Research Article |
Corresponding author: Amanda Kaatz ( aekaatz@gmail.com ) Academic editor: Uwe Fritz
© 2021 Amanda Kaatz, Jesse L. Grismer, 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:
Kaatz A, Grismer JL, Grismer LL (2021) Convergent evolution of karst habitat preference and its ecomorphological correlation in three species of Bent-toed Geckos (Cyrtodactylus) from Peninsular Malaysia. Vertebrate Zoology 71: 367-386. https://doi.org/10.3897/vz.71.e66871
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Abstract
By studying ecomorophology in the context of phylogeny, researchers can parse out similarity due to common ancestry versus that due to convergence. This is especially true among relatively closely related species where both phylogenetic and environmental constraints may be operating simultaneously. We explored these issues among three karst-associated species from two lineages of Cyrtodactylus—the sworderi group from Peninsular Malaysia and the swamp clade from Peninsular Malaysia and western Indonesia of the agamensis group. A stochastic character mapping analysis using five different habitat preferences corroborated a larger previous analysis in recovering a general habitat preference as an ancestral condition for all habitat preferences and a karst habitat preference in C. guakanthanensis and C. gunungsenyumensis of the sworderi group and C. metropolis of the swamp clade as convergently evolved. Multivariate and univariate analyses of 10 morphometric characters revealed that the ecomorphological similarity between C. guakanthanensis and C. gunungsenyumensis of the sworderi group was also convergent. The ecomorphology of C. metropolis of the swamp clade was intermediate between a karst-adapted ecomorphology and a swamp-generalists ecomorphology. Of the 10 morphometric characters employed in this analysis, only three—head length, head width, and forelimb width—showed any signs of phylogenetic signal. Cyrtodactylus metropolis is hypothesized to be a recently refuged swamp-dwelling species that frequented the Batu Caves environments prior to urbanization of the surrounding swamp habitat to which it is now confined.
ecomorph, Gekkonidae, Sundaland, swamp clade, sworderi group
The concept that an animal’s form has evolved in response to the way it navigates its habitat underpins the study of ecomorphology—the intersection of organismal morphology, life history, and adaptation (
Only a few such studies (e.g.,
We ran a Bayesian Evolutionary Analysis by Sampling Trees (BEAST) (see below) using the genetic data set of
Data were taken from all species in the sworderi group (Cyrtodactylus sworderi (n=5), C. gunungsenyumensis (n=10), C. guakanthanensis (n=22), C. tebuensis (n=12), C quadrivirgatus (n=10)) and four available species in the swamp clade (Cyrtodactylus majulah (n=3), C. metropolis (n=6), C. payacola (n=7), and C. pantiensis (n=13)). Measurements were taken on the left side of the body when possible to the nearest 0.1 mm using Mitutoyo dial calipers under a Nikon SMZ 1500 dissecting microscope and follow
Analyses of variance (ANOVA) were conducted on characters (see below) with normalized data and statistically similar variances (i.e., p values ≤ 0.05 in a Levene’s test) to search for the presence of statistically significant mean differences (p < 0.05) among species across the data set. Characters bearing statistical differences were subjected to a TukeyHSD test to ascertain which species pairs differed significantly from each other for those particular characters. Boxplots were generated in order to visualize the range, mean, median, and degree of differences between pairs of species bearing statistically different mean values. All statistical analyses were performed in R [v3.4.3].
The morphospatial clustering of the sampled individuals was visualized using principal component analysis (PCA) from the ADEGENET package in R (
A discriminant analysis of principal components (DAPC) from the ADEGENET package in R was also performed. DAPC relies on scaled data calculated from its own PCA as a prior step to ensure that variables analyzed are not correlated and number fewer than the sample size. Dimension reduction of the DAPC prior to plotting, is accomplished by retaining the first set of PCs that account for approximately 90% of the variation in the data set (
Related species in a clade can resemble one another because of common ancestry (phylogenetic signal), evolutionary convergence (no phylogenetic signal), random evolution, or various degrees of each. To test for the presence or absence of phylogenetic signal in the morphological data set, a mean value for each scaled trait was calculated and tested independently by employing the phylosig () command in the R package phytools (
A BEAST 2 analysis version 2.4.6 (
Habitat preference (general, granite, karst, arboreal, and terrestrial, sec.
Museum acronyms are
The BEAST 2 analysis recovered the same relationships among the species groups as those in the ML and BI analyses of
A Maximum clade credibility BEAST tree based on 1469 bp of ND2 and its flanking tRNAs. BPP support values are at the nodes B Stochastic character map showing probability estimates of the ancestral states of habitat preference at each node in the tree. GenBank accession numbers are appended to each species.
The morphological analyses of the sworderi group demonstrated that the karst-associated species C. gunungsenyumensis and C. guakanthanensis and the habitat generalists C. quadrivirgatus, C. sworderi, and C. tebuensis only overlapped slightly in the PCA and did not overlap in the DAPC (Fig.
