Research Article |
Corresponding author: Siriwadee Chomdej ( siriwadee.submission@gmail.com ) Corresponding author: L. Lee Grismer ( lgrismer@lasierra.edu ) Academic editor: Thomas Ziegler
© 2023 Siriwadee Chomdej, Chatmongkon Suwannapoom, Waranee Pradit, Apichaya Phupanbai, L. Lee Grismer.
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:
Chomdej S, Suwannapoom C, Pradit W, Phupanbai A, Grismer LL (2023) A new species of the Cyrtodactylus brevipalmatus group (Squamata, Gekkonidae) from Tak Province, northwestern Thailand. ZooKeys 1164: 63-88. https://doi.org/10.3897/zookeys.1164.101263
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An integrative taxonomic analysis was used to delimit and diagnose a new species of the Cyrtodactylus brevipalmatus group from Tak Province in western Thailand. Although Bayesian phylogenetic analyses place C. denticulatus sp. nov. within the brevipalmatus group, the new species is neither nested within nor is it the sister species of any other species in the brevipalmatus group. Furthermore, based on the mitochondrial NADH dehydrogenase subunit 2 gene (ND2) and adjacent tRNAs, it bears an uncorrected pairwise sequence divergence of 7.87–21.94% from all other species in the brevipalmatus group. Cyrtodactylus denticulatus sp. nov. is differetiated from all other species in the brevipalmatus group by having a number of unique charateristics such as denticulate ventrolateral body folds and ventrolateral subcaudal ridges, characters not seen in any other species of the group (n = 51 individuals). Additionally, based on a multiple factor anlaysis, C. denticulatus sp. nov. does not overlap with any other species in multivariate space. The discovery of C. denticulatus sp. nov. underscores the unrealized diversity of upland ecosystems across Thailand and the urgent need for increased exploration and conservation of these unique imperiled montane refugia, especially in this era of climate change.
Bent-toed gecko, conservation, Indochina, integrative taxonomy, Southeast Asia, systematics
The Cyrtodactylus brevipalmatus group (sec.
We present herein the description of another new species of the brevipalmatus group from the Chao Doi waterfall, Mae Meoi, Tha Song Yang District, Tak Province, Thailand (Fig.
Under the general lineage concept (GLC:
Methods for DNA extraction, sequencing, and editing followed
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.
brevipalmatus | cf. ngati1 | cf. ngati2 | elok | fluvicavus | interdigitalis | kochangensis | ngati | rivularis | rukhadeva | thongphaphumensis. | denticulatus sp. nov. | sp. 11 | cf. brevipalmatus | uthaiensis | |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
brevipalmatus | n/a | ||||||||||||||
n = 1 | |||||||||||||||
cf. ngati1 | 21.03 | n/a | |||||||||||||
n = 1 | |||||||||||||||
cf. ngati2 | 21.68 | 4.39 | n/a | ||||||||||||
n = 1 | |||||||||||||||
elok | 20.77 | 22.58 | 21.42 | n/a | |||||||||||
n = 1 | |||||||||||||||
fluvicavus | 18.86 | 10.64 | 11.02 | 20.15 | 0.10 | ||||||||||
n = 7 | (18.84–18.97) | (10.58–10.84) | (10.97–11.23) | (20.13–20.26) | (0.00–0.26) | ||||||||||
interdigitalis | 20.77 | 6.97 | 9.16 | 22.84 | 12.02 | n/a | |||||||||
n = 1 | (12.00–12.13) | ||||||||||||||
kochangensis | 19.35 | 14.58 | 14.71 | 20.90 | 12.31 | 15.23 | n/a | ||||||||
n = 1 | (12.26–12.31) | ||||||||||||||
ngati | 20.70 | 3.30 | 3.71 | 21.11 | 11.34 | 8.13 | 14.58 | 0.84 | |||||||
n = 7 | (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) | |||||||
rivularis | 20.00 | 15.87 | 15.03 | 21.61 | 12.57 | 15.48 | 12.26 | 15.03 | 0.52 | ||||||
n = 2 | (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) | |||||||
rukhadeva | 20.65 | 15.42 | 15.48 | 21.61 | 12.25 | 16.00 | 13.10 | 15.23 | 4.65 | 1.55 | |||||
n = 2 | (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) | ||||||
thongphaphumensis | 20.34 | 7.93 | 9.51 | 22.02 | 9.75 | 8.96 | 13.22 | 8.81 | 13.12 | 13.25 | 0.22 | ||||
n = 9 | (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) | ||||
denticulatus sp. nov. | 19.87 | 9.29 | 10.84 | 21.94 | 10.12 | 10.19 | 13.68 | 10.21 | 13.94 | 14.32 | 8.06 | n/a | |||
n = 1 | (10.06–10.32) | (10.06–10.45) | (13.68–14.19) | (13.68–14.97) | (7.87–8.13) | ||||||||||
sp. 11 | 20.39 | 7.23 | 8.90 | 22.19 | 11.12 | 3.87 | 14.58 | 8.28 | 15.35 | 15.61 | 8.96 | 10.45 | n/a | ||
n = 1 | (11.10–11.23) | (8.00–8.65) | (15.10–15.61) | (14.97–16.26) | (8.77–9.03) | ||||||||||
