Extinction of lizards in Sri Lanka’s geographically isolated locations

Climate warming is projected to drive the extinction of approximately one-fifth of the world’s lizard species within the next 60 years(by 2080).However,it remains unclear to what extent and how the global lizard population will be affected by this phenomenon on a geographical scale.Furthermore,there is a need to establish a connection between extinction risk and the prioritization of conservation efforts.In this context,we aim to address concerns related to geographically isolated and critically small lizard populations,highlighting why they face an exceptionally high risk of extinction.We also explore the potential for entire populations to vanish without notice.A comprehensive understanding of these factors is essential to inform the prioritization of conservation initiatives aimed at safeguarding lizard populations in tropical countries under future climate change scenarios.

Differences in Incubation Length and Hatchling Morphology among Five Species of Oviparous Phrynocephalus Lizards (Agamidae) from China

We incubated eggs of five Phrynocephalus species （P. albolineatus, P. axillaries, P. grumgrzimailoi, P. helioscopus and P. przewalskii） at three constant temperatures （24℃, 28℃ and 32℃） to examine differences in incubation length and hatchling morphology among species and among temperature treatments. We combined data from this study with those reported previously for P. frontalis and P. versicolor to examine whether embryonic stage at laying is a causal factor for interspecific variation in incubation length, and whether the phylogenetic relationship inferred from hatchling morphology is consistent with the relationship based on mitochondrial DNA data. Mean values for incubation length differed among the five species studied herein and, in all these five species, incubation length decreased at a decreasing rate as temperature increased. In none of the five species did hatchling size （snout-vent length and body mass） and other morphological variables differ among the three temperature treatments. The seven oviparous Phrynocephalus lizards found in China differ from each other in hatchling morphology, and embryonic stage at laying is a causal factor of inter- and intra-specific variation in incubation length. The phylogenetic relationship inferred from hatchling morphology is not always consistent with the currently known relationship based on mitochondrial DNA data. Data from this study and those reported previously allow the conclusion that any Phrynocephalus species may have its unique position along the axis defined by hatchling morphology.

Differences in Thermal Preference and Tolerance among Three Phrynocephalus Lizards (Agamidae) with Different Body Sizes and Habitat Use

We acclimated adults of two viviparous （Phrynocephalus guinanens＆ and P. vlangalii） and one oviparous （P. versicolor） species of toad-headed lizards （Agamidae） to 28 ℃, 33 ℃ and 38 ℃ to examine whether thermal preference （preferred body temperature, Tp） and thermal tolerance （critical thermal minimum, CTMin; critical thermal maximum, CTMax） were affected by acclimation temperature, and correlate with body size and habitat use. Both Tp and CTMax were highest in P. versicolor and lowest in P. vlangalii, with P. guinanensis in between. The two viviparous species did not differ in CTMin and thermal tolerance range, and they both were more resistant to low temperatures and had a wider range of thermal tolerance than the oviparous species. Both CTMin and CTMax shifted upward as acclimation temperature increased in all the three species. Tp was higher in the lizards acclimated to 33 ℃ than in those to 28 ℃ or 38 ℃. The range of thermal tolerance was wider in the lizards acclimated to 28 ℃ than in those to 33 ℃ or 38 ℃. The data showed that： 1） thermal preference and tolerance were affected by acclimation temperature, and differed among the three species of Phrynocephalus lizards with different body sizes and habitat uses; 2） both Tp and CTMax were higher in the species exchanging heat more rapidly with the environment, and CTMin was higher in the species using warmer habitats during the active season; and 3） thermal preference and tolerance might correlat with body size and habitat use in Phrynocephalus lizards.