Species pairs bearing statistically different mean values based on ANOVA and TukeyHSD tests of adjusted morphometric characters and their adjusted p values.
p adj | |
---|---|
axilla-groin length | |
quadrivirgatus-sworderi | 0.0087833 |
forelimb length | |
guakanthanensis-majulah | 0.0188355 |
guakanthanensis-pantiensis | 0.0000102 |
guakanthanensis-payacola | 0.0000163 |
guakanthanensis-quadrivirgatus | 0.0000503 |
guakanthanensis-sworderi | 0.0059021 |
guakanthanensis-tebuensis | 0.0190158 |
gunungsenyumensis-majulah | 0.0445273 |
gunungsenyumensis-pantiensis | 0.0006199 |
gunungsenyumensis-payacola | 0.0003075 |
gunungsenyumensis-quadrivirgatus | 0.0013346 |
gunungsenyumensis-sworderi | 0.0243946 |
metropolis-pantiensis | 0.0454672 |
metropolis-payacola | 0.0145310 |
forelimb width | |
pantiensis-quadrivirgatus | 0.0049576 |
hindlimb length | |
guakanthanensis-quadrivirgatus | 0.0000006 |
guakanthanensis-sworderi | 0.0000007 |
guakanthanensis-tebuensis | 0.0003022 |
gunungsenyumensis-quadrivirgatus | 0.0020836 |
gunungsenyumensis-sworderi | 0.0003406 |
sworderi-metropolis | 0.0523656 |
sworderi-pantiensis | 0.0070785 |
head length | |
guakanthanensis-majulah | 0.0000340 |
guakanthanensis-pantiensis | 0.0000045 |
guakanthanensis-quadrivirgatus | 0.0003565 |
guakanthanensis-sworderi | 0.0211978 |
gunungsenyumensis-majulah | 0.0014134 |
gunungsenyumensis-pantiensis | 0.0075965 |
tebuensis-majulah | 0.0002370 |
tebuensis-pantiensis | 0.0004053 |
tebuensis-quadrivirgatus | 0.0080412 |
head width | |
guakanthanensis-metropolis | 0.0532702 |
guakanthanensis-pantiensis | 0.0177667 |
guakanthanensis-payacola | 0.0001369 |
gunungsenyumensis-payacola | 0.0021279 |
tebuensis-pantiensis | 0.0495110 |
tebuensis-payacola | 0.0005296 |
pelvic height | |
metropolis-guakanthanensis | 0.0213616 |
metropolis-pantiensis | 0.0043248 |
metropolis-payacola | 0.0011999 |
metropolis-sworderi | 0.0003470 |
metropolis-tebuensis | 0.0033463 |
pelvic width | |
guakanthanensis-majulah | 0.0068838 |
guakanthanensis-pantiensis | 0.0000032 |
guakanthanensis-tebuensis | 0.0104445 |
gunungsenyumensis-majulah | 0.0052087 |
gunungsenyumensis-pantiensis | 0.0000216 |
gunungsenyumensis-tebuensis | 0.0136552 |
sworderi-majulah | 0.0525812 |
pantiensis-metropolis | 0.0276788 |
pantiensis-quadrivirgatus | 0.0176241 |
pantiensis-sworderi | 0.0081379 |
Summary statistics and principal component scores for the sworderi group’s scaled mensural data.
PC1 | PC2 | PC3 | PC4 | PC5 | PC6 | PC7 | PC8 | PC9 | PC10 | |
Standard deviation | 1.750 | 1.277 | 1.169 | 1.035 | 0.949 | 0.758 | 0.721 | 0.665 | 0.481 | 0.445 |
Proportion of Variance | 0.306 | 0.163 | 0.137 | 0.107 | 0.090 | 0.057 | 0.052 | 0.044 | 0.023 | 0.020 |
Cumulative Proportion | 0.306 | 0.470 | 0.606 | 0.713 | 0.803 | 0.861 | 0.913 | 0.957 | 0.980 | 1.000 |
Eigen | 3.064 | 1.631 | 1.367 | 1.071 | 0.900 | 0.575 | 0.520 | 0.442 | 0.232 | 0.198 |
PW | 0.012 | –0.451 | 0.063 | –0.267 | 0.788 | –0.087 | 0.020 | 0.067 | 0.280 | –0.095 |
PH | 0.058 | –0.304 | –0.647 | –0.079 | –0.274 | –0.238 | 0.397 | 0.397 | 0.171 | 0.049 |
AXG | 0.026 | 0.530 | –0.397 | 0.167 | 0.414 | –0.241 | 0.218 | –0.355 | 0.052 | 0.357 |
HL | –0.451 | –0.080 | –0.059 | 0.232 | 0.178 | –0.344 | –0.364 | 0.410 | –0.474 | 0.238 |
HW | –0.236 | –0.210 | –0.566 | 0.064 | 0.075 | 0.635 | –0.299 | –0.259 | –0.091 | –0.051 |
SNT | –0.475 | 0.226 | –0.113 | 0.070 | –0.079 | –0.314 | –0.193 | –0.086 | 0.429 | –0.610 |
ED | –0.104 | 0.130 | –0.118 | –0.886 | –0.127 | –0.182 | –0.191 | –0.177 | –0.217 | 0.094 |
HDL | –0.456 | 0.041 | 0.116 | –0.080 | 0.111 | 0.176 | 0.696 | –0.033 | –0.411 | –0.270 |
FLW | –0.262 | –0.527 | 0.143 | 0.144 | –0.234 | –0.301 | 0.093 | –0.607 | 0.072 | 0.292 |
FLL | –0.472 | 0.159 | 0.188 | –0.139 | –0.069 | 0.321 | 0.072 | 0.263 | 0.502 | 0.513 |
Similar to the karst-adapted species in the sworderi group, the PCA and DAPC of the swamp clade species indicate that the karst-associated Cyrtodactylus metropolis is well-differentiated from all other members of that clade (Fig.
Summary statistics and principal component scores for the swamp clade’s scaled mensural data.
