Research Article |
Corresponding author: L. Lee Grismer ( lgrismer@lasierra.edu ) Academic editor: Uwe Fritz
© 2022 L. Lee Grismer, Lelani del Pinto, Evan S. H. Quah, Shahrul Anuar, Michael Cota, Jimmy A. McGuire, Djoko T. Iskandar, Perry L. Wood Jr, Jesse L. 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:
Grismer LL, del Pinto L, Quah ESH, Anuar S, Cota M, McGuire JA, Iskandar DT, Wood PL Jr, Grismer JL (2022) Phylogenetic and multivariate analyses of Gekko smithii Gray, 1842 recover a new species from Peninsular Malaysia and support the resurrection of G. albomaculatus (Giebel, 1861) from Sumatra. Vertebrate Zoology 72: 47-80. https://doi.org/10.3897/vz.72.e77702
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Abstract
An integrative taxonomic analysis of Sundaic populations of Gekko smithii from the Thai-Malaya Peninsula, Sumatra, and Borneo recovered four deeply divergent mitochondrial lineages that are separated by major geographic barriers (mountains and seaways). Furthermore, they bear a number of concordant statistically significant differences in meristic and morphometric features, morphospatial separation in multivariate space, and discrete differences in color pattern. Gekko smithii sensu stricto is restricted to southern Thailand south of the Isthmus of Kra and Peninsular Malaysia west of the Banjaran (mountain range) Titiwangsa, being that the type locality is on Penang Island, Penang. Gekko hulk sp. nov. is a new species from extreme southern Thailand and Peninsular Malaysia east of the Banjaran Titiwangsa and five east coast islands—the type locality being Pulau (island) Tioman, Pahang. Gekko cf. albofasciolatus is tentatively used to include Bornean populations west of the Iran Mountains in Sabah and Sarawak which, in the absence of molecular data, cannot unequivocally be separated morphologically from G. albofasciolatus from the type locality at Banjarmasin, Kalimantan, Indonesia east of the Iran Mountains. In the absence of molecular data, G. albomaculatus is resurrected to include mainland Sumatran, Nias Island, and Banyak Islands populations which, based on their morphology, cannot be separated from descriptions of G. albomaculatus from the type locality of Bangka Island, 15 km off the southeast coast of mainland Sumatra. Further integrative analyses of all Sumatran and Bornean populations are currently underway as well as the enigmatic Wallacean populations from Sulawesi. Data are presented that strongly suggest all references to G. smithii from Java stem from a 151 year-old misidentification of a specimen of G. gecko of unknown provenance. Additionally, there are no vouchered records of G. smithii from Myanmar. The phylogeographic patterns of Sundaic populations of the G. smithii complex are concordant with those of a plethora of other Sundaic lineages.
Gekkonidae, integrative taxonomy, phylogenetics, Sundaland, systematics
The genus Gekko Laurenti, 1768 is an ecologically and morphologically diverse radiation of scansorial, nocturnal lizards comprising at least 82 species that collectively range throughout Southeast and East Asia to western Melanesia (
Distribution of the species of the Gekko smithii complex from throughout their respective Sundaic land masses. Circles denote the localities of specimens that were both examined first hand and are represented in the phylogeny (Fig.
Integrative taxonomic analyses of many other species of Sundaic amphibians and reptiles were recovered as species complexes and whose pronounced phylogeographic sub-structuring across well-defined geographic features necessitated their taxonomic partitioning into multiple species (e.g.,
The general lineage concept (GLC:
We obtained 1267 base pairs of NADH dehydrogenase subunit 2 gene (ND2) and its flanking tRNAs from six specimens from GenBank and 60 newly sequenced specimens (Table
Taxon sampling, locality data, and ND2 GenBank accession numbers from specimens used in the phylogenetic analyses.
Species | Cat no. | Locality | Accession no. |
---|---|---|---|
Gekko gecko | LSUHC 6813 | Pulau Langkawi, Kedah, West Malaysia | OM420678 |
Gekko cf. albofasciolatus | FMNH 267868 | Bintulu Division, Sarawak, East Malaysia | OM420649 |
Gekko cf. albofasciolatus | FMNH 269014 | Bintulu Division, Sarawak, East Malaysia | OM420650 |
Gekko cf. albofasciolatus | FMNH 269015 | Bintulu Division, Sarawak, East Malaysia | OM420651 |
Gekko cf. albomaculatus | MVZ 271122 | Pulau Tuangku, Aceh, Sumatra | OM420692 |
Gekko cf. albomaculatus | MVZ 271123 | Pulau Tuangku, Aceh, Sumatra | OM420693 |
Gekko cf. albomaculatus | MVZ 271125 | Pulau Nias, North Sumatra, Sumatra | OM420677 |
Gekko hulk sp. nov. | LSUHC 8696 | Pulau Perhentian Besar, Teregganu, West Malaysia | OM420680 |
Gekko hulk sp. nov. | LSUHC 8690 | Pulau Perhentian Besar, Teregganu, West Malaysia | OM420679 |
Gekko hulk sp. nov. | LSUHC 6749 | Endau Rompin, Johor, West Malaysia | OM420652 |
Gekko hulk sp. nov. | LSUHC 7651 | Endau Rompin, Johor, West Malaysia | OM420654 |
Gekko hulk sp. nov. | LSUHC 7694 | Endau Rompin, Johor, West Malaysia | OM420655 |
Gekko hulk sp. nov. | LSUHC 7702 | Endau Rompin, Johor, West Malaysia | OM420656 |
Gekko hulk sp. nov. | LSUHC 7649 | Endau Rompin, Johor, West Malaysia | OM420652 |
Gekko hulk sp. nov. | LSUHC 7650 | Endau Rompin, Johor, West Malaysia | OM420653 |
Gekko hulk sp. nov. | LSUHC 9959 | Gunung Lambak, Johor, West Malaysia | OM420657 |
Gekko hulk sp. nov. | LSUHC 6748 | Endau Rompin, Johor, West Malaysia | JN019056 |
Gekko hulk sp. nov. | LSUHC 10585 | Gunung Ledang, Johor, West Malaysia | OM420659 |
Gekko hulk sp. nov. | LSUHC 14015 | Gunung Ledang, Johor, West Malayia | OM420658 |
Gekko hulk sp. nov. | LSUHC 7251 | Pulau Tulai, Johor, West Malaysia | OM420705 |
Gekko hulk sp. nov. | LSUHC 5062 | Pulau Tulai, Johor, West Malaysia | OM420696 |
Gekko hulk sp. nov. | LSUHC 5063 | Pulau Tulai, Johor, West Malaysia | OM420697 |
Gekko hulk sp. nov. | LSUHC 3891 | Pulau Tulai, Johor, West Malaysia | OM420694 |
Gekko hulk sp. nov. | LSUHC 6265 | Pulau Tulai, Johor, West Malaysia | OM420699 |
Gekko hulk sp. nov. | LSUHC 5064 | Pulau Tulai, Johor, West Malaysia | OM420698 |
Gekko hulk sp. nov. | LSUHC 6277 | Pulau Tulai, Johor, West Malaysia | OM420701 |
Gekko hulk sp. nov. | LSUHC 6278 | Pulau Tulai, Johor, West Malaysia | OM420702 |
Gekko hulk sp. nov. | LSUHC 7024 | Pulau Tulai, Johor, West Malaysia | OM420703 |
Gekko hulk sp. nov. | LSUHC 7257 | Pulau Tulai, Johor, West Malaysia | OM420706 |
Gekko hulk sp. nov. | LSUHC 6268 | Pulau Tulai, Johor, West Malaysia | OM420700 |
Gekko hulk sp. nov. | LSUHC 7025 | Pulau Tulai, Johor, West Malaysia | OM420704 |
Gekko hulk sp. nov. | LSUHC 5061 | Pulau Tulai, Johor, West Malaysia | OM420695 |
Gekko hulk sp. nov. | LSUHC 6095 | Pekan, Pahang, West Malaysia | JQ173534 |
Gekko hulk sp. nov. | LSUHC 7264 | Pulau Tioman, Pahang, West Malaysia | OM420690 |
Gekko hulk sp. nov. | LSUHC 7263 | Pulau Tioman, Pahang, West Malaysia | OM420689 |
Gekko hulk sp. nov. | LSUHC 4681 | Pulau Tioman, Pahang, West Malaysia | OM420681 |
Gekko hulk sp. nov. | LSUHC 6278 | Pulau Tioman, Pahang, West Malaysia | OM420702 |
Gekko hulk sp. nov. | LSUHC 5390 | Pulau Tioman, Pahang, West Malaysia | OM420683 |
Gekko hulk sp. nov. | LSUHC 7299 | Pulau Tioman, Pahang, West Malaysia | OM420691 |
Gekko hulk sp. nov. | LSUHC 6284 | Pulau Tioman, Pahang, West Malaysia | OM420688 |
Gekko hulk sp. nov. | LSUHC 6890 | Pulau Tioman, Pahang, West Malaysia | JN019055 |
Gekko hulk sp. nov. | LSUHC 5152 | Pulau Tioman, Pahang, West Malaysia | OM420682 |
Gekko hulk sp. nov. | LSUHC 6283 | Pulau Tioman, Pahang, West Malaysia | OM420687 |
Gekko hulk sp. nov. | LSUHC 5849 | Pulau Tioman, Pahang, West Malaysia | OM420685 |
Gekko hulk sp. nov. | LSUHC 5399 | Pulau Tioman, Pahang, West Malaysia | OM420684 |
Gekko hulk sp. nov. | LSUHC 6260 | Pulau Tioman, Pahang, West Malaysia | OM420686 |
Gekko hulk sp. nov. | LSUHC 11977 | Sungai Bubu, Terengganu, West Malaysia | OM420711 |
Gekko hulk sp. nov. | LSUHC 11976 | Sungai Bubu, Terengganu, West Malaysia | OM420710 |
Gekko hulk sp. nov. | LSUHC 11206 | Lata Belatan, Terengganu, West Malaysia | OM420663 |
Gekko cf. albofasciolatus | ID 8774 | Gunung Mulu, Sarawak, East Malaysia | JN019054 |
Gekko smithii | LSUHC 10596 | Sedim, Kedah, West Malaysia | OM420708 |
Gekko smithii | LSUHC 9626 | Sedim, Kedah, West Malaysia | OM420707 |
Gekko smithii | LSUHC 12690 | The Gap, Pahang, West Malaysia | OM420712 |
Gekko smithii | LSUHC 14005 | The Gap, Pahang, West Malaysia | OM420713 |
Gekko smithii | LSUHC 13624 | Penang Island, Penang, West Malaysia | OM420669 |
Gekko smithii | LSUHC 13625 | Penang Island, Penang, West Malaysia | OM420670 |
Gekko smithii | LSUHC 13626 | Penang Island, Penang, West Malaysia | OM420671 |
Gekko smithii | LSUHC 13627 | Penang Island, Penang, West Malaysia | OM420672 |
Gekko smithii | LSUHC 9157 | Lata Iskandar, Pahang, West Malaysia | OM420667 |
Gekko smithii | LSUHC 9158 | Lata Iskandar, Pahang, West Malaysia | OM420668 |
Gekko smithii | LSUHC 9153 | Lata Iskandar, Pahang, West Malaysia | OM420664 |
Gekko smithii | LSUHC 9154 | Lata Iskandar, Pahang, West Malaysia | OM420665 |
Gekko smithii | LSUHC 9155 | Lata Iskandar, Pahang, West Malaysia | OM420666 |
Gekko smithii | LSUHC 15041 | Perlis State Park, Perlis, West Malaysia | OM420673 |
Gekko smithii | LSUHC 15042 | Perlis State Park, Perlis, West Malaysia | OM420674 |
Gekko smithii | LSUHC 15052 | Perlis State Park, Perlis, West Malaysia | OM420675 |
Gekko smithii | LSUHC 15053 | Perlis State Park, Perlis, West Malaysia | OM420676 |
Gekko smithii | LSUHC 6542 | Kepong, Selangor, West Malaysia | JQ173535 |
Gekko smithii | LSUHC 15084 | Kepong, Selangor, West Malaysia | OM420662 |
Gekko smithii | none | Ulu Gombok, Selangor, West Malaysia | FJ487868 |
Gekko smithii | LSUHC 6606 | Kepong, Selangor, West Malaysia | OM420661 |
Gekko smithii | LSUHC 6564 | Kepong, Selangor, West Malaysia | OM420660 |
Gekko smithii | none | Captive specimen no data | JN019057 |
Gekko smithii | LSUHC 15085 (JAM 1712) | Selangor, West Malaysia | OM420709 |
Primers used for amplification and sequencing reactions for the ND2 gene and the flanking tRNA’s.