cf. brevipalmatus | 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 | |
n = 1 | (18.32–18.45) | (20.26–20.65) | (19.48–20.00) | (19.48–20.52) | (19.48–19.87) | ||||||||||
uthaiensis | 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 |
n = 1 | (10.06–10.32) | (6.58–7.61) | (13.68–14.19) | (13.29–14.58) | (7.61–7.87) |
The morphological data included 15 meristic, 16 normalized morphometric, and eight categorical characters. The data were taken using the protocol of
Evaluated meristic characters 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; number of femoral pores (FP) in males from each thigh combined as a single metric; number postcloacal tubercles (PCT) on each side of the base of the tail (this character was not used in the analyses); 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); ventrolateral body fringe denticulate (VFD); slightly enlarged medial subcaudals (SC1); single enlarged, unmodified, medial subcaudal scales (SC2); enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginated (SC3); small longitudinal ventrolateral subcaudal ridge (SC4); 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
Morphospatial clustering and positioning among the species/populations and individuals was analyzed using multiple factor analysis (MFA) on a concatenated data set comprised of 15 meristic characters, 16 normalized morphometric characters, and eight categorical characters (Suppl. material
The BEAST analysis recovered C. sp. 10 as being deeply nested within the brevipalmatus group on a long branch that was not embedded within, nor sister to any other species (Fig.
The MFA recovered C. sp. 10 as well-separated from all other species of the brevipalmatus group along the ordination of the first two dimensions (Dim) in that the specimen did not cluster near or within the convex hull of any other species (Fig.
MFA analyses of the Cyrtodactylus brevipalmatus group A MFA of the species-level lineages based on the BEAST phylogeny (Fig.
Based on the phylogenetic position of C. sp. 10, its high percentage value of pairwise sequence divergence from all other species, and its unique morphology, we hypothesize C. sp. 10 represents a diagnosable evolutionarily distinct population at the northwestern extent of the range of the brevipalmatus group that should be recognized as a new species. As such, it is described below.
Cyrtodactylus
sp. 10:
Holotype. Adult male AUP-00680 collected on 8 March 2019 by Parinya Pawangkhanant and Chatmongkon Suwannapoom from a bamboo forest near a rocky stream at the Chao Doi waterfall, Mae Meoi, Tha Song Yang District, Tak Province, Thailand at 17°30.000'N, 98°03.000'E (DDM) and 610 m a.s.l.
Cyrtodactylus denticulatus sp. nov. is tentatively separated (see below) from all other species of the brevipalmatus group by the combination of having nine supralabials, nine infralabials, 20 paravertebral tubercles, 19 rows of longitudinally arranged tubercles, 42 transverse rows of ventrals, 158 longitudinal rows of ventrals, nine expanded subdigital lamellae on the fourth finger, 11 unexpanded subdigital lamellae on the fourth toe, 19 total subdigital lamellae on the fourth toe; nine expanded subdigital lamellae on the fourth finger, ten unexpanded subdigital lamellae on the fourth finger, 19 total subdigital lamellae on the fourth finger; 16 total number of enlarged femoral scales, 20 total number of femoral pores in the male specimen; 13 precloacal pores in the male specimen; 16 enlarged precloacals; enlarged femorals and enlarged precloacals not continuous; proximal femorals smaller than distal femorals; tubercles on forelimbs and flanks nearly same size as those on body; ventrolateral body fold weakly denticulate; spinose paravertebral rows; row of large dorsolateral caudal tubercles; wide ventrolateral caudal fringes; ventrolateral caudal fringes composed scales of different size; small longitudinal ventrolateral subcaudal ridges; tail square in cross-section; no slightly enlarged unpaired medial subcaudals; subcaudals not posteromedially furrowed; SVL 69.5 mm; three dark transverse body bands (Tables
Sex, raw meristic, categorical, and morphometric data used in the analyses of specimens in the Cyrtodactylus brevipalmatus group. ♂ = male; ♀ = female; R/L = right/left; / = data unavailable. Shaded cells denote characters that potentially differentiate C. denticulatus sp. nov. from the other species.