PC1 | PC2 | PC3 | PC4 | PC5 | PC6 | PC7 | PC8 | PC9 | PC10 | |
Standard deviation | 1.859 | 1.251 | 1.134 | 1.069 | 0.851 | 0.762 | 0.664 | 0.618 | 0.554 | 0.339 |
Proportion of Variance | 0.346 | 0.157 | 0.129 | 0.114 | 0.072 | 0.058 | 0.044 | 0.038 | 0.031 | 0.011 |
Cumulative Proportion | 0.346 | 0.502 | 0.631 | 0.745 | 0.818 | 0.876 | 0.920 | 0.958 | 0.989 | 1.000 |
Eigen | 3.458 | 1.565 | 1.286 | 1.143 | 0.724 | 0.581 | 0.440 | 0.382 | 0.307 | 0.115 |
PW | –0.211 | 0.382 | –0.313 | 0.491 | –0.305 | 0.082 | –0.398 | 0.438 | –0.013 | 0.143 |
PH | –0.284 | 0.382 | –0.400 | –0.147 | 0.205 | 0.426 | 0.120 | –0.411 | –0.425 | –0.062 |
AXG | 0.144 | 0.505 | 0.445 | 0.243 | 0.482 | –0.061 | 0.174 | 0.033 | –0.021 | 0.446 |
HL | –0.342 | –0.163 | –0.083 | –0.377 | 0.523 | –0.288 | –0.380 | 0.380 | –0.234 | 0.086 |
HW | –0.392 | 0.156 | 0.357 | 0.320 | 0.009 | –0.381 | –0.045 | –0.194 | –0.157 | –0.619 |
SNT | –0.399 | –0.278 | 0.231 | 0.042 | –0.396 | –0.064 | 0.325 | 0.036 | –0.480 | 0.457 |
ED | –0.405 | 0.157 | 0.176 | –0.246 | –0.001 | 0.352 | 0.425 | 0.474 | 0.376 | –0.222 |
HDL | –0.448 | 0.040 | 0.155 | –0.148 | –0.096 | 0.022 | –0.368 | –0.473 | 0.521 | 0.334 |
FLW | –0.232 | –0.219 | –0.509 | 0.382 | 0.291 | –0.322 | 0.436 | –0.081 | 0.308 | 0.113 |
FLL | –0.083 | –0.501 | 0.205 | 0.451 | 0.327 | 0.589 | –0.195 | –0.006 | –0.037 | –0.055 |
To test whether or not the karst-adapted species—Cyrtodactylus metropolis of the swamp clade and C. gunungsenyumensis and C. guakanthanensis of the sworderi group converged on each other morphologically, a PCA and DAPC were performed on a concatenated data set composed of all the species of each group. The analyses showed that the karst-adapted C. gunungsenyumensis and C. guakanthanensis grouped together as before and that the swamp-adapted species (C. majulah, C. payacola, and C. pantiensis) and habitat generalist (C. quadrivirgatus, C. sworderi, and C. tebuensis) also grouped together but separately from the karst-adapted species (Fig.
Summary statistics and principal component scores for the scaled mensural concatenated data.
PC1 | PC2 | PC3 | PC4 | PC5 | PC6 | PC7 | PC8 | PC9 | PC10 | |
Standard deviation | 1.744 | 1.277 | 1.178 | 0.950 | 0.901 | 0.754 | 0.698 | 0.685 | 0.609 | 0.573 |
Proportion of Variance | 0.304 | 0.163 | 0.139 | 0.090 | 0.081 | 0.057 | 0.049 | 0.047 | 0.037 | 0.033 |
Cumulative Proportion | 0.304 | 0.467 | 0.606 | 0.696 | 0.777 | 0.834 | 0.883 | 0.930 | 0.967 | 1.000 |
eigen | 3.040 | 1.632 | 1.389 | 0.902 | 0.812 | 0.569 | 0.488 | 0.469 | 0.371 | 0.328 |
PW | 0.106 | –0.526 | 0.144 | –0.592 | 0.058 | –0.431 | 0.134 | –0.217 | 0.200 | –0.216 |
PH | –0.078 | –0.314 | 0.628 | 0.238 | –0.154 | 0.392 | –0.381 | –0.124 | 0.248 | –0.213 |
AXG | 0.052 | 0.465 | 0.331 | –0.576 | –0.323 | 0.242 | 0.138 | 0.211 | 0.249 | 0.227 |
HL | –0.442 | –0.022 | 0.036 | 0.233 | –0.201 | –0.003 | 0.566 | –0.509 | 0.243 | 0.264 |
HW | –0.370 | 0.058 | 0.327 | 0.068 | –0.400 | –0.591 | –0.211 | 0.174 | –0.388 | 0.116 |
SNT | –0.479 | 0.094 | –0.058 | –0.066 | 0.003 | 0.129 | 0.304 | 0.313 | –0.035 | –0.739 |
ED | –0.272 | –0.009 | 0.404 | –0.006 | 0.769 | –0.045 | 0.124 | 0.256 | 0.009 | 0.298 |
HDL | –0.403 | –0.106 | –0.158 | –0.440 | 0.073 | 0.392 | –0.269 | –0.351 | –0.485 | 0.132 |
FLW | –0.135 | –0.601 | –0.243 | –0.004 | –0.256 | 0.205 | 0.109 | 0.562 | 0.080 | 0.345 |
FLL | –0.405 | 0.154 | –0.342 | –0.069 | 0.081 | –0.195 | –0.510 | 0.010 | 0.619 | 0.046 |
K and λ statistics for the mean values of each trait mirrored one another and indicated there was no phylogenetic signal in PW, PH, AXG, SNT, ED, HDL, and FLL but both statistics indicated that HL showed phylogenetic signal in both groups and FLW and HW were phylogenetically influenced in only the swamp clade (Table
Results from the phylosig analysis testing for the presence or absence of phylogenetic signal in the morphological data set of the swamp clade and sworderi group.