Primer name | Primer reference | Sequence | |
L4437 | ( |
EXTERNAL | 5’ -AAGCTTTCGGGCCCATACC- 3’ |
H5934 | ( |
EXTERNAL | 5’ -AGRGTGCCAATGTCTTTGTGRTT- 3’ |
We used both maximum likelihood (ML) and Bayesian inference (BI) to estimate the phylogenetic relationships among the sampled geckos in our sequence alignment. A ML phylogeny was estimated using the IQ-TREE webserver (
Morphometric data were taken with Mitutoyo dial calipers to the nearest 0.1 mm under a Nikon SMZ 1500 dissecting microscope on the left side of the body where appropriate. Data taken were snout-vent length (SVL), taken from the tip of the snout to the vent; tail width (TW), measured at the base of the tail; axillia-groin length (AG), taken from the posterior margin of the forelimb at its insertion point on the body to the anterior margin of the hind limb at its insertion point on the body; head length (HL), measured from the posterior margin of the retroarticular process of the lower jaw to the tip of the snout; head width (HW), measured at the angle of the jaws; head depth (HD), measured from the top of the head posterior to the eyes to the bottom of the lower jaw; internarial distance (IN), measured across the snout between the dorsal margins of the external nares; orbital diameter (OD), the horizontal diameter of the bony orbit; eye to ear distance (EE), measured from the anterior margin of the auricular opening to the posterior margin of the eyeball; snout-eye length (SE), measured from anteriormost margin of the eyeball to the tip of snout; nares-eye length (NE), measured from the anterior margin of the eyeball to the posterior margin of the external nares; auricular opening (TD), measured as the horizontal distance of the ear opening; interorbital distance (IO), measured across the narrowest part of the frontal bone between the orbits; forearm length (FL), measured from the distal edge of the elbow when flexed 90° to the wrist; and crus length (CL), measured from the distal edge of the knee when flexed 90° to the ankle. The tail width (TW) measured at the base of the tail where it contacts the body.
Meristic characters evaluated were scales across frontal bone (FS), counted as the number of scales across the frontal bone at the midline of the orbits; supralabials (SL) and infralabials (IL) counted from the angle of the jaw to the rostral and mental scales, respectively; chin scales (CS), counted as the number of enlarged scales medial to and contacting the infralabials; ventral scales counted across the belly between the ventrolateral body folds midway between the limb insertions (VS); midbody scales counted across the dorsum between the ventrolateral body folds (MB); paravertebral tubercles (PVT) counted as the number of longitudinally arranged tubercles between an imaginary line between the middle of the limb insertions; longitudinal rows of tubercles (LRT) counted across the dorsum between the ventrolateral body folds midway between the limb insertions; subdigital lamellae on the first (TL1) and fourth (TL4) toes; precloacal pore-bearing scales in males (PP); and cloacal spurs (CSP). Codes for natural history collections follow
Seven of the 86 preserved specimens of Gekko smithii from the Thai-Malay Peninsula used in this study were not represented in the molecular phylogenetic analyses. These speciemens were place in a specific mitochondrial lineage based on their geographic proximity to that lineage. Following this, all individuals were subjected to a Discriminant Function Analysis (DFA) using the MASS Package in R (
All characters conformed to parametric test assumptions of normality (Shapiro-Wilk test, p < 0.05) and homogeneity of variances (F-test, p > 0.05). One-way analyses of variance (ANOVA) were conducted on characters to search for the presence of statistically significant mean differences (p < 0.05) across the data set. Characters bearing statistical differences were subjected to a TukeyHSD post hoc test to ascertain which population pairs differed significantly from each other for those particular characters. Boxplots were generated for discrete meristic characters in order to visualize the range, mean, median, and degree of differences between species pairs bearing statistically different mean values and violin plots with embedded boxplots were generated for continuous morphometric characters to visualize the same plus the distribution frequency of the data. All statistical analyses were performed in R [v3.4.3].
The morphospatial relationships of each species relative to one another and the clustering of the sampled individuals were visualized using multivariate ordination analyses on the appropriate combinations of data (meristic and morphometric). These analyses were employed as a multivariate assessment to determine if the respective data sets were morphospatially consistent with one another and the putative species boundaries delimited by the molecular phylogenetic analyses and diagnosed and defined by the univariate analyses (see below). It should be made clear these are not species delimitation analyses. Principal component analysis (PCA) implemented by the prcomp() command in R was employed to analyze the morphometric data. PCA is a dimension reducing analysis that decreases the complexity of a data set by finding a subset of input variables that contain the most relevant information (i.e. the greatest variance in the data) while de-emphasizing those characters that do not, thus increasing the overall accuracy of the model by eliminating noise and the potential of overfitting (
Discriminant analysis of principal components (DAPC) from the ADEGENET package in R (Jombart et al. 2015) was performed on the morphometric data set. The DAPC is a supervised analysis that places individuals from each predefined population into separate clusters (i.e. plots of points) bearing the smallest within-group variance that produce linear combinations of centroids having the greatest between-group variance (i.e. linear distance;
A principal coordinate analysis (PCoA)—conceptually similar to a PCA—using a Gower (dis)similarity index was employed on a concatenated meristic and adjusted morphometric data set. The dissimilarity matrix is used as the input to the analysis, not the original variables themselves. Therefore, information concerning the original variables cannot be recovered. Because this unsupervised multivariate analysis is based on a (dis)similarity index constructed from Euclidean distances between data points, it is appropriate for data sets containing discrete characters (scale counts) because it does not require the data to fulfill the assumptions of linearity (as does PCA) or unimodality (
Non-parametric permutation multivariate analyses of variance (PERMANOVA) from the VEGAN package in R (
The ML and BI analyses recovered trees with identical strongly supported topologies delimiting four major lineages separated by relatively long branch lengths that correspond to distinct geographic regions (Fig.
Percent uncorrected pairwise sequence divergence for species of the Gekko smithii complex calculated from 1267 base pairs of the mitochondrial gene ND2 and flanking tRNAs.
cf. albofasciolatus | hulk sp. nov. | cf. albomaculatus | smithii | |
cf. albofasciolatus | 0.012 | 0.128 | 0.133 | 0.133 |
hulk sp. nov. | 0.128 | 0.007 | 0.025 | 0.048 |
cf. albomaculatus | 0.133 | 0.025 | 0.015 | 0.133 |
smithii | 0.133 | 0.048 | 0.133 | 0.023 |
The DFA and DAPC placed nearly all Thai individuals for which there were no sequence data with all the sequenced individuals from the west peninsular lineage with a 99–100% posterior probability (PsP). The only exception was THNHM 01841 from the Hala-Bala Wildlife Sanctuary, Narathiwat in southeastern Thailand which was placed in the Sumatran lineage with a 79.5% PsP and in the west peninsular lineage with a 20.2% PsP. The DAPC placed it in the west peninsular lineage with a 100% PsP. All other individuals from the east peninsular lineage, Sumatra, and Borneo grouped together with a 98.9–99.9%, 83.0–99.6%, and 88.2–99.1% PsP, respectively. These data were taken as statistical justification for downstream statistical comparisons among the four major lineages. Given the results of both analyses, we consider the placement of THNHM 01841 in the Sumatran lineage as an anomaly. For downstream analyses, it was considered part of the west peninsular lineage and its placement therein is discussed below.
The PCA and DAPC (Fig.
Summary statistics and principal component analysis scores for the Gekko smithii complex. Abbreviations are listed in the Materials and methods.
PC1 | PC2 | PC3 | PC4 | PC5 | PC6 | PC7 | PC8 | PC9 | PC10 | PC11 | PC12 | PC13 | PC14 | PC15 | |
Standard deviation | 2.7774 | 1.2686 | 1.0521 | 0.9805 | 0.8919 | 0.8035 | 0.6850 | 0.6636 | 0.6195 | 0.5293 | 0.4392 | 0.3830 | 0.3579 | 0.2928 | 0.2008 |
Proportion of variance | 0.5143 | 0.1073 | 0.0738 | 0.0641 | 0.0530 | 0.0430 | 0.0313 | 0.0294 | 0.0256 | 0.0187 | 0.0129 | 0.0098 | 0.0085 | 0.0057 | 0.0027 |
Cumulative proportion | 0.5143 | 0.6216 | 0.6954 | 0.7594 | 0.8125 | 0.8555 | 0.8868 | 0.9162 | 0.9417 | 0.9604 | 0.9733 | 0.9831 | 0.9916 | 0.9973 | 1.0000 |
Eigenvalue | 7.7139 | 1.6094 | 1.1069 | 0.9613 | 0.7955 | 0.6456 | 0.4693 | 0.4403 | 0.3838 | 0.2802 | 0.1929 | 0.1467 | 0.1281 | 0.0858 | 0.0403 |
SVL | –0.2670 | –0.0713 | –0.2095 | 0.0708 | –0.1368 | 0.1959 | –0.0262 | 0.7912 | –0.4339 | 0.0327 | –0.0134 | –0.0011 | –0.0360 | –0.0101 | 0.0056 |
HH | –0.3108 | 0.1252 | 0.0566 | –0.2202 | 0.0012 | –0.1208 | 0.1317 | 0.0136 | –0.0030 | –0.5180 | 0.6112 | 0.2228 | 0.0683 | 0.2174 | –0.2458 |
HL | –0.3130 | –0.1084 | –0.1615 | 0.0913 | –0.0211 | 0.2363 | 0.0117 | –0.3320 | –0.1645 | –0.2507 | –0.3411 | –0.1758 | –0.5291 | 0.3518 | –0.2199 |
HW | –0.3019 | 0.2881 | 0.0488 | –0.1983 | 0.0018 | 0.2507 | –0.1829 | –0.1517 | –0.0452 | –0.0191 | 0.0033 | –0.0115 | –0.1359 | –0.7773 | –0.1990 |
IN | –0.2455 | 0.3503 | 0.1525 | –0.0695 | –0.2712 | –0.0030 | 0.0768 | 0.3192 | 0.6867 | 0.1069 | –0.1851 | –0.2018 | –0.0935 | 0.1732 | –0.1042 |
IO | –0.1120 | –0.4619 | 0.5028 | –0.2884 | –0.1288 | 0.1457 | 0.5758 | –0.0181 | –0.0831 | 0.2170 | 0.0125 | –0.0307 | –0.0673 | –0.0875 | –0.0331 |
TD | –0.1209 | –0.2948 | 0.5368 | 0.5117 | –0.2618 | 0.0651 | –0.4826 | –0.0036 | 0.0627 | –0.1438 | 0.0921 | 0.0542 | 0.0626 | –0.0239 | –0.0473 |
EE | –0.2132 | –0.0887 | 0.1304 | 0.2195 | 0.8088 | 0.0081 | 0.0194 | 0.1704 | 0.2056 | 0.1073 | 0.1866 | –0.0653 | –0.2876 | –0.0312 | 0.1461 |
NE | –0.3204 | 0.0005 | 0.0143 | 0.0732 | 0.3104 | 0.1812 | 0.0833 | –0.0722 | –0.0044 | –0.0081 | –0.3876 | 0.1053 | 0.7122 | 0.1017 | –0.2674 |
SE | –0.3431 | 0.0153 | –0.0478 | –0.0375 | –0.1196 | 0.1176 | 0.0811 | –0.1184 | 0.0847 | –0.2592 | –0.1184 | 0.1768 | 0.0788 | –0.0428 | 0.8378 |
OD | –0.0831 | –0.5050 | –0.1428 | –0.6109 | 0.0846 | –0.0621 | –0.5166 | 0.0734 | 0.2085 | –0.0195 | –0.0570 | –0.0893 | 0.0512 | 0.0591 | 0.0027 |
FL | –0.2209 | –0.2934 | –0.2339 | 0.2314 | –0.0827 | –0.6963 | 0.2218 | 0.0365 | 0.1004 | –0.1775 | –0.1879 | –0.0441 | –0.0185 | –0.3491 | –0.1168 |
CL | –0.3220 | 0.0301 | –0.1229 | 0.0794 | –0.1117 | –0.0752 | –0.0393 | –0.2255 | –0.1487 | 0.3058 | 0.3715 | –0.6873 | 0.2391 | 0.0775 | 0.1193 |
AG | –0.3023 | –0.0817 | –0.2673 | 0.0982 | –0.1522 | –0.0131 | –0.0620 | –0.1804 | 0.1052 | 0.5943 | 0.1739 | 0.5806 | –0.1111 | 0.1024 | –0.0816 |
TW | –0.1972 | 0.3236 | 0.4231 | –0.2267 | 0.0878 | –0.5115 | –0.2066 | –0.0064 | –0.4049 | 0.1756 | –0.2471 | 0.0935 | –0.0992 | 0.1771 | 0.0898 |
PERMANOVA summary statistics for the centroid placement between all species pairs from the PCA and PCoA analyses. Shaded cells denote insignificant adjusted p-values.