Species | denticulatus sp. nov. | brevipalmatus | cf. brevipalmatus | cf. brevipalmatus | brevipalmatus | brevipalmatus | elok | elok | elok | elok | interdigitalis | interdigitalis | interdigitalis | interdigitalis | sp. 11 |
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
Institutional catalog number | AUP-00680 | 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 | THNMH 20227 paratype | ZMMU R-16492 |
Sex | ♂ | ♂ | ♀ | ♀ | ♀ | ♀ | ♀ | ♂ | ♂ | ♀ | ♀ | ♀ | ♀ | ♀ | ♂ |
Meristic data | |||||||||||||||
supralabials (SL) | 9 | 11 | 12 | 10 | 14 | 12 | 11 | 8 | 13 | 9 | 14 | 12 | 11 | 12 | 11 |
infralabials (IL) | 9 | 8 | 10 | 9 | 11 | 11 | 11 | 8 | 11 | 9 | 9 | 8 | 8 | 7 | 9 |
paravertebral tubercles (PVT) | 20 | 39 | 37 | 38 | 37 | 37 | 0 | 0 | 0 | 0 | 32 | 33 | 33 | 33 | 30 |
longitudinal rows of tubercles (LRT) | 19 | 15 | 16 | 17 | 16 | 14 | 6 | 7 | 4 | 4 | 19 | 20 | 19 | 19 | 18 |
ventral scales (VS) | 42 | 38 | 38 | 38 | 36 | 39 | 45 | 45 | 47 | 36 | 42 | 40 | 42 | 43 | 34 |
ventral scales along middle of the body (VSM) | 158 | 176 | 170 | 182 | 154 | 160 | 190 | 225 | 234 | 192 | 187 | 170 | 187 | 178 | 160 |
expanded subdigital lamellae on 4th toe (TL4E) | 9 | 7 | 8 | 9 | 8 | 8 | 10 | 9 | 9 | 9 | 12 | 10 | 10 | 11 | 9 |
unmodified subdigital lamellae on 4th toe (TL4U) | 11 | 13 | 11 | 11 | 11 | 12 | 11 | 10 | 11 | 9 | 14 | 13 | 12 | 14 | 10 |
total subdigital lamellae 4th toe (TL4T) | 20 | 20 | 19 | 20 | 19 | 20 | 21 | 19 | 20 | 18 | 26 | 23 | 22 | 23 | 19 |
expanded subdigital lamellae on 4th finger (FL4E) | 9 | 8 | 8 | 8 | 7 | 8 | 8 | 9 | 9 | 9 | 9 | 8 | 9 | 9 | 10 |
unmodified subdigital lamellae on 4th finger (FL4U) | 10 | 9 | 11 | 10 | 10 | 10 | 12 | 13 | 9 | 8 | 12 | 11 | 12 | 13 | 9 |
total subdigital lamellae 4th finger (FL4T) | 19 | 17 | 19 | 18 | 17 | 18 | 20 | 22 | 18 | 17 | 21 | 21 | 21 | 22 | 19 |
enlarged femoral scales (R/L) | 8 | 0 | 0 | 0 | 8R/8L | 7R/7L | 0 | 0 | 0 | 0 | 11R/8L | 10R/9L | 8R/8L | 9R/10L | 9R/8L |
total enlarged femoral scales (FS) | 16 | 16 | 10 | 11 | 16 | 14 | 0 | 0 | 0 | 0 | 14 | 19 | 19 | 19 | 17 |
total femoral pores in males (FP) | 20 | 7 | / | / | / | / | / | 0 | 0 | / | / | / | / | / | 17 |
enlarged precloacal scales (PCS) | 13 | 7 | 7 | 7 | 8 | 7 | 8 | 8 | 8 | 7 | 14 | 15 | 13 | 19 | 13 |
precloacal pores in males (PP) | 13 | 7 | / | / | / | / | / | 8 | 8 | / | 0 | / | 0 | 0 | 13 |
postcloacal tubercles (PCT) | 3 | 3 | 3 | 2 | 3 | 3 | 3 | 2 | 3 | 3 | 3 | 2 | 3 | 3 | 3 |
Body bands (BB) | 3 | 4 | 6 | 3 | 5 | 5 | 5 | 5 | 3 | 3 | 5 | 5 | 5 | 5 | 3 |
Categorical data | |||||||||||||||
small tubercles on flank (FKT) | present | present | present | present | present | present | absent | absent | absent | absent | present | present | present | present | present |
denticulate ventrolateral body folds (VFD) | present | absent | absent | absent | absent | absent | absent | absent | absent | absent | absent | absent | absent | absent | absent |
dorsolateral caudal tubercles (DCT) | small | small | small | small | / | small | large | large | large | large | small | / | small | small | large |
ventrolateral caudal fringe narrow or wide (VLF1) | wide | narrow | narrow | narrow | / | narrow | wide | wide | wide | wide | narrow | / | narrow | narrow | wide |
ventrolateral caudal fringe scales generally homogenous (VLF2) | no | no | no | no | / | no | no | no | no | no | yes | yes | yes | yes | yes |
tail cross-section (TLcross) | square | circular | circular | circular | / | circular | square | square | square | square | circular | / | circular | circular | square |