Swamp clade | ||||||
Character | lambda | p lambda | Phylogenetic signal | K | p K | Phylogenetic signal |
PW | 1.11E-01 | 0.768 | no | 0.4541 | 0.563 | no |
PH | 6.61E-05 | 1.000 | no | 0.3069 | 0.896 | no |
AXG | 6.61E-05 | 1.000 | no | 0.7180 | 0.115 | no |
HL | 1.00E+00 | 0.104 | yes | 1.1424 | 0.013 | yes |
HW | 1.00E+00 | 0.005 | yes | 1.9345 | 0.001 | yes |
SNT | 6.61E-05 | 1.000 | no | 0.5563 | 0.338 | no |
ED | 2.91E-01 | 0.575 | no | 0.6460 | 0.191 | no |
HDL | 6.61E-05 | 1.000 | no | 0.5285 | 0.404 | no |
FLW | 1.00E+00 | 0.132 | yes | 1.0381 | 0.014 | yes, but weak |
FLL | 9.24E-04 | 0.998 | no | 0.5677 | 0.33 | no |
sworderi group | ||||||
PW | 2.00E-01 | 0.699 | no | 0.6565 | 0.190 | no |
PH | 6.61E-05 | 1.000 | no | 0.3311 | 0.853 | no |
AXG | 6.61E-05 | 1.000 | no | 0.5786 | 0.305 | no |
HL | 1.00E+00 | 0.085 | yes | 1.1759 | 0.005 | yes |
HW | 7.60E-01 | 0.097 | no | 0.9712 | 0.045 | no |
SNT | 6.61E-05 | 1.000 | no | 0.4962 | 0.450 | no |
ED | 0.09342955 | 0.852 | no | 0.5509 | 0.399 | no |
HDL | 6.61E-05 | 1.000 | no | 0.5231 | 0.427 | no |
FLW | 6.61E-05 | 1.000 | no | 0.5231 | 0.389 | no |
FLL | 6.61E-05 | 1.000 | no | 0.5464 | 0.348 | no |
The analyses demonstrated that the evolution of a karst-adapted ecomorphology evolved independently in three species of Cyrtodactylus from Peninsular Malaysia from two unrelated lineages, and that it co-evolved with a karst habitat preference.
Interestingly however, in the concatenated analysis, Cyrtodactylus metropolis does not unequivocally group with the karst ecomorphs of the sworderi group nor with the swamp-dwellers or generalist of the swamp clade but instead, is somewhat intermediate among them—although distinctively closer to the karst-adapted species (Fig.
Color pattern of karst-adapted species showing the wide, darkly colored, straight-edged interspaces on the body in A Cyrtodactylus guakanthanensis of the sworderi group from Peninsular Malaysia B C. metropolis of the swamp clade from Peninsular Malaysia C C. gunungsenyumensis of the sworderi group from Peninsular Malaysia.
Color pattern of the swamp-adapted and habitat generalist taxa showing the irregularly shaped broken bands, blotches, or stripes on the body in A Cyrtodactylus majulah of the swamp clade from Singapore B C. payacola of the swamp clade from Peninsular Malaysia C C. quadrivirgatus from the sworderi group from Peninsular Malaysia and Sumatra D C. jarakensis of the swamp clade from Peninsular Malaysia E C. pantiensis of the swamp clade from Peninsular Malaysia F C. tebuensis of the sworderi group from Peninsular Malaysia G C. sworderi of the sworderi group from Peninsular Malaysia H C. rosichonarieforum of the swamp clade from Natuna Besar, Indonesia.
We thank Perry L. Wood, Jr. for sequencing and the College of Arts and Sciences for funding for field work. We thanks Evan S. H. Quah and two anonymous reviewers for helpful comments on the manuscript.