Lineage pairs | F ratio | R2 | p-value | adjusted p-value |
PCA statistics | ||||
cf. albofasciolatus vs hulk sp. nov. | 5.53231 | 0.11399 | 0.00014 | 0.00084 |
cf. albofasciolatus vs cf. albomaculatus | 3.38702 | 0.23542 | 0.00411 | 0.02471 |
cf. albofasciolatus vs smithii | 14.8605 | 0.26602 | 2.00E-05 | 0.00011 |
hulk sp. nov. vs cf. albomaculatus | 13.74370 | 0.24655 | 2.00E-05 | 0.00012 |
hulk sp. nov. vs smithii | 102.01761 | 0.58624 | 2.00E-05 | 0.00012 |
cf. albomaculatus vs smithii | 8.29619 | 0.17177 | 6.00E-05 | 0.00036 |
PCoA statistics | ||||
cf. albofasciolatus vs hulk sp. nov. | 1.83536 | 0.04387 | 0.04194 | 0.25163 |
cf. albofasciolatus vs cf. albomaculatus | 1.11739 | 0.21835 | 0.50000 | 1.00000 |
cf. albofasciolatus vs smithii | 6.80440 | 0.22089 | 0.00032 | 0.00191 |
hulk sp. nov. vs cf. albomaculatus | 5.34969 | 0.11796 | 0.00018 | 0.00107 |
hulk sp. nov. vs smithii | 53.62436 | 0.47194 | 2.00E-05 | 0.00011 |
cf. albomaculatus vs smithii | 3.93881 | 0.14098 | 0.00046 | 0.00275 |
The ANOVA analyses and subsequent TukeyHSD post hoc tests of the meristic and morphometric data sets recovered various combinations of statistically different mean values between various combinations of lineage pairs at varying levels of significance (Fig.
A. Box plots of meristic characters showing the range, mean (blue dot), and 50% quartile (rectangle) for each character. White dots are y-axis values. B. Violin plots of mensural characters embedded with boxplots showing the range, frequency, mean (white dot), and 50% quartile (black rectangle) for each character. Violin plots are vertically oriented mirror-imaged frequency diagrams.
Summary statistics from ANOVA and TukeyHSD tests of meristic data. difference = the average difference between species. lower and upper = confidence interval of the average differences. Shaded cells denoted species pairs bearing significantly different mean values.
Infralabials (IL) | difference | lower | upper | p adj | Longitudinal tubercle rows (LRT) | diff | lower | upper | p adj | |
cf. albofasciolatus-hulk sp. nov. | –0.5952 | –1.7145 | 0.5241 | 0.5074 | cf. albofasciolatus-hulk sp. nov. | –0.2857 | –1.0020 | 0.4305 | 0.7238 | |
cf. albomaculatus-hulk sp. nov. | –0.1429 | –1.6084 | 1.3227 | 0.9941 | cf. albomaculatus-hulk sp. nov. | –0.1429 | –1.0806 | 0.7949 | 0.9784 | |
smithii-hulk sp. nov. | –0.0309 | –1.1610 | 1.0992 | 0.9999 | smithii-hulk sp. nov. | –0.6178 | –1.3409 | 0.1054 | 0.1213 | |
cf. albomaculatus-cf. albofasciolatus | 0.4524 | –0.6669 | 1.5717 | 0.7157 | cf. albomaculatus-cf. albofasciolatus | 0.1429 | –0.5734 | 0.8591 | 0.9535 | |
smithii-cf. albofasciolatus | 0.5644 | –0.0538 | 1.1825 | 0.0863 | smithii-cf. albofasciolatus | –0.3320 | –0.7276 | 0.0635 | 0.1318 | |
smithii-cf. albomaculatus | 0.1120 | –1.0181 | 1.2420 | 0.9938 | smithii-cf. albomaculatus | –0.4749 | –1.1980 | 0.2482 | 0.3198 | |
Internasals (IS) | Ventral scales (VS) | |||||||||
cf. albofasciolatus-hulk sp. nov. | –0.5952 | –1.7145 | 0.5241 | 0.5074 | cf. albofasciolatus-hulk sp. nov. | –1.8333 | –5.5633 | 1.8967 | 0.5736 | |
cf. albomaculatus-hulk sp. nov. | –0.1429 | –1.6084 | 1.3227 | 0.9941 | cf. albomaculatus-hulk sp. nov. | 2.2857 | –2.5980 | 7.1694 | 0.6125 | |
smithii-hulk sp. nov. | –0.0309 | –1.1610 | 1.0992 | 0.9999 | smithii-hulk sp. nov. | 1.7104 | –2.0554 | 5.4762 | 0.6352 | |
cf. albomaculatus-cf. albofasciolatus | 0.4524 | –0.6669 | 1.5717 | 0.7157 | cf. albomaculatus-cf. albofasciolatus | 4.1190 | 0.3891 | 7.8490 | 0.0244 | |
smithii-cf. albofasciolatus | 0.5644 | –0.0538 | 1.1825 | 0.0863 | smithii-cf. albofasciolatus | 3.5438 | 1.4837 | 5.6038 | 0.0001 | |
smithii-cf. albomaculatus | 0.1120 | –1.0181 | 1.2420 | 0.9938 | smithii-cf. albomaculatus | –0.5753 | –4.3411 | 3.1905 | 0.9782 | |
Frontal bone scales (FS) | Subdigital lamellae on first toe (TL1) | |||||||||
cf. albofasciolatus-hulk sp. nov. | –2.4048 | –4.5951 | –0.2144 | 0.0256 | cf. albofasciolatus-hulk sp. nov. | 0.9048 | –0.6002 | 2.4098 | 0.3986 | |
cf. albomaculatus-hulk sp. nov. | –0.4286 | –3.2965 | 2.4393 | 0.9795 | cf. albomaculatus-hulk sp. nov. | 0.5714 | –1.3991 | 2.5419 | 0.8725 | |
smithii-hulk sp. nov. | 1.0618 | –1.1496 | 3.2732 | 0.5925 | smithii-hulk sp. nov. | 2.0965 | 0.5771 | 3.6160 | 0.0028 | |
cf. albomaculatus-cf. albofasciolatus | 1.9762 | –0.2142 | 4.1666 | 0.0920 | cf. albomaculatus-cf. albofasciolatus | –0.3333 | –1.8383 | 1.1717 | 0.9379 | |
smithii-cf. albofasciolatus | 3.4665 | 2.2568 | 4.6763 | 0.0000 | smithii-cf. albofasciolatus | 1.1918 | 0.3606 | 2.0229 | 0.0017 | |
smithii-cf. albomaculatus | 1.4903 | –0.7211 | 3.7018 | 0.2972 | smithii-cf. albomaculatus | 1.5251 | 0.0056 | 3.0445 | 0.0488 | |
Chin scales (CS) | Subdigital lamellae on fourth toe (TL4) | |||||||||
cf. albofasciolatus-hulk sp. nov. | –0.7619 | –1.7015 | 0.1777 | 0.1538 | cf. albofasciolatus-hulk sp. nov. | 1.3095 | –0.1772 | 2.7963 | 0.1043 | |
cf. albomaculatus-hulk sp. nov. | 0.7143 | –0.5160 | 1.9446 | 0.4299 | cf. albomaculatus-hulk sp. nov. | 0.2857 | –1.6609 | 2.2323 | 0.9806 | |
smithii-hulk sp. nov. | –0.0116 | –0.9603 | 0.9371 | 1.0000 | smithii-hulk sp. nov. | 2.1660 | 0.6650 | 3.6670 | 0.0016 | |
cf. albomaculatus-cf. albofasciolatus | 1.4762 | 0.5365 | 2.4158 | 0.0005 | cf. albomaculatus-cf. albofasciolatus | –1.0238 | –2.5105 | 0.4629 | 0.2788 | |
smithii-cf. albofasciolatus | 0.7503 | 0.2314 | 1.2693 | 0.0016 | smithii-cf. albofasciolatus | 0.8565 | 0.0354 | 1.6776 | 0.0375 | |
smithii-cf. albomaculatus | –0.7259 | –1.6745 | 0.2228 | 0.1945 | smithii-cf. albomaculatus | 1.8803 | 0.3793 | 3.3813 | 0.0079 | |
Midbody scales (MB) | Precloacal pores (PP) | |||||||||
cf. albofasciolatus-hulk sp. nov. | 0.0238 | –7.8616 | 7.9092 | 1.0000 | cf. albofasciolatus-hulk sp. nov. | –4.6912 | –7.8552 | –1.5271 | 0.0015 | |
cf. albomaculatus-hulk sp. nov. | 2.2857 | –8.0387 | 12.6102 | 0.9379 | cf. albomaculatus-hulk sp. nov. | 0.2500 | –4.0986 | 4.5986 | 0.9987 | |
smithii-hulk sp. nov. | 13.9614 | 6.0002 | 21.9226 | 0.0001 | smithii-hulk sp. nov. | –0.6136 | –3.7085 | 2.4812 | 0.9512 | |
cf. albomaculatus-cf. albofasciolatus | 2.2619 | –5.6235 | 10.1473 | 0.8760 | cf. albomaculatus-cf. albofasciolatus | 4.9412 | 1.3757 | 8.5067 | 0.0033 | |
smithii-cf. albofasciolatus | 13.9376 | 9.5826 | 18.2926 | 0.0000 | smithii-cf. albofasciolatus | 4.0775 | 2.2389 | 5.9161 | 0.0000 | |
smithii-cf. albomaculatus | 11.6757 | 3.7145 | 19.6369 | 0.0013 | smithii-cf. albomaculatus | –0.8636 | –4.3678 | 2.6406 | 0.9117 | |
Supralabials (SL) | Paravertebral tubercles (PVT) | |||||||||
cf. albofasciolatus-hulk sp. nov. | –0.7143 | –1.9074 | 0.4789 | 0.4024 | cf. albofasciolatus-hulk sp. nov. | 0.0952 | –1.6545 | 1.8450 | 0.9990 | |
cf. albomaculatus-hulk sp. nov. | 0.0000 | –1.5622 | 1.5622 | 1.0000 | cf. albomaculatus-hulk sp. nov. | 0.4286 | –1.8624 | 2.7195 | 0.9612 | |
smithii-hulk sp. nov. | –0.1351 | –1.3398 | 1.0695 | 0.9911 | smithii-hulk sp. nov. | 0.4170 | –1.3496 | 2.1835 | 0.9261 | |
cf. albomaculatus-cf. albofasciolatus | 0.7143 | –0.4789 | 1.9074 | 0.4024 | cf. albomaculatus-cf. albofasciolatus | 0.3333 | –1.4164 | 2.0831 | 0.9591 | |
smithii-cf. albofasciolatus | 0.5792 | –0.0798 | 1.2381 | 0.1055 | smithii-cf. albofasciolatus | 0.3218 | –0.6446 | 1.2881 | 0.8194 | |
smithii-cf. albomaculatus | –0.1351 | –1.3398 | 1.0695 | 0.9911 | smithii-cf. albomaculatus | –0.0116 | –1.7781 | 1.7550 | 1.0000 |
Summary statistics from ANOVA and TukeyHSD tests of morphometric data. difference = the average difference between species. lower and upper = confidence interval of the average differences. Shaded cells denoted species pairs bearing significantly different mean values.
