1 Introduction

Climate change, driven by anthropogenic activities, has emerged as one of the most pressing environmental challenges of the 21st century. The rapid increase in global temperatures, changes in precipitation patterns, and the frequency of extreme weather events are altering ecosystems and affecting biodiversity worldwide. These climatic shifts have profound impacts on natural systems, influencing species distributions, phenology, and interactions within ecosystems (Diele-Viegas and Rocha, 2018). The Earth’s warming is projected to exceed 4.8°C by the end of the 21st century, posing significant threats to biodiversity and ecosystem services. Understanding the extent and nature of these impacts is crucial for developing effective conservation strategies.

Reptiles, as ectothermic organisms, are particularly sensitive to changes in their thermal environment, making them excellent bioindicators of climate change (Zhang et al., 2023; Li et al., 2024). Their physiological and behavioral traits are closely tied to environmental temperatures, and even slight changes can have significant effects on their survival, reproduction, and distribution (Noble et al., 2018). Reptiles exhibit a range of adaptive responses, including phenotypic plasticity and genetic adaptation, which can provide insights into the mechanisms of resilience and vulnerability to climate change (Urban et al., 2013; Catullo et al., 2019). Moreover, reptiles occupy diverse ecological niches and are integral components of many ecosystems, thus their responses to climate change can have cascading effects on ecosystem structure and function (Du et al., 2023).

This study seeks to identify patterns and trends in how reptiles respond to changing climatic conditions. By evaluating the role of phenotypic plasticity and genetic adaptation in mediating reptile responses to climate change, assess the impacts of developmental environments on thermal physiological traits and survival, and investigate the influence of climate change on reptile behavior, reproductive strategies, and population dynamics, the study will provide a comprehensive overview of the adaptive capacities of reptiles, highlighting the importance of integrating multiple ecological and evolutionary factors, and inform conservation strategies aimed at mitigating the impacts of climate change on reptile populations.

2 Overview of Reptilian Physiology and Ecology

2.1 Diversity of Reptiles and Their Ecological Roles

Reptiles represent one of the most diverse and ecologically significant groups of vertebrates, having successfully colonized a wide range of habitats across the globe. This group includes lizards, snakes, turtles, crocodilians, and the tuatara, each playing unique roles in their respective ecosystems. Lizards and snakes, which belong to the order Squamata, are particularly diverse, with lizards alone comprising over 6 650 species (Meiri, 2018). These species have adapted to various ecological niches, from deserts to rainforests, showcasing a remarkable array of morphological and physiological traits that enable them to thrive in different environments (Pincheira-Donoso et al., 2013).

Turtles (order Testudines) and crocodilians (order Crocodylia) are less diverse but equally important. Turtles are found in both aquatic and terrestrial habitats, contributing to nutrient cycling and seed dispersal, while crocodilians are apex predators in many freshwater ecosystems, playing a crucial role in maintaining the balance of these environments. The tuatara, a unique reptile from New Zealand, represents the order Rhynchocephalia and provides valuable insights into the evolutionary history of reptiles. Despite their ecological importance, many reptile species are under threat due to habitat loss, climate change, and other anthropogenic factors, highlighting the need for comprehensive conservation efforts (Nordstrom et al., 2022).

2.2 Climate Sensitivity in Reptiles

Reptiles, being ectothermic animals, are particularly sensitive to changes in their thermal environment. Temperature, humidity, and precipitation significantly influence their behavior, physiology, and overall fitness. For instance, the thermal physiology of reptiles, including traits like critical thermal maximum (CTmax) and thermal preference (Tpref), is crucial for their survival and reproductive success. However, studies have shown that developmental environments do not significantly affect these thermal physiological traits, suggesting that reptiles may rely more on behavioral or evolutionary adaptations to cope with changing temperatures (Zhang et al., 2023).

The ability of reptiles to adapt to climate change through phenotypic plasticity and genetic adaptation is a subject of ongoing research. While phenotypic plasticity allows for immediate responses to environmental changes, genetic adaptation may provide long-term solutions. However, the rate of thermal trait evolution in reptiles is often slow, particularly at the warm end of the thermal performance curve, which may limit their ability to keep pace with rapid climate changes (Bodensteiner et al., 2020). Understanding the interactions between plasticity, behavior, and genetic adaptation is essential for predicting how reptiles will respond to future climate scenarios (Urban et al., 2013; Valero et al., 2021). Additionally, the use of advanced techniques like environmental DNA (eDNA) can enhance our ability to monitor and conserve reptile populations in the face of climate change.