Species | PW | PH | AXG | HL | HW | SNT | ED | HDL | FLW | FLL | |
---|---|---|---|---|---|---|---|---|---|---|---|
|
C. sworderi | 1.786 | 1.752 | 3.251 | 2.941 | 2.366 | 1.935 | 1.479 | 3.380 | 0.930 | 3.183 |
|
C. sworderi | 1.827 | 1.948 | 3.259 | 2.878 | 2.446 | 1.880 | 1.480 | 3.528 | 0.892 | 3.229 |
|
C. sworderi | 1.768 | 1.865 | 3.247 | 2.867 | 2.427 | 1.938 | 1.510 | 3.354 | 0.770 | 3.175 |
|
C. sworderi | 1.786 | 1.826 | 3.287 | 2.940 | 2.395 | 1.937 | 1.441 | 3.421 | 0.990 | 3.140 |
|
C. sworderi | 1.725 | 1.878 | 3.332 | 2.913 | 2.436 | 1.990 | 1.415 | 3.313 | 0.961 | 3.009 |
|
C. gunungsenyumensis | 1.745 | 1.815 | 3.343 | 2.934 | 2.478 | 2.007 | 1.425 | 3.467 | 0.968 | 3.226 |
|
C. gunungsenyumensis | 1.724 | 1.749 | 3.338 | 2.926 | 2.397 | 1.993 | 1.467 | 3.522 | 0.908 | 3.246 |
|
C. gunungsenyumensis | 1.795 | 1.751 | 3.303 | 2.972 | 2.460 | 2.025 | 1.415 | 3.487 | 0.993 | 3.320 |
|
C. gunungsenyumensis | 1.758 | 1.737 | 3.406 | 2.938 | 2.422 | 1.982 | 1.376 | 3.420 | 0.767 | 3.223 |
|
C. gunungsenyumensis | 1.752 | 1.782 | 3.333 | 2.941 | 2.430 | 2.040 | 1.497 | 3.572 | 0.889 | 3.201 |
|
C. gunungsenyumensis | 1.802 | 1.751 | 3.380 | 2.962 | 2.447 | 2.010 | 1.565 | 3.501 | 0.745 | 3.213 |
|
C. gunungsenyumensis | 1.828 | 1.732 | 3.265 | 2.983 | 2.427 | 2.032 | 1.457 | 3.638 | 1.264 | 3.334 |
|
C. gunungsenyumensis | 1.737 | 1.696 | 3.338 | 2.966 | 2.417 | 2.019 | 1.443 | 3.605 | 0.712 | 3.409 |
|
C. gunungsenyumensis | 1.685 | 1.792 | 3.359 | 2.942 | 2.453 | 1.995 | 1.549 | 3.571 | 0.786 | 3.306 |
|
C. gunungsenyumensis | 1.789 | 1.688 | 3.345 | 2.931 | 2.386 | 1.981 | 1.535 | 3.479 | 0.724 | 3.262 |
|
C. guakanthanensis | 1.810 | 1.792 | 3.261 | 2.997 | 2.457 | 2.058 | 1.556 | 3.537 | 1.043 | 3.484 |
|
C. guakanthanensis | 1.773 | 1.810 | 3.392 | 2.950 | 2.431 | 1.993 | 1.569 | 3.502 | 0.906 | 3.250 |
|
C. guakanthanensis | 1.807 | 1.830 | 3.312 | 2.965 | 2.512 | 2.034 | 1.563 | 3.574 | 1.054 | 3.244 |
|
C. guakanthanensis | 1.844 | 1.744 | 3.324 | 2.966 | 2.405 | 2.022 | 1.367 | 3.583 | 1.039 | 3.292 |
|
C. guakanthanensis | 1.866 | 1.820 | 3.274 | 2.939 | 2.483 | 1.980 | 1.411 | 3.623 | 1.005 | 3.254 |
|
C. guakanthanensis | 1.809 | 1.830 | 3.363 | 2.957 | 2.465 | 2.073 | 1.475 | 3.532 | 0.910 | 3.211 |
|
C. guakanthanensis | 1.725 | 1.779 | 3.327 | 2.973 | 2.443 | 2.019 | 1.449 | 3.525 | 1.053 | 3.302 |
|
C. guakanthanensis | 1.766 | 1.642 | 3.305 | 2.970 | 2.388 | 2.016 | 1.378 | 3.550 | 0.898 | 3.273 |
|
C. guakanthanensis | 1.781 | 1.756 | 3.299 | 2.963 | 2.394 | 2.019 | 1.463 | 3.485 | 1.076 | 3.249 |
|
C. guakanthanensis | 1.722 | 1.727 | 3.336 | 2.967 | 2.386 | 2.008 | 1.457 | 3.499 | 0.861 | 3.201 |
|
C. guakanthanensis | 1.751 | 1.739 | 3.349 | 2.996 | 2.401 | 2.035 | 1.399 | 3.506 | 1.170 | 3.242 |
|
C. guakanthanensis | 1.763 | 1.746 | 3.384 | 2.970 | 2.426 | 2.019 | 1.346 | 3.603 | 0.854 | 3.286 |
|
C. guakanthanensis | 1.738 | 1.794 | 3.358 | 2.944 | 2.418 | 2.033 | 1.482 | 3.582 | 0.789 | 3.296 |
|
C. guakanthanensis | 1.757 | 1.782 | 3.398 | 2.953 | 2.406 | 2.007 | 1.552 | 3.529 | 0.958 | 3.304 |
|
C. guakanthanensis | 1.782 | 1.815 | 3.326 | 3.009 | 2.457 | 2.008 | 1.442 | 3.556 | 0.820 | 3.273 |