Snout-vent length (SVL) | difference | lower | upper | p adj | Tympanum diameter (TD) | difference | lower | upper | p adj | |
cf. albofasciolatus-hulk sp. nov. | -0.0317 | -0.0854 | 0.0219 | 0.4086 | cf. albofasciolatus-hulk sp. nov. | 0.0239 | -0.0594 | 0.1073 | 0.8730 | |
cf. albomaculatus-hulk sp. nov. | -0.0031 | -0.0762 | 0.0700 | 0.9995 | cf. albomaculatus-hulk sp. nov. | 0.0139 | -0.0997 | 0.1275 | 0.9883 | |
smithii-hulk sp. nov . | 0.0490 | -0.0060 | 0.1040 | 0.0971 | smithii-hulk sp. nov. | 0.0622 | -0.0232 | 0.1476 | 0.2294 | |
cf. albomaculatus-cf. albofasciolatus | 0.0287 | -0.0250 | 0.0823 | 0.4979 | cf. albomaculatus-cf. albofasciolatus | -0.0100 | -0.0934 | 0.0733 | 0.9888 | |
smithii-cf. albofasciolatus | 0.0807 | 0.0572 | 0.1043 | 0.0000 | smithii-cf. albofasciolatus | 0.0383 | 0.0017 | 0.0748 | 0.0370 | |
smithii-cf. albomaculatus | 0.0521 | -0.0029 | 0.1070 | 0.0697 | smithii-cf. albomaculatus | 0.0483 | -0.0371 | 0.1337 | 0.4481 | |
Head height (HH) | Eye-ear distance (EE) | |||||||||
cf. albofasciolatus-hulk sp. nov. | -0.0195 | -0.0673 | 0.0283 | 0.7054 | cf. albofasciolatus-hulk sp. nov. | -0.0188 | -0.0744 | 0.0369 | 0.8101 | |
cf. albomaculatus-hulk sp. nov. | 0.0257 | -0.0394 | 0.0908 | 0.7258 | cf. albomaculatus-hulk sp. nov. | 0.0118 | -0.0640 | 0.0876 | 0.9764 | |
smithii-hulk sp. nov. | 0.0490 | 0.0000 | 0.0979 | 0.0499 | smithii-hulk sp. nov. | 0.0494 | -0.0076 | 0.1064 | 0.1123 | |
cf. albomaculatus-cf. albofasciolatus | 0.0452 | -0.0026 | 0.0930 | 0.0703 | cf. albomaculatus-cf. albofasciolatus | 0.0306 | -0.0251 | 0.0862 | 0.4736 | |
smithii-cf. albofasciolatus | 0.0685 | 0.0475 | 0.0894 | 0.0000 | smithii-cf. albofasciolatus | 0.0681 | 0.0437 | 0.0925 | 0.0000 | |
smithii-cf. albomaculatus | 0.0233 | -0.0257 | 0.0722 | 0.5957 | smithii-cf. albomaculatus | 0.0376 | -0.0194 | 0.0946 | 0.3127 | |
Head length (HL) | Nares-eye distance (NE) | |||||||||
cf. albofasciolatus-hulk sp. nov. | -0.0129 | -0.0348 | 0.0089 | 0.4091 | cf. albofasciolatus-hulk sp. nov. | -0.0143 | -0.0464 | 0.0178 | 0.6436 | |
cf. albomaculatus-hulk sp. nov. | 0.0216 | -0.0083 | 0.0514 | 0.2354 | cf. albomaculatus-hulk sp. nov. | 0.0144 | -0.0293 | 0.0580 | 0.8216 | |
smithii-hulk sp. nov. | 0.0574 | 0.0350 | 0.0798 | 0.0000 | smithii-hulk sp. nov. | 0.0534 | 0.0206 | 0.0862 | 0.0004 | |
cf. albomaculatus-cf. albofasciolatus | 0.0345 | 0.0126 | 0.0564 | 0.0006 | cf. albomaculatus-cf. albofasciolatus | 0.0287 | -0.0034 | 0.0607 | 0.0957 | |
smithii-cf. albofasciolatus | 0.0704 | 0.0608 | 0.0800 | 0.0000 | smithii-cf. albofasciolatus | 0.0677 | 0.0536 | 0.0818 | 0.0000 | |
smithii-cf. albomaculatus | 0.0359 | 0.0135 | 0.0583 | 0.0004 | smithii-cf. albomaculatus | 0.0390 | 0.0062 | 0.0719 | 0.0135 | |
Head width (HW) | Snout-eye distance (SE) | |||||||||
cf. albofasciolatus-hulk sp. nov. | -0.0164 | -0.0570 | 0.0242 | 0.7117 | cf. albofasciolatus-hulk sp. nov. | -0.0102 | -0.0329 | 0.0126 | 0.6415 | |
cf. albomaculatus-hulk sp. nov. | 0.0176 | -0.0377 | 0.0730 | 0.8349 | cf. albomaculatus-hulk sp. nov. | 0.0266 | -0.0044 | 0.0576 | 0.1171 | |
smithii-hulk sp. nov. | 0.0461 | 0.0045 | 0.0877 | 0.0243 | smithii-hulk sp. nov. | 0.0614 | 0.0381 | 0.0847 | 0.0000 | |
cf. albomaculatus-cf. albofasciolatus | 0.0340 | -0.0066 | 0.0746 | 0.1313 | cf. albomaculatus-cf. albofasciolatus | 0.0368 | 0.0141 | 0.0596 | 0.0004 | |
smithii-cf. albofasciolatus | 0.0625 | 0.0446 | 0.0803 | 0.0000 | smithii-cf. albofasciolatus | 0.0716 | 0.0616 | 0.0816 | 0.0000 | |
smithii-cf. albomaculatus | 0.0284 | -0.0132 | 0.0700 | 0.2819 | smithii-cf. albomaculatus | 0.0347 | 0.0114 | 0.0581 | 0.0012 | |
Internarial distance (IN) | Orbit diameter (OD) | |||||||||
cf. albofasciolatus-hulk sp. nov. | -0.0133 | -0.0739 | 0.0474 | 0.9387 | cf. albofasciolatus-hulk sp. nov. | 0.0181 | -0.0218 | 0.0580 | 0.6323 | |
cf. albomaculatus hulk sp. nov. | 0.0413 | -0.0413 | 0.1240 | 0.5541 | cf. albomaculatus-hulk sp. nov. | 0.0249 | -0.0295 | 0.0792 | 0.6253 | |
smithii-hulk sp. nov. | 0.0426 | -0.0195 | 0.1048 | 0.2784 | smithii-hulk sp. nov. | 0.0380 | -0.0028 | 0.0789 | 0.0772 | |
cf. albomaculatus-cf. albofasciolatus | 0.0546 | -0.0060 | 0.1152 | 0.0923 | cf. albomaculatus-cf. albofasciolatus | 0.0068 | -0.0331 | 0.0467 | 0.9696 | |
smithii-cf. albofasciolatus | 0.0559 | 0.0293 | 0.0825 | 0.0000 | smithii-cf. albofasciolatus | 0.0200 | 0.0025 | 0.0375 | 0.0192 | |
smithii-cf. albomaculatus | 0.0013 | -0.0608 | 0.0634 | 0.9999 | smithii-cf. albomaculatus | 0.0132 | -0.0277 | 0.0541 | 0.8302 | |
Interorbital distance (IO) | Forearm length (FL) | |||||||||
cf. albofasciolatus-hulk sp. nov. | 0.0258 | -0.0298 | 0.0815 | 0.6135 | cf. albofasciolatus-hulk sp. nov. | -0.0063 | -0.0553 | 0.0427 | 0.9864 | |
cf. albomaculatus-hulk sp. nov. | 0.0201 | -0.0557 | 0.0959 | 0.8967 | cf. albomaculatus-hulk sp. nov. | 0.0094 | -0.0574 | 0.0762 | 0.9822 | |
smithii-hulk sp. nov. | 0.0457 | -0.0113 | 0.1028 | 0.1588 | smithii-hulk sp. nov. | 0.0764 | 0.0262 | 0.1266 | 0.0009 | |
cf. albomaculatus-cf. albofasciolatus | -0.0057 | -0.0614 | 0.0499 | 0.9930 | cf. albomaculatus-cf. albofasciolatus | 0.0157 | -0.0333 | 0.0648 | 0.8318 | |
smithii-cf. albofasciolatus | 0.0199 | -0.0045 | 0.0443 | 0.1482 | smithii-cf. albofasciolatus | 0.0827 | 0.0612 | 0.1042 | 0.0000 | |
smithii-cf. albomaculatus | 0.0256 | -0.0314 | 0.0826 | 0.6380 | smithii-cf. albomaculatus | 0.0670 | 0.0167 | 0.1172 | 0.0044 | |
Crus length (CL) | Axilla-groin distance (AG) | |||||||||
cf. albofasciolatus-hulk sp. nov. | -0.0236 | -0.0595 | 0.0123 | 0.3136 | cf. albofasciolatus-hulk sp. nov. | 0.0094 | -0.0186 | 0.0374 | 0.8128 | |
cf. albomaculatus-hulk sp. nov. | 0.0105 | -0.0384 | 0.0594 | 0.9416 | cf. albomaculatus-hulk sp. nov. | 0.0317 | -0.0065 | 0.0699 | 0.1365 | |
smithii-hulk sp. nov. | 0.0597 | 0.0229 | 0.0964 | 0.0004 | smithii-hulk sp. nov. | 0.0821 | 0.0534 | 0.1108 | 0.0000 | |
cf. albomaculatus-cf. albofasciolatus | 0.0341 | -0.0018 | 0.0700 | 0.0681 | cf. albomaculatus-cf. albofasciolatus | 0.0223 | -0.0057 | 0.0503 | 0.1634 | |
smithii-cf. albofasciolatus | 0.0833 | 0.0676 | 0.0990 | 0.0000 | smithii-cf. albofasciolatus | 0.0727 | 0.0604 | 0.0850 | 0.0000 | |
smithii-cf. albomaculatus | 0.0492 | 0.0124 | 0.0859 | 0.0042 | smithii-cf. albomaculatus | 0.0504 | 0.0217 | 0.0791 | 0.0001 | |
Tail width (TW) | ||||||||||
cf. albofasciolatus-hulk sp. nov. | 0.0094 | -0.0186 | 0.0374 | 0.8128 | ||||||
cf. albomaculatus-hulk sp. nov. | 0.0317 | -0.0065 | 0.0699 | 0.1365 | ||||||
smithii-hulk sp. nov. | 0.0821 | 0.0534 | 0.1108 | 0.0000 | ||||||
cf. albomaculatus-cf. albofasciolatus | 0.0223 | -0.0057 | 0.0503 | 0.1634 | ||||||
smithii-cf. albofasciolatus | 0.0727 | 0.0604 | 0.0850 | 0.0000 | ||||||
smithii-cf. albomaculatus | 0.0504 | 0.0217 | 0.0791 | 0.0001 |
Summary statistics of the adjusted morphometric data. sd = 1 standard deviation. N = sample size. Character abbreviations are in the Materials and methods.
