Thermal stress and developmental disruption in the Neotropical lizard Tropidurus torquatus under climate change scenarios

1 Introduction

Climate change and global warming are among the main threats to Earth’s ecosystems (Sinervo et al., 2010), with documented impacts across a wide range of biological systems (Hughes et al., 2018; Warren et al., 2018) (Warren et al., 2018). Over the last century, the global mean temperature has risen by 0.74°C, and predictions indicate an additional increase of 4.4 ± 1.1°C over the next 100 years (Kirtman et al., 2013). Moreover, warming has impacts on many environmental aspects, such as precipitation (Hughes, 2000) and atmospheric CO₂ concentration (Sabine et al., 2004), which can amplify the frequency and intensify extreme weather events (Grimmond, 2007). Because temperature directly constrains multiple biological processes in ectothermic organisms, rising environmental temperatures are expected to have pervasive consequences across life stages (Booth, 2006; Angilletta et al., 2009).

As a direct result of climate change, many species have shown alterations from distribution and occurrence (Thomas et al., 2006) to genetic and phenotypic diversity (Booth, 2006; Habel et al., 2011). Understanding and predicting how phenotypic traits respond to ongoing climate change remains one of the central challenges in contemporary biology (Bonino et al., 2015; Ficetola et al., 2016; Campbell-Staton et al., 2017). Such climate-driven responses have been documented across multiple levels of biological organization and taxonomic groups, ranging from ecosystem-scale impacts on coral reefs to population-level responses in terrestrial vertebrates under sustained warming (Hughes et al., 2018; Warren et al., 2018).

Research on the biological consequences of global warming has expanded considerably in recent decades, with most studies focusing on adult stages including investigations of thermal physiology, behavior, and performance in lizards (Angilletta et al., 2002; Huey et al., 2012; Campbell-Staton et al., 2017; Diele-Viegas et al., 2018), statistical modeling of species distributions and extinction risk (Sinervo et al., 2010; Pontes-da-Silva et al., 2018) and migration patterns (La Sorte et al., 2017). In contrast, a growing number of experimental studies have demonstrated that incubation temperature can profoundly affect development, morphology, and physiology in reptiles, particularly lizards (Goodman and Walguarnery, 2007; Ji et al., 2007; Goodman, 2008; De-Lima et al., 2022; Rossigalli-Costa and Kohlsdorf, 2022).

The influence of temperature on development is well documented in the literature, as incubation temperature also plays a pivotal role in sex determination in some ectothermic vertebrates, including several lineages of fishes, amphibians, and reptiles (Conover and Kynard, 1981; Valenzuela et al., 2019; Tomillo and Spotila, 2020; Krueger and Janzen, 2023). Moreover, elevated temperatures are consistently associated with shorter incubation durations, but concurrently increase mortality, reduce hatchling size, skew sex ratios, and alter thermal physiology (Goodman, 2008; Scott and Johnston, 2012; Vargesson and Fraga, 2017; De-Lima et al., 2022).

Temperature has also been linked to the occurrence of developmental anomalies across vertebrates (Haghighi et al., 2021; Dillenburg et al., 2024). Thermally induced developmental anomalies have been reported for many lineages from fish to birds and non-avian reptiles (Noiva et al., 2014; Idrisova, 2018; Ashaf-Ud-Doulah et al., 2021). In lizards, a recent study has shown pronounced thermal sensitivity during embryogenesis, where even moderate increases in incubation temperature can disrupt axial patterning, limb formation, and craniofacial symmetry (Sanger et al., 2021).

Lizards’ reproductive modes range from viviparous, ovoviviparous to oviparous species (Blackburn, 2016). In oviparous lizards, embryonic development occurs outside the maternal body and is therefore more directly influenced by environmental factors such as humidity, oxygen availability, and temperature (Scott and Johnston, 2012; Flewelling and Parker, 2015). In contrast to free-living juveniles and adults, which can partially buffer environmental variation through behavioral thermoregulation (Bogert, 1949; Muñoz, 2022) embryos developing within eggs experience incubation temperatures largely passively and cannot behaviorally avoid thermal stress (Shine and Harlow, 1993; Refsnider and Janzen, 2010). Hence, considering that developmental anomalies can be influenced by extrinsic factors, species with oviparous development are expected to show a higher incidence of environmentally induced malformations during incubation, which represents a particularly vulnerable life stage due to the absence of behavioral thermoregulation (Hopkins et al., 2013).