slightly enlarged medial subcaudals (SC1) | absent | present | present | present | / | absent | absent | absent | absent | absent | absent | / | absent | absent | present |
single enlarged medial subcaudal (SC2) | absent | absent | absent | absent | / | 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 | no | yes | / | yes | yes | no |
small ventrolateral subcaudal ridge of scales (SC4) | yes | no | no | no | no | no | no | no | no | no | no | no | no | no | no |
Morphometric data | |||||||||||||||
SVL | 69.5 | 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 |
AG | 83.0 | 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 |
HumL | 9.1 | 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 |
ForL | 10.2 | 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 |
FemL | 12.8 | 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 |
TibL | 6.5 | 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 |
HL | 18.8 | 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 |
HW | 12.7 | 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 |
HD | 7.2 | 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 |
ED | 4.0 | 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 |
EE | 5.4 | 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 |
ES | 7.9 | 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 |
EN | 5.8 | 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 |
IO | 7.1 | 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 |
EL | 0.7 | 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 |
IN | 2.4 | 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 |
Sex, raw meristic, categorical, and morphometric data used in the analyses of specimens in the Cyrtodactylus brevipalmatus group. ♂ = male; ♀ = female; R/L = right/left; / = data unavailable. Shaded cells denote characters that potentially differentiate C. denticulatus sp. nov. from the other species.
Species | denticulatus sp. nov. | ngati | ngati3 | ngati4 | cf. ngati1 | cf. ngati2 | ||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|
Institutional catalog number | AUP-00680 | HNUE-R00111 | IEBR 4829 | VNUF R.2020.12 | HNUE-R00112 | FMNH 255454 | FMNH 270493 | FMNH 270492 | FMNH 265806 | NCSM 79472 | ZMMU R-14917 | NCSM 80100 |
Sex | ♂ | ♂ | ♀ | ♀ | ♀ | ♀ | ♂ | ♂ | ♂ | ♀ | ♀ | ♀ |
Meristic data | ||||||||||||
supralabials (SL) | 9 | 10 | 10 | 10 | 10 | 13 | 13 | 13 | 10 | 14 | 9 | 12 |
infralabials (IL) | 9 | 9 | 9 | 9 | 9 | 10 | 9 | 11 | 8 | 11 | 10 | 12 |
paravertebral tubercles (PVT) | 20 | 39 | 40 | 38 | 40 | 28 | 27 | 26 | 27 | 28 | 32 | 29 |
longitudinal rows of tubercles (LRT) | 19 | 18 | 18 | 17 | 22 | 19 | 18 | 17 | 19 | 18 | 24 | 19 |
ventral scales (VS) | 42 | 38 | 36 | 35 | 32 | 37 | 36 | 36 | 33 | 33 | 36 | 35 |
ventral scales along middle of the body (VSM) | 158 | 168 | 164 | 178 | 158 | 159 | 166 | 156 | 158 | 164 | 166 | 165 |
expanded subdigital lamellae on 4th toe) TL4E( | 9 | 8 | 10 | 9 | 9 | 10 | 10 | 8 | 10 | 9 | 8 | 10 |
unmodified subdigital lamellae on 4th toe) TL4U( | 11 | 11 | 10 | 11 | 10 | 11 | 11 | 11 | 11 | 12 | 10 | 10 |
total subdigital lamellae 4th toe (TL$T) | 20 | 13 | 16 | 17 | 16 | 21 | 21 | 19 | 21 | 21 | 18 | 20 |
expanded subdigital lamellae on 4th finger (FL4E) | 9 | 6 | 6 | 7 | 6 | 8 | 8 | 8 | 8 | 9 | 7 | 9 |
unmodified subdigital lamellae on 4th finger (FL4U) | 10 | 9 | 9 | 9 | 9 | 10 | 10 | 10 | 10 | 8 | 9 | 10 |
total subdigital lamellae 4th finger (FL4T) | 19 | 15 | 15 | 18 | 15 | 18 | 18 | 18 | 18 | 17 | 16 | 19 |
