Morphometric variation of the Asian Water Monitor Lizard (Varanus salvator) in different habitats in Peninsular Malaysia: a model analysis with machine learning

A tail of evolution: evaluating body length, weight and locomotion as potential drivers of tail length scaling in Australian marsupial mammals

The present study investigates relationships among size, shape and speed in the Australian agamid lizard Amphibolurus nuchalis. Maximal running speed, body mass, snout‐vent length, tail length, fore‐ and hind limb spans and thigh muscle mass were measured in 68 field‐fresh individuals spanning the entire ontogenetic size range (1.3 48 g). Relative lengths of both foreand hind limbs decrease with increasing body mass (= negative allometry), whereas relative tail length and thigh muscle mass increase with body mass (= positive allometry). Repeatable and significant differences in maximal running speed exist among individuals. Maximal running speed scales as (body mass)0.161, and 59% of the variation in maximal speed was related to body mass. Based on the results of the present and previous studies, data on scaling of body proportions alone appear inadequate to infer scaling relationships of functional characters such as top speed.Surprisingly, individual variation in maximal speed is not related to individual variation in shape (relative limb, tail and body lengths). These components of overall shape are not independent; individuals tended to have either relatively long or relatively short limbs, tails and bodies for their body mass. Even the significant difference in multivariate shape between adult males and females has no measurable consequences for maximal speed. Speeds of field‐fresh animals did not vary on a seasonal basis, and eight weeks of captivity had no effect on maximal running speeds. Gravid females and long‐term (obese) captive lizards were both approximately 12% slower than field‐fresh lizards.

Living primates vary considerably in tail length-body size relation, ranging from tailless species to those where the tail is more than twice as long as the body. Because the general pattern and determinants of tail evolution remain incompletely known, we reconstructed evolutionary changes in relative tail length across all primates and sought to explain interspecific variation in this trait. We combined data on tail length, head-body length, intermembral index (IMI), habitat use, locomotion type, and range latitude for 340 species from published sources. We reconstructed the evolution of relative tail length to identify all independent cases of regime shifts on a primate phylogeny, using several methods based on Ornstein-Uhlenbeck (OU) models. Accounting for phylogeny, we also examined the effects of habitat, locomotion type, distance from the equator and IMI on interspecific variation in tail length-body size relation. Primate tail length is not sexually dimorphic. A phylogenetic reconstruction allowing multiple optima explains the observed regime shifts best. During the evolutionary history of primates, relative tail length changed 50 times under an OU model. Specifically, relative tail length increased 26 and decreased 24 times. Most of these changes occurred among Old World primates. Among the variables tested here, interspecific variation in IMI and the difference between leaping and non-leaping locomotion explained interspecific variation in relative tail length: Evolutionary decreases in relative tail length are generally associated with an increase in IMI and an absence of leaping behavior. Regime shifts for relative tail length in living primates occurred in concert with fundamental changes in IMI and a change from leaping to non-leaping locomotion, or vice versa. Exceptions from this general pattern are linked to the presence of a prehensile tail or specialized foraging strategies. Thus, the primate tail appears to have evolved in functional coordination with limb proportions, presumably to assist body balance.

The common lizard (Lacerta vivipara) is a small, nonterritorial, live-bearing lacertid that is sexually dimorphic in several morphological traits (e.g., tail length, snout–vent length, head size). Using microsatellites, we examined paternity in a wild population and investigated whether sexual dimorphism could be the result of intra- or intersexual selection. We found multiple paternity in 65.4% of 26 clutches. There was no evidence of assortative mating. Successfully reproducing males were larger and heavier and had longer tail regenerates or intact tails compared to those that did not reproduce. Tail length and body condition of males were related to the number of offspring sired. However, we found no evidence that head width was related to male reproductive success. We conclude that (1) males with higher body condition index might be more successful in male-male interactions or might be able to search more effectively for females, (2) sex divergence in relative tail length in common lizards reflects the action of sexual selection for male reproductive success, and (3) intersexual dietary divergence could be an alternative hypothesis for head size difference between sexes rather than intrasexual selection.