The Cerrado biome, a Neotropical savanna, is among the biomes most severely impacted by climate change and global warming, as well as by the widespread loss of native vegetation driven by agricultural expansion (Hofmann et al., 2021; Rodrigues et al., 2022). This environmental vulnerability, combined with the limited amount of research focused on Neotropical herpetological fauna, creates a concerning scenario of multiple knowledge gaps regarding the effects of climate change on Neotropical species (Urbina-Cardona, 2008; Pio Caetano Machado et al., 2023).

Tropidurus torquatus is a South American lizard widely distributed across the Cerrado biome (Carvalho, 2013). Recent studies have documented population-level shifts in growth rate, body size, and age at sexual maturity associated with a 1–2°C increase in ambient temperature over the last four decades in this species (Piantoni et al., 2019). T. torquatus produces clutches averaging 5–10 eggs (Wiederhecker et al., 2002; Ortiz et al., 2014), has a well-established embryonic staging table (Py‐Daniel et al., 2017), and can be easily maintained in captivity (Rossigalli‐Costa et al., 2021), making it suitable for experimental and embryological studies. Furthermore, recent experimental evidence shows that incubation temperature influences neonatal morphology and thermal physiology in this species, supporting its use as a model organism for studies on temperature-dependent development in Neotropical reptiles (De-Lima et al., 2022). Here, we provide the first quantitative assessment of developmental anomalies in T. torquatus under different incubation temperature regimes that reflect thermal conditions potentially experienced in natural nests under ongoing climate warming scenarios.

2 Materials and methods

2.1 Animal collection and husbandry

We collected 159 eggs from 27 pregnant adult females of T. torquatus during the reproductive season of the species from October to December (Wiederhecker et al., 2002) in two field campaigns conducted in 2017 and 2018. Females were captured by noose in urban areas of Brasilia, Distrito Federal, Brazil (15°50′20.2″S, 47°58′50.2″W; 15°45′49.5″S, 47°52′05.5″W; and 15°56′57.0″S, 47°56′05.4″W). Once captured, females were visually marked using non-toxic, water-based paint for identification and housed in terraria, with a maximum of 5 females per terrarium. They were provided ad libitum access to food and water, as well as heat plates for thermoregulation. Captures were licensed by the Brazilian Ministry of the Environment (ICMBio/SISBIO permits n° #63226–1 and #63226-2). The experimental procedure described herein was fully approved and performed in accordance with the University of Brasilia Ethics Committee (CEUA-UnB permit n° 116/2017). Also, the experimental design is in accordance with the ARRIVE guidelines 2.0 (Percie du Sert et al., 2020).

2.2 Egg collection and treatments

Terrariums were checked daily for eggs. Upon detection of a clutch, the eggs were collected and individually placed in labeled plastic containers filled with moist vermiculite, maintained a saturated humidity environment, following the same protocol and substrate hydration ratio described in De-Lima et al. (2022). To minimize the genetic effects, eggs were randomly assigned to one of five incubation treatments. Only eggs considered viable at oviposition were included in the incubation experiment. Egg viability was assessed by visual inspection under ambient light, where eggs exhibiting a clearly visible embryonic pole through shell translucency, characterized by a distinct reddish embryonic disc, were selected for incubation. Clutches compromised by desiccation or laid under maternal stress were not used in the experiment.

Treatments and sample numbers were as follows: (i) an artificial nest constructed in the field beneath rocks and logs to simulate natural nesting conditions (n = 30; mean temperature = 28.6°C); and four incubator-based treatments maintained under controlled laboratory conditions: (ii) 30.0°C (n=33), constant temperature; (iii) 32.5°C (n=28), constant temperature; (iv) 32.5°C (n=31), with daily thermal variation reaching a peak of 39.0°C for 2 h; and (v) 35°C (n=22), constant temperature.