smithii | SVL | HH | HL | HW | IN | IO | TD | EE | NE | SE | OD | FL | CL | AG | TW |
mean (±sd) | 2.21 (±0.036) | 1.22 (±0.023) | 1.66 (±0.018) | 1.45 (±0.03) | 0.63 (±0.04 | 0.95 (±0.027) | 0.74 (±0.03) | 1.09 (±0.023) | 1.13 (±0.014) | 1.27 (±0.013) | 1.05 (±0.032) | 1.33 (±0.043) | 1.41 (±0.021) | 1.91 (±0.019) | 1.08 (±0.054) |
range | 2.13–2.25 | 1.19–1.27 | 1.63–1.68 | 1.39–1.5 | 0.59–0.74 | 0.91–0.99 | 0.69–0.79 | 1.06–1.14 | 1.11–1.15 | 1.24–1.29 | 0.99–1.1 | 1.26–1.41 | 1.37–1.44 | 1.88–1.93 | 1.01–1.15 |
N | 11 | 11 | 11 | 11 | 11 | 11 | 11 | 11 | 11 | 11 | 11 | 11 | 11 | 11 | 11 |
hulk sp. nov. | |||||||||||||||
mean (±sd) | 2.14 (±0.038) | 1.17 (±0.026) | 1.59 (±0.009) | 1.41 (±0.019) | 0.58 (±0.037) | 0.93 (±0.036) | 0.72 (±0.056) | 1.04 (±0.038) | 1.08 (±0.019) | 1.21 (±0.013) | 1.02 (±0.019) | 1.25 (±0.026) | 1.34 (±0.017) | 1.83 (±0.015) | 1.1 (±0.041) |
range | 2.07–2.21 | 1.11–1.23 | 1.57–1.61 | 1.37–1.46 | 0.52–0.66 | 0.84–1.0 | 0.61–0.82 | 0.96–1.13 | 1.05–1.12 | 1.18–1.23 | 0.98–1.05 | 1.15–1.28 | 1.31–1.39 | 1.8–1.86 | 0.95–1.18 |
N | 38 | 38 | 38 | 38 | 38 | 38 | 38 | 38 | 38 | 38 | 38 | 38 | 38 | 38 | 38 |
cf. albomaculatus | |||||||||||||||
mean (±sd) | 2.17 (±0.012) | 1.21 (±0.011) | 1.63 (±0.011) | 1.44 (±0.007) | 0.63 (±0.036) | 0.91 (±0.051) | 0.7 (±0.059) | 1.07 (±0.011) | 1.11 (±0.019) | 1.25 (±0.009) | 1.01 (±0.032) | 1.28 (±0.025) | 1.37 (±0.035) | 1.85 (±0.026) | 1.09 (±0.029) |
range | 2.16–2.2 | 1.2–1.22 | 1.61–1.64 | 1.43–1.44 | 0.59–0.68 | 0.8–0.94 | 0.6–0.76 | 1.05–1.08 | 1.07–1.12 | 1.23–1.26 | 0.95–1.04 | 1.25–1.31 | 1.33–1.43 | 1.82–1.88 | 1.05–1.13 |
N | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 | 6 |
cf. albofasciolatus | |||||||||||||||
mean (±sd) | 2.16 (±0.02 | 1.18 (±0.019 | 1.62 (±0.015 | 1.42 (±0.013 | 0.61 (±0.014 | 0.93 (±0.036 | 0.71 (±0.034 | 1.07 (±0.027 | 1.10 (±0.014 | 1.23 (±0.013 | 1.00 (±0.03 |
1.26 (±0.031 | 1.37 (±0.014 | 1.83 (±0.016 | 1.07 (±0.034 |
range | 2.13–2.18 | 1.16–1.21 | 1.6–1.64 | 1.41–1.44 | 0.59–0.62 | 0.87–0.99 | 0.63–0.73 | 1.03–1.11 | 1.08–1.12 | 1.21–1.25 | 0.95–1.04 | 1.22–1.32 | 1.35–1.39 | 1.81–1.85 | 1.03–1.13 |
N | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 |
Summary statistics of the meristic data. sd = 1 standard deviation. N = sample size. Character abbreviations are in the Materials and methods.
smithii | SL | IL | IN | FS | CS | MB | PVT | LRT | VS | TL1 | TL4 | PP |
mean (±sd) | 13.7 (±1.48) | 11.6 (±1.28) | 4.5 (±0.75) | 22.6 (±2.06) | 6.7 (±1.03) | 112.9 (±9.73) | 19.1 (±1.94) | 9.7 (±0.61) | 29.2 (±3.17) | 17.6 (±1.39) | 21.4 (±1.5) | 13.2 (±1.95) |
range | 11–17 | 9–14 | 3–6 | 19–26 | 4–9 | 94–137 | 17–23 | 8–11 | 23–35 | 15–20 | 19–24 | 13–15 |
N | 27 | 27 | 27 | 27 | 27 | 27 | 27 | 27 | 27 | 27 | 27 | 23 |
hulk sp. nov. | ||||||||||||
mean (±sd) | 13.3 (±0.94) | 11 (±0.92) | 4.2 (±0.55) | 19.5 (±2.1) | 6 (±0.91) | 98.7 (±6.82) | 18.4 (±1.31) | 10.1 (±0.57) | 26.0 (3.74) | 16.6 (±1.27) | 20.7 (±1.23) | 8.7 (±1.85) |
range | 10–15 | 9–13 | 3–5 | 13–24 | 4–8 | 84–110 | 16–21 | 9–11 | 22–28 | 14–19 | 18–24 | 6–13 |
N | 42 | 42 | 42 | 42 | 42 | 42 | 42 | 42 | 42 | 42 | 42 | 16 |
cf. albomaculatus | ||||||||||||
mean (±sd) | 14 (±1.00) | 11.4 (±1.27) | 4.7 (±0.49) | 21.4 (±2.7) |
7.4 (±0.53) | 101 (±5.69) |
18.7 (±1.8) |
10.3 (±0.76) | 30.1 (±2.41) | 16.3 (±1.5) |
19.7 (±1.5) |
14.0 (±3.00) |
range | 13–16 | 10–14 | 4–5 | 17–26 | 7–8 | 90–109 | 16–21 | 9–11 | 26–33 | 14–18 | 17–21 | 11–17 |
N | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 3 |
cf. albofasciolatus | ||||||||||||
mean (±sd) | 14 (±0.82) | 11.6 (±0.98 |
5.3 (±0.49) | 21.9 (±1.77) | 6.7 (±0.49) | 98.7 (±3.35) |
18.3 (±1.5) | 10.4 (±0.98) | 27.9 (±4.1) |
15.7 (±1.8) |
19.4 (±1.99) | 13.6 (±0.96) |
range | 13–15 | 11–13 | 5 or 6 | 19–24 | 6 or 7 | 94–103 | 16–20 | 9–12 | 23–33 | 14–19 | 17–22 | 13–15 |
N | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 7 | 4 |
Geckos from the west peninsular lineage have a considerable amount of dark dorsal blotching in the interspaces between the transversely aranged white spots which often tend to form bands (Fig.
Distribution and color pattern variation in Gekko smithii and G. hulk sp. nov. on the Thai-Malay Peninsula. A. Gekko smithii (LSUHC 15052) from Perlis State Park, Perlis, Peninsular Malaysia. B. Gekko smithii (LSUDPC 12884) from Puncak Janing, Kedah, Peninsualr Malaysia. C. Gekko smithii (LSUHC 9626) from Sedim, Kedah, Peninsular Malaysia. D. Gekko smithii (LSUDPC 12869) from Penang Hill (type locality), Penang, Peninsular Malaysia. E. Gekko smithii (LSUDPC 12886) from Lata Kedondong, Selangor, Peninsular Malaysia. F. Gekko smithii (LSUDPC 12887) from Kota Damansara, Selangor, Peninsular Malaysia. G. Gekko smithii (LSUDPC 11174) from Khaopu Khaoya, Nakhon Si Thamarat Province, Thailand. H. Gekko smithii (LSUDPC 11173) from Bacho Falls, Narathiwat Province, Thailand. I. Gekko smithii (LSUDPC 12878) from the Weang District, Narathiwat Province, Thailand. J. Gekko hulk sp. nov. (LSUDPC 12880) from the Hala-Bala Wildlife Sanctuary, Narathiwat Province, Thailand. K. Gekko hulk sp. nov. (LSUHC 8696) from Pulau Perhentian Besar, Terengganu, Peninsular Malaysia. L. Gekko hulk sp. nov. (LSUDPC 7991) from Gunung Tebu, Terengganu, Peninsular Malaysia. M. Gekko hulk sp. nov. (LSUDPC 12897) from Merapoh, Pahang, Peninsular Malaysia. N. Gekko hulk sp. nov. (LSUDPC 12897) from Pulau Tioman, Pahang, Peninsular Malaysia. Photographs by L. Lee Grismer (A, K, L, N), Evan S. H. Quah (B, C, D), Kurt H. P. Guek (E), Steven Wong (F), Henrick Bringsøe (G, H); Parinya Pawangkhanant (I), Ian Dugdale (J), and Nick Baker (M). Unannotated circles denote localities of specimens or photographs examined here or verified from other publications as well as vouchered samples in the literature.
