail bearing hemipenial swellings; 2R/2L conical postcloacal tubercles at base of hemipenial swellings; and postcloacal scales flat, imbricate.
(Fig.
(Fig.
Cyrtodactylus rivularis sp. nov. is currently known from the type locality at Pa La-U Waterfall, Kaeng Krachan National Park, Huai Sat Yai Subdistrict, Hua Hin District Prachuap Khiri Khan Province, Thailand.
The specific epithet rivularis is derived from the Latin rivus, meaning stream, brook, or creek refers to rocky brook or stream habitat of the new species.
Cyrtodactylus rivularis sp. nov. is the sister species to C. rukhadeva (Fig.
The holotype and paratype were collected at night (1900–2055 h) on granite boulders by a rocky stream dry evergreen forest at 368 m in elevation (Fig.
Cyrtodactylus rivularis sp. nov. and C. cf. rukhadeva occur on separate branches of the phylogeny, thus neither is embedded within one another’s branch indicating there is no evidence of gene flow between them. Both species are reported to occur within the boundaries of Kaeng Krachan National Park (
Sex and raw meristic, categorical, and morphometric data used in the analyses from specimens in the Cyrtodactylus brevipalmatus group. m = male; f = female; R/L = right/left; / = data unavailable.
Species | C. brevipalmatus | C. cf. brevipalmatus | C. cf. brevipalmatus | C. brevipalmatus | C. brevipalmatus | C. elok | C. elok | C. elok | C. elok | C. interdigitalis | C. interdigitalis | C. interdigitalis | C. interdigitalis | C. sp.11 | C. ngati | C. ngati | C. ngati | C. ngati | C. ngati3 | C. ngati3 | C. ngati3 | C. ngati4 | C. cf. ngati1 | C. cf. ngati2 | C. cf. ngati2 | C. rukhadeva | C. rukhadeva | C. rukhadeva | C. cf. rukhadeva | C. cf. rukhadeva | C. cf. rukhadeva | C. cf. rukhadeva | C. cf. rukhadeva | C. cf. rukhadeva | C. cf. rukhadeva | Cyrtodactylus fluvicavus sp. nov. | Cyrtodactylus fluvicavus sp. nov. | Cyrtodactylus fluvicavus sp. nov. | Cyrtodactylus fluvicavus sp. nov. | Cyrtodactylus fluvicavus sp. nov. | Cyrtodactylus fluvicavus sp. nov. | Cyrtodactylus fluvicavus sp. nov. | Cyrtodactylus kochangensis sp. nov | C. cf. kochangensis | Cyrtodactylus uthaiensis sp. nov. | Cyrtodactylus rivularis sp. nov. | Cyrtodactylus rivularis sp. nov. | C. sp.9 Thong Pha Phum | C. sp.9 Thong Pha Phum | C. sp.9 Thong Pha Phum | C. sp.9 Thong Pha Phum | C. sp.9 Thong Pha Phum | C. sp.9 Thong Pha Phum | C. sp.9 Thong Pha Phum | C. sp.9 Thong Pha Phum | C. sp.13 | C. sp.13 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Institutional catalog number | LSUHC 1899 | LSUHC 15076 | LSUHC 11788 | THNHM 10670 | THNHM 14112 | LSUHC 8238 | LSUHC 12180 | LSUHC 12181 | ZMMU R-16144 | THNHM 20226 paratype | THNHM 20228 paratype | THNHM 20229 paratype | THNHM 20227 paratype | ZMMU R-16492 | HNUE-R00111 holotype | IEBR 4829 paratype | VNUF R.2020.12 paratype | HNUE-R00112 paratype | FMNH 255454 | FMNH 270493 | FMNH 270492 | FMNH 265806 | NCSM 79472 | ZMMU R-14917 | NCSM 80100 | ZMMU R-16851 holotype | ZMMU R-16852 paratype | ZMKU R 00948 topotype | THNHM 24622 | THNHM 24838 | THNHM 03251 | THNHM 03252 | THNHM 03253 | THNHM 03254 | THNHM 01807 | ZMKU R 00959 holotype | ZMKU R 00958 paratype | ZMKU R 00960 paratype | ZMKU R 00961 paratype | ZMKU R 00962 paratype | ZMKU R 00963 paratype | ZMKU R 00964 paratype | ZMKU R 00945 holotype | THNHM 01667 | ZMKU R 00949 holotype | ZMKU R 00947 holotype | ZMKU R 00946 paratype | ZMKU R 00950 | ZMKU R 00951 | ZMKU R 00952 | ZMKU R 00953 | ZMKU R 00954 | ZMKU R 00955 | ZMKU R 00956 | ZMKU R 00957 | THNHM 00104 | THNHM 27821 |
Sex | m | f | f | f | f | f | m | m | f | f | f | f | f | m | m | f | f | f | f | m | m | m | f | f | f | m | f | f | m | f | m | m | f | m | m | m | m | m | f | f | f | f | f | m | m | f | f | f | m | f | m | m | f | m | f | f | f |
Meristic data | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Supralabials (SL) | 11 | 12 | 10 | 14 | 12 | 11 | 8 | 13 | 9 | 14 | 12 | 11 | 12 | 11 | 10 | 10 | 10 | 10 | 13 | 13 | 13 | 10 | 14 | 9 | 12 | 11 | 9 | 14 | 11 | 13 | 13 | 11 | 12 | 13 | 12 | 12R/12L | 13R/12L | 13R/12L | 11R/12L | 12R/12L | 13R/12L | 12R/11L | 12R/13L | 12 | 13R/15L | 13R/12L | 13R/12L | 12 | 13 | 13 | 14 | 13 | 13 | 13 | 13 | 12 | 15 |
Infralabials (IL) | 8 | 10 | 9 | 11 | 11 | 11 | 8 | 11 | 9 | 9 | 8 | 8 | 7 | 9 | 9 | 9 | 9 | 9 | 10 | 9 | 11 | 8 | 11 | 10 | 12 | 10 | 11 | 9 | 10 | 10 | 10 | 10 | 10 | 11 | 10 | 10R/10L | 10R/10L | 9R/10L | 10R/10L | 10R/10L | 10R/10L | 10R/10L | 9R/9L | 10 | 10R/11L | 11R/10L | 10R/9L | 8 | 8 | 10 | 10 | 9 | 10 | 10 | 9 | 10 | 11 |