3 Phenotypic Plasticity in Response to Climate Change

3.1 Definition and Mechanisms of Phenotypic Plasticity

Phenotypic plasticity refers to the ability of an organism to alter its morphology, physiology, or behavior in response to environmental changes. This adaptability is crucial for survival in rapidly changing climates, as it allows organisms to maintain functionality and fitness without requiring genetic changes. Phenotypic plasticity can manifest as temporary or reversible changes, such as alterations in body size, coloration, or metabolic rates, which can be adjusted back when environmental conditions revert to their original state. For instance, reptiles often exhibit changes in their morphology and physiology in response to varying thermal environments, which can significantly impact their survival and reproductive success (Noble et al., 2018).

The mechanisms underlying phenotypic plasticity are diverse and complex. They include changes in gene expression, hormonal regulation, and cellular processes that enable organisms to respond to environmental cues. For example, reptiles may adjust their metabolic rates or alter their developmental pathways in response to temperature fluctuations, which can affect their growth rates, survival, and reproductive output (Fox et al., 2019). These plastic responses are often mediated by specific genes that are sensitive to environmental conditions, allowing for rapid and reversible changes in phenotype. Understanding these mechanisms is essential for predicting how reptiles and other organisms will cope with ongoing climate change.

3.2 Behavioral Adjustments and Shifts in Activity Patterns

Behavioral plasticity is another critical aspect of phenotypic plasticity, enabling organisms to modify their behavior in response to environmental changes. Reptiles, for instance, may alter their basking behavior, feeding times, and seasonal activity cycles to cope with temperature variations and other climatic factors. These behavioral adjustments can help mitigate the adverse effects of climate change by optimizing energy use and reducing exposure to extreme conditions. For example, changes in basking behavior can help reptiles regulate their body temperature more effectively, thereby maintaining optimal physiological functions (Canale and Henry, 2010; Urban et al., 2013).

Shifts in activity patterns are also common among reptiles as a response to climate change. These shifts can include changes in the timing of daily activities, such as feeding and mating, as well as alterations in seasonal behaviors, such as migration and hibernation. By adjusting their activity patterns, reptiles can avoid the most extreme temperatures and exploit more favorable conditions for survival and reproduction. For instance, some reptiles may extend their active periods during cooler parts of the day or season to avoid overheating, while others may enter a state of torpor or hibernation during unfavorable conditions (Boutin and Lane, 2013). These behavioral adaptations are crucial for maintaining population stability and resilience in the face of climate change.

4 Morphological and Physiological Adaptations

4.1 Morphological Changes

Reptiles exhibit a range of morphological changes in response to climate change, including alterations in body size, scale pattern, and coloration. These changes are often driven by the need to adapt to new environmental conditions that affect their survival and reproductive success. For instance, phenotypic plasticity allows reptiles to modify their physical traits in response to varying climatic conditions, which can be crucial for their survival (Urban et al., 2013). Studies have shown that early thermal environments, such as nest temperatures, can significantly impact the morphology of reptile offspring, influencing traits like body size and scale pattern (Noble et al., 2018). These morphological adaptations are essential for coping with the changing climate, as they can affect a reptile's ability to thermoregulate, find food, and avoid predators.

Moreover, the concept of countergradient variation (CnGV) highlights how genetic variation can counteract environmental variation, leading to morphological adaptations that are beneficial in specific climates. Most published studies show evidence for CnGV between development time and environmental temperature (Figure 1) (Pettersen, 2020). These morphological changes are not just limited to body size but also include alterations in coloration and scale patterns, which can provide camouflage and reduce predation risks in different habitats. Overall, morphological adaptations play a critical role in enabling reptiles to survive and thrive in the face of climate change.