Allometry in size and shape between sexes was investigated in preserved Bearded Dragon Pogona barbata museum specimens. Measurements for snout-vent, head, leg and tail lengths were obtained from 236 individuals ranging from hatchlings to large adults. Juveniles and adults were sexed, and size at onset of maturity was determined. Geographic variation in P. barbata was also studied by comparing lizards from three regions in New South Wales (NSW), which differ in annual rainfall. Relative to snout-vent length, head and leg lengths were in negative allometry in all specimens, and tail length was in positive allometry in juveniles, but negative in adults. Sexual dimorphism was evident in body size and shape. Males grew larger and reached maturity at larger sizes than females. While juvenile shape did not differ between the sexes, mature males had proportionately longer heads, legs and tails than adult females. This generalisation in sexual shape dimorphism did not hold true for animals from different NSW regions. Western males had proportionately longer legs and heads than western females, but relative tail lengths were not significantly different. Sexual dimorphism in body shape was not seen in the central animal group. In the east, heads and tails were longer in males than in females. Both sexes showed lower relative head and leg lengths in more arid regions. We suggest that a combination of genetic drift and phenotypic responses are likely causes of these variations.

In primates, tail length is subject to wide variation, and the tail may even be absent. Tail length varies greatly between each species group of the genus Macaca, which is explained by climatic factors and/or phylogeographic history. Here, tail length variability was studied in hybrids of the Japanese (M. fuscata) and Taiwanese (Macaca cyclopis) macaque, with various degrees of hybridization being evaluated through autosomal allele typing. Relative tail length (percent of crown-rump length) correlated well with the number of caudal vertebrae. Length profiles of caudal vertebrae of hybrids and parent species revealed a common pattern: the length of several proximal-most vertebrae do not differ greatly; then from the third or fourth vertebra, the length rapidly increases and peaks at around the fifth to seventh vertebra; then the length plateaus for several vertebrae and finally shows a gentle decrease. As the number of caudal vertebrae and relative tail length increase, peak vertebral length and lengths of proximal vertebrae also increase, except that of the first vertebra, which only shows a slight increase. Peak vertebral length and the number of caudal vertebrae explained 92 % of the variance in the relative tail length of hybrids. Relative tail length correlated considerably well with the degree of hybridization, with no significant deviation from the regression line being observed. Thus, neither significant heterosis nor hybrid depression occurred.

Sexual dimorphism is widespread in animals, including snakes, and has impor- tant implication in both ecology and behaviour. I studied a grass snake (Natrix natrix) population from Stawy Milickie nature reserve. Mean snout-vent length (SVL) for fe- males was significantly greater than SVL for males, but males had proportionally longer tails. However, relative tail length (TL) in males decreased with increasing SVL whereas in females it was constant. Larger tails in males have frequently been associated with in- creasing mating success (e.g. tail wrestling behaviour between males). However, it is pos- sible this is less important in the population that I studied; instead, male snakes can allocate more energy to body growth.

The influence of natural selection and sexual selection on the tails of sea-snakes (Laticauda colubrina)

Many phenotypic traits perform more than one function, and so can influence organismal fitness in more than one way. Sexually dimorphic traits offer an exceptional opportunity to clarify such complexity, especially if the trait involved is subject to natural as well as sexual selection, and if the sexes differ in ecology as well as reproductive behaviour. Relative tail length in sea-snakes fulfils these conditions. Our field studies on a Fijian population of yellow-lipped sea kraits ( Laticauda colubrina ) show that relative tail lengths in male sea kraits have strong consequences for individual fitness, both via natural and sexual selection. Males have much longer tails (relative to snout–vent length) than do females. Mark-recapture studies revealed a trade-off between growth and survival: males with relatively longer tails grew more slowly, but were more likely to survive, than were shorter-tailed males. A male snake's tail length relative to body length influenced not only his growth rate and probability of survival, but also his locomotor ability and mating success. Relative tail length in male sea kraits was thus under a complex combination of selective forces. These forces included directional natural selection (through effects on survival, growth and swimming speed) as well as stabilizing natural selection (males with average-length tails swam faster) and stabilizing sexual selection (males with average-length tails obtained more matings). In contrast, our study did not detect significant selection on relative tail length in females. This sex difference may reflect the fact that females use their tails primarily for swimming, whereas males also must frequently use the tail in terrestrial locomotion and in courtship as well as for swimming.