Paravertebral tubercles (PVT) | 39 | 37 | 38 | 37 | 37 | 0 | 0 | 0 | 0 | 32 | 33 | 33 | 33 | 30 | 39 | 40 | 38 | 40 | 28 | 27 | 26 | 27 | 28 | 32 | 29 | 27 | 30 | 30 | 26 | 28 | 27 | 27 | 30 | 30 | 26 | 30 | 28 | 27 | 27 | 28 | 26 | 28 | 34 | 29 | 33 | 34 | 33 | 32 | 33 | 34 | 34 | 36 | 36 | 30 | 30 | 33 | 29 |
Longitudinal rows of tubercles (LRT) | 15 | 16 | 17 | 16 | 14 | 6 | 7 | 4 | 4 | 19 | 20 | 19 | 19 | 18 | 18 | 18 | 17 | 22 | 19 | 18 | 17 | 19 | 18 | 24 | 19 | 19 | 20 | 19 | 18 | 19 | 18 | 18 | 19 | 19 | 19 | 17 | 17 | 14 | 16 | 17 | 18 | 16 | 14 | 19 | 17 | 20 | 18 | 21 | 19 | 20 | 20 | 21 | 21 | 19 | 19 | 18 | 20 |
Ventral scales (VS) | 38 | 38 | 38 | 36 | 39 | 45 | 45 | 47 | 36 | 42 | 40 | 42 | 43 | 34 | 38 | 36 | 35 | 32 | 37 | 36 | 36 | 33 | 33 | 36 | 35 | 34 | 43 | 38 | 38 | 36 | 37 | 37 | 39 | 34 | 35 | 34 | 37 | 33 | 30 | 36 | 37 | 39 | 35 | 34 | 36 | 34 | 37 | 34 | 33 | 33 | 34 | 30 | 33 | 32 | 33 | 37 | 36 |
Ventral scales along middle of the body (VSM) | 176 | 170 | 182 | 154 | 160 | 190 | 225 | 234 | 192 | 187 | 170 | 187 | 178 | 160 | 168 | 164 | 178 | 158 | 159 | 166 | 156 | 158 | 164 | 166 | 165 | 154 | 152 | 165 | 162 | 158 | 157 | 159 | 168 | 160 | 161 | 155 | 154 | 155 | 172 | 164 | 175 | 170 | 172 | 159 | 159 | 160 | 166 | 173 | 158 | 156 | 166 | 159 | 159 | 150 | 169 | 159 | 165 |
Expanded subdigital lamellae on 4th toe (TL4E) | 7 | 8 | 9 | 8 | 8 | 10 | 9 | 9 | 9 | 12 | 10 | 10 | 11 | 9 | 8 | 10 | 9 | 9 | 10 | 10 | 8 | 10 | 9 | 8 | 10 | 9 | 9 | 9 | 8 | 9 | 9 | 10 | 9 | 10 | 10 | 9R/9L | 10R/10L | 9R/9L | 9R/9L | 10R/11L | 9R/10L | 9R/9L | 9R/8L | 8 | 8R/(broken)L | 9R/9L | 9R/9L | 9 | 10 | 9 | 8 | 10 | 8 | 9 | 9 | 9 | 7 |
Unmodified subdigital lamellae on 4th toe (TL4U) | 13 | 11 | 11 | 11 | 12 | 11 | 10 | 11 | 9 | 14 | 13 | 12 | 14 | 10 | 11 | 10 | 11 | 10 | 11 | 11 | 11 | 11 | 12 | 10 | 10 | 11 | 11 | 12 | 11 | 13 | 12 | 12 | 15 | 13 | 13 | 11R/11L | 12R/11L | 10R/10L | 12R/12L | 11R/11L | 10R/10L | 12R/13L | 12R/11L | 13 | 12R/(broken)L | 13R/13L | 12R/13L | 12 | 14 | 13 | 12 | 13 | 12 | 11 | 13 | 12 | 12 |
Total subdigital lamellae 4th toe (TL4T) | 20 | 19 | 20 | 19 | 20 | 21 | 19 | 20 | 18 | 26 | 23 | 22 | 23 | 19 | 13 | 16 | 17 | 16 | 21 | 21 | 19 | 21 | 21 | 18 | 20 | 20 | 18 | 21 | 19 | 22 | 21 | 22 | 14 | 23 | 23 | 20R/20L | 22R/21L | 19R/19L | 21R/21L | 21R/22L | 19R/20L | 22R/22L | 21R/19L | 21 | 20 | 22R/22L | 21R/22L | 21 | 24 | 22 | 20 | 23 | 20 | 20 | 22 | 21 | 19 |
Expanded subdigital lamellae on 4th finger (FL4E) | 8 | 8 | 8 | 7 | 8 | 8 | 9 | 9 | 9 | 9 | 8 | 9 | 9 | 10 | 6 | 6 | 7 | 6 | 8 | 8 | 8 | 8 | 9 | 7 | 9 | 9 | 8 | 8 | 7 | 8 | 8 | 8 | 8 | 8 | 8 | 8R/8L | 8R/8L | 8R/8L | 8R/8L | 7R/7L | 8R/9L | 7R/7L | 8R/8L | 8 | 7R/7L | 8R/8L | 8R/8L | 8 | 7 | 7 | 8 | 8 | 8 | 8 | 8 | 8 | 8 |
Unmodified subdigital lamellae on 4th finger (FL4U) | 9 | 11 | 10 | 10 | 10 | 12 | 13 | 9 | 8 | 12 | 11 | 12 | 13 | 9 | 9 | 9 | 9 | 9 | 10 | 10 | 10 | 10 | 8 | 9 | 10 | 10 | 9 | 11 | 10 | 11 | 10 | 10 | 12 | 12 | 12 | 10R/10L | 10R/10L | 10R/9L | 11R/11L | 10R/10L | 9R/9L | 10R/10L | 10R/10L | 12 | 11R/11L | 11R/10L | 12R/12L | 10 | 12 | 12 | 11 | 12 | 12 | 11 | 12 | 11 | 10 |
Total subdigital lamellae 4th finger (FL4T) | 17 | 19 | 18 | 17 | 18 | 20 | 22 | 18 | 17 | 21 | 21 | 21 | 22 | 19 | 15 | 15 | 18 | 15 | 18 | 18 | 18 | 18 | 17 | 16 | 19 | 19 | 17 | 19 | 17 | 17 | 18 | 18 | 20 | 20 | 20 | 18R/18L | 18R/18L | 18R/17L | 19R/19L | 17R/17L | 17R/18L | 17R/17L | 18R/18L | 20 | 18R/18L | 19R/18L | 20R/20L | 18 | 19 | 19 | 19 | 20 | 20 | 19 | 20 | 19 | 18 |
Enlarged femoral scales (R/L) | 0 | 0 | 0 | 8R/8L | 7R/7L | 0 | 0 | 0 | 0 | 11R/8L | 10R/9L | 8R/8L | 9R/10L | 9R/8L | 10R/10L | 9R/8L | 10R/9L | 8R/9L | 9R/7L | 8R/9L | 9R/9L | 8R/8L | 9R/8L | 7R/8L | 7R/8L | 9R/8L | 8R/8L | 9R/8L | 9R/L | 9R/9L | 9R/7L | 7R/7L | 6R/7L | 5R/8L | 7R/7L | 5R/6L | 4R/5L | 5R/6L | 6R/6L | 5R/6L | 5R/6L | 6R/6L | 6R/6L | 7R/7L | 8R/8L | 8R/8L | 6R/8L | 5R/7L | 8R/8L | 8R/8L | 7R/8L | 8R/8L | 7R/8L | 7R/6L | 8R/8L | 9R/9L | 7R/10L |
Total enlarged femoral scales (FS) | 16 | 10 | 11 | 16 | 14 | 0 | 0 | 0 | 0 | 14 | 19 | 19 | 19 | 17 | 20 | 17 | 19 | 17 | 16 | 17 | 18 | 16 | 17 | 15 | 15 | 17 | 16 | 17 | 18 | 18 | 16 | 14 | 13 | 13 | 14 | 11 | 9 | 11 | 12 | 11 | 11 | 12 | 12 | 14 | 16 | 16 | 14 | 12 | 16 | 16 | 15 | 16 | 15 | 13 | 16 | 18 | 17 |
Total femoral pores in males (FP) | 7 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 17 | 14 | 0 | 0 | 0 | 0 | 14 | 15 | 13 | 0 | 0 | 0 | 17 | 0 | 0 | 14 | 0 | 12 | 13 | 0 | 11 | 13 | 11 | 8 | 10 | 0 | 0 | 0 | 0 | 0 | 14 | 12 | 0 | 0 | 0 | 16 | 0 | 14 | 15 | 14 | 12 | 0 | 0 | 0 |