Figure 1 Effect sizes (Hedges’ g) for differences in the thermal sensitivity of development time (time from oviposition until hatching) and metabolic (heart) rate across cold and warm-adapted populations for 15 species of reptiles across 8 families (± variance) (Adopted from Pettersen, 2020) Image caption: For development time (D; green data points and variance bars), positive Hedges’ g values indicate positive Cov(G,E), or cogradient variation, where cold-adapted populations have longer D, relative to warm-adapted populations. Negative values of D indicate negative Cov(G,E), or countergradient variation, where genotypic differences oppose environmental temperature effects – in these instances, cold-adapted populations develop faster than warm-adapted populations. For metabolic rates (MR; orange data points and variance bars), negative Hedges’ g values indicate positive Cov(G,E), or cogradient variation, where cold-adapted populations have lower MR, relative to warm-adapted populations, while positive values of MR indicate negative Cov(G,E), or countergradient variation – here cold-adapted populations maintain higher MR, relative to warm-adapted populations (Adopted from Pettersen, 2020)

4.2 Physiological Adaptations

Physiological adaptations in reptiles are equally important for coping with the challenges posed by climate change. These adaptations include adjustments in metabolism, thermal tolerance, and reproductive strategies. Reptiles, being ectothermic animals, rely heavily on external temperatures to regulate their body functions. As global temperatures rise, reptiles must adapt their thermal physiology to maintain homeostasis and ensure their survival. Studies have shown that reptiles can exhibit significant plasticity in their thermal physiological traits, such as critical thermal maximum (CTmax) and thermal preference (Tpref), allowing them to cope with varying thermal environments (Zhang et al., 2023).

One of the key physiological adaptations observed in reptiles is the adjustment of metabolic rates. For instance, reptiles in cooler climates may exhibit slower metabolic rates to conserve energy, while those in warmer climates may have faster metabolic rates to support increased activity levels (Figure 1) (Pettersen, 2020). Additionally, thermal tolerance is a crucial aspect of physiological adaptation, with some reptiles evolving to withstand higher temperatures through behavioral thermoregulation and physiological adjustments (Muñoz et al., 2014). Reproductive strategies also play a vital role in physiological adaptation, as changes in nesting behavior and timing of oviposition can influence the thermal environment of developing embryos, thereby affecting their survival and fitness (Du et al., 2023).

5 Evolutionary Adaptations to Climate Change

5.1 Genetic Mechanisms and Natural Selection

The role of genetic mechanisms and natural selection in shaping the evolutionary adaptations of reptiles to climate change is a critical area of study. Natural selection acts on genetic variation within populations, leading to changes in gene frequencies that enhance survival and reproduction in changing environments (Burggren and Mendez-Sanchez, 2023). This process is fundamental to the adaptive evolution of species facing new climatic conditions. For instance, the study by Radchuk et al. highlights that while phenotypic changes in response to climate change are well-documented, the adaptive nature of these changes remains uncertain, particularly in terms of maintaining a good match between phenotype and environment. This underscores the importance of understanding the genetic basis of these adaptations.

Moreover, the research on lacertid lizards and other vertebrates by the functional genomics approach reveals that specific subsets of genes are under positive diversifying selection, which are involved in physiological and morphological adaptations to climate (Bairos-Novak et al., 2021; Valero et al., 2021). This indicates that certain genes play a pivotal role in enabling reptiles to cope with environmental stressors induced by climate change. The identification of these genes and their functions can provide insights into the genetic mechanisms that facilitate adaptation and highlight the selective pressures that shape gene frequencies in reptile populations.

5.2 Evidence of Adaptive Evolution in Reptiles

Evidence of adaptive evolution in reptiles in response to climate change is emerging, although it is often challenging to distinguish between genetic adaptations and phenotypic plasticity. The study by Telemeco et al. on the impact of developmental temperatures on reptile phenotypes demonstrates that early thermal environments can have significant and lasting effects on offspring phenotype and survival, suggesting a potential for adaptive responses to changing climates (Noble et al., 2018; Putman and Tippie, 2020). This highlights the role of phenotypic plasticity as a mechanism that can buffer populations against climate change, although it also points to the need for more research on genetic changes over time.

Historical perspectives also provide valuable insights into adaptive evolution. The research on the early evolution and radiation of reptiles during successive climate crises in the deep past shows that climatic shifts have historically driven significant evolutionary changes in reptiles (Bodensteiner et al., 2020; Simões et al., 2022). This long-term view underscores the potential for reptiles to undergo adaptive evolution in response to current and future climate change. However, the pace of contemporary climate change poses a unique challenge, as the rate of environmental change may outstrip the ability of species to adapt through natural selection alone.