The artificial nest treatment was designed to reproduce nesting microhabitats commonly used by females in the study population, based on repeated field observations of oviposition occurring at wall corners beneath concrete slabs. Accordingly, a wire-mesh enclosure was installed at an external wall corner and covered with concrete slabs, creating a sheltered nesting structure. A temperature datalogger was placed within the structure and programmed to record temperatures at 10-min intervals throughout incubation. Across 20,069 temperature records, the artificial nest exhibited a mean incubation temperature of 28.7°C. Thermal variation was asymmetric, with daily maxima contributing disproportionately to overall variability: the mean temperature was approximately 0.7°C higher than the median recorded value (28.0°C). Mean daily maximum and minimum temperatures were 39.1°C and 22.7°C, respectively, while absolute extremes ranged from 20.9°C to 45.8°C.

Artificial incubation temperatures were selected to encompass the range of mean and extreme thermal conditions documented for natural nests of open-habitat lizards, as well as values projected to become more frequent under near-term climate-warming scenarios. Experimental rooms housing the incubators were maintained under a standardized 12:12 h light–dark cycle, ensuring consistent photoperiod conditions across all incubator-based treatments (ii–v). Throughout the entire female oviposition period (November to February), all incubation treatments were simultaneously available, such that each egg was randomly assigned to a treatment immediately after oviposition, regardless of laying date, ensuring effective randomization and preventing confounding effects between treatment and oviposition timing.

Incubator-based treatments were maintained so that the last egg introduced remained under incubation for up to 90 days, based on the species-specific embryonic staging framework for Tropidurus torquatus, which indicates an incubation duration of approximately 75 days at 30°C (Py‐Daniel et al., 2017). This extended period ensured that all embryos had sufficient time to complete development across treatments. For the artificial nest treatment, the end of incubation was defined as the mean hatching time of the first hatchlings, followed by a conservative additional incubation period exceeding the maximum hatching interval observed, to ensure that no viable embryos were prematurely excluded. No incubation-related inconsistencies were detected, and no eggs or hatchlings were removed prior to the completion of their expected developmental period.

2.3 Hatchling and embryo collection and anomaly classification

Once hatched, the individuals were removed from their respective treatments and taken from the containers. Hatchlings were euthanized using an overdose of anesthetic consisting of pentobarbital sodium (100 mg/ml) and lidocaine hydrochloride (20 mg/ml) mixed in a 1:1 ratio. After euthanasia, specimens were fixed in Baker’s solution for 48 hours and then preserved in 70% ethanol.

Eggs that did not hatch in the average developmental time for the species – between three and five months (Wiederhecker et al., 2002; Py‐Daniel et al., 2017) – or were visibly dead by the exterior aspect of the shell were removed from the incubators, fixed in Baker’s solution for 48 hours, and preserved in 70% ethanol. To facilitate embryo extraction, we rehydrated the eggs in water at 40°C for 24 hours. We extracted the embryos using scissors and tweezers under a stereomicroscope and stored them in 70% ethanol. Eggs that failed to hatch and contained embryos at very early developmental stages could not be reliably assessed for malformations. Therefore, these eggs were classified as embryonic mortality rather than as malformed individuals.

Neonates were radiographed with a Micro X-ray machine (Faxitron DX) to assess skeletal morphology and screen for possible anomalies and photographed using a Canon^® EOS 40D camera. The embryos were staged using the available staging table for the species (Py‐Daniel et al., 2017).

We identified and described the anomalies, following the nomenclature classification system proposed by Dillenburg et al. (2024). We also classified the anomalies according to Bell et al. (2006) as mild, when their presence created little to nonfunctional impairment to the individual; moderate, when the anomalies could generate some competitive disadvantage to the animal in natural conditions; and severe, when the presence of the anomaly greatly impacted the individual’s ability to survive and often resulted in the animal’s death.