enlarged femoral scales (R/L) | 8 | 10R/10L | 9R/8L | 10R/9L | 8R/9L | 9R/7L | 8R/9L | 9R/9L | 8R/8L | 9R/8L | 7R/8L | 7R/8L |
total enlarged femoral scales (FS) | 16 | 20 | 17 | 19 | 17 | 16 | 17 | 18 | 16 | 17 | 15 | 15 |
total femoral pores (FP) | 20 | 14 | / | / | / | / | 14 | 15 | 13 | / | / | / |
enlarged precloacal scales (PCS) | 13 | 13 | 13 | 13 | 13 | 15 | 13 | 13 | 13 | 12 | 13 | 13 |
precloacal pores (PP) | 13 | 13 | / | / | / | 13 | 13 | 13 | 13 | / | / | / |
postcloacal tubercles (PCT) | 3 | 3 | 2 | 1 | 2 | / | / | / | / | 2 | 3 | 4 |
Body bands (BB) | 3 | 6 | 6 | 6 | 6 | 3 | 4 | 3 | 3 | 3 | 3 | 3 |
Categorical data | ||||||||||||
small tubercles on flank (FKT) | present | present | present | present | present | present | present | present | present | present | present | present |
denticulate ventrolateral body folds (VFD) | present | absent | absent | absent | absent | absent | absent | absent | absent | absent | absent | absent |
dorsolateral caudal tubercles (DCT) | small | small | small | small | small | small | small | small | small | small | small | small |
ventrolateral caudal fringe narrow or wide (VLF1) | wide | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow | narrow |
ventrolateral caudal fringe scales generally homogenous (VLF2) | no | no | no | no | no | yes | yes | yes | yes | yes | yes | yes |
tail cross-section (TLcross) | square | circular | circular | circular | circular | circular | circular | circular | circular | circular | circular | circular |
slightly enlarged medial subcaudals (SC1) | absent | present | 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 |
enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginate (SC3) | no | no | no | no | no | / | no | no | no | no | no | no |
small ventrolateral subcaudal ridge of scales (SC4) | yes | no | no | no | no | / | no | no | no | no | no | no |
Morphometric data | ||||||||||||
SVL | 69.5 | 66.5 | 68.1 | 69.3 | 46.6 | 83.6 | 70.2 | 74.1 | 73.8 | 78.0 | 87.1 | 77.7 |
AG | 83.0 | 28.8 | 29.8 | 30.2 | 19.7 | 41.3 | 35.4 | 37.0 | 31.3 | 38.2 | 41.9 | 36.8 |
HumL | 9.1 | 7.9 | 8.1 | 8.5 | 5.6 | 8.6 | 8.7 | 8.6 | 6.9 | 8.7 | 11.5 | 9.2 |
ForL | 10.2 | 9.2 | 10.0 | 10.1 | 6.5 | 10.2 | 9.3 | 10.4 | 10.0 | 10.3 | 10.4 | 10.7 |
FemL | 12.8 | 11.5 | 11.5 | 11.5 | 7.6 | 13.7 | 12.7 | 13.0 | 13.1 | 13.1 | 15.2 | 14.2 |
TibL | 6.5 | 10.8 | 11.1 | 11.8 | 7.8 | 12.5 | 11.8 | 11.2 | 11.1 | 12.8 | 12.6 | 12.7 |
HL | 18.8 | 20.1 | 20.4 | 20.7 | 16.1 | 21.7 | 20.6 | 20.3 | 20.7 | 21.2 | 22.1 | 21.4 |
HW | 12.7 | 12.6 | 12.0 | 11.8 | 8.8 | 13.8 | 12.5 | 13.0 | 12.3 | 12.7 | 14.8 | 13.5 |
HD | 7.2 | 7.4 | 7.2 | 6.6 | 5.1 | 9.2 | 8.4 | 9.1 | 7.6 | 8.3 | 8.7 | 9.2 |
ED | 4.0 | 3.8 | 4.1 | 3.4 | 2.6 | 4.9 | 4.9 | 4.9 | 4.8 | 6.5 | 4.6 | 6.0 |
EE | 5.4 | 5.8 | 5.5 | 5.9 | 4.4 | 6.9 | 6.1 | 6.2 | 5.7 | 5.3 | 6.5 | 6.2 |
ES | 7.9 | 7.5 | 7.6 | 6.9 | 5.0 | 9.0 | 8.3 | 8.3 | 8.2 | 8.7 | 8.8 | 8.4 |
EN | 5.8 | 6.7 | 6.3 | 6.2 | 4.5 | 6.5 | 6.2 | 6.1 | 6.2 | 6.2 | 6.6 | 6.0 |
IO | 7.1 | 5.6 | 5.4 | 5.6 | 4.2 | 6.6 | 5.6 | 5.4 | 5.1 | 4.9 | 3.5 | 5.7 |
EL | 0.7 | 0.8 | 0.8 | 0.7 | 0.3 | 1.3 | 1.1 | 1.2 | 1.0 | 1.5 | 1.2 | 0.9 |
IN | 2.4 | 2.8 | 2.6 | 2.6 | 2.0 | 2.8 | 2.5 | 2.5 | 2.3 | 2.7 | 2.2 | 2.5 |
Sex, raw meristic, categorical, and morphometric data used in the analyses of specimens in the Cyrtodactylus brevipalmatus group. ♂ = male; ♀ = female; R/L = right/left; / = data unavailable. Shaded cells denote characters that potentially differentiate C. denticulatus sp. nov. from the other species.