6 Case Study: Evolutionary Adaptation in the Sand Lizard (Lacerta agilis)

6.1 Background and Relevance of the Sand Lizard

The sand lizard (Lacerta agilis) is a species of lizard that is widely distributed across Europe and parts of Asia (Zakharov et al., 2022). It inhabits a variety of environments, including heathlands, sand dunes, and grasslands, which are characterized by their sandy soils and sparse vegetation (Valero et al., 2021; Zakharov et al., 2022). This species is particularly notable for its adaptability to different climatic conditions, which makes it an excellent model for studying the impacts of climate change on reptilian species. The Swedish populations of Lacerta agilis, for instance, are considered relict populations from the post-glacial warmth period, indicating their long-term persistence in varying climatic conditions.

The genetic diversity within these populations has been shaped by historical climatic events and more recent anthropogenic disturbances. Studies have shown that Swedish sand lizards have lower genetic variability compared to their Hungarian counterparts, likely due to a population bottleneck during post-glacial colonization. Despite this reduced genetic diversity, these populations exhibit a high level of observed heterozygosity, suggesting that they have maintained a degree of genetic health that could be crucial for their adaptability to ongoing climate changes.

6.2 Response to Warming Temperatures

The response of Lacerta agilis to warming temperatures has been a subject of extensive research, revealing significant changes in their phenology, morphology, and reproductive cycles. One of the most notable phenological changes observed is the advancement of oviposition dates in response to higher spring temperatures. This shift is attributed to both population-level plasticity and individual-level variation, which together enhance the fitness and survival of offspring in a warming climate (Ljungström et al., 2015). Additionally, studies have shown that moderate warming can have beneficial effects on embryonic and hatchling development, particularly in low-latitude margin populations. These populations exhibit higher hatching success, growth rates, and survival rates under moderate warming scenarios, although extreme temperatures can have detrimental effects (Figure 2) (Cui et al., 2022).

Figure 2 Natural habitats of (A) and thermal environments (B) for sand lizards (Lacerta agilis) from Burqin population (Adopted from Cui et al., 2022) Image caption: (A) The photographs indicate the natural habitats and demographic photographs for Lacerta agilis. (B) The blue solid, green long dash and red short dash lines indicate the temperatures of nest at depth of 5 cm underground, surface and air. The shade column indicates the reproductive season mainly for embryonic development. The data of temperatures were from China Meteorological Data Service Centre (Adopted from Cui et al., 2022)

Morphological changes have also been documented, such as variations in dorsal patterns and coloration, which are influenced by local temperature conditions. For instance, in the common lizard (Lacerta vivipara), warmer climates have been associated with changes in the frequency of different morphotypes, suggesting that similar mechanisms could be at play in Lacerta agilis (Lepetz et al., 2009). These morphological adaptations, along with behavioral changes such as altered basking patterns and habitat use, are crucial for thermoregulation and overall fitness in a changing climate.

6.3 Implications of the Case Study

The case study of Lacerta agilis provides valuable insights into the broader mechanisms of reptile adaptation to climate change. The observed phenotypic plasticity and potential for evolutionary adaptation highlight the importance of both short-term and long-term responses in mitigating the impacts of climate change. Phenotypic plasticity allows for immediate adjustments to changing environmental conditions, while genetic variation and evolutionary processes enable populations to track and adapt to these changes over longer timescales (Urban et al., 2013; Ljungström et al., 2015).

Moreover, the findings from Lacerta agilis underscore the significance of maintaining genetic diversity within populations to enhance their adaptive potential. The high level of observed heterozygosity in Swedish populations, despite historical bottlenecks, suggests that even small, isolated populations can retain the genetic resources necessary for adaptation. This has important implications for conservation strategies, emphasizing the need to preserve genetic diversity and facilitate gene flow between fragmented populations (Schlindwein et al., 2022; Bestion et al., 2023).