Enlarged precloacal scales (PCS) | 7 | 7 | 7 | 8 | 7 | 8 | 8 | 8 | 7 | 14 | 15 | 13 | 19 | 13 | 13 | 13 | 13 | 13 | 15 | 13 | 13 | 13 | 12 | 13 | 13 | 17 | 13 | 15 | 15 | 15 | 14 | 13 | 15 | 15 | 14 | 15 | 14 | 14 | 15 | 14 | 15 | 15 | 12 | 16 | 14 | 15 | 15 | 17 | 15 | 15 | 15 | 15 | 15 | 15 | 15 | 14 | 16 |
Precloacal pores in males (PP) | 7 | 0 | 0 | 0 | 0 | 0 | 8 | 8 | 0 | 0 | 0 | 0 | 0 | 13 | 0 | 0 | 0 | 0 | 13 | 13 | 13 | 13 | 0 | 0 | 0 | 17 | 13 | 0 | 15 | 0 | 14 | 13 | 0 | 15 | 14 | 15 | 14 | 14 | 0 | 0 | 0 | 0 | 0 | 16 | 14 | 0 | 0 | 0 | 15 | 0 | 15 | 15 | 15 | 15 | 0 | 0 | 0 |
Postcloacal tubercles (PCT) | 3 | 3 | 2 | 3 | 3 | 3 | 2 | 3 | 3 | 3 | 2 | 3 | 3 | 3 | 3 | 2 | 1 | 2 | 0 | 0 | 0 | 0 | 2 | 3 | 4 | 3 | 2 | 2R/3L | 3 | 2 | 3 | 2 | 2 | 3 | 2 | 3R/2L | 3R/2L | 3R/3L | 1R/1L | 3R/2L | 3R/3L | 2R/2L | 1R/1L | 3 | 3R/3L | 2R/2L | 3R/3L | 2 | 2R/3L | 3 | 3 | 2R/3L | 2R/3L | 3 | 2 | 3 | 3 |
Body bands (BB) | 4 | 6 | 3 | 5 | 5 | 5 | 5 | 3 | 3 | 5 | 5 | 5 | 5 | 3 | 6 | 6 | 6 | 6 | 3 | 4 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 4 | 4 | / | / | 5 | 3 | 3 | 3 | 3 | 3 | 3 | 3 | 5 | 5 | 6 | 3 | 4 | 3 | 4 | 3 | 4 | 3 | 5 | 4 | 4 | 3 | / |
Categorical data | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Small tubercles on flank (FKT) | present | present | present | present | present | absent | absent | absent | absent | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present | present |
Dorsolateral caudal tubercles (DCT) | small | small | small | / | small | large | large | large | large | small | / | small | small | large | small | small | small | small | small | small | small | small | small | small | small | small | small | small | small | small | small | small | small | small | / | small | small | small | small | small | small | small | large | large | large | large | large | large | large | large | large | large | large | / | large | small | small |
Ventrolateral caudal fringe narrow or wide (VLF1) | narrow | narrow | narrow | / | narrow | wide | wide | wide | wide | narrow | / | narrow | narrow | wide | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | / | narrow | narrow | narrow | narrow | narrow | narrow | narrow | wide | wide | wide | wide | wide | wide | wide | wide | wide | wide | wide | / | wide | narrow | narrow |
Ventrolateral caudal fringe scales generally homogenous (VLF2) | no | no | no | / | no | no | no | no | no | yes | yes | yes | yes | yes | no | no | no | no | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | yes | / | no | no | no | no | no | no | no | no | no | no | yes | yes | no | no | no | no | no | no | / | no | yes | yes |
Tail cross-section (TLcross) | circular | circular | circular | / | circular | square | square | square | square | circular | / | circular | circular | square | circular | circular | circular | circular | circular | circular | circular | circular | circular | circular | circular | square | square | square | square | square | square | square | square | square | / | circular | circular | circular | circular | circular | circular | circular | square | / | circular | square | square | square | square | square | square | square | square | / | square | circular | circular |
Slightly enlarged medial subcaudals (SC1) | present | present | present | / | absent | absent | absent | absent | absent | absent | / | absent | absent | present | present | present | present | present | / | present | present | present | present | present | present | absent | absent | absent | absent | absent | absent | absent | absent | absent | / | present | present | present | present | present | present | present | present | present | present | absent | absent | present | present | present | present | present | present | / | present | present | present |
Single enlarged medial subcaudal (SC2) | absent | absent | absent | / | absent | absent | absent | absent | absent | absent | / | absent | absent | absent | absent | absent | absent | absent | / | absent | absent | absent | absent | absent | absent | present | present | present | present | present | present | present | present | present | / | absent | absent | absent | absent | absent | absent | absent | absent | absent | absent | present | present | absent | absent | absent | absent | absent | absent | / | absent | absent | absent |
Enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginate (SC3) | no | no | no | / | no | no | no | no | no | yes | / | yes | yes | no | no | no | no | no | / | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | no | yes | no | no | no | no | no | no | yes | no | / | no | no | no |
Morphometric data | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||
SVL | 68.8 | 70.8 | 64.1 | 65.95 | 63.79 | 80.2 | 78.2 | 84.8 | 78.6 | 81.19 | 74.80 | 78.56 | 59.70 | 68.1 | 66.5 | 68.1 | 69.3 | 46.6 | 83.6 | 70.2 | 74.1 | 73.8 | 78.0 | 87.1 | 77.7 | 74.9 | 71.7 | 71.6 | 68.3 | 71.8 | 73.6 | 75.3 | 74.7 | 73.2 | 61.5 | 72.5 | 72.0 | 69.6 | 68.4 | 76.8 | 65.7 | 78.2 | 60.1 | 70.2 | 58.1 | 73.9 | 68.1 | 73.1 | 73.5 | 73.7 | 73.2 | 64.4 | 76.6 | 76.6 | 74.2 | 63.7 | 72.9 |
AG | 35.7 | 33.4 | 30.1 | 30.0 | 26.5 | 39.7 | 37.8 | 41.5 | 36.2 | 34.5 | 33.7 | 32.7 | 24.6 | 34.6 | 28.8 | 29.8 | 30.2 | 19.7 | 41.3 | 35.4 | 37.0 | 31.3 | 38.2 | 41.9 | 36.8 | 34.6 | 32.6 | 33.9 | 27.3 | 29.9 | 30.9 | 31.3 | 32.2 | 30.3 | 26.2 | 33.4 | 33.6 | 32.0 | 30.4 | 35.6 | 30.6 | 38.1 | 29.0 | 31.5 | 26.6 | 34.8 | 33.2 | 34.8 | 33.9 | 35.4 | 33.6 | 28.5 | 37.1 | 33.2 | 35.1 | 25.8 | 30.6 |
HumL | 9.7 | 9.3 | 8.0 | 9.6 | 9.5 | 10.2 | 9.1 | 10.1 | 1.7 | 9.8 | 10.2 | 11.2 | 7.4 | 10.3 | 7.9 | 8.1 | 8.5 | 5.6 | 8.6 | 8.7 | 8.6 | 6.9 | 8.7 | 11.5 | 9.2 | 10.7 | 10.4 | 7.9 | 9.8 | 8.3 | 12.2 | 11.3 | 11.8 | 11.0 | 10.1 | 9.1 | 8.8 | 9.0 | 8.0 | 10.0 | 7.5 | 10.1 | 6.5 | 10.2 | 7.0 | 8.1 | 7.6 | 8.4 | 7.2 | 9.0 | 9.0 | 7.2 | 8.0 | 8.1 | 8.6 | 7.6 | 10.1 |
ForL | 9.9 | 9.8 | 8.9 | 8.2 | 8.7 | 11.5 | 11.7 | 11.8 | 10.2 | 10.6 | 10.5 | 11.1 | 8.4 | 8.5 | 9.2 | 10.0 | 10.1 | 6.5 | 10.2 | 9.3 | 10.4 | 10.0 | 10.3 | 10.4 | 10.7 | 8.6 | 7.9 | 9.6 | 8.7 | 8.5 | 9.0 | 10.6 | 9.6 | 9.2 | 7.9 | 10.5 | 10.3 | 10.5 | 10.1 | 11.1 | 8.8 | 10.8 | 7.6 | 8.6 | 8.3 | 9.7 | 9.1 | 9.5 | 9.1 | 9.2 | 9.8 | 9.2 | 10.0 | 8.6 | 9.8 | 8.1 | 9.6 |
FemL | 12.0 | 12.6 | 11.5 | 11.7 | 9.8 | 12.9 | 14.2 | 14.6 | 13.1 | 14.7 | 13.2 | 12.7 | 10.2 | 12.6 | 11.5 | 11.5 | 11.5 | 7.6 | 13.7 | 12.7 | 13.0 | 13.1 | 13.1 | 15.2 | 14.2 | 12.6 | 11.8 | 10.5 | 10.8 | 10.9 | 11.5 | 10.2 | 11.9 | 12.1 | 9.5 | 13.1 | 12.5 | 12.5 | 13.5 | 14.1 | 11.5 | 13.9 | 10.4 | 12.1 | 10.0 | 11.4 | 10.4 | 12.8 | 11.6 | 12.3 | 12.5 | 10.9 | 13.7 | 10.8 | 12.5 | 10.7 | 12.8 |
TibL | 11.6 | 12.2 | 10.5 | 9.7 | 8.2 | 13.5 | 14.0 | 13.8 | 12.3 | 13.1 | 11.9 | 12.9 | 10.2 | 11.4 | 10.8 | 11.1 | 11.8 | 7.8 | 12.5 | 11.8 | 11.2 | 11.1 | 12.8 | 12.6 | 12.7 | 10.1 | 9.3 | 11.2 | 9.7 | 10.7 | 10.9 | 11.7 | 11.3 | 11.1 | 9.1 | 11.3 | 10.6 | 10.2 | 9.9 | 11.2 | 9.4 | 12.3 | 8.4 | 11.8 | 8.4 | 11.2 | 10.3 | 10.5 | 10.1 | 10.6 | 10.6 | 9.9 | 11.1 | 10.0 | 11.4 | 10.1 | 10.2 |
HL | 19.3 | 19.3 | 19.0 | 17.9 | 18.2 | 21.8 | 21.6 | 21.9 | 21.7 | 20.8 | 19.9 | 21.7 | 16.7 | 18.4 | 20.1 | 20.4 | 20.7 | 16.1 | 21.7 | 20.6 | 20.3 | 20.7 | 21.2 | 22.1 | 21.4 | 20.2 | 19.2 | 19.7 | 19.7 | 19.9 | 20.8 | 21.3 | 20.8 | 21.5 | 17.9 | 20.1 | 20.5 | 19.7 | 20.1 | 21.2 | 18.6 | 21.3 | 17.3 | 18.3 | 16.1 | 20.3 | 19.3 | 19.9 | 20.9 | 20.1 | 20.0 | 17.6 | 20.4 | 19.3 | 20.0 | 17.6 | 19.9 |
HW | 13.2 | 13.8 | 12.3 | 12.3 | 12.0 | 15.6 | 16.1 | 15.9 | 15.1 | 14.0 | 13.4 | 14.2 | 11.4 | 13.1 | 12.6 | 12.0 | 11.8 | 8.8 | 13.8 | 12.5 | 13.0 | 12.3 | 12.7 | 14.8 | 13.5 | 14.6 | 13.4 | 14.0 | 13.1 | 13.9 | 14.9 | 15.0 | 13.1 | 14.1 | 11.8 | 14.0 | 13.4 | 12.9 | 13.0 | 14.9 | 13.0 | 15.4 | 11.6 | 12.1 | 10.9 | 14.9 | 13.7 | 14.5 | 14.3 | 15.7 | 13.9 | 12.8 | 14.7 | 14.4 | 14.1 | 11.9 | 13.8 |
HD | 8.0 | 7.6 | 7.6 | 7.3 | 7.0 | 9.6 | 9.8 | 10.4 | 9.8 | 3.4 | 8.6 | 8.7 | 6.6 | 8.3 | 7.4 | 7.2 | 6.6 | 5.1 | 9.2 | 8.4 | 9.1 | 7.6 | 8.3 | 8.7 | 9.2 | 9.2 | 8.5 | 8.3 | 7.3 | 8.9 | 8.2 | 8.2 | 8.1 | 8.9 | 7.5 | 8.5 | 8.1 | 8.3 | 7.9 | 8.1 | 7.8 | 8.3 | 6.5 | 7.8 | 6.3 | 8.2 | 8.2 | 7.8 | 7.7 | 7.9 | 7.7 | 7.0 | 8.2 | 7.8 | 7.6 | 7.7 | 8.4 |
ED | 5.2 | 4.5 | 4.3 | 5.3 | 4.4 | 4.8 | 5.0 | 5.7 | 5.0 | 5.3 | 5.5 | 5.9 | 4.4 | 4.4 | 3.8 | 4.1 | 3.4 | 2.6 | 4.9 | 4.9 | 4.9 | 4.8 | 6.5 | 4.6 | 6.0 | 4.6 | 4.3 | 5.5 | 4.9 | 5.1 | 5.8 | 5.4 | 5.0 | 5.5 | 4.7 | 5.0 | 5.0 | 4.9 | 4.7 | 5.1 | 4.5 | 5.3 | 4.2 | 5.2 | 4.6 | 5.8 | 5.6 | 5.0 | 5.1 | 5.0 | 5.0 | 4.8 | 5.6 | 5.3 | 4.9 | 4.1 | 5.3 |
EE | 5.7 | 5.9 | 4.9 | 5.7 | 5.7 | 6.4 | 7.1 | 7.0 | 6.8 | 5.8 | 6.2 | 6.4 | 4.8 | 6.2 | 5.8 | 5.5 | 5.9 | 4.4 | 6.9 | 6.1 | 6.2 | 5.7 | 5.3 | 6.5 | 6.2 | 6.2 | 6.2 | 5.8 | 5.1 | 6.2 | 5.6 | 5.7 | 5.4 | 6.2 | 4.3 | 6.5 | 5.9 | 5.7 | 5.8 | 6.1 | 5.4 | 6.5 | 5.0 | 4.9 | 4.7 | 6.5 | 6.2 | 5.9 | 5.9 | 6.0 | 5.9 | 5.3 | 6.1 | 6.0 | 6.0 | 4.9 | 6.3 |