Species | denticulatus sp. nov. | thongphaphumensis | sp.13 | ||||||||
---|---|---|---|---|---|---|---|---|---|---|---|
Institutional catalog number | AUP-00680 | ZMKU R 00950 paratype | ZMKU R 00951 paratype | ZMKU R 00952 paratype | ZMKU R 00953 holotype | ZMKU R 00954 paratype | ZMKU R 00955 pratype | ZMKU R 00956 paratype | ZMKU R 00957 paratype | THNHM 00104 | THNHM 27821 |
Sex | ♂ | ♀ | ♂ | ♀ | ♂ | ♂ | ♀ | ♂ | ♀ | ♀ | ♀ |
Meristic data | |||||||||||
supralabials (SL) | 9 | 12 | 13 | 13 | 14 | 13 | 13 | 13 | 13 | 12 | 15 |
infralabials (IL) | 9 | 8 | 8 | 10 | 10 | 9 | 10 | 10 | 9 | 10 | 11 |
paravertebral tubercles (PVT) | 20 | 32 | 33 | 34 | 34 | 36 | 36 | 30 | 30 | 33 | 29 |
longitudinal rows of tubercles (LRT) | 19 | 21 | 19 | 20 | 20 | 21 | 21 | 19 | 19 | 18 | 20 |
ventral scales (VS) | 42 | 34 | 33 | 33 | 34 | 30 | 33 | 32 | 33 | 37 | 36 |
ventral scales along middle of the body (VSM) | 158 | 173 | 158 | 156 | 166 | 159 | 159 | 150 | 169 | 159 | 165 |
expanded subdigital lamellae on 4th toe (TL4E) | 9 | 9 | 10 | 9 | 8 | 10 | 8 | 9 | 9 | 9 | 7 |
unmodified subdigital lamellae on 4th toe (TL4U) | 11 | 12 | 14 | 13 | 12 | 13 | 12 | 11 | 13 | 12 | 12 |
total subdigital lamellae 4th toe (TL$T) | 20 | 21 | 24 | 22 | 20 | 23 | 20 | 20 | 22 | 21 | 19 |
expanded subdigital lamellae on 4th finger (FL4E) | 9 | 8 | 7 | 7 | 8 | 8 | 8 | 8 | 8 | 8 | 8 |
unmodified subdigital lamellae on 4th finger (FL4U) | 10 | 10 | 12 | 12 | 11 | 12 | 12 | 11 | 12 | 11 | 10 |
total subdigital lamellae 4th finger (FL4T) | 19 | 18 | 19 | 19 | 19 | 20 | 20 | 19 | 20 | 19 | 18 |
enlarged femoral scales (R/L) | 8 | 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 | 12 | 16 | 16 | 15 | 16 | 15 | 13 | 16 | 18 | 17 |
total femoral pores (FP) | 20 | 0 | 16 | 0 | 14 | 15 | 14 | 12 | 0 | 0 | 0 |
enlarged precloacal scales (PCS) | 13 | 17 | 15 | 15 | 15 | 15 | 15 | 15 | 15 | 14 | 16 |
precloacal pores (PP) | 13 | 0 | 15 | 0 | 15 | 15 | 15 | 15 | 0 | 0 | 0 |
postcloacal tubercles (PCT) | 3 | 2 | 2R/3L | 3 | 3 | 2R/3L | 2R/3L | 3 | 2 | 3 | 3 |
Body bands (BB) | 3 | 3 | 4 | 3 | 4 | 3 | 5 | 4 | 4 | 3 | / |
Categorical data | |||||||||||
small tubercles on flank (FKT) | present | present | present | present | present | present | present | present | present | present | present |
denticulate ventrolateral body folds (VFD) | present | absent | absent | absent | absent | absent | absent | absent | absent | absent | absent |
dorsolateral caudal tubercles (DCT) | small | large | large | large | large | large | large | / | large | small | small |
ventrolateral caudal fringe narrow or wide (VLF1) | 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 |