7 Conservation Implications and Challenges

7.1 Threats to Reptile Populations

Reptile populations are facing significant threats due to climate change, which manifests in various forms such as habitat fragmentation, increased temperatures, and resource depletion. Habitat fragmentation, often exacerbated by human activities, disrupts the natural habitats of reptiles, making it difficult for them to find suitable environments for survival and reproduction. This fragmentation can also impede their ability to adapt to changing climates, as seen in other species where habitat fragmentation has disrupted climate adaptation and led to diminished reproductive success (Van Daele et al., 2023). Increased temperatures, a direct consequence of global warming, have profound effects on reptile physiology and behavior. For instance, changes in nest temperatures can significantly impact the phenotype and survival rates of reptile offspring, with warmer temperatures generally reducing incubation time but potentially leading to maladaptive traits (Noble et al., 2018). Additionally, resource depletion, driven by both climate change and human exploitation, further threatens reptile populations. The crocodile lizard, for example, is experiencing a significant reduction in its habitat due to climate change, which is compounded by over-exploitation for traditional medicine and the pet trade.

The combined effects of these threats create a challenging environment for reptile conservation. The sensitivity of reptiles to temperature changes, due to their ectothermic nature, makes them particularly vulnerable to climate-induced habitat alterations. Studies have shown that reptiles exhibit a range of responses to climate change, from shifts in geographic ranges to changes in reproductive behaviors (Diele-Viegas and Rocha, 2018). However, the ability of reptiles to adapt to these changes is often limited by the rapid pace of environmental change and the additional pressures of habitat fragmentation and resource depletion. This underscores the urgent need for comprehensive conservation strategies that address these multifaceted threats.

7.2 Conservation Strategies

To mitigate the threats posed by climate change and other anthropogenic factors, several conservation strategies have been proposed and implemented. Assisted migration, which involves relocating species to more suitable habitats, is one such strategy. This approach can help species like the crocodile lizard, whose current habitats are shrinking due to climate change, by moving them to areas with more favorable conditions (Zhang et al., 2022). Habitat management is another critical strategy, focusing on preserving and restoring natural habitats to ensure they remain viable for reptile populations. Effective habitat management can help maintain the ecological balance and provide the necessary resources for reptiles to thrive. For instance, managing nesting sites to ensure optimal thermal conditions can enhance the survival rates of reptile offspring (Putman and Tippie, 2020; Du et al., 2023).

Breeding programs also play a vital role in reptile conservation. These programs aim to increase population sizes and genetic diversity, which are crucial for the long-term survival of species. By breeding reptiles in controlled environments and then reintroducing them into the wild, conservationists can bolster populations that are at risk of extinction. Additionally, the use of environmental DNA (eDNA) has emerged as a valuable tool for monitoring reptile populations and informing conservation efforts. eDNA techniques can detect the presence of elusive or rare species, providing critical data for conservation planning and management (Simões et al., 2022; Nordstrom et al., 2022). Overall, a combination of assisted migration, habitat management, breeding programs, and advanced monitoring techniques like eDNA can help address the conservation challenges faced by reptiles in the face of climate change.

8 Future Directions and Research Gaps

8.1 Need for Longitudinal Studies

Longitudinal studies are crucial for understanding the long-term impacts of climate change on reptile populations. These studies involve tracking the same populations over extended periods, allowing researchers to observe changes in phenotypic traits, population dynamics, and genetic adaptations in response to shifting environmental conditions. The importance of such studies is underscored by the current gaps in our understanding of how reptiles adapt over time. For instance, while phenotypic plasticity has been shown to play a significant role in immediate responses to climate variation, the long-term evolutionary consequences remain unclear (Noble et al., 2018). Longitudinal studies can provide insights into whether observed phenotypic changes are temporary adjustments or indicative of more permanent genetic adaptations.

Moreover, longitudinal data can help identify critical periods in the life cycle of reptiles that are most sensitive to climate change. This information is vital for developing effective conservation strategies. For example, studies have shown that early developmental environments, such as nest temperatures, can have lasting effects on reptile phenotypes and survival rates. By tracking these effects over multiple generations, researchers can better predict how future climate scenarios might impact reptile populations and identify which species or populations are most at risk. This approach can also help in understanding the role of genetic accommodation and assimilation in facilitating or hindering adaptive responses to climate change (Kelly 2019).