Given that the multivariate and univariate diagnoses corroborate the geographically structured phylogenetic delimitation of distinct allopatric mitochondrial lineages within Gekko smithii across prominent biogeographic barriers (i.e. mountain ranges and seaways), we hypothesize these are non-reticulating lineages that should be designated as separate species. Because Penang Island is the type locality for Gekko smithii (
Gecko Smithii:
Platydactylus Stentor:
Gecko Smithii: Duméril 1856:449;
Gecko Stentor:
Gecko Stentor:
Gekko stentor:
Gekko smithi:
Gecko smithii:
Gekko gecko:
Gekko smithii:
Gekko albofasciolatus:
Gekko (Gekko) smithii:
Gekko cf. smithii:
Non-technical books, field guides, and pockets guides are not listed
Gekko smithii sensu stricto herein after referred to as G. smithii unless noted otherwise can be separated from all other species of Gekko in the G. smithii species complex by having the combination of a maximum SVL of 191.1 mm, 11–17 supralabials, 9–14 infralabials, 3–6 internarial scales, 19–26 frontal scales, 4–9 chin scales, 94–137 midbody scales, 17–23 paravertebral tubercles, 8–11 longitudinal rows of tubercles, 23–35 ventral scales, 15–20 1st toe subdigital lamellae, 19–24 4th toe lamellae; 13–15 precloacal pores in males (absent in females); enlarged subcaudal scales; thin, white nuchal band at base of occiput composed of closely spaced spots; thin dark nuchal band contacting the eyes; large white ocelli surrounding dorsal tubercles or bordering them posteriorly in six or seven transverse rows; and a thick dark reticulum to diffuse banded dorsal pattern (Tables
Diagnostic character states (shaded cells) separating Gekko hulk sp. nov. from the other species of G. (Gekko). Data on G. gecko, G. nutaphandi, G. reevesi, G. siamensis, G. stoliczkai, and G. verreauxi come from
albofasciolatus | alblomaculatus | gecko | hulk sp. nov. | nutaphandi | reevesi | siamensis | smithii | stoliczkai | verreauxi | |
max SVL | 165.1 | 157.4 | 185 | 161.3 | 117 | 173 | 150 | 191 | 128.4 | 155 |
precloacal pores (males) | 13–15 | 11–17 | 12–16 | 6–13 | 17–22 | 13–20 | 10–13 | 13–15 | 13–15 | 11–13 |
rostral scale contacting nares | no | no | no | no | no | no | no | no | no | yes |
occiput scales smaller than snout scales | yes | yes | no | yes | yes | no | yes | yes | yes | yes |
longitudinal rows of tubercles | 9–12 | 9–11 | 9–18 | 9–11 | 14 | 12–18 | 16–19 | 8–11 | 10–12 | 10–14 |
subcaudals enlarged | yes | yes | no | yes | yes | no | yes | yes | yes | yes |
iris color | green | green | gold/copper/ olive/brown | turquoise/green | red | gold/copper/ olive/brown | green | turquoise/green | green | golden |
solid, thin, white nuchal band | no | yes | no | no | no | no | no | no | no | no |
Gecko smithii ranges from southern Thailand south of the Isthmus of Kra from at least Khao Phanom Bencha National Park in Krabi and Khao Nan National Park in Nakhon Si Thammarat Province to the northern border of the Banjaran Titiwangsa in southeastern Thailand and northwestern Peninsular Malaysia. Its range continues southward along the west side of the Banjaran Titiwangsa to at least the state of Selangor but very likely farther as well (Figs
The dorsal body tubercles of Gekko smithii from northernmost Peninsular Malaysia and Thailand north of the Kangar-Pattani Line (generally running from Gunung Jerai, Kedah, Peninsular Malaysia to Songkhla, Songkhla, Thailand) are surrounded by a large white ocellus whereas in southern populations, the ocelli are smaller and may only border the tubercles posteriorly (Figs
Geographic variation in Gekko smithii (A–G) and G. hulk sp. nov. (H–M). A. Gekko smithii (LSUDPC 12899) from Ulu Yam, Selangor, Peninsular Malaysia. B. Gekko smithii (LSUHC 4959) from Pulau Pangkor, Perak, Peninsular Malaysia. C. Gekko smithii (LSUDPC 12898) from Ulu Yam, Selangor, Peninsular Malaysia. D. Gekko smithii (LSUDPC 12895) from Kota Damansara, Selangor, Peninsular Malaysia. E. Gekko smithii (LSUDPC 12903) from Bukit Panchor, Penang, Peninsular Malaysia. F. Gekko smithii (LSUDPC 12902) from Ulu Yam, Selangor, Peninsular Malaysia. G. Gekko smithii (THNHM 01844) from Phu Pha Phet, Satun, Province, Thailand. H. Gekko hulk sp. nov. (LSUDPC 12912) from Lata Kekabu, Setiu, Terengganu, Peninsular Malaysia. I. Gekko hulk sp. nov. (LSUDPC 8696) from Endau-Rompin, Johor, Peninsular Malaysia. J. Gekko hulk sp. nov. (LSUDPC 12953) from the Hala-Bala Wildlife Sanctuary, Narathiwat Province, Thailand. K. Gekko hulk sp. nov. (LSUHC 5152) from the Tekek-Juara Trail, Pulau Tioman, Pahang, Peninsular Malaysia. L. Gekko hulk sp. nov. (LSUHC 5873) from Kota Tinggi, Peninsular Malaysia. M. Gekko hulk sp. nov. (LSUDPC 12955) from Hutan Lipur Sekayu, Terengganu, Peninsular Malaysia. Photographs by Kurt H. P. Guek (A, C, D, F), L. Lee Grismer (B, K, L), Evan S. H. Quah (E, H), Michael Cota (G, J), M. A. Muin (I) and Syed A. Rizal (M).
Gekko smithii is an arboreal nocturnal species that is well-established in all types of primary and secondary forests. It is particularly well-suited for inhabiting occupied and abandoned human dwellings along forest edges where it is most easily observed. In forested habitats, geckos occur on the trunks of large trees and boulders from where males are commonly heard calling during the day and early in the evening.
Distribution and color pattern variation in Gekko albofasciolatus and G. cf. albofasciolatus in Borneo A. LSUDPC 12916 from Gunung Mulu, Sarawak, East Malaysia. B. LSUDPC 12918 from Gunung Mulu, Sarawak, East Malaysia. C. LSUDPC 12919) from Kelay Subdistrict, Kalimantan, Indonesia. D. LSUDPC 1293 from Kalimantan, Indonesia. E. LSUDPC 12949 from Kinabatangan River, Sabah, East Malaysia. F. LSUDPC 12924 from Lambir, Sarawak, East Malaysia. G. LSUDPC 12925 from Lambir, Sarawak, East Malaysia. H. LSUDPC 12929 from Gunung Gading, Sarawak, East Malaysia. Unannotated circles denote localities of specimens or photographs examined here or verified from other publications as well as vouchered samples in the literature. Photographs by Alan Watson www.alanwatsonfeatherstone.com (A), Wolfgang Grossmann (B, H), C. J. Franklin (C), Mistar Kamsi (D), Chien C. Lee (E), and Nick Baker (F,G).
Platydactylus albomaculatus:
Gecko stentor:
Gecko fascoilatus:
Gekko stentor:
Gekko smithii: Ota, Hikida, and Matsui 1991:150 (in part);
Non-technical books, field guides, and pockets guides are not listed
Comparing the short description of the two syntypes of Platydactylus albomaculatus by
Distribution and color pattern variation in Gekko albomaculatus and G. cf. albomaculatus in Sumatra A. LSUDPC 12933 from Gunung Leuser NP, North Sumatra, Sumatra, Indonesia. B. LSUDPC 12934 (JAM 10261) from Pulau Nias, Sumatra, Indonesia. C. LSUDPC 12940 from Andalas University, Sumatra Barat, Sumatra, Indonesia. D. LSUDPC 12944 from Kecamatan Ngambur, Lampung, Sumatra, Indonesia. E. LSUDPC 12952 from Sum, Sumatra, Indonesia. F. LSUDPC 12924 from Sumber Rejio, Bengkuat Belimbing, Sumatra, Indonesia. G. Syntypes of G. albomaculatus from Banka Island, Sumatra, Indonesia. Unannotated circles denote localities of specimens or photographs examined here or verified from other publications as well as vouchered samples in the literature. Photographs by Andrea Molyneaux (A), Jimmy A. McGuire (B), Eric N. Smith (C, E), C. J. Franklin (D), Photo from Creative Commons Attribution Share Alike (F), and Frank Tillack (G).
Gecko Stentor:
Gekko smithi:
Gekko smithii:
Gekko albofasciolatus:
Gekko albomaculatus:
Gekko
sp.
Gekko (Gekko) smithii:
Non-technical books, field guides, and pockets guides are not listed
Adult male LSUHC 6284 from the Tekek-Juara trail on Pulau Tioman, Pahang, Peninsular Malaysia (2.821021°N 104.179596°E; 462 m) collected by Jesse L. Grismer, Perry L. Wood, Jr., and L. Lee Grismer on 2 July 2004.
All paratypes are from Peninsular Malaysia and were collected by various personnel from La Sierra University, Universiti Sains Malaysia, Universiti Kabangsaan Malaysia, and the Department of Wildlife and National Parks Malaysia. Adult female LSUHC 6283 bears the same data as the holotype. Adult females LSUHC 5152 and 5399 from the upper Tekek-Juara trail on Pulau Tioman, Pahang (2.821021°N 104.179596°E; 462 m) collected on 3 March 2003. Adult female LSUHC 7026 from Pulau Tulai, Johor (2.909920°N 104.105315°E; 11 m) collected on 14 September 2004. Adult male LSUHC 5062 from Pulau Tulai, Johor (2.909920°N 104.105315°E; 11 m) collected on 17 August 2002. Adult male LSUHC 7648 from Endau-Rompin, Peta, Visitor center, Johor (2.530818°N 103.414191°E; 42 m) collected on 25 August 2005 by Perry L. Wood, Jr., Kin Onn Chan, and L. Lee Grismer. Adult female LSUHC 7649 and 7651 from Endau-Rompin, Peta, Visitor center, Johor (2.530818°N 103.414191°E; 42 m) collected on 25 August 2005 by Perry L. Wood, Jr., Kin Onn Chan, and L. Lee Grismer. Adult female LSUHC 7694 from Endau-Rompin, Peta, Sungai Semawak, Johor (2.529150°N 103.401173°E; 36 m) collected on 29 August 2005 by Perry L. Wood, Jr., Kin Onn Chan, Norhayati Ahmad, and L. Lee Grismer. Juvenile female LSUHC 10585 from Gunung Ledang, Johor (2.529150°N 103.401173°E; 36 m) collected on 31 May 2008, collector unknown 2008. Juvenile female LSUHC 9959 from Gunung Lambak, (2.029934°N 103.353323°E; 255 m). Juvenile male ZRC 2.6014 from FRIM, Pasoh, Negeri Sembian (2.968545°N 102.297043°E; 255 m) collected on 27 November 2008. Adult male LSUHC 1197 from Sungai Bubu, Terengganu (4.997105°N 102.953106°E; 174 m) collected on 1 September 2009 by L. Lee Grismer and Kin Onn Chan.
Gekko hulk sp. nov. can be separated from all other species of Gekko in the G. smithii species complex by having the combination of a maximum SVL of 161.3 mm, 10–15 supralabials, 9–13 infralabilas, 3–5 internarial scales, 13–24 frontal scales, 4–8 chin scales, 84–110 midbody scales, 16–21 paravertebral tubercles, 9–11 longitudinal rows of tubercles, 22–28 ventral scales, 14–19 1st toe subdigital lamellae, 18–24 4th toe lamellae; 6–13 precolacal pores in males (absent in females); subcaudals enlarged; thin, white nuchal band at base of occiput composed of closely spaced spots; thin dark nuchal band absent or faded and never contacting eyes; small white ocelli confined to dorsal tubercles or their anterior margin in six or seven transverse rows; and no thick dark reticulum on body (Tables
Adult male SVL 147.5 mm; head moderate in length (HL/SVL 0.28), width (HW/HL 0.65), somewhat flattened, distinct from neck, triangular in dorsal profile; lores concave slightly anteriorly, weakly inflated posteriorly; prefrontal region concave; canthus rostralis rounded; snout elongate (SE/HL 0.41), rounded in dorsal profile; eye large (OD/HL 0.26), pupil vertical, margins crenulated; ear opening elliptical, obliquely oriented, moderate in size; eye to ear distance slightly less than diameter of eye; rostral rectangular, bordered posteriorly by large left and right supranasals and smaller azygous postrostral, bordered laterally by first supralabials; external nares seated anteriorly in nasal scale bordered anteriorly by rostral, dorsally by large anterior and smaller posterior supranasals, posteriorly by three small postnasals, ventrally by first and second supralabials; five internasals; 12 (R,L) rectangular supralabials, first supralabial slightly larger than second; 12(R) 10(L) infralabials tapering smoothly to angle of jaw; scales of rostrum and lores flat, larger than flat scales on top of head and occiput; scales of occiput intermixed with distinct, small tubercles; superciliaries elongate, largest dorsally; mental not enlarged, subtriangular, bordered laterally by first infralabials and posteriorly by left and right trapezoidal postmentals contacting medially for 60% of their length posterior to mental; one row of slightly enlarged, elongate chin scales extending posteriorly to fifth (R) and sixth (L) infralabial; gular and throat scales small, flat, juxtaposed, grading posteriorly into larger, smooth, imbricate, pectoral scales which grade into larger ventral scales.
Body slightly flattened, relatively long (AG/SVL 0.51) with well-defined ventrolateral folds; dorsal scales small, flat, juxtaposed, interspersed with larger, smooth subconical, regularly arranged tubercles; 110 longitudinal rows of scales at midbody; 17 paravertebral tubercles, those at midbody surrounded by 9–12 smaller scales; 10 longitudinal rows of tubercles at midbody; body tubercles extend from occiput onto base of tail forming transverse rows, terminating near end of original tail; smaller tubercles in temporal and postocular regions; 25 longitudinal rows of flat, imbricate, ventral scales much larger than dorsal scales between body folds; and eight large, pore-bearing, precloacal scales.