tail cross-section (TLcross) | square | square | square | square | square | square | square | / | square | circular | circular |
slightly enlarged medial subcaudals (SC1) | 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 |
enlarged medial subcaudals intermittent, medially furrowed, posteriorly emarginate (SC3) | no | no | no | no | no | yes | no | / | no | no | no |
small ventrolateral subcaudal ridge of scales (SC4) | yes | no | no | no | no | no | no | / | no | no | no |
Morphometric data | |||||||||||
SVL | 69.5 | 73.1 | 73.5 | 73.7 | 73.2 | 64.4 | 76.6 | 76.6 | 74.2 | 63.7 | 72.9 |
AG | 83.0 | 34.8 | 33.9 | 35.4 | 33.6 | 28.5 | 37.1 | 33.2 | 35.1 | 25.8 | 30.6 |
HumL | 9.1 | 8.4 | 7.2 | 9.0 | 9.0 | 7.2 | 8.0 | 8.1 | 8.6 | 7.6 | 10.1 |
ForL | 10.2 | 9.5 | 9.1 | 9.2 | 9.8 | 9.2 | 10.0 | 8.6 | 9.8 | 8.1 | 9.6 |
FemL | 12.8 | 12.8 | 11.6 | 12.3 | 12.5 | 10.9 | 13.7 | 10.8 | 12.5 | 10.7 | 12.8 |
TibL | 6.5 | 10.5 | 10.1 | 10.6 | 10.6 | 9.9 | 11.1 | 10.0 | 11.4 | 10.1 | 10.2 |
HL | 18.8 | 19.9 | 20.9 | 20.1 | 20.0 | 17.6 | 20.4 | 19.3 | 20.0 | 17.6 | 19.9 |
HW | 12.7 | 14.5 | 14.3 | 15.7 | 13.9 | 12.8 | 14.7 | 14.4 | 14.1 | 11.9 | 13.8 |
HD | 7.2 | 7.8 | 7.7 | 7.9 | 7.7 | 7.0 | 8.2 | 7.8 | 7.6 | 7.7 | 8.4 |
ED | 4.0 | 5.0 | 5.1 | 5.0 | 5.0 | 4.8 | 5.6 | 5.3 | 4.9 | 4.1 | 5.3 |
EE | 5.4 | 5.9 | 5.9 | 6.0 | 5.9 | 5.3 | 6.1 | 6.0 | 6.0 | 4.9 | 6.3 |
ES | 7.9 | 7.9 | 8.5 | 7.9 | 7.9 | 7.3 | 8.2 | 7.9 | 7.9 | 7.2 | 8.0 |
EN | 5.8 | 6.0 | 6.1 | 6.0 | 5.8 | 5.4 | 6.1 | 6.0 | 5.9 | 5.6 | 5.9 |
IO | 7.1 | 5.4 | 5.5 | 5.8 | 5.5 | 4.9 | 5.7 | 5.6 | 5.3 | 4.8 | 6.1 |
EL | 0.7 | 1.1 | 1.5 | 1.5 | 1.2 | 1.2 | 1.0 | 1.2 | 1.3 | 1.4 | 1.4 |
IN | 2.4 | 2.3 | 2.4 | 2.2 | 2.0 | 2.0 | 2.3 | 2.2 | 2.2 | 2.1 | 2.3 |
(Figs
Holotype of Cyrtodactylus denticulatus sp. nov. AUP-00680 from the Chao Doi waterfall, Mae Meoi, Tha Song Yang District, Tak Province, Thailand A ventral view of the cloacal and femoral regions B ventral view of the left hand C ventral view of the left foot D dorsal view of the anterior section of the tail E ventral view of the tail showing the ventrolateral subcaudal ridge.
Body relatively short (AG/SVL 0.46) with well-defined denticulate 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 19 longitudinal rows of tubercles at midbody; approximately 20 paravertebral tubercles; small tubercles on flanks; 42 longitudinal rows of flat, imbricate, ventral scales much larger than dorsal scales; 158 transverse rows of ventral scales; 13 large, pore-bearing, precloacal scales; no deep precloacal groove or depression; and three rows of enlarged post-precloacal scales on midline.