Population Dynamics Following the Last Glacial Maximum in Two Sympatric Lizards in Northern China

Phylogeographic studies of Eremias lizards （Lacertidae） in East Asia have been limited, and the impact of major climatic events on their population dynamics remains poorly known. This study aimed to investigate population histories and refugia during the Last Glacial Maximum of two sympatric Eremias lizards （E. argus and E. brenchleyi） inhabiting northern China. We sequenced partial mitochondrial DNA from the ND4 gene for 128 individuals of E. argus from nine localities, and 46 individuals of E. brenchleyi from five localities. Forty-four ND4 haplotypes were determined from E. argus samples, and 33 from E. brenchleyi samples. Population expansion events began about 0.0044 Ma in E. argus, and 0.031 Ma in E. brenchleyi. The demographic history of E. brenchleyi indicates a long-lasting population decline since the most recent common ancestor, while that of E. argus indicates a continuous population growth. Among-population structure was significant in both species, and there were multiple refugia across their range. Intermittent gene flow occurred among expanded populations across multiple refugia during warmer phases of the glacial period, and this may explain why the effective population size has remained relatively stable in E. brenchleyi and grown in E. argus.

Tail regeneration reduction in lizards after repetitive amputation or cauterization reflects an increase of immune cells in blastemas

Lizards are key amniote models for studying organ regeneration. During tail regeneration in lizards, blastemas contain sparse granulocytes, macrophages, and lymphocytes among the prevalent mesenchymal cells. Using transmission electron microscopy to examine scarring blastemas after third and fourth sequential tail amputations, the number of granulocytes, macrophages, and lymphocytes increased at 3-4 weeks in comparison to the first regeneration. An increase in granulocytes and agranulocytes also occurred within a week after blastema cauterization during the process of scarring Blood at the third and fourth regeneration also showed a significant increase in white blood cells compared with that under normal conditions and at the first regeneration. The extracellular matrix of the scarring blastema, especially after cauterization, was denser than that in the normal blastema and numerous white blood cells and fibroblasts were surrounded by electron-pale, fine fibrinoid material mixed with variable collagen fibrils. In addition to previous studies, the present observations support the hypothesis that an increase in inflammation and immune reactions determine scarring rather than regeneration. These new findings verify that an immune reaction against mesenchymal and epidermal cells of the regenerative blastema is one of the main causes for the failure of organ regeneration in amniotes.

Mating Ethogram of a Video-aided Study of Mating and Parturition in Captive Chinese Crocodile Lizards (Shinisaurus crocodilurus)

Mating behavior and parturition are important components in the life history of any species. The endangered Chinese crocodile lizard(Shinisaurus crocodilurus) is suffering dramatic population decreases in the wild. Successful captive breeding can provide individuals to be reintroduced into their natural habitat and help to restore the wild populations. Studying the mating behavior and parturition of these lizards in captivity will enable us understand how is this ancient species reproduction and optimize artificial reproduction protocol. We studied the mating behavior and parturition of Chinese crocodile lizards in captivity using video recordings at the Daguishan National Nature Reserve, China, from 2015 to 2017. We analyzed pre-copulation and copulation, which occurred in mid-March to late May from all studied years. Pre-copulation and copulation lasted on average 7.51(n=11) and 27.45(n=11) minutes, respectively. We found that large-bodied male lizards have longer copulation periods, a pattern seen in other lizard species. In the mating event, males always initiated sex, while the females were passive. The duration of parturition lasted 153.8 ± 26.8(n=46) minutes, and larger females had heavier clutches. In addition, even though Chinese crocodile lizards represent an ancient lizard clade with primitive characteristics, they still abide by the reproduction patterns seen in most other lizard species.

Regeneration of adhesive tail pad scales in the New Zealand gecko (Hoplodactylus maculatus) (Reptilia, Squamata; Lacertilia) can serve as an experimental model to analyze setal formation in lizards generally

During the regeneration of the tail in the arboreal New Zealand gecko （Hoplodactylus maculatus） a new set of tail scales, modified into pads bearing setae 5-20 μm long, is also regenerated. Stages of the formation of these specialized scales from epidermal pegs that invaginate the dermis of the regenerating tail are described on the basis of light and electron microscopic images. Within the pegs a differentiating clear layer interfaces with the spinulae and setae of the Oberh^utchen according to a process similar to that described for the digital pads. A layer of clear cytoplasm surrounds the growing tiny setae and eventually comities around them and their spatular ends, later leaving the new setae free- standing on the epidermal surface. The fresh adhesive pads help the gecko to maintain the prehensile function of its regenerated tail as together with the axial skeleton （made of a cylinder of elastic cartilage） the pads allow the regenerated tail to curl around twigs and small branches just like the original tail. The regeneration of caudal adhesive pads represents an ideal system to study the cellular processes that determine setal formation under normal or experimental manipulation as the progressive phases of the formation of the setae can be sequentially analyzed.