2.4 Statistical analysis

To evaluate the effects of incubation temperature on embryonic outcomes, we used generalized linear mixed models (GLMMs) with binomial error distribution and a logit link function. Three biologically distinct models were fitted: (1) egg mortality, (2) neonate malformations (among hatchlings only), and (3) undesirable outcomes (eggs that either died or produced malformed hatchlings). Because all severe malformations occurred in embryos that did not hatch, severe outcomes were fully contained within mortality and therefore did not constitute an independent response variable.

For each model, the response consisted of the number of affected and unaffected eggs per temperature treatment, with temperature (°C) as the continuous predictor. To account for non-independence among eggs originating from the same female, clutch identity (each clutch corresponding to a single female) was included as a random intercept in all models. Each fitted model was formally compared against an intercept-only null model, which assumes a constant probability of the response independent of incubation temperature. Statistical significance of temperature effects was assessed using both Wald tests on fixed-effect coefficients and likelihood ratio tests comparing full models against intercept-only null models with identical random-effects structure. Model performance was evaluated by likelihood-based model fit, and effect size was expressed as regression coefficients (β), odds ratios (OR=eβ), Wald z statistics, and 95% confidence intervals.

Model assumptions were evaluated using diagnostic procedures appropriate for binomial generalized linear mixed models. Overdispersion was assessed using Pearson residual deviance ratios and simulation-based dispersion tests. Dispersion ratios were close to or below 1 for all models (mortality: 0.91; hatchling malformations: 0.96; undesirable outcomes: 0.98), indicating no evidence of overdispersion. Simulation-based residual diagnostics implemented in the DHARMa package (Hartig, 2024) further confirmed the adequacy of model fit, showing no evidence of zero inflation for any response (all p > 0.4). These results support the appropriateness of the binomial error structure and model specification used in the analyses.

Analyses and plots were performed in R version 4.4.0 (R Core Team, 2024).

3 Results

3.1 Occurrence of morphological anomalies

Twelve distinct types of anomalies were identified and classified as mild, moderate and severe according to Bell et al. (2006) and Dillenburg et al. (2024) (Table 1).

Table 1. Morphological anomalies observed in embryos (E) and hatchlings (H) of Tropidurus torquatus incubated under five thermal treatments.

Anomalies are grouped according to severity class (Mild, Moderate, Severe). Numbers represent the count of individuals exhibiting each anomaly within each treatment.

Mild anomalies included abnormal pigmentation, axial sacro-caudal asymmetry, caudal amputation, deformed caudal vertebrae, hemivertebrae, and twisted tail. Moderate anomalies included hemimelia, a congenital reduction or absence of part of a limb, and oligodactyly, characterized by a reduced number of digits. Severe anomalies comprised four distinct types: axial torsion, involving twisting of the vertebral column; displacements affecting multiple axial and appendicular regions; exencephaly, a severe cranial defect in which the developing brain is exposed due to failure of skull formation; and extreme general anomalies, characterized by widespread malformations throughout the body, encompassing both axial and limb structures (Table 1). Skeletal anomalies in hatchlings are illustrated in the radiographs presented in Figure 1, whereas embryonic malformations are shown in the photographic documentation in Figure 2.

3.2 Frequency of morphological anomalies

From the total 159 eggs incubated, 84 hatchlings were obtained; 30 of these (35.7%) exhibited morphological anomalies (Table 2). Seventy-five eggs did not hatch, and among these non-hatched eggs it was possible to locate embryos at intermediate to advanced developmental stages in 30 cases (embryonic stages 36–42; Py‐Daniel et al., 2017), 11 of which presented malformations. In the remaining eggs (n = 45), embryos were either at very early developmental stages and/or could not be reliably located. Because these eggs showed no evidence of hatching and could not be morphologically assessed, they were conservatively classified as embryonic mortality.

Table 2. Incubation outcomes and frequency of morphological anomalies in Tropidurus torquatus across thermal treatments.

Numbers refer to affected embryos (E) or hatchlings (H).