ES | 7.4 | 7.6 | 7.0 | 7.0 | 7.2 | 8.6 | 8.7 | 9.5 | 8.6 | 8.3 | 7.8 | 9.1 | 6.8 | 7.7 | 7.5 | 7.6 | 6.9 | 5.0 | 9.0 | 8.3 | 8.3 | 8.2 | 8.7 | 8.8 | 8.4 | 8.3 | 7.7 | 7.9 | 7.4 | 8.1 | 8.4 | 8.8 | 8.1 | 8.6 | 7.3 | 8.5 | 8.3 | 8.2 | 8.1 | 9.2 | 7.3 | 9.3 | 6.9 | 7.5 | 6.4 | 8.3 | 7.9 | 7.9 | 8.5 | 7.9 | 7.9 | 7.3 | 8.2 | 7.9 | 7.9 | 7.2 | 8.0 |
EN | 5.7 | 5.4 | 4.9 | 5.3 | 5.4 | 6.0 | 6.2 | 6.5 | 6.2 | 6.0 | 5.5 | 6.8 | 5.1 | 5.5 | 6.7 | 6.3 | 6.2 | 4.5 | 6.5 | 6.2 | 6.1 | 6.2 | 6.2 | 6.6 | 6.0 | 6.3 | 5.7 | 5.8 | 5.4 | 6.0 | 6.2 | 6.4 | 5.8 | 6.2 | 5.3 | 6.5 | 6.2 | 5.9 | 6.1 | 6.6 | 5.6 | 6.5 | 5.2 | 5.5 | 4.9 | 6.1 | 5.8 | 6.0 | 6.1 | 6.0 | 5.8 | 5.4 | 6.1 | 6.0 | 5.9 | 5.6 | 5.9 |
IO | 5.4 | 4.7 | 4.7 | 4.2 | 5.2 | 5.7 | 5.4 | 5.4 | 3.9 | 4.8 | 4.7 | 5.5 | 4.3 | 2.9 | 5.6 | 5.4 | 5.6 | 4.2 | 6.6 | 5.6 | 5.4 | 5.1 | 4.9 | 3.5 | 5.7 | 3.3 | 3.1 | 5.6 | 4.5 | 4.7 | 5.6 | 5.7 | 5.7 | 5.6 | 4.2 | 5.5 | 5.4 | 5.3 | 5.1 | 5.6 | 5.0 | 5.6 | 4.2 | 4.0 | 4.3 | 5.8 | 5.5 | 5.4 | 5.5 | 5.8 | 5.5 | 4.9 | 5.7 | 5.6 | 5.3 | 4.8 | 6.1 |
EL | 1.0 | 1.4 | 1.1 | 1.3 | 1.0 | 1.9 | 1.4 | 1.5 | 1.4 | 1.3 | 1.3 | 1.6 | 1.2 | 0.9 | 0.8 | 0.8 | 0.7 | 0.3 | 1.3 | 1.1 | 1.2 | 1.0 | 1.5 | 1.2 | 0.9 | 1.2 | 1.0 | 1.4 | 1.6 | 1.5 | 1.2 | 1.3 | 1.2 | 1.2 | 0.9 | 1.4 | 1.5 | 1.7 | 1.4 | 1.8 | 1.6 | 1.8 | 1.0 | 1.3 | 1.5 | 1.1 | 1.1 | 1.1 | 1.5 | 1.5 | 1.2 | 1.2 | 1.0 | 1.2 | 1.3 | 1.4 | 1.4 |
IN | 1.7 | 2.1 | 2.3 | 2.1 | 2.2 | 2.7 | 2.6 | 2.5 | 3.1 | 2.1 | 2.2 | 2.5 | 1.8 | 2.3 | 2.8 | 2.6 | 2.6 | 2.0 | 2.8 | 2.5 | 2.5 | 2.3 | 2.7 | 2.2 | 2.5 | 2.2 | 2.1 | 2.1 | 2.0 | 2.2 | 2.4 | 2.5 | 2.4 | 2.3 | 2.0 | 2.3 | 2.4 | 2.5 | 2.3 | 2.3 | 2.3 | 2.6 | 1.9 | 2.2 | 1.8 | 2.3 | 2.0 | 2.3 | 2.4 | 2.2 | 2.0 | 2.0 | 2.3 | 2.2 | 2.2 | 2.1 | 2.3 |
Adult female ZMKU R 00945 from Ko Chang Island, Ko Phayam Subdistrict, Mueang Ranong District, Ranong Province, Thailand (9.82411°N, 98.43999°E, 36 m a.s.l.), collected by Siriporn Yodthong, Natee Ampai, Attapol Rujirawan, and Piyawan Puanprapai on 8 July 2017.
Cyrtodactylus cf. kochangensis sp. nov. adult male THNHM 01667 from Khlong Naka Wildlife Sanctuary, Suk Samran District, Ranong Province, Thailand (~N 9.4596, E 98.5044, elevation unknown), collected by Yodchaiy Chuaynkern between 28 December 2000 and 2 January 2001.
(based on the holotype). Cyrtodactylus kochangensis sp. nov. can be separated from all other species of the brevipalmatus group by the combination of having 12 or 13 supralabials, nine infralabials, 34 paravertebral tubercles, 14 rows of longitudinally arranged tubercles, 35 transverse rows of ventrals, 172 longitudinal rows of ventrals, 8 or 9 expanded subdigital lamellae on the fourth toe, 11 or 12 unexpanded subdigital lamellae on the fourth toe, 19–21 total subdigital lamellae on the fourth toe; eight expanded subdigital lamellae on the fourth finger, ten unexpanded subdigital lamellae on the fourth finger, 18 total subdigital lamellae on the fourth finger; 12 total enlarged femoral scales; 12 enlarged precloacal scales; enlarged femoral and enlarged precloacal scales not continuous; proximal femoral scales less than one-half the size of the distal femorals; small tubercles on forelimbs and flanks; large dorsolateral caudal tubercles and a wide ventrolateral caudal fringe not composed homogeneous scales; tail square in cross-section; slightly enlarged paired medial subcaudals not posteromedially furrowed; maximum SVL 60.1 mm; five dark transverse body bands (Tables
(Fig.
Holotype of Cyrtodactylus kochangensis sp. nov. ZMKU R 00945 (field no. AA 04439) from Thailand, Ranong Province, Mueng Ranong District, Ko Phayam Subdistrict, Ko Chang A dorsal view B ventral view C dorsal view of head D ventral view of femoral and precloacal regions E dorsal view of tail F ventral view of tail G holotype in life.
Body relatively short (AG/SVL 0.48) with well-defined ventrolateral folds; dorsal scales small, granular, interspersed with larger conical, semi-regularly arranged, weakly keeled tubercles; tubercles extend from occipital region onto base of tail and slightly beyond as paravertebral rows; tubercles of nape and occiput smaller than those on body; approximately 14 longitudinal rows of tubercles at midbody; approximately 34 paravertebral tubercles; tubercles on upper flanks smaller than those on dorsum; approximately 35 longitudinal rows of flat, imbricate, ventral scales much larger than dorsal scales; approximately 172 transverse rows of ventral scales; no pore-bearing, precloacal scales; 12 enlarged precloacal scales; no deep precloacal groove or depression; and approximately three rows of post-precloacal scales on midline.