Forelimbs moderately robust, relatively short (FL/SVL 0.13); large, imbricate scales of upper arm and anterior surface of forearm larger than those on posterior surface which are interspersed with large tubercles; palmar scales flat, subimbricate; digits well-developed, inflected at penultimate interphalangeal joints, arising from digital pad; subdigital lamellae wide, transversely expanded forming digital pad; claws well-developed, claw base sheathed by a dorsal and lateral scale; digit I clawless; hind limbs more robust than forelimbs, moderate in length (CL/SVL 0.16), covered anteriorly by large, flat, imbricate scales, dorsally and posteriorly by much smaller flat, juxtaposed scales interspersed with large, subconical tubercles; ventral scales of hind limbs large, flat, imbricate, abruptly contacting small postfemoral scales; femoral pores absent; plantar scales flat, subimbricate; digits well-developed, inflected at penultimate interphalangeal joints, arising from digital pad; distal subdigital lamellae wide, transversely expanded forming digital pad; 15 transverse lamellae beneath digit I, 19 transverse lamellae beneath digit IV; claws well-developed, claw base sheathed by a dorsal and lateral scale; digit I clawless; small amount of webbing between digits I–IV.
Tail original, 139.0 mm in length, tapering to a point; dorsal scales flat, square, bearing transverse rows of six large, subconical tubercles; tubercle rows separated by six or seven transverse rows of dorsal scales; large, paired, median, transversely expanded subcaudal scales; and base of tail bearing hemipenal swellings, each with two conical postcloacal tubercles.
Ground color of all dorsal surfaces yellowish-brown bearing slightly darker faint mottling; top of head bearing small white spots and dark-colored, diffuse Y-marking; thin white nuchal band composed of closely spaced spots extends from one ear opening to other, edged anteriorly by faint, dark-colored nuchal band running between postocular regions; incomplete series of obliquely aligned, small white spots immediately anterior to forelimb insertions parallel the white nuchal spots; series of five rows of transversely arranged, widely separated, small white spots between limb insertions, another between hind limb insertions; white spots on body generally confined to tubercles or border them anteriorly; limbs bearing incomplete, thin white bands; white spots on base of each digit and one or two more on each digit; venter beige, mottled with faint dark-colored markings, weakest in gular region, most dense in subcaudal region; and center of iris gold, transitioning distally to green then turquoise.
Color pattern variation in Gekko hulk sp. nov. is not as extensive as that in G. smithii. The hue and intensity of the ground color changes from day to night and is generally lighter and less bold during evening hours. The description here is of the daytime coloration. The dorsal ground color in life can be dark brownish green, light-green, tan or yellowish brown. There are no thick, dark-colored reticulations on the dorsum although some specimens have faint, brown markings. The white dorsal spots are small and typically confined to the tubercles although in some specimens they may border the tubercles anteriorly or both. The light-colored caudal bands can vary in both width and boldness. In one specimen from Pulau Tioman (LSUHC 5152; Fig.
Raw morphometric data of the type series of Gekko hulk sp. nov. from Peninsular Malaysia. All measurements are in millimeters. Abbreviations are in the Materials and methods.
Locality | LSUHC cat. no. | Sex | SVL | HH | HL | HW | IN | IO | TD | EE | NE | SE | OD | FL | CL | AG | TW |
Pulau Tioman | holotype 6284 | M | 147.5 | 15.9 | 41.7 | 27.2 | 3.7 | 8.2 | 6.3 | 10.8 | 12.2 | 16.9 | 11.0 | 19.8 | 23.8 | 75.8 | 12.8 |
Pulau Tioman | 5399 | F | 149.2 | 14.3 | 42.4 | 27.8 | 4.3 | 8.7 | 6.2 | 11.8 | 13.0 | 18.0 | 10.8 | 20.0 | 22.9 | 77.5 | 15.2 |
Pulau Tioman | 6283 | F | 161.3 | 15.0 | 44.2 | 28.1 | 4.4 | 9.7 | 7.0 | 13.8 | 13.4 | 18.4 | 10.3 | 21.0 | 25.3 | 81.3 | 12.9 |
Pulau Tioman | 5152 | F | 157.2 | 16.7 | 41.1 | 28.6 | 4.4 | 9.0 | 4.5 | 10.9 | 12.6 | 17.9 | 11.4 | 20.2 | 25.6 | 75.7 | 16.7 |
Pulau Tioman | 7263 | F | 141.2 | 15.3 | 39.9 | 26.2 | 3.6 | 9.4 | 5.1 | 10.0 | 13.0 | 16.4 | 11.2 | 14.3 | 22.3 | 67.8 | 13.1 |
Pulau Tioman | 6890 | M | 158.4 | 16.9 | 44.0 | 28.3 | 3.7 | 9.6 | 5.2 | 12.4 | 14.7 | 18.1 | 11.3 | 19.7 | 25.2 | 77.4 | 14.5 |
Pulau Tioman | 5151 | M | 146.5 | 16.3 | 39.8 | 27.2 | 3.6 | 9.7 | 5.0 | 10.0 | 12.2 | 17.3 | 11.3 | 19.6 | 24.2 | 71.1 | 15.3 |
Pulau Tioman | 4681 | M | 149.0 | 16.6 | 42.4 | 25.9 | 3.6 | 8.9 | 5.3 | 12.2 | 12.4 | 16.7 | 10.3 | 18.7 | 24.8 | 70.0 | 13.7 |
Pulau Tioman | 5390 | F | 117.0 | 11.9 | 33.3 | 21.1 | 3.3 | 7.3 | 4.5 | 11.9 | 10.8 | 13.6 | 9.0 | 15.4 | 19.1 | 58.8 | 10.2 |
Pulau Tioman | 7299 | F | 149.9 | 15.9 | 41.3 | 27.1 | 4.3 | 8.3 | 5.3 | 12.9 | 12.3 | 16.9 | 10.5 | 19.3 | 26.2 | 72.7 | 14.6 |
Pulau Tioman | 6260 | F | 121.4 | 12.9 | 35.7 | 20.8 | 3.8 | 7.2 | 5.3 | 10.2 | 10.9 | 15.1 | 9.2 | 16.8 | 21.1 | 60.9 | 11.3 |
Pulau Tioman | 5849 | M | 122.6 | 13.8 | 36.2 | 23.1 | 3.3 | 8.3 | 5.0 | 10.2 | 11.9 | 15.4 | 9.6 | 17.6 | 20.8 | 60.3 | 10.8 |
Pulau Tioman | 7264 | F | 133.4 | 14.5 | 39.5 | 25.0 | 3.7 | 8.4 | 6.1 | 12.4 | 12.5 | 16.7 | 10.9 | 18.8 | 22.8 | 67.5 | 12.1 |
Pulau Tulai | 4694 | F | 127.5 | 12.9 | 36.5 | 23.9 | 3.9 | 7.7 | 6.2 | 9.7 | 10.9 | 14.9 | 10.1 | 15.9 | 20.2 | 58.6 | 11.6 |
Pulau Tulai | 6282 | M | 134.5 | 13.6 | 39.1 | 25.8 | 4.5 | 9.7 | 6.4 | 10.9 | 12.0 | 16.6 | 10.3 | 17.6 | 19.8 | 68.8 | 12.7 |
Pulau Tulai | 7026 | F | 143.0 | 15.3 | 39.4 | 27.3 | 4.4 | 8.7 | 4.8 | 11.0 | 13.1 | 16.7 | 10.3 | 17.9 | 23.1 | 69.2 | 13.6 |
Pulau Tulai | 4697 | F | 138.1 | 14.7 | 38.7 | 27.0 | 4.5 | 8.6 | 4.2 | 11.0 | 11.8 | 15.8 | 10.4 | 18.8 | 21.7 | 66.4 | 12.7 |
Pulau Tulai | 6278 | F | 117.1 | 11.8 | 33.4 | 21.5 | 3.5 | 7.6 | 5.4 | 8.5 | 11.0 | 13.9 | 10.3 | 15.2 | 18.8 | 54.0 | 11.7 |
Pulau Tulai | 5063 | F | 133.0 | 14.7 | 38.3 | 24.4 | 3.9 | 8.6 | 5.2 | 9.8 | 11.0 | 16.1 | 10.2 | 18.1 | 20.8 | 68.0 | 11.7 |
Pulau Tulai | 5062 | M | 139.6 | 16.4 | 39.5 | 27.7 | 4.4 | 9.2 | 5.2 | 11.8 | 12.7 | 17.1 | 11.4 | 17.2 | 21.9 | 65.4 | 12.5 |
Pulau Tulai | 6265 | F | 128.5 | 13.7 | 37.0 | 25.7 | 3.8 | 7.5 | 5.2 | 10.3 | 11.1 | 15.3 | 10.5 | 15.9 | 20.9 | 64.6 | 12.2 |
Pulau Tulai | 6277 | F | 141.1 | 15.6 | 38.8 | 29.2 | 3.8 | 8.8 | 4.1 | 10.7 | 12.1 | 16.5 | 10.7 | 18.2 | 21.9 | 68.6 | 14.1 |
Pulau Tulai | 7025 | M | 136.6 | 15.1 | 38.1 | 25.9 | 3.7 | 8.9 | 6.1 | 10.5 | 11.6 | 16.1 | 10.2 | 17.4 | 21.3 | 62.6 | 12.4 |
Pulau Tulai | 7257 | F | 137.6 | 14.3 | 39.4 | 25.4 | 3.6 | 8.4 | 5.3 | 10.5 | 11.8 | 16.1 | 10.7 | 17.1 | 22.4 | 66.6 | 12.9 |
Pulau Tulai | 3891 | F | 127.1 | 13.4 | 37.2 | 23.5 | 3.6 | 8.1 | 4.8 | 10.6 | 11.1 | 14.9 | 9.9 | 17.8 | 20.6 | 64.1 | 10.5 |
Pulau Tulai | 5059 | F | 121.5 | 14.2 | 36.5 | 23.2 | 3.5 | 9.3 | 5.3 | 10.2 | 10.8 | 15.4 | 10.4 | 17.0 | 20.8 | 60.5 | 11.1 |
Pulau Tulai | 7024 | M | 135.6 | 14.7 | 40.1 | 26.5 | 3.9 | 9.8 | 5.1 | 11.1 | 11.9 | 16.7 | 9.8 | 16.6 | 21.6 | 65.6 | 12.4 |
Pulau Tulai | 5061 | F | 137.3 | 15.8 | 38.5 | 24.8 | 4.0 | 9.0 | 4.9 | 10.9 | 11.6 | 16.2 | 10.1 | 17.2 | 22.2 | 70.5 | 13.2 |
Pulau Tulai | 3990 | M | 148.8 | 16.2 | 42.9 | 26.4 | 4.8 | 9.3 | 5.7 | 12.2 | 13.0 | 17.6 | 10.8 | 17.9 | 22.2 | 67.5 | 11.4 |
Pulau Tulai | 5060 | M | 124.7 | 15.4 | 37.0 | 25.2 | 3.6 | 9.3 | 5.5 | 12.3 | 11.2 | 15.5 | 10.6 | 16.6 | 20.9 | 60.5 | 13.7 |
Pulau Tulai | 7251 | F | 143.9 | 15.0 | 38.8 | 25.4 | 3.9 | 9.2 | 6.2 | 11.1 | 11.6 | 16.6 | 10.3 | 17.5 | 21.8 | 72.1 | 12.0 |
Endau Rompin | 7649 | F | 143.5 | 15.9 | 40.2 | 27.1 | 3.7 | 8.6 | 5.7 | 11.4 | 12.1 | 16.6 | 11.0 | 16.9 | 21.7 | 72.0 | 11.8 |
Endau Rompin | 7694 | F | 143.0 | 14.9 | 39.5 | 25.4 | 3.7 | 8.2 | 4.5 | 10.8 | 11.7 | 16.3 | 9.7 | 18.9 | 22.2 | 70.5 | 13.3 |
Endau Rompin | 7651 | M | 116.4 | 11.9 | 34.3 | 21.2 | 3.4 | 7.2 | 4.1 | 8.9 | 11.2 | 13.5 | 9.7 | 15.3 | 19.1 | 58.4 | 10.5 |
Endau Rompin | 7702 | F | 67.1 | 7.3 | 20.5 | 12.6 | 2.3 | 4.5 | 2.7 | 4.9 | 6.2 | 8.3 | 4.7 | 7.8 | 9.8 | 33.4 | 4.8 |
Endau Rompin | 7650 | F | 61.2 | 6.1 | 19.5 | 11.8 | 2.3 | 4.9 | 2.4 | 4.4 | 5.1 | 7.4 | 5.1 | 8.0 | 9.6 | 32.0 | 4.6 |
Endau Rompin | 7648 | M | 151.5 | 16.8 | 43.0 | 30.7 | 3.5 | 8.0 | 4.3 | 12.9 | 13.7 | 17.8 | 10.5 | 20.0 | 23.5 | 75.3 | 14.3 |
Gunung Lambak | 9959 | M | 56.9 | 6.4 | 17.9 | 11.3 | 1.1 | 3.7 | 2.0 | 3.6 | 4.9 | 6.7 | 5.3 | 7.0 | 9.0 | 27.9 | 3.1 |
Gunung Ledang | 10585 | F | 54.6 | 5.3 | 17.1 | 10.6 | 1.8 | 4.0 | 2.2 | 3.8 | 4.4 | 6.7 | 4.9 | 7.1 | 8.9 | 26.1 | 3.0 |
Meristic data of the type series of Gekko hulk sp. nov. from Peninsular Malaysia. Abbreviations are in the Materials and methods.