Among malformed individuals, 9 embryos and 23 hatchlings exhibit only one malformation; 2 embryos exhibited two simultaneous malformation types, while 4 hatchlings showed two malformations; 2 hatchlings exhibited three malformations, and 1 additional hatchling exhibited four malformations, totaling 54 malformation occurrences distributed across 41 individuals (30 hatchlings and 11 embryos) (Tables 1, 2).

3.3 Influence of incubation temperature

Egg mortality was significantly influenced by incubation temperature with a positive fixed effect of temperature (Table 3; GLMM; β = 0.580 ± 0.121 SE; z = 4.84; p< 0.001), corresponding to a 79.4% increase in the odds of mortality per 1 °C (OR = 1.79; Figure 3). This effect was further supported by a likelihood ratio test comparing the full model against an intercept-only null model (χ² = 30.98, df = 1, p< 0.001). Clutch identity (female), modeled as a random intercept, showed substantial variation in baseline mortality risk (variance = 2.957; SD = 1.720).

Table 3. Summary of generalized linear mixed models (GLMMs) testing the effects of incubation temperature on developmental outcomes in Tropidurus torquatus.

Models were fitted with a binomial error distribution and a logit link function. Incubation temperature (°C) was included as a fixed effect, and clutch identity (female) was included as a random intercept. For each model, effect sizes are reported as regression coefficients (β), odds ratios (OR = e^β), and 95% confidence intervals. Statistical support for temperature effects is shown both by Wald tests (Wald p-value) and by likelihood ratio tests comparing each full model to its corresponding intercept-only null model (ΔDeviance and LRT p-value).

In contrast, malformations detected among hatchlings did not show a significant relationship with temperature (Table 3; GLMM; β = −0.033 ± 0.154 SE; z = −0.218; p = 0.828; Figure 3), and the temperature effect did not improve model fit relative to the null model (likelihood ratio test: χ² = 0.046, df = 1, p = 0.83). Variation among clutches was detected as a random intercept effect (variance = 1.600; SD = 1.265). However, this pattern does not indicate reduced developmental sensitivity, as many of the most severe anomalies occurred in embryos that failed to hatch and are therefore incorporated into the mortality component rather than into the hatchling model.

When mortality and any type of hatchling malformation were combined into a single category of undesirable outcomes (i.e., events reflecting any developmental failure), temperature again exerted a significant positive effect (Table 3; GLMM; β = 0.311 ± 0.120 SE; z = 2.60; p = 0.009), resulting in a 36.5% increase in the odds of such outcomes per 1 °C (OR = 1.37; Figure 3). This effect was also supported by a significant improvement over the null model (likelihood ratio test: χ² = 7.23, df = 1, p = 0.007). Female identity also showed high variance in baseline risk (variance = 4.380; SD = 2.093). A direct comparison between fitted curves for mortality and for mortality plus malformations is shown in Figure 3.

4 Discussion

4.1 Thermal effects on mortality and congenital anomalies

To date, this is the first systematic experimental estimate of the natural rates of animals with anomalies in a Neotropical lizard species. This is also the first record of multiple congenital anomalies in Tropidurus torquatus, and all anomalies identified were previously unreported for the species. This represents a relevant contribution to the understanding of temperature-dependent developmental instability in Neotropical squamates, a group for which embryological and teratological data remain severely limited (Dillenburg et al., 2024; Horváth, 2025). However, the scarcity of comparable datasets from South American reptiles continues to limit broader inferences about how environmental warming affects early developmental trajectories (Bodensteiner et al., 2021; Hall et al., 2022).

It was evident that the direct effect of high incubation temperatures was primarily manifested as increased egg mortality. A clear positive association between rising incubation temperature and embryonic lethality has been repeatedly documented in lizards, with upper thermal limits between 33–36°C commonly inducing near-complete mortality (Hall and Sun, 2021; De-Lima et al., 2022; Rossigalli-Costa and Kohlsdorf, 2022). Field studies show that lizards frequently nest in highly exposed microhabitats where soil temperatures at nesting depth can reach or exceed ~36°C (Angilletta et al., 2009; Telemeco et al., 2009), indicating that nests constructed in open environments may periodically experience thermal conditions capable of producing similar developmental outcomes.