Higher Body Temperatures and Earlier Parturition in Response to Hypoxia Experienced by Pregnant Lizards

Many species are shifting towards higher altitudes in response to global wa rming, but how these upslope-shifting species will respond to hypoxic environments at high altitudes remains unclear. Hypoxia can be especially challenging for viviparous reproduction because of the limitation of oxygen supply to the female and her developing embryos. To investigate the effect of hypoxia on viviparous females and their offspring we acclimated pregnant females of a high-altitude dwelling vivipa rous liza rd(Phr ynocephalus vlangalii) to local oxygen and hypoxia conditions, respectively. We then recorded maternal body temperatures, postpartum body condition, as well as offspring morphology and locomotor performance. We found that pregnant females had higher body temperatures and advanced their parturition under hypoxic acclimation. However, maternal body condition, offspring morphology and locomotor performance were unaffected by the hypoxic conditions during gestation. Our study suggests that upslope-shifting viviparous lizards respond to hypoxic environments by plastically adjusting their body tempera tures to reduce parturition time, without short-term costs to offspring traits.

High-elevation Adaptation of Motion Visual Display Modifications in the Toad-Headed Agamid Lizards(Phrynocephalus)

Understanding the process of adaptation is a key mission in modern evolutionary biology.Animals living at high elevations face challenges in energy meta bolism due to several environmental constraints(e.g., oxygen supply, food availa bility,and movement time). Animal behavioral processes are intimately related to energy meta bolism, and therefore, behavioral modifica tions are expected to be an important mechanism for high-elevation adaptation. We tested this behavioral adaptation hypothesis using va ria tions of motion visual displays in toad-headed agamid lizards of the genus Phr ynocephalus. We predicted tha t complexity of visual motion displays would decrease with the increase of elevation, because motion visual displays are energetically costly. Displays of 12 Phr ynocephalus species were collected with elevations ranging from sea level to 4600 m. We quantified display complexity using the number of display components, display duration, pathways of display components, as well as display speed for each species. Association between display complexity and elevation was analyzed using the phylogenetic generalized least squares(PGLS)model. We found that both the number of display components and the average value of tail coil speed were negatively correlated with elevation, suggesting that toad-headed lizards living at high-elevation areas reduced their display complexity to cope with the environmental constraints. Our research provides direct evidence for high-elevation adaptation from a behavioral aspect and illustrates the potential impacts of environment heterogeneity on motion visual display diversification.

Body Size and Reproductive Tactics in Varanid lizards

Body size and female reproduction in the water adult females larger than 500 mm SVL and 32 adult males Ledong, Hainan under permit to our laboratory in Hainan monitor lizard （Varanus salvator） were studied. Forty-two larger than 400 mm SVL were donated by local people in in 2013 and 2014. The largest male and female measured 745 and 755 mm SVL, respectively. The mean SVL was greater in adult females than in adult males. Males had larger heads （head width） than females of the same SVL. The smallest reproductive female in our sample was 565 mm SVL. Females produced a single clutch of 17.1 （10-23） pliable-shelled eggs per breeding season stretching from mid-June and mid-September. Clutch size and clutch mass were all positively related to female SVL. However, there was no significant linear relationship between egg mass and female SVL. Larger females generally produced more eggs, and thus heavier clutches than did smaller ones. There was no significant linear relationship between relative clutch mass and female SVL. Phylogenetic generalized least squares （PGLS） analysis, accounting for phylogenetic relationships, showed that clutch size was positively correlated with mean maternal SVL in varanid lizards. PGLS analysis showed that phylogenetic relationships did not affect clutch （or/and egg） mass and the SVL although there were significant linear relationship between clutch （or/and egg） mass and mean maternal SVL. Therefore, we could draw some general conclusions about the body size and reproductive tactics in varanid lizards that larger females generally produced more eggs, larger eggs and thus heavier clutches than did smaller ones.