Forelimbs moderate in stature, relatively short (ForL/SVL 0.13); granular scales of forearm slightly larger than those on body, interspersed with tubercles; palmar scales rounded, slightly raised; digits well-developed, relatively short, inflected at basal interphalangeal joints; digits narrower distal to inflections; subdigital lamellae wide, transversely expanded proximal to joint inflections, narrower transverse lamellae distal to joint inflections; claws well-developed, claw base sheathed by a dorsal and ventral scale; 8R/8L expanded and 10R/10L unexpanded lamellae beneath the fourth finger; hind limbs larger and thicker than forelimbs, moderate in length (TibL/SVL 0.14), covered dorsally by granular scales interspersed with moderately sized, conical tubercles dorsally and posteriorly and anteriorly by flat, slightly larger, subimbricate scales; ventral scales of thigh flat, subimbricate, larger than dorsals; subtibial scales flat, imbricate; no pore-bearing femoral scales; 6R/6L enlarged femoral scales; enlarged femoral scales not contiguous with enlarged precloacal scales, terminating distally at knee; proximal femoral scales smaller than distal femorals, the former forming an abrupt union with much smaller, rounded, ventral scales of posteroventral margin of thigh; plantar scales flat; digits relatively long, well-developed, inflected at basal interphalangeal joints; 9R/8L wide, transversely expanded subdigital lamellae on the fourth toe proximal to joint inflection that extend onto sole, and 12R/11L unexpanded lamellae beneath the fourth toe; and claws well-developed, sheathed by a dorsal and ventral scale at base.
Tail original (but in two pieces), long 80.3 mm (TL/SVL 1.34), 4.0 mm in width at base, tapering to a point; nearly square in cross-section; dorsal scales flat, square bearing tubercles forming paravertebral rows and large tubercles forming a dorsolateral longitudinal row; slightly larger, posteriorly directed, semi-spinose tubercles forming wide distinct ventrolateral caudal fringe; scales of ventrolateral fringe generally interspersed at regular intervals with larger spinose scales; medial subcaudal scales paired, slightly enlarged; subcaudals, larger than dorsal caudal scales; base of tail bearing hemipenial swellings; one conical postcloacal tubercle at base of hemipenial swellings; and postcloacal scales flat, imbricate.
(Fig.
The additional specimen (THNHM 01667) closely approximates the holotype in overall coloration and pattern except that it is more boldly marked. It has four dark-colored body bands as opposed to three and a complete original tail bearing eight dark-colored and seven pale-colored bands. The pale-colored postorbital stripe is slightly thinner and less distinct. Meristic and morphometric differences are listed in Table
Cyrtodactylus kochangensis sp. nov. is currently known only from the type locality at Ko Chang Island, Ko Phayam Subdistrict, Mueang Ranong District, Ranong Province, Thailand. The additional population of pending species status occurs in the Khlong Naka Wildlife Sanctuary, Suk Samran District Ranong Province.
The specific epithet kochangensis is in reference to the type locality, Ko Chang, Ranong Province, Thailand
(based on the holotype). Cyrtodactylus kochangensis sp. nov. forms a clade with the sister species Cyrtodactylus rivularis sp. nov. and C. rukhadeva (Fig.
The holotype (ZMKU R 00945) was collected at night (2107 h) among branches of a small tree approximately 100 cm above the ground at 36 m elevation with a temperature of 28.6 °C and relative humidity of 83.9%. The surrounding habitat was dry evergreen forest with a rocky stream nearby (Fig.
Adult male ZMKU R 00949 from Thung Na Ngam Subdistrict, Lan Sak District, Uthai Thani Province, Thailand (15.37649°N, 99.63426°E, 106 m a.s.l.), collected by Attapol Rujirawan, Siriporn Yodthong, Korkhwan Termprayoon, and Natee Ampai on 18 June 2018.
Cyrtodactylus uthaiensis sp. nov. can be separated from all other species of the brevipalmatus group by the combination of having 13–15 supralabials, 10–11 infralabials, 33 paravertebral tubercles, 17 rows of longitudinally arranged tubercles, 36 transverse rows of ventrals, 159 longitudinal rows of ventrals, eight expanded subdigital lamellae on the fourth toe, 12 unexpanded subdigital lamellae on the fourth toe, 20 total subdigital lamellae on the fourth toe; seven expanded subdigital lamellae on the fourth finger, 11 unexpanded subdigital lamellae on the fourth finger, 18 total subdigital lamellae on the fourth finger; 16 total enlarged femoral scales, 12 total femoral pores; 14 enlarged pore-bearing precloacals; enlarged femorals and enlarged precloacals not continuous; proximal femorals less than one-half the size of the distal femorals; small tubercles on forelimbs and flanks; large dorsolateral caudal tubercles and wide ventrolateral caudal fringe; ventrolateral caudal fringe composed scales of different size; tail circular in cross-section; slightly enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginated; maximum SVL 58.1 mm; and six dark transverse body bands (Tables
(Fig.
Significant p-values from the results of the ANOVA analyses comparing all combinations of OTU pairs of the Cyrtodactylus brevipalmatus group. SVL and TL4U are not listed because no species pairs differed significantly from one another. * = Results based on a Games-Howell post hoc test. Character abbreviations are listed in the Materials and methods.