Forelimbs moderate in stature, relatively short (HumL/SVL 0.13; ForL/SVL 14.7); granular scales of forearm larger than those on body, interspersed with slightly larger tubercles; palmar scales rounded, slightly raised, subimbricate; 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 lamellae distal to inflections; claws well-developed, claw base sheathed by a dorsal and ventral scale; 9R/9L expanded and 11R/11L unexpanded lamellae beneath the fourth finger; hind limbs longer and thicker than forelimbs, moderate in length (FemL/SVL 18.4; TibL/SVL 0.09), covered dorsally by granular scales interspersed with moderately sized, conical tubercles dorsally and posteriorly and anteriorly by flat, larger, subimbricate scales; ventral scales of thigh flat, imbricate, larger than dorsals; subtibial scales flat, imbricate; one row of 8R/8L 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 raised, subimbricate; 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 distal to joint inflection; and claws well-developed, claw base sheathed by a dorsal and ventral scale.
Posterior one-fifth of tail regenerated, 83.0 mm long (TL/SVL 1.19), 6.5 mm in width at base, tapering to a point; nearly square in cross-section; dorsal scales flat, intermixed with large tubercles forming spinose paravertebral rows; row of large dorsolateral caudal tubercles; large, posteriorly directed, spinose tubercles forming wide ventrolateral caudal fringe; much larger scales of ventrolateral fringe occur at regular intervals; medial subcaudals slightly enlarged, no enlarged single medial subcaudal longitudinal row; subcaudals, larger than dorsal caudals; small longitudinal ventrolateral subcaudal ridges; base of tail bearing hemipenal swellings; 3R/3L conical postcloacal tubercles at base of hemipenal swellings; and postcloacal scales flat, imbricate.
(Fig.
Cyrtodactylus denticulatus sp. nov. represents the northwestern-most species of the brevipalmatus group. At present, it is known only from the type locality at Chao Doi waterfall, Tha Song Yang District, Tak Province, western Thailand (Fig.
The specific epithet denticulatus is given as a noun in apposition, meaning “denticulate” or with small teeth, a reference to bearing small tooth-like dorsolateral and ventrolateral caudal tubercles and denticulate ventrolateral body folds.
(Tables
The holotype was collected at night between 19.00–23.00 hours in a bamboo forest near a rocky stream. The lizard was found on a bamboo branch 4 m above a large granite boulder (Fig.
The data above clearly demonstrate the unique phylogenetic and morphological properties of the individual described above as Cyrtodactylus denticulatus sp. nov. However, describing a species on the basis of a single specimen can be potentially misleading, especially in the absence of genetic data. Here we provide compelling genetic evidence for the unique phylogenetic position of this specimen wherein it resides on its own long branch (pairwise sequence divergence of 7.87–21.94%) that is not embedded within that of any other species nor sister to any other species as with C. kochangensis Grismer, Aowphol, Yodthong, Ampai, Temprayon, Aksornneam & Rujirawan, 2022 and C. uthaiensis Grismer, Aowphol, Yodthong, Ampai, Temprayon, Aksornneam & Rujirawan, 2022. Its phylogenetic position is what delimits it as a distinct evolutionarily independent lineage, not its morphology. The morphological diagnosis of a species provides evidence of how different or similar a species may be to other closely related species but has no bearing on whether or not it is a new species (see discussion in
The addition of Cyrtodactylus denticulatus sp. nov. brings the total number of described species in the brevipalmatus group to ten. Based on photographs in social media, there are as many as five potentially undescribed species from unsampled areas (Fig.
This work was supported by the National Research Council of Thailand (167951), the Thailand Science Research and Innovation (TSRI) (DBG6180025), and Chiang Mai University to Siriwadee Chomdej. We would like to thank the Laboratory Animal Research Center, University of Phayao, and Parinya Pawangkhanant for sample collection and field trips. Specimen collection protocols were approved by the Institutional Ethical Committee of Animal Experimentation of the University of Phayao (certificate number 640204005 issued to Chatmongkon Suwannapoom). Our thanks to the Thailand Research Fund 2019 (MRG6280203) and the Unit of Excellence 2023 on Biodiversity and Natural Resources Management, University of Phayao (FF66-UoE003). We also thank Anchalee Aowphol and Attapol Rujirawan for their assistance in data checking of Cyrtodactylus species in Thailand.
Dataset of meristic, normalized morphometric, and categorical characters
Data type: table (.csv file)