The Role of Environmental Stress in Determining Gut Microbiome:Case Study of Two Sympatric Toad-headed Lizards

The gut microbiota has gained attention because of its importance in facilitating host survival and evolution.However,it is unclear whether gut microbial communities are determined by the host(heritable factor) or environment(environmental factor).In this study,we investigated the gut microbial communities and potential functional signatures of two sympatric species distributed along an elevation gradient,the toadheaded lizards Phrynocephalus axillaris and P.forsythii.Our results indicated that at high elevations,the gut microbial communities of P.axillaris and P.forsythii did not significantly differ,and the phylogenetic relationships of gut microbial communities contradicted their hosts.At low altitudes,the two lizards could be distinguished based on their significantly different gut microbial communities.Compared to low-altitude populations,Kyoto Encyclopedia of Genes and Genomes(KEGG)pathway analysis showed that at higher altitudes,energy metabolism,such as carbohydrate,lipid,and amino acids metabolism were higher in both lizards.While a larger number of pathogenic bacteria were found in the lowaltitude population of P.forsythii.This suggests that the convergence of gut microbiota of two lizards at highaltitude stem from environmental factors,as they were exposed to the same environmental stress,whereas the divergence at low-altitude stemmed from heritable factors,as they were exposed to different environmental stresses.These results provide a new perspective regarding whether heritable or environmental factors dominate the gut microbiota during exposure to environmental stress.

The functional significance of residual yolk in lizards

Residual yolk is assumed to be an important source of energy and nutrients during early life in nonmammalian amniotes.Available data show that the mean size of residual yolk is far smaller in lizards than in turtles,snakes,crocodiles,and birds,raising a question of whether residual yolk is of functional significance in lizards.Here,we compared data from 26 lizard species with those from other nonmammalian amniotes to test the hypothesis that residual yolk is functionally less significant in species producing more fully developed offspring.In our sample,species mean offspring water contents ranged from 73%to 84%of body wet mass;species mean proportions of carcass dry mass,fat-body dry mass,and residual yolk dry mass to offspring dry mass ranged from 84%to 99%,0%to 5.0%,and 0%to 14.4%,respectively.Lizards are,on average,more fully developed at hatching or birth than snakes,as revealed by the fact that the mean proportion of carcass dry mass to body dry mass and offspring water contents were both higher in lizards than in snakes.We conclude that the functional significance of residual yolk during early life is generally less evident in lizards.Even in the lizards where residual yolk is of potential functional significance,this portion of yolk contributes little,if any,to postembryonic growth.Future work could usefully collect data across a wider spectrum of reptile taxa to establish a precocial-altricial continuum and test the hypothesis that species with a smaller amount of residual yolk are closer to the precocial end of the continuum.

Climate change,thermal niches,extinction risk and maternal-effect rescue of toad-headed lizards,Phrynocephalus,in thermal extremes of the Arabian Peninsula to the Qinghai-Tibetan Plateau

Determining the susceptibility of species to changing thermal niches is a major goal for biologists.In this paper we develop an eco-physiological model of extinction risk under climate change premised on behavioral thermoregulation.Our method downscales operative environmental temperatures,which restrict hours of activity of lizards,hr,for present-day climate(1975)and future climate scenarios(2070).We apply our model using occurrence records of 20 Phrynocephalus lizards(or taxa in species complexes)drawn from literature and museum records.Our analysis is phylogenetically informed,because some clades may be more sensitive to rising temperatures.The limits for computed hr predict local extirpations among Phrynocephalus lizards at continental scales and delineate upper boundaries of thermal niches as defined by Extreme Value Distributions.Under the 8.5 Representative Concentration Pathway scenario,we predict extirpation of 64%of local populations by 2070 across 20 Phrynocephalus species,and 12 are at high risk of total extinction due to thermal limits being exceeded.In tandem with global strategies of lower CO_(2) emissions,we propose regional strategies for establishing new national parks to protect extinction-prone taxa by preserving high-elevation climate refugia within existing sites of species occurrence.We propose that evolved acclimatization–maternal plasticity–may ameliorate risk,but is poorly studied.Previous studies revealed that adaptive maternal plasticity by thermoregulating gravid females alter progeny thermal preferences by±1℃.We describe plasticity studies for extinction-prone species that could assess whether they might be buffered from climate warming–a self-rescue.We discuss an epigenetic framework for studying such maternal-effect evolution.