Tail reduction/loss independently evolved in a number of mammalian lineages, including hominoid primates. One prerequisite to appropriately contextualizing its occurrence and understanding its significance is the ability to track evolutionary changes in tail length throughout the fossil record. However, to date, the bony correlates of tail length variation among living taxa have not been comprehensively examined. This study quantifies postsacral vertebral morphology among living primates and other mammals known to differ in relative tail length (RTL). Linear and angular measurements with known biomechanical significance were collected on the first, mid-, and transition proximal postsacral vertebrae, and their relationship with RTL was assessed using phylogenetic generalized least-squares regression methods. Compared to shorter-tailed primates, longer-tailed primates possess a greater number of postsacral vertebral features associated with increased proximal tail flexibility (e.g., craniocaudally longer vertebral bodies), increased intervertebral body joint range of motion (e.g., more circularly shaped cranial articular surfaces), and increased leverage of tail musculature (e.g., longer spinous processes). These observations are corroborated by the comparative mammalian sample, which shows that distantly related short-tailed (e.g., Phascolarctos, Lynx) and long-tailed (e.g., Dendrolagus, Acinonyx) nonprimate mammals morphologically converge with short-tailed (e.g., Macaca tonkeana) and long-tailed (e.g., Macaca fascicularis) primates, respectively. Multivariate models demonstrate that the variables examined account for 70% (all mammals) to 94% (only primates) of the variance in RTL. Results of this study may be used to infer the tail lengths of extinct primates and other mammals, thereby improving our understanding about the evolution of tail reduction/loss.

Background:Varanus salvator is one of the reptiles being hunted by human beings for several purposes, including traditional medicine. The studies about reproductive biology aspects were limited. Aim:This study aimed to determine the morphology, histology, and histometry of V. salvator paryphasmata and hemibaculum based on Snout-Vent Length (SVL) as an indicator of sexual maturity.Methods:This study examined 18 pairs of hemipenis of V. salvator with SVL more and less than 40 cm in equal number. Paryphasmata and hemibaculum parts were observed visually and micro-sliced, then stained with Hematoxylin-Eosin (HE). The histological observation was conducted under a 40×, 100×, and 400× magnification of a light microscope. The histometry of the paryphasmata was examined using 13 Megapixels Coolpad and OptiLab Plus for microscopic pictures. The chondrocyte cell area was measured using the Optilab Plus and Image Raster three applications.Results:The sizes of glans of hemipenis, paryphasmata, and hemibaculum increased according to the increasing of SVL. The average paryphasmata row number, epidermis, and loose connective tissue thickness were not significantly different (p > 0.05). However, dense connective tissue was thicker (p < 0.05), which corresponds to SVL. Hemibaculum was composed of fibrous and hyaline cartilage characterized by chondrocyte cells. The SVL also affects (p < 0.05) the ossification of hyaline in hemipenis, while the chondrocyte cell area followed the equation −1.87E7 + 7.09E5* SVL.Conclusion:The SVL size of V. salvator affects the paryphasmata, hemibaculum, thickness of dense connective tissue of paryphasmata, and the area of chondrocyte cells.

According to the hypothesis of parasite-mediated sexual selection, for a communication system to work reliably, parasites should reduce the showiness of sexual signals of their host. In this study, we examined whether the expression of breeding coloration in free-ranging adult European Green Lizards ( Lacerta viridis (Laurenti, 1768)) is linked with infestation by their common ectoparasite Ixodes ricinus (L., 1758) (Acari: Ixodidae). We found that tick infestation was higher in males than in females. Males showing relatively heavier body for their tail length (predominantly males with regenerated tails) and relatively thinner tail base experienced higher infestation rates. In turn, relatively heavier females for their snout–vent length were less tick infested. Although some components of throat and chest coloration varied significantly with relative tail length, tail-base thickness, body mass, and head size, a measure of male throat and female chest color saturation seemed independent of lizard morphology. After correcting for the effects of morphology on skin coloration and tick load, the saturation of blue throat color in male lizards decreased with increasing level of tick infestation. In contrast, yellow chest color saturation increased with residual tick numbers in females. Considering presumably different signaling functions of male and female lizard coloration, our work suggests that tick infestation might represent a handicap for Green Lizards.