|
C. guakanthanensis | 1.653 | 1.824 | 3.424 | 2.967 | 2.456 | 2.002 | 1.426 | 3.546 | 0.826 | 3.244 |
|
C. guakanthanensis | 1.756 | 1.751 | 3.399 | 2.924 | 2.392 | 1.997 | 1.485 | 3.593 | 0.808 | 3.291 |
|
C. guakanthanensis | 1.750 | 1.799 | 3.349 | 2.950 | 2.456 | 2.014 | 1.420 | 3.545 | 0.856 | 3.264 |
|
C. guakanthanensis | 1.775 | 1.778 | 3.354 | 2.978 | 2.460 | 2.030 | 1.474 | 3.552 | 0.909 | 3.305 |
|
C. guakanthanensis | 1.839 | 1.794 | 3.374 | 2.952 | 2.440 | 2.045 | 1.536 | 3.556 | 0.920 | 3.255 |
|
C. guakanthanensis | 1.780 | 1.767 | 3.337 | 2.973 | 2.449 | 2.023 | 1.515 | 3.587 | 1.002 | 3.304 |
|
C. guakanthanensis | 1.802 | 1.802 | 3.370 | 2.925 | 2.435 | 2.002 | 1.427 | 3.574 | 1.038 | 3.275 |
|
C. tebuensis | 1.802 | 1.828 | 3.364 | 2.958 | 2.514 | 1.933 | 1.441 | 3.423 | 1.028 | 3.211 |
|
C. tebuensis | 1.936 | 1.781 | 3.283 | 2.979 | 2.366 | 1.995 | 1.475 | 3.485 | 0.962 | 3.219 |
|
C. tebuensis | 1.807 | 1.844 | 3.332 | 2.951 | 2.427 | 1.944 | 1.282 | 3.481 | 0.894 | 3.133 |
|
C. tebuensis | 1.874 | 1.761 | 3.291 | 2.946 | 2.488 | 1.913 | 1.382 | 3.455 | 1.121 | 3.124 |
|
C. tebuensis | 1.830 | 1.772 | 3.279 | 2.921 | 2.356 | 1.912 | 1.445 | 3.570 | 0.854 | 3.245 |
|
C. tebuensis | 1.735 | 1.839 | 3.301 | 2.949 | 2.421 | 1.948 | 1.377 | 3.446 | 0.975 | 3.126 |
|
C. tebuensis | 1.881 | 1.722 | 3.269 | 2.956 | 2.433 | 1.877 | 1.391 | 3.463 | 0.874 | 3.159 |
|
C. tebuensis | 1.843 | 1.851 | 3.360 | 2.947 | 2.433 | 1.988 | 1.370 | 3.463 | 0.963 | 3.207 |
|
C. tebuensis | 1.957 | 1.790 | 3.328 | 2.955 | 2.421 | 1.960 | 1.514 | 3.503 | 1.011 | 3.251 |
|
C. tebuensis | 1.714 | 1.864 | 3.338 | 2.975 | 2.411 | 1.977 | 1.428 | 3.343 | 0.624 | 3.198 |
|
C. tebuensis | 1.957 | 1.759 | 3.391 | 2.947 | 2.464 | 1.962 | 1.335 | 3.457 | 0.873 | 3.199 |
|
C. tebuensis | 1.921 | 1.840 | 3.336 | 3.012 | 2.494 | 2.013 | 1.476 | 3.525 | 0.984 | 3.236 |
|
C. quadrivirgatus | 1.812 | 1.717 | 3.282 | 2.888 | 2.437 | 1.913 | 1.554 | 3.381 | 0.754 | 3.135 |
|
C. quadrivirgatus | 1.817 | 1.833 | 3.357 | 2.943 | 2.381 | 1.942 | 1.481 | 3.542 | 1.191 | 3.117 |
|
C. quadrivirgatus | 1.896 | 1.687 | 3.381 | 2.924 | 2.375 | 1.956 | 1.499 | 3.454 | 0.668 | 3.207 |
|
C. quadrivirgatus | 1.695 | 1.759 | 3.399 | 2.888 | 2.376 | 1.909 | 1.332 | 3.381 | 0.678 | 3.147 |
|
C. quadrivirgatus | 1.825 | 1.790 | 3.372 | 2.888 | 2.364 | 1.914 | 1.451 | 3.428 | 0.638 | 3.088 |
|
C. quadrivirgatus | 1.828 | 1.827 | 3.314 | 2.894 | 2.398 | 1.905 | 1.507 | 3.470 | 0.858 | 3.124 |
|
C. quadrivirgatus | 1.756 | 1.599 | 3.370 | 2.862 | 2.406 | 1.928 | 1.428 | 3.414 | 0.800 | 3.167 |
|
C. quadrivirgatus | 1.763 | 1.830 | 3.389 | 2.897 | 2.402 | 1.964 | 1.450 | 3.369 | 0.888 | 3.196 |
|
C. quadrivirgatus | 1.832 | 1.800 | 3.342 | 2.923 | 2.356 | 1.951 | 1.482 | 3.437 | 0.988 | 3.139 |
|
C. quadrivirgatus | 1.862 | 1.869 | 3.523 | 2.950 | 2.467 | 1.981 | 1.523 | 3.460 | 0.555 | 3.118 |
|
C. majulah | 1.911 | 1.743 | 3.389 | 2.849 | 2.357 | 1.927 | 1.382 | 3.464 | 0.866 | 3.144 |
|
C. majulah | 1.922 | 1.765 | 3.373 | 2.868 | 2.373 | 1.941 | 1.358 | 3.454 | 0.676 | 3.119 |
|
C. majulah | 1.917 | 1.755 | 3.381 | 2.858 | 2.365 | 1.934 | 1.370 | 3.459 | 0.776 | 3.131 |