The movement dynamics of autotomized lizards and their tails reveal functional costs of caudal autotomy

Autotomy has evolved independently several times in different animal lineages.It frequently involves immediate functional costs,so regeneration evolved in many instances to restore the functionality of that body part.Caudal autotomy is a widespread antipredator strategy in lizards,although it may affect energy storage,locomotion dynamics,or survival in future encounters with predators.Here,we assessed the effect of tail loss on the locomotor performance of wall lizards(Podarcis muralis),as well as the recovery of locomotor functionality of lizards with regenerated tails,and the movement dynamics of shed tails that were either intact or having regenerated portions.Tail loss had no effect on locomotion over unhindered spaces,possibly due to compensation between a negative effect on the stride of front limbs,and a positive effect of losing mass and friction force.We found a clear negative impact of tail loss on locomotion in spaces with interspersed obstacles,in which tailed lizards jumped larger distances when leaving the obstacles.Besides,lizards that used the tail to push off the ground were able to approach the obstacles from further,so that the tail seemed to be useful when used during jumping.Regeneration fully restores lizard’s locomotor capacities,but tail antipredator value,as indicated by the intensity of post-autotomic movements,is only partially retrieved.From these results,we propose that,together with the recovery of post-autotomy antipredator capacities,the restoration of the organismal locomotor performance may have been an important,yet frequently neglected factor in the evolution of lizard’s regeneration ability.

Do Larger and Older Individuals Occupy Better Areas in the Colony of Qinghai Toad-headed Agamas?

Group living reduces the chances of predation.This is also true for burrowing animals,which use burrows as refuges to escape predators.Living in the center of a colony is the safest because it offers access to the greatest number of burrows;however,it is stressful due to frequent aggressive interactions there.We predicted that larger and older animals should occupy areas of a colony with a higher density of burrows than smaller and younger ones.To test this prediction,we observed individually marked Qinghai toad-headed agamas(Phynocephalus vlangalii)from a single colony late in the breeding season when females were gravid.Density of burrows was the highest and distance between them the shortest in the center of the colony.Aggressive encounters were more frequent in the central part of the colony than on its periphery.We found a negative correlation between body size and distance to the center of the colony for females,but not for males.There was no such correlation between age of individuals and distance to the center of the colony.Females in this species likely compete with other females and males for central burrows in the colony even after mating season finishes.We suggest that gravid females likely compete for the dense burrows within the central part of the colony mainly due to their relatively lower sprint speed and higher probability of being caught by a predator if they are not refuged in a burrow.

A Glimpse into the Natural History of Leiolepis rubritaeniata Mertens,1961(Squamata:Leiolepididae)

During the months of April through July 2020 we studied aspects of the natural history of Leiolepis rubritaeniata,a species of butterfly lizard that occurs on the Khorat Plateau in Thailand and adjacent regions of Laos and Cambodia as well as in south-central Vietnam.We present data on population size and structure,as well as location,size,compass orientation,and structure of the lizard’s burrows.Also,we present climatic data(ambient air temperature and precipitation)at the study site for the duration of our field work as well as temperature data for the complete year 2021.Furthermore,we provide data on egg laying,incubation conditions and characteristics of the hatchlings.Finally,we present an easy to use and reliable non-invasive method for the long-term recognition of individual butterfly lizards based on their unique dorsal patterning.Butterfly lizards are utilized as a food source for the local human population.So far,there are no farming projects in Thailand involving this species and all individuals are collected from the wild populations.No data are available on the population dynamics of L.rubritaeniata but it can be assumed that habitat destruction due to land use change as well as its utilization for human consumption have negative effects on the long-term survival of the local populations of this lizard species.The baseline data presented here are essential for any meaningful conservation strategy for these lizards.