Locality | LSUHC cat. no. | Sex | SL | IL | FS | CS | MB | PVT | LRT | VS | TL1 | TL4 | PP | CSP |
Pulau Tioman | holotype 6284 | M | 12 | 10 | 18 | 6 | 110 | 17 | 10 | 25 | 15 | 19 | 8 | 4 |
Pulau Tioman | 5399 | F | 14 | 12 | 21 | 5 | 104 | 18 | 10 | 22 | 17 | 21 | 0 | 4 |
Pulau Tioman | 6283 | F | 14 | 11 | 18 | 6 | 95 | 18 | 10 | 26 | 16 | 20 | 0 | 4 |
Pulau Tioman | 5152 | F | 12 | 10 | 18 | 5 | 91 | 17 | 10 | 21 | 16 | 21 | 0 | 4 |
Pulau Tioman | 7263 | F | 15 | 13 | 18 | 6 | 90 | 16 | 9 | 22 | 18 | 22 | 0 | 2 |
Pulau Tioman | 6890 | M | 14 | 12 | 19 | 7 | 98 | 19 | 11 | 27 | 17 | 21 | 7 | 4 |
Pulau Tioman | 5151 | M | 14 | 12 | 21 | 6 | 109 | 19 | 10 | 23 | 19 | 24 | 8 | 4 |
Pulau Tioman | 4681 | M | 15 | 12 | 20 | 5 | 91 | 20 | 10 | 21 | 16 | 20 | 7 | 6 |
Pulau Tioman | 5390 | F | 13 | 11 | 17 | 7 | 106 | 21 | 11 | 26 | 17 | 21 | 0 | 4 |
Pulau Tioman | 7299 | F | 14 | 11 | 18 | 6 | 105 | 19 | 10 | 27 | 15 | 19 | 0 | 2 |
Pulau Tioman | 6260 | F | 14 | 11 | 21 | 7 | 108 | 21 | 11 | 32 | 17 | 21 | 0 | 4 |
Pulau Tioman | 5849 | M | 13 | 11 | 22 | 7 | 91 | 21 | 11 | 22 | 16 | 21 | 6 | 4 |
Pulau Tioman | 7264 | F | 12 | 9 | 20 | 5 | 93 | 17 | 11 | 26 | 16 | 21 | 0 | 4 |
Pulau Tulai | 4694 | F | 13 | 11 | 19 | 8 | 104 | 17 | 9 | 27 | 19 | 23 | 0 | 2 |
Pulau Tulai | 6282 | M | 13 | 11 | 18 | 6 | 101 | 19 | 10 | 24 | 16 | 20 | 10 | 4 |
Pulau Tulai | 7026 | F | 12 | 10 | 21 | 6 | 98 | 18 | 10 | 25 | 17 | 21 | 0 | 2 |
Pulau Tulai | 4697 | F | 12 | 10 | 17 | 6 | 98 | 18 | 11 | 27 | 16 | 20 | 0 | 4 |
Pulau Tulai | 6278 | F | 13 | 11 | 18 | 6 | 102 | 17 | 10 | 24 | 17 | 21 | 0 | 2 |
Pulau Tulai | 5063 | F | 13 | 11 | 17 | 6 | 99 | 17 | 10 | 25 | 16 | 20 | 0 | 4 |
Pulau Tulai | 5062 | M | 13 | 11 | 20 | 5 | 106 | 18 | 10 | 27 | 15 | 19 | 6 | 2 |
Pulau Tulai | 6265 | F | 13 | 11 | 21 | 7 | 109 | 19 | 10 | 28 | 17 | 21 | 0 | 2 |
Pulau Tulai | 6277 | F | 14 | 12 | 19 | 6 | 106 | 17 | 10 | 26 | 17 | 22 | 0 | 2 |
Pulau Tulai | 7025 | M | 14 | 12 | 23 | 6 | 96 | 17 | 10 | 24 | 19 | 22 | 9 | 2 |
Pulau Tulai | 7257 | F | 13 | 11 | 20 | 7 | 95 | 18 | 9 | 23 | 16 | 20 | 0 | 2 |
Pulau Tulai | 3891 | F | 13 | 11 | 20 | 5 | 102 | 18 | 10 | 26 | 18 | 22 | 0 | 2 |
Pulau Tulai | 5059 | F | 14 | 12 | 22 | 6 | 104 | 18 | 10 | 25 | 16 | 21 | 0 | 2 |
Pulau Tulai | 7024 | M | 14 | 12 | 21 | 6 | 100 | 20 | 10 | 24 | 17 | 21 | 8 | 4 |
Pulau Tulai | 5061 | F | 14 | 11 | 19 | 6 | 98 | 19 | 10 | 27 | 17 | 21 | 0 | 2 |
Pulau Tulai | 3990 | M | 13 | 10 | 22 | 6 | 104 | 21 | 10 | 23 | 17 | 20 | 9 | 3 |
Pulau Tulai | 5060 | M | 13 | 10 | 15 | 7 | 92 | 18 | 10 | 26 | 19 | 23 | 9 | 4 |
Pulau Tulai | 7251 | F | 13 | 11 | 21 | 5 | 101 | 19 | 11 | 29 | 19 | 22 | 0 | 4 |
Endau Rompin | 7649 | F | 14 | 9 | 20 | 8 | 191 | 19 | 11 | 32 | 16 | 20 | 0 | 4 |
Endau Rompin | 7694 | F | 13 | 11 | 18 | 6 | 102 | 18 | 10 | 29 | 15 | 19 | 0 | 4 |
Endau Rompin | 7651 | M | 13 | 11 | 19 | 5 | 105 | 16 | 11 | 26 | 18 | 22 | 10 | 4 |
Endau Rompin | 7702 | F | 10 | 10 | 13 | 4 | 89 | 19 | 9 | 23 | 16 | 20 | 0 | 0 |
Endau Rompin | 7650 | F | 13 | 10 | 20 | 5 | 87 | 19 | 10 | 26 | 15 | 19 | 0 | 2 |
Endau Rompin | 7648 | M | 13 | 10 | 21 | 7 | 95 | 19 | 11 | 30 | 14 | 18 | 11 | 6 |
Gunung Lambak | 9959 | M | 13 | 10 | 18 | 6 | 84 | 18 | 10 | 24 | 16 | 20 | 10 | 0 |
Gunung Ledang | 10585 | F | 14 | 11 | 24 | 5 | 89 | 18 | 10 | 23 | 16 | 21 | 0 | 2 |
Gekko hulk sp. nov. ranges from at least the southeastern corner of southernmost Thailand in the Hala-Bala Wildlife Sanctuary, Narathiwat Province, southward east of the Banjaran Titiwangsa through Peninsular Malaysia to Singapore. It approaches the west coast of Peninsular Malaysia south the Banjaran Titiwangsa at Gunung Ledang, Johor. It is known from the east coast islands of Perhentian Besar and Redang, Terengganu in the north and from the islands of Tulai and Jahat, Johor and Tioman, Pahang in the south (Figs
As noted above, the DFA placed THNHM 01841 from the Hala-Bala Wildlife Sanctuary, Waeng District, Narathiwat, Province, Thailand in Gekko cf. albomaculatus with a 79.5% PsP and in G. smithii with a 20.2% PsP. The DAPC placed it in G. smithii with a 100% PsP. THNHM 01841 bears the color pattern of the northern populations of G. smithii having white ocelli surrounding the tubercles (e.g. Figs
The specific epithet “hulk” is a noun in apposition in reference to ‘‘The Incredible Hulk’’, who is a fictional character and superhero created by Stan Lee and artist Jack Kirby in 1962 and appears in the Marvel Comics publications. When angry, The Incredible Hulk becomes a large, green-skinned, muscular beast possessing great physical strength and a very aggressive temperament—all characteristics of Gekko hulk sp. nov.
Gekko hulk sp. nov. can be separated from other species of the subgenus G. (Gekko) by a number of discrete characters (Table
Much like other species of Gekko (Gekko), G. hulk sp. nov. is an arboreal nocturnal species that is well-established in all types of primary and secondary forests as well as buildings on forest edges. We observed lizards 3–4 m above the ground on the trunks of large trees on Gunung Tebu (Fig.
Microhabitats of Gekko hulk sp. nov. on Peninsular Malaysia. A. Granite boulder microhabitat on Pulau Perhentian Besar, Terengganu. B. Forest microhabitat of the type locality along the Tekek-Juara Trail, Pulau Tioman, Pahang. Strangler Fig. microhabitat along Sungai Mentawak, Pulau Tioman, Pahang. Photographs by L. Lee Grismer.
Several integrative taxonomic analyses of Peninsular Malaysian genera and species across a number of taxonomic groups have consistently found the Banjaran Titiwangsa to be an effective geographic barrier separating sister clades and sister species (
We wish to thank Esther Dondrop and Wendy von Bohemen of Naturalis Biodiversity Center, Leiden for photographs and the loan of specimens. For the loan of specimens, we thank Alan Resetar of the Field Museum of Natural History. We thank Sunchai Makchai of the National Science Museum, Thailand for allowing LLG to examine specimens in his care. We thank Carol Spencer of the Museum of Vertebrate Zoology for processing the loan of specimens. For photographs and literature, we thank Wolfgang Grossmann, Herbert Rösler, and Perry L. Wood Jr. For specimen photographs and translated literature we thank Frank Tillack, Hendrick Muller, and Eric N. Smith. For granting Indonesian research and export permits, JAM thanks RISTEK and the Museum Zoologicum Bogoriense, Indonesian Institute of Sciences (LIPI). Mistar Kamsi (LMU Leuser) provided us with images from Sumatra and Kalimantan. We are grateful to Chien C. Lee, Ian Dugdale, Nick Baker, Kurt H.P. Guek, Steven S.P. Wong, Henrik Bringsøe, Parinya Pawangkhanant, M.A. Muin, Syed A. Rizal, Alan Watson, C.J. Franklin, Andrea Molyneaux and Eric N. Smith for sharing photographs that aided in our work. We are grateful to Mr. Sahir bin Othman, former director of the Department of Wildlife, Jabatan Perlindungan Hidupan Liar dan Taman Negara (PERHILITAN), for permission to conduct fieldwork in the Seribuat Archipelago as part of the La Sierra University Biology 487E field course with personnel from the Universiti Sains Malaysia, Universiti Kabangsaan Malaysia, and the Department of Wildlife and National Parks Malaysia. A research pass (40/200/19SJ.1105) was issued to LLG by the Economic Planning Unit, Prime Minister’s Department. Aaron M. Bauer and Herbert Rösler provided many helpful comments on the manuscript.