|
C. payacola | 1.677 | 1.723 | 3.353 | 2.894 | 2.284 | 1.881 | 1.366 | 3.413 | 0.933 | 3.093 |
|
C. payacola | 1.922 | 1.840 | 3.390 | 2.991 | 2.423 | 1.945 | 1.452 | 3.480 | 0.988 | 3.153 |
|
C. payacola | 1.786 | 1.716 | 3.290 | 2.832 | 2.292 | 1.880 | 1.245 | 3.403 | 0.880 | 3.025 |
|
C. payacola | 1.833 | 1.842 | 3.433 | 2.879 | 2.378 | 1.918 | 1.390 | 3.513 | 0.884 | 3.217 |
|
C. payacola | 1.874 | 1.936 | 3.381 | 3.025 | 2.412 | 2.001 | 1.574 | 3.548 | 1.067 | 3.211 |
|
C. payacola | 1.838 | 1.884 | 3.243 | 2.979 | 2.329 | 1.958 | 1.471 | 3.477 | 0.998 | 3.113 |
|
C. payacola | 1.904 | 1.857 | 3.274 | 2.894 | 2.374 | 2.014 | 1.404 | 3.515 | 1.085 | 3.009 |
|
C. metropolis | 1.813 | 1.574 | 3.296 | 2.899 | 2.380 | 2.008 | 1.353 | 3.464 | 1.092 | 3.413 |
|
C. metropolis | 1.816 | 1.643 | 3.284 | 2.919 | 2.388 | 2.007 | 1.392 | 3.499 | 1.029 | 3.308 |
|
C. metropolis | 1.816 | 1.746 | 3.262 | 2.906 | 2.352 | 1.936 | 1.274 | 3.480 | 1.138 | 3.252 |
|
C. metropolis | 1.813 | 1.748 | 3.351 | 2.894 | 2.428 | 1.989 | 1.499 | 3.504 | 1.092 | 3.243 |
|
C. metropolis | 1.776 | 1.681 | 3.282 | 2.975 | 2.382 | 2.029 | 1.502 | 3.547 | 0.710 | 3.082 |
|
C. metropolis | 1.752 | 1.689 | 3.329 | 2.924 | 2.358 | 1.973 | 1.428 | 3.468 | 0.945 | 3.215 |
|
C. pantiensis | 1.820 | 1.796 | 3.368 | 2.926 | 2.357 | 1.917 | 1.442 | 3.495 | 0.967 | 3.192 |
|
C. pantiensis | 1.888 | 1.815 | 3.348 | 2.877 | 2.382 | 1.994 | 1.411 | 3.444 | 1.060 | 3.223 |
|
C. pantiensis | 1.829 | 1.738 | 3.397 | 2.836 | 2.346 | 1.944 | 1.557 | 3.481 | 0.783 | 3.189 |
|
C. pantiensis | 1.871 | 1.838 | 3.284 | 2.937 | 2.438 | 1.988 | 1.452 | 3.523 | 1.032 | 3.144 |
|
C. pantiensis | 1.942 | 1.821 | 3.396 | 2.878 | 2.374 | 1.937 | 1.271 | 3.458 | 1.199 | 3.127 |
|
C. pantiensis | 1.849 | 1.714 | 3.389 | 2.836 | 2.385 | 1.916 | 1.380 | 3.505 | 0.880 | 3.087 |
|
C. pantiensis | 1.913 | 1.641 | 3.381 | 2.903 | 2.413 | 1.937 | 1.355 | 3.503 | 0.960 | 3.186 |
|
C. pantiensis | 2.110 | 1.825 | 3.279 | 2.893 | 2.372 | 1.981 | 1.568 | 3.485 | 1.271 | 3.174 |
|
C. pantiensis | 1.841 | 1.908 | 3.319 | 2.896 | 2.442 | 2.063 | 1.588 | 3.552 | 1.042 | 3.161 |
|
C. pantiensis | 1.918 | 1.907 | 3.325 | 2.879 | 2.398 | 1.941 | 1.469 | 3.513 | 0.913 | 3.185 |
|
C. pantiensis | 1.854 | 1.753 | 3.405 | 2.920 | 2.396 | 1.919 | 1.438 | 3.450 | 1.001 | 3.092 |
|
C. pantiensis | 1.905 | 1.765 | 3.364 | 2.934 | 2.408 | 1.944 | 1.503 | 3.520 | 1.068 | 2.899 |
|
C. pantiensis | 1.933 | 1.886 | 3.232 | 2.958 | 2.349 | 1.930 | 1.460 | 3.555 | 1.116 | 3.233 |
Summary statistics of the scaled morphometric data of swamp clade and sworderi group species. SD = standard deviation, n = sample size.
Cyrtodactylus sworderi | Cyrtodactylus gunungsenyumensis | Cyrtodactylus guakanthanensis | Cyrtodactylus tebuensis | Cyrtodactylus quadrivirgatus | Cyrtodactylus majulah | Cyrtodactylus payacola | Cyrtodactylus metropolis | Cyrtodactylus pantiensis | |
---|---|---|---|---|---|---|---|---|---|
pelvic width (PW) | |||||||||
Mean | 1.78 | 1.76 | 1.77 | 1.85 | 1.81 | 1.92 | 1.83 | 1.8 | 1.9 |
SD(±) | 0.037 | 0.042 | 0.046 | 0.081 | 0.057 | 0.006 | 0.083 | 0.027 | 0.075 |