Morphometric characters | AG* | HumL* | ForL | FemL | TibL | HL | HW | HD* | ED* | EE* | ES | EN* | IO | EL | IN |
Cyrtodactylus fluvicavus sp. nov. vs. C. brevipalmatus | 0.00 | 0.01 | < 0.001 | 0.001 | 0.048 | < 0.001 | 0.004 | 0.002 | 0.004 | ||||||
C. interdigitalis vs. C. brevipalmatus | 0.00 | 0.00 | 0.00 | < 0.001 | |||||||||||
C. ngati vs. C. brevipalmatus | 0.00 | 0.003 | 0.001 | < 0.001 | |||||||||||
C. ngati3 vs. C. brevipalmatus | 0.014 | 0.03 | 0.01 | < 0.001 | 0.008 | 0.038 | < 0.001 | 0.004 | 0.000 | ||||||
C. rukhadeva vs. C. brevipalmatus | < 0.001 | 0.001 | 0.022 | < 0.001 | 0.021 | ||||||||||
C. sp.9 vs. C. brevipalmatus | 0.01 | < 0.001 | < 0.001 | 0.03 | |||||||||||
C. interdigitalis vs. Cyrtodactylus fluvicavus sp. nov. | 0.01 | ||||||||||||||
C. ngati vs. Cyrtodactylus fluvicavus sp. nov. | < 0.001 | 0.002 | 0.04 | < 0.001 | 0.005 | < 0.001 | < 0.001 | ||||||||
C. ngati3 vs. Cyrtodactylus fluvicavus sp. nov. | < 0.001 | 0.025 | 0.038 | ||||||||||||
C. rukhadeva vs. Cyrtodactylus fluvicavus sp. nov. | 0.049 | < 0.001 | < 0.001 | 0.004 | |||||||||||
C. sp.9 vs. Cyrtodactylus fluvicavus sp. nov. | 0.0 | 0.007 | 0.013 | 0.023 | 0.007 | ||||||||||
C. ngati vs. C. interdigitalis | 0.007 | 0.031 | 0.010 | < 0.001 | 0.000 | ||||||||||
C. ngati3 vs. C. interdigitalis | 0.011 | 0.003 | |||||||||||||
C. rukhadeva vs. C. interdigitalis | 0.0 | 0.01 | 0.00 | 0.044 | |||||||||||
C. sp.9 vs. C. interdigitalis | 0.00 | 0.007 | |||||||||||||
C. ngati3 vs. C. ngati | < 0.001 | 0.019 | 0.006 | < 0.001 | 0.001 | ||||||||||
C. rukhadeva vs. C. ngati | 0.003 | < 0.001 | < 0.001 | 0.046 | < 0.001 | 0.001 | < 0.001 | < 0.001 | 0.001 | ||||||
C. sp.9 vs. C. ngati | < 0.001 | < 0.001 | 0.042 | 0.007 | < 0.001 | < 0.001 | 0.000 | ||||||||
C. rukhadeva vs. C. ngati3 | < 0.001 | 0.021 | 0.0 | 0.00 | 0.007 | 0.02 | 0.006 | ||||||||
C. sp.9 vs. C. ngati3 | 0.001 | 0.01 | 0.03 | 0.003 | 0.043 | 0.001 | 0.019 | 0.019 | 0.003 | ||||||
C. sp.9 vs. C. rukhadeva | 0.02 | 0.004 | 0.02 | 0.033 | |||||||||||
Meristic characters | SL | IL* | PVT* | LRT | VS | VSM | TL4E | TL4T | FL4E | FL4U* | FL4T* | FS | PCS* | BB* | |
Cyrtodactylus fluvicavus sp. nov. vs. C. brevipalmatus | 0.037 | < 0.001 | 0.05 | ||||||||||||
C. interdigitalis vs C. brevipalmatus | < 0.001 | 0.000 | 0.002 | 0.044 | |||||||||||
C. ngati vs C. brevipalmatus | < 0.001 | 0.028 | < 0.001 | ||||||||||||
C. ngati3 vs C. brevipalmatus | 0.017 | 0.029 | |||||||||||||
C. rukhadeva vs C. brevipalmatus | < 0.001 | < 0.001 | |||||||||||||
C. sp.9 vs C. brevipalmatus | < 0.001 | 0.003 | 0.022 | < 0.001 | 0.05 | ||||||||||
C. interdigitalis vs Cyrtodactylus fluvicavus sp. nov. | 0.000 | 0.000 | 0.021 | < 0.001 | 0.000 | < 0.001 | |||||||||
C. ngati vs Cyrtodactylus fluvicavus sp. nov. | 0.037 | 0.002 | < 0.001 | < 0.001 | 0.033 | 0.000 | 0.002 | < 0.001 | |||||||
C. ngati3 vs Cyrtodactylus fluvicavus sp. nov. | 0.005 | ||||||||||||||
C. rukhadeva vs Cyrtodactylus fluvicavus sp. nov. | 0.000 | 0.001 | 0.001 | ||||||||||||
C. sp.9 vs Cyrtodactylus fluvicavus sp. nov. | < 0.001 | < 0.001 | 0.001 | 0.004 | 0.020 | ||||||||||
C. ngati vs C. interdigitalis | 0.001 | 0.029 | < 0.001 | < 0.001 | 0.029 | ||||||||||
C. ngati3 vs C. interdigitalis | 0.029 | 0.025 | 0.005 | ||||||||||||
C. rukhadeva vs C. interdigitalis | 0.004 | 0.011 | 0.001 | 0.044 | 0.001 | 0.002 | |||||||||
C. sp.9 vs C. interdigitalis | 0.003 | < 0.001 | 0.005 | 0.043 | 0.01 | < 0.001 | |||||||||
C. ngati3 vs C. ngati | 0.0267 | 0.001 | < 0.001 | ||||||||||||
C. rukhadeva vs C. ngati | < 0.001 | < 0.001 | 0.008 | < 0.001 | 0.008 | 0.03 | < 0.001 | ||||||||
C. sp.9 vs C. ngati | 0.003 | 0.016 | 0.011 | 0.000 | < 0.001 | < 0.001 | < 0.001 | < 0.001 | |||||||
C. rukhadeva vs C. ngati3 | |||||||||||||||
C. sp.9 vs C. ngati3 | 0.001 | 0.042 | < 0.001 | 0.001 | |||||||||||
C. sp.9 vs C. rukhadeva | 0.029 | < 0.001 | 0.002 | 0.001 |
Holotype of Cyrtodactylus uthaiensis sp. nov. ZMKU R 00949 (field no. AA 06298) from Thailand, Uthai Thani Province, Lan Sak District, Thung Na Ngam Subdistrict A dorsal view B ventral view C dorsal view of head D ventral view of femoral and precloacal regions E dorsal view of tail F ventral view of tail G holotype in life.
Body relatively short (AG/SVL 0.46) with well-defined ventrolateral folds; dorsal scales small, granular, interspersed with larger, conical, semi-regularly arranged, weakly keeled tubercles; tubercles extend from occipital region onto base of tail and slightly beyond as paravertebral rows; smaller tubercles extend anteriorly onto nape and occiput, diminishing in size anteriorly; approximately 17 longitudinal rows of tubercles at midbody; approximately 33 paravertebral tubercles; small tubercles on flanks; 36 longitudinal rows of flat, imbricate, ventral scales much larger than dorsal scales; 159 transverse rows of ventral scales; 16 total large femoral scales; 12 total femoral pores; 14 enlarged pore-bearing precloacals; no deep precloacal groove or depression; and two rows of post-precloacal scales on midline.
Forelimbs moderate in stature, relatively short (ForL/SVL 0.14); granular scales of forearm slightly larger than those on body, interspersed with large tubercles; palmar scales rounded, slightly raised; digits well-developed, relatively short, inflected at basal interphalangeal joints; digits narrower distal to inflections; subdigital lamellae wide, transversely expanded proximal to joint inflections, narrower transverse lamellae distal to joint inflections; claws well-developed, claw base sheathed by a dorsal and ventral scale; 7R/7L expanded and 11R/11L unexpanded lamellae beneath the fourth finger; hind limbs larger and thicker than forelimbs, moderate in length (TibL/SVL 0.14), covered dorsally by granular scales interspersed with moderately sized, conical tubercles dorsally and posteriorly and anteriorly by flat, slightly larger, subimbricate scales; ventral scales of thigh flat, subimbricate, larger than dorsals; subtibial scales flat, imbricate; one row of 6R/6L of enlarged pore-bearing femoral scales not continuous with enlarged pore bearing precloacal scales, terminating distally at knee; 8R/8L enlarged femoral scales; proximal femoral scales smaller than distal femorals, the former forming an abrupt union with much smaller, rounded, ventral scales of posteroventral margin of thigh; plantar scales flat; digits relatively long, well-developed, inflected at basal interphalangeal joints; 8R/(broken)L wide, transversely expanded subdigital lamellae on fourth toe proximal to joint inflection that extend onto sole, 12R/(broken)L unexpanded lamellae beneath first toe; and claws well-developed, sheathed by a dorsal and ventral scale at base.