Context Each year, between 50 000 and 120 000 Asian water monitors (Varanus salvator, to &amp;gt;2 m total length) are harvested from the wild in Peninsular Malaysia for their skins. Under the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES), international trade is allowable only if it is sustainable. Aims To assess the sustainability of Malaysia’s harvest of water monitors by quantifying the abundance and demography of V. salvator in the wild, and to develop cost-effective methods for estimating the parameters needed to evaluate sustainability. Methods We conducted trapping surveys to determine the abundance, population demography and density of V. salvator in four habitat types in five states in Peninsular Malaysia in 2010, 2011, 2012 and 2018. Key results Of 1025 lizards captured, only 63% (mostly females) were within the preferred body-size range exploited for commercial trade. Densities were high (37–372 lizards km–2 based on estimated population sizes; 1–35 lizards km–2 based on number of animals captured). Anthropogenic habitats (e.g. oil palm plantations) contained denser populations of monitors than did natural habitats where no hunting occurs, but mean body sizes were smaller. Conclusions Despite intensive harvesting for many decades, V. salvator remains abundant and widespread. Harvesting alters the demographic structure of lizard populations, but harvests of V. salvator in Malaysia are likely to be sustainable because a significant proportion of the population is not exploited. Implications Ongoing monitoring is required to continually reassess harvest sustainability. For this purpose, relatively simple population approaches, such as line-trapping transects to elucidate relative abundances, can provide important data on the makeup of hunted populations of water monitors more cost-effectively than can mark–recapture studies for assessments of sustainable use of these economically important lizards.

Tail movement is an important component of vertebrate locomotion and likely contributes to dynamic stability during steady-state locomotion. Previous results suggest that the tail plays a significant role in lizard locomotion, but little data are available on tail motion during locomotion and how it differs with morphological, ecological, and phylogenetic parameters. We collected high-speed vertical climbing and horizontal locomotion video data from 43 lizard species from four taxonomic groups (Agamidae, Gekkota, Scincidae, and Varanidae) across four habitats. We introduce a new semi-automated and generalizable analysis pipeline for tail and spine motion analysis including markerless pose-estimation, semi-automated kinematic recognition, and muti-species data analysis. We found that step length relative to snout-vent length (SVL) increased with tail length relative to SVL. Examining spine cycles agnostic to limb stride phase, we found that ranges of inter-tail bending compared with inter-spine bending increased with relative tail length, while ranges of tail deflection relative to spine deflection increased with relative speed. Considering stepwise strides, we found the angular velocity and acceleration of the tail center of mass increased with relative speed. These results will provide general insights into the biomechanics of tails in sprawling locomotion enabling biomimetic applications in robotics, and a better understanding of vertebrate form and function. We look forward to adding more species, behaviors, and locomotor speeds to our analysis pipeline through collaboration with other research groups.

SummaryGravity imposes potentially important constraints on blood circulation in tall or elongate animals during upright posture or climbing. Upright postures create vertical gradients of gravitational (=hydrostatic) pressures within circulatory vessels. In terrestrial animals, this pressure potentially induces blood pooling and oedema in dependent (inferior) tissues and, secondarily, decreases blood volume reaching the head and vital organs. Arboreal snakes exhibit a suite of adaptations for countering the effects of gravity on blood circulation, including relatively non‐compliant tissue compartments in the tail. However, patterns of tail length related to arboreal habitats and gravity have not been previously studied. Here, we test the hypothesis that arboreal snakes have relatively longer tails than non‐climbing species, and we interpret the results within the context of adaptation to gravity stress.Length data were obtained for 226 snake species, 139 genera and 15 snake families assigned to three broad gravitational habitat categories (G‐habitats: stenotopically arboreal, eurytopically arboreal/terrestrial and non‐scansorial). We constructed a composite phylogenetic tree and quantified the relationships between snake morphology and G‐habitat using conventional regression analysis of species data and regressions of phylogenetically independent contrasts.Mean relative tail length (RTL) increased and mean relative snout‐vent length decreased with increasing arboreality among the three G‐habitats. However, mean total body length (TL) was not different between the two arboreal G‐habitats.Snakes with longerRTLs have a higher percentage of elongate blood vessels contained within the relatively non‐compliant integument of the tail, which mitigates postural blood pooling experienced during climbing.TLin adult female arboreal snakes is hypothesized to be an evolutionary trade‐off between demands for a longer tail to enhance cardiovascular performance and the length of body cavity to provide space for young.Our results complement previous studies correlatingRTLwith arboreal habitat use, in addition to physiological experiments quantifying the effects of gravity on blood flow in snakes. These results provide evidence that longRTLs of arboreal snakes function, at least in part, as an adaptive response to cardiovascular stresses on blood circulation imposed by gravity. Selection on tail length related to other factors seems less likely, but is in need of further investigation.