Thermal-physiological Strategies Underlying the Sympatric Occurrence of Three Desert Lizard Species

Sympatric reptiles are the ideal system for investigating temperature-driven coexistence. Understanding thermally physiological responses of sympatric lizards is necessary to reveal the physiological mechanisms that underpin the sympatric occurrence of reptiles. In this study, we used three lizard species, Eremias argus, E. multiocellata, and Phrynocephalus przewalskii, which are sympatric in the Inner Mongolia desert steppe, as a study system. By comparing their resting metabolic rates(RMR) and locomotion at different body temperatures, we aimed to better understand their physiological responses to thermal environments, which may explain the sympatric occurrence of these lizards. Our results showed that E. argus had significantly higher RMR and sprint speed than E. multiocellata, and higher RMR than P. przewalskii. In addition, the optimal temperature that maximized metabolic rates and locomotion for E. argus and E. multiocellata was 36°C, whereas for P. przewalskii it was 39°C. Our study revealed the physiological responses to temperatures that justify the sympatric occurrence of these lizards with different thermal and microhabitat preferences and active body temperatures. Eremias argus and E. multiocellata, which have lower body temperatures than P. przewalskii, depend on higher RMR and locomotion to compensate for their lower body temperatures in field conditions. Our study also highlights the importance of using an integrative approach, combining behavior and physiology, to explore the basis of sympatric occurrence in ectothermic species.

基于Lizard框架Ctrip Wireless H5数据存储方式的优化

Lizard是针对目前前端开源框架中存在的不足,由携程研发出的适用于无线移动端开发的框架。Lizard Web App是运行在浏览器上的单页应用,所有加载资源都放在一个html文件上,所有的用户交互都交给Java Script,所有的通信都交给Ajax。该文基于Lizard框架实现Hybrid App(混合模式移动应用)的开发,并针对框架中本地存储方式进行改进,由此提出了扩展store的存储方式。以多态方式封装store,由Local Storage、Index DB、SQLite by App替换cookie,进而支持多种本地存储方式,该文只是针对其中一种方式即Local Storage进行封装,以提高携程业务开发效率。

Kinematic Analysis and Experimental Verification on the Locomotion of Gecko

This paper presents a kinematic analysis of the locomotion of a gecko,and experimental verification of the kinematic model.Kinematic analysis is important for parameter design,dynamic analysis,and optimization in biomimetic robot research. The proposed kinematic analysis can simulate,without iteration,the locomotion of gecko satisfying the constraint conditions that maintain the position of the contacted feet on the surface.So the method has an advantage for analyzing the climbing motion of the quadruped mechanism in a real time application.The kinematic model of a gecko consists of four legs based on 7-degrees of freedom spherical-revolute-spherical joints and two revolute joints in the waist.The motion of the kinematic model is simulated based on measurement data of each joint.The motion of the kinematic model simulates the investigated real gecko's motion by using the experimental results.The analysis solves the forward kinematics by considering the model as a combination of closed and open serial mechanisms under the condition that maintains the contact positions of the attached feet on the ground. The motions of each joint are validated by comparing with the experimental results.In addition to the measured gait,three other gaits are simulated based on the kinematic model.The maximum strides of each gait are calculated by workspace analysis.The result can be used in biomimetic robot design and motion planning.

Effects of Constant versus Fluctuating Incubation Temperatures on Hatching Success, Incubation Length, and Hatchling Morphology in the Chinese Skink (Plestiodon chinensis)

We incubated eggs ofPlestiodon chinensis under five constant （24, 26, 28, 30, and 32 ℃） and one fluctuating thermal regimes to examine the effects of constant versus fluctuating incubation temperatures on hatching success, incubation length, and hatchling morphology. The duration of incubation varied considerably among the six temperature treatments, whereas hatching success did not. The mean incubation length decreased as temperature increased in a nonlinear way, and increased as the thermal variance increased. Incubation temperature affected the body size （linear length and mass） and shape of hatchlings, with eggs incubated at 26, 28, and 30 ℃ producing larger and heavier hatchlings than did those incubated at 24 ℃, 32 ℃, or fluctuating temperatures. Our results showed that exposure of P. chinensis eggs to extreme temperatures for brief periods of time did not increase embryonic mortality and, in the fluctuating-temperature treatment, the thermal variance affected hatchling morphology more evidently than the thermal mean. Our results highlight the importance of the thermal variance in affecting embryonic development and hatchling morphology, and add further evidence that temperatures within the range of 26-30 ℃ are optimal for P. chinensis embryos.