8.2 Role of Genomics in Studying Adaptations

The advent of genomic technologies has opened new avenues for studying the adaptive responses of reptiles to climate change. Genomic studies can uncover the molecular pathways and genetic variations that underlie phenotypic adaptations, providing a deeper understanding of how reptiles cope with changing environments. For instance, functional genomics approaches have identified specific genes and regulatory regions that are under selection in response to environmental stressors, such as temperature and salinity (Li et al., 2021; Valero et al., 2021). These findings highlight the potential of genomics to reveal the genetic basis of adaptive traits and the evolutionary processes driving these adaptations.

Furthermore, integrating genomic data with ecological and phenotypic information can enhance our ability to predict species' responses to climate change. Eco-evolutionary models that incorporate genomic data can provide more accurate projections of species distributions and adaptive potential under different climate scenarios. However, there is a need for standardized and replicable genomic methodologies to facilitate large-scale studies across diverse reptile species. This will require collaborative efforts to develop comprehensive genomic databases and analytical tools that can be used to study the adaptive potential of reptiles on a global scale (Sun et al., 2020).

8.3 Integrating Climate Models into Conservation Planning

Integrating climate models into conservation planning is essential for developing proactive strategies to mitigate the impacts of climate change on reptile populations. Predictive tools that combine climate projections with ecological and evolutionary data can guide conservation actions by identifying areas and species at greatest risk. For example, species distribution models that incorporate both ecological and genetic data can provide more accurate predictions of how reptile ranges might shift in response to climate change (Waldvogel et al., 2020). These models can also help identify potential refugia and corridors that facilitate species migration and gene flow, which are critical for maintaining genetic diversity and adaptive potential.

Moreover, climate models can inform the design of conservation interventions, such as habitat restoration and assisted migration. By predicting how different climate scenarios might affect habitat suitability and connectivity, conservationists can prioritize actions that enhance the resilience of reptile populations. For instance, understanding the impact of climate change on nesting sites and thermal environments can help in the development of strategies to protect critical habitats and ensure the survival of vulnerable species (Noble et al., 2018). Integrating climate models into conservation planning thus provides a science-based framework for making informed decisions that can help safeguard reptile biodiversity in the face of ongoing climate change.

9 Concluding Remarks

This study explored the phenotypic and evolutionary adaptations of reptiles to climate change, revealing their ability to survive in changing environments through diverse adaptive strategies. Phenotypic plasticity, as a short-term response mechanism, allows reptiles to adjust their morphology, behavior, and physiology to adapt to environmental temperatures and climate fluctuations. Additionally, behavioral shifts, changes in body size and coloration, as well as reproductive adjustments, demonstrate the flexibility of reptiles in responding to climate change. Long-term evolutionary adaptations are facilitated by genetic variation and natural selection, enabling reptiles to adapt over longer time scales. Studies indicate that certain genes are under positive selection for coping with environmental stress, playing a crucial role in the success of populations in future climates.

Reptiles play essential ecological roles in ecosystems, and their resilience is critical for maintaining ecological balance. They contribute to nutrient cycles and species interactions, while their behaviors influence habitat structure and function. The adaptive capacity of reptiles is directly linked to biodiversity and the long-term stability of ecosystems. However, as climate change intensifies, the environmental pressures on reptiles will increase, affecting their survival and the overall health of ecosystems. Thus, the resilience of reptiles is not only essential for the conservation of their populations but also integral to maintaining ecological balance and addressing the challenges posed by climate change.

To ensure the continued survival of reptiles under future climate scenarios, adaptive policies and conservation frameworks are necessary. Assisted migration and habitat management strategies should be implemented to relocate threatened species to suitable new environments and restore connectivity among critical habitats. Additionally, the use of environmental DNA (eDNA) monitoring should be promoted to track population dynamics in real time, providing essential data for conservation planning. Conservation policies must prioritize maintaining genetic diversity within populations and facilitating gene flow between isolated populations. Furthermore, climate models and eco-evolutionary models should be integrated into conservation decision-making to guide future efforts with scientific rigor. These comprehensive strategies will not only support the long-term survival of reptiles but also contribute to global biodiversity and the stability of ecosystems.

Acknowledgments

The authors are grateful to Dr. Jin for critically reading the manuscript and providing valuable feedback.

Conflict of Interest Disclosure

The author affirms that this research was conducted without any commercial or financial relationships that could be construed as a potential conflict of interest.

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