Tail original, long 76.7 mm (TL/SVL 1.32), 4.1 mm in width at base, tapering to a point; sub-circular or nearly round in cross-section; dorsal scales flat, square bearing tubercles forming paravertebral rows and large tubercles forming a dorsolateral longitudinal row; slightly larger, posteriorly directed, semi-spinose tubercles forming small but distinct ventrolateral caudal fringe; larger scales of ventrolateral fringe occur at regular intervals; slightly enlarged medial subcaudals intermittent, medially furrowed, posteriorly; single enlarged medial subcaudals absent; subcaudal scales, larger than dorsal caudal scales; base of tail bearing hemipenial swellings; 3R/3L conical postcloacal tubercles at base of hemipenial swellings; and postcloacal scales flat, imbricate.
(Fig.
Cyrtodactylus uthaiensis sp. nov. is currently known from the type locality at Thung Na Ngam Subdistrict, Lan Sak District, Uthai Thani Province, Thailand.
The specific epithet uthaiensis refers to the type locality, Uthai Thani Province, Thailand.
Cyrtodactylus uthaiensis sp. nov. is the sister species to a clade comprised the sister species C. interdigitalis and C. sp.11 (Fig.
Cyrtodactylus uthaiensis sp. nov. is the only species of the brevipalmatus group that occurs in an isolated hilly area within the Chao Phraya River Basin (Fig.
Summary statistics from the PERMANOVA analysis of the OTUs and proposed morphogroups of the Cyrtodactylus brevipalmatus group.
OTU pairs | F model | R2 | p-value | p-adjusted |
---|---|---|---|---|
C. rukhadeva vs. C. cf. ngati2 | 6.8474 | 0.4064 | 0.015 | 0.544 |
C. rukhadeva vs. C. ngati3 | 8.8824 | 0.4467 | 0.003 | 0.122 |
C. rukhadeva vs. C. interdigitalis | 3.3630 | 0.2189 | 0.006 | 0.201 |
C. rukhadeva vs. C. ngati | 10.4580 | 0.4874 | 0.003 | 0.114 |
C. rukhadeva vs. C. brevipalmatus | 6.5983 | 0.3367 | 0.000 | 0.012 |
C. rukhadeva vs. Cyrtodactylus fluvicavus sp. nov. | 6.6357 | 0.3067 | 0.000 | 0.004 |
C. rukhadeva vs. C. sp.9 | 3.8646 | 0.1945 | 0.001 | 0.033 |
C. cf. ngati2 vs. C. brevipalmatus | 15.4818 | 0.7559 | 0.048 | 1.000 |
C. cf. ngati2 vs. Cyrtodactylus fluvicavus sp. nov. | 15.9186 | 0.6946 | 0.027 | 0.967 |
C. cf. ngati2 vs. C. sp.9 | 19.0130 | 0.7038 | 0.022 | 0.804 |
C. ngati3 vs. C. interdigitalis | 4.4753 | 0.4723 | 0.029 | 1.000 |
C. ngati3 vs. C. brevipalmatus | 14.9425 | 0.7135 | 0.018 | 0.643 |
C. ngati3 vs. Cyrtodactylus fluvicavus sp. nov. | 8.7953 | 0.5237 | 0.009 | 0.317 |
C. ngati3 vs. C. sp.9 | 14.7978 | 0.6218 | 0.006 | 0.226 |
C. interdigitalis vs. C. ngati | 9.8976 | 0.6644 | 0.029 | 1.000 |
C. interdigitalis vs. C. brevipalmatus | 4.5646 | 0.3947 | 0.008 | 0.278 |
C. interdigitalis vs. Cyrtodactylus fluvicavus sp. nov. | 6.7120 | 0.4272 | 0.003 | 0.124 |
C. interdigitalis vs. C. sp.9 | 5.6585 | 0.3614 | 0.002 | 0.067 |
C. ngati vs. C. brevipalmatus | 7.4818 | 0.5550 | 0.018 | 0.643 |
C. ngati vs. Cyrtodactylus fluvicavus sp. nov. | 22.8234 | 0.7405 | 0.008 | 0.283 |
C. ngati vs. C. sp.9 | 17.0146 | 0.6540 | 0.006 | 0.227 |
C. brevipalmatus vs. Cyrtodactylus fluvicavus sp. nov. | 17.8585 | 0.6410 | 0.001 | 0.048 |
C. brevipalmatus vs. C. sp.9 | 9.3960 | 0.4607 | 0.001 | 0.025 |
Cyrtodactylus fluvicavus sp. nov. vs. C. sp.9 | 8.2047 | 0.3869 | 0.000 | 0.005 |
A taxonomy consistent with evolutionary history is paramount to any downstream comparative analyses employed to address the evolution of other features of the group be they ecology, behavior, or habitat preference. By delimiting new and existing species using a phylogenetic analysis and then diagnosing those species using univariate and multivariate statistical analyses of their morphological data, phylogenetic history will not conflated with convergent evolution (Fig.
Comparison of a morphology-based taxonomy pre-2020 and the current phylogenetic-based taxonomy illustrating the misrepresentation of the diversity of the Cyrtodactylus brevipalmatus group by the former. Colored lines connecting species on the left to species on the right indicate under which name the former were masquerading as the latter.
The description of the four new species, Cyrtodactylus fluvicavus sp. nov., Cyrtodactylus kochangensis sp. nov., Cyrtodactylus rivularis sp. nov., and Cyrtodactylus uthaiensis sp. nov. adds to the rapidly growing list of new species of Cyrtodactylus from Thailand. Many of the poorly sampled populations reported here such as C. sp.9 from Thong Pha Phum National Park, Kanchanaburi Province; C. sp.10 from Chao Doi Waterfall, Tak Province; C. sp.11 from Phu Hin Rong Kla National Park, Phitsanulok Province; C. sp.13 from Ban Saphan Lao, Kanchanaburi Province and Thung Yai Naresuan Wildlife Sanctuary, Tak Provinces; C. sp.14 from Langkawi Island, Peninsular Malaysia; C. cf. ngati1 and C. cf. ngati2 from Xaignabouli and Vientiane Provinces, respectively, Laos (Figs
This work was financially supported by Office of the Permanent Secretary, Ministry of Higher Education, Science, Research and Innovation (grant no. RGNS 64-038). NA was supported by Srinakharinwirot University Research Grant (no. 596/2564). This research was reviewed and approved by the Institutional Animal Care and Use Committee of Faculty of Science, Kasetsart University (ACKU64-SCI-006) and the Department of National Parks, Wildlife and Plant Conservation, Thailand provided the research permission. We would like to thank Suchai Horradee (Chaloem Rattanakosin National Park), Charoen Jaichon (Thong Pha Phum National Park) and Mana Phermpool (Kaeng Krachan National Park) for facilitating the fieldwork. Wachara Sanguansombat and Sunchai Makchai (Thailand Natural History Museum) made specimens in their care available for study. Piyawan Puanprapai assisted with fieldwork. We would like to thank Bryan L. Stuart for suggestions on this research and Truong Nguyen and an anonymous reviewer improved the manuscript.
Data frame for the multiple factor analysis of the putative species of the Cyrtodactylus brevipalmatus group
Data type: morphological data
Explanation note: Data frame for the multiple factor analysis of the putative species of the Cyrtodactylus brevipalmatus group.