Range | 1.73–1.83 | 1.69–1.83 | 1.65–1.87 | 1.71–1.96 | 1.69–1.90 | 1.91–1.92 | 1.68–1.92 | 1.75–1.82 | 1.82–2.11 |
n | 5 | 10 | 22 | 12 | 10 | 3 | 7 | 6 | 13 |
pelvic height (PH) | |||||||||
Mean | 1.85 | 1.75 | 1.78 | 1.8 | 1.77 | 1.75 | 1.83 | 1.68 | 1.8 |
SD(±) | 0.072 | 0.04 | 0.043 | 0.046 | 0.082 | 0.011 | 0.081 | 0.066 | 0.078 |
Range | 1.75–1.95 | 1.69–1.81 | 1.64–1.83 | 1.72–1.86 | 1.60–1.87 | 1.74–1.77 | 1.72–1.94 | 1.57–1.75 | 1.64–1.91 |
n | 5 | 10 | 22 | 12 | 10 | 3 | 7 | 6 | 13 |
axilla-groin length (AXG) | |||||||||
Mean | 3.28 | 3.34 | 3.35 | 3.32 | 3.37 | 3.38 | 3.34 | 3.3 | 3.35 |
SD(±) | 0.035 | 0.038 | 0.041 | 0.038 | 0.064 | 0.008 | 0.07 | 0.033 | 0.054 |
Range | 3.25–3.33 | 3.27–3.41 | 3.26–3.42 | 3.27–3.39 | 3.28–3.52 | 3.37–3.39 | 3.24–2.42 | 3.26–3.35 | 3.23–3.40 |
n | 5 | 10 | 22 | 12 | 10 | 3 | 7 | 6 | 13 |
head length (HL) | |||||||||
Mean | 2.91 | 2.95 | 2.96 | 2.96 | 2.91 | 2.86 | 2.93 | 2.92 | 2.9 |
SD(±) | 0.035 | 0.02 | 0.021 | 0.023 | 0.028 | 0.01 | 0.071 | 0.03 | 0.037 |
Range | 2.87–2.94 | 2.93–2.98 | 2.92–3.01 | 2.92–3.01 | 2.86–2.95 | 2.85–2.87 | 2.83–3.02 | 2.89–2.98 | 2.84–2.96 |
n | 5 | 10 | 22 | 12 | 10 | 3 | 7 | 6 | 13 |
head width (HW) | |||||||||
Mean | 2.41 | 2.43 | 2.43 | 2.44 | 2.4 | 2.37 | 2.36 | 2.38 | 2.39 |
SD(±) | 0.033 | 0.029 | 0.033 | 0.048 | 0.034 | 0.008 | 0.056 | 0.027 | 0.031 |
Range | 2.37–2.45 | 2.39–2.48 | 2.39–2.51 | 2.36–2.51 | 2.36–2.47 | 2.36–2.37 | 2.28–2.42 | 2.35–2.43 | 2.35–2.44 |
n | 5 | 10 | 22 | 12 | 10 | 3 | 7 | 6 | 13 |
snout length (SNT) | |||||||||
Mean | 1.94 | 2.01 | 2.02 | 1.95 | 1.94 | 1.93 | 1.94 | 1.99 | 1.95 |
SD(±) | 0.039 | 0.021 | 0.021 | 0.04 | 0.026 | 0.007 | 0.053 | 0.033 | 0.042 |
Range | 1.88–1.99 | 1.98–2.04 | 1.98–2.01 | 1.88–2.01 | 1.90–1.98 | 1.93–1.94 | 1.88–2.01 | 1.94–2.03 | 1.92–2.06 |
n | 5 | 10 | 22 | 12 | 10 | 3 | 7 | 6 | 13 |
eye diameter (ED) | |||||||||
Mean | 1.47 | 1.47 | 1.46 | 1.41 | 1.47 | 1.37 | 1.41 | 1.42 | 1.45 |
SD(±) | 0.037 | 0.062 | 0.065 | 0.066 | 0.061 | 0.012 | 0.102 | 0.088 | 0.089 |
Range | 1.42–1.51 | 1.38–1.56 | 1.35–1.57 | 1.28–1.51 | 1.33–1.55 | 1.36–1.38 | 1.25–1.57 | 1.27–1.50 | 1.27–1.59 |
n | 5 | 10 | 22 | 12 | 10 | 3 | 7 | 6 | 13 |
hindlimb length (HDL) | |||||||||
Mean | 3.4 | 3.53 | 3.55 | 3.47 | 3.43 | 3.46 | 3.48 | 3.49 | 3.5 |
SD(±) | 0.082 | 0.068 | 0.036 | 0.056 | 0.052 | 0.005 | 0.054 | 0.031 | 0.035 |
Range | 3.31–3.53 | 3.42–3.64 | 3.49–3.62 | 3.34–3.57 | 3.37–3.54 | 3.45–3.46 | 3.40–3.55 | 3.46–3.55 | 3.44–3.56 |
n | 5 | 10 | 22 | 12 | 10 | 3 | 7 | 6 | 13 |
forelimb width (FLW) | |||||||||
Mean | 0.91 | 0.88 | 0.95 | 0.93 | 0.8 | 0.77 | 0.98 | 1 | 1.02 |
SD(±) | 0.086 | 0.17 | 0.104 | 0.123 | 0.188 | 0.095 | 0.0818 | 0.158 | 0.13 |
Range | 0.77–0.99 | 0.71–1.26 | 0.79–1.17 | 0.62–1.12 | 0.56–1.19 | 0.68–0.87 | 0.88–1.08 | 0.71–1.14 | 0.78–1.27 |
n | 5 | 10 | 22 | 12 | 10 | 3 | 7 | 6 | 13 |
forelimb length (FLL) | |||||||||
Mean | 3.15 | 3.27 | 3.28 | 3.19 | 3.14 | 3.13 | 3.12 | 3.25 | 3.15 |
SD(±) | 0.084 | 0.066 | 0.055 | 0.046 | 0.037 | 0.012 | 0.083 | 0.109 | 0.086 |
Range | 3.01–3.23 | 3.20–3.41 | 3.20–3.48 | 3.12–3.25 | 3.09–3.21 | 3.12–3.14 | 3.01–3.22 | 3.08–3.41 | 2.90–3.23 |
n | 5 | 10 | 22 | 12 | 10 | 3 | 7 | 6 | 13 |