The frequency of individuals exhibiting congenital anomalies ranged from 21% to 44% across treatments, values that exceed the typical ranges reported for other reptile species under natural or experimental conditions (Idrisova, 2018; Horváth, 2025). Experimental studies on Natrix natrix and Lacerta agilis have documented that deviations and malformations typically occur in 8–24% of hatchlings, with the highest values associated with elevated incubation temperatures (Idrisova, 2018). Similar ranges were identified across multiple reptile taxa in a broad survey of congenital abnormalities, where malformation frequencies generally remained below 10–20% in natural populations (Horváth, 2025). Because all anomalies in the present dataset were observed directly in embryos and hatchlings, these values likely represent the full spectrum of developmental outcomes rather than a subset filtered by post-hatching survival. Previous studies indicate that elevated incubation temperatures substantially increase both the incidence and severity of congenital defects, particularly those affecting axial and caudal structures (Telemeco et al., 2009; Idrisova, 2018). In our dataset, however, a temperature-dependent effect was not detected among hatchlings only because embryos with the most severe defects failed to hatch and were therefore incorporated into the mortality component rather than into the malformation model.

Tail anomalies are commonly found in lizards and have been described in many families (Passos et al., 2016; Kolenda et al., 2017; Koleska and Jablonski, 2018; Handal and Christopoulos, 2022). In many cases, however, the anomalies found in the tails result from a failure of tail regeneration and do not constitute a developmental anomaly proper (Barr et al., 2019). In the present dataset, tail anomalies accounted for 33 out of 54 malformation occurrences (61.1%). When these caudal anomalies are excluded, the remaining 21 non-caudal malformations represent 38.9% of all recorded defects, a proportion that still lies within the upper range reported for reptiles (Idrisova, 2018) and remains substantially higher than most field-based estimates (Bárcenas-Ibarra et al., 2015; Horváth, 2025). A major source of underestimation is an “anomaly paradox,” in which the probability of detection is inversely related to the severity of the malformation. Mild anomalies often affect structures that are expendable or capable of regeneration – most notably the tail – which means that such defects can persist into adulthood with minimal fitness consequences and may go unnoticed in wild individuals (Bateman and Fleming, 2009). In contrast, moderate and severe malformations, especially those involving cranial, vertebral, or limb development, are strongly associated with reduced viability and high embryonic or neonatal mortality (Shine and Harlow, 1993; Idrisova, 2018). As a result, these more consequential anomalies rarely appear in adult populations, creating a natural sampling bias that gives the false impression that congenital defects are rare in lizards (Lowenborg et al., 2011; Bateman et al., 2022).

4.2 Developmental vulnerability under climate-change scenarios

Experimental temperatures that increase mortality rate and developmental anomalies fall within the range of extreme thermal conditions already documented for open microhabitats in open dry savannas (Elphick, 1998; Telemeco et al., 2009). Climate projections indicate that the Cerrado biome will experience higher mean temperatures and more frequent and intense heat waves (Hofmann et al., 2021), a pattern that is likely to raise nest temperatures to levels comparable to those associated with the highest rates of lethal and sublethal outcomes in experimental studies. Climate change is also expected to increase thermal variability within nests, not only mean temperatures. Experimental studies using ecologically realistic, fluctuating thermal regimes show that broader thermal variance alone can elevate embryonic mortality and intensify developmental disruption under warming scenarios (Valenzuela et al., 2019).

Temperature-driven embryonic mortality is a well-established mechanism through which climate warming may erodes oviparous vertebrate populations, and the strong temperature dependence observed here reinforces that even modest increases in nest temperature can push embryos beyond their developmental limits (Du and Shine, 2015; Noble et al., 2018; Sanger et al., 2021). This thermal sensitivity is consistent with a growing body of experimental evidence demonstrating that elevated incubation temperatures disrupt axial patterning, skeletal formation, and neural development in reptiles (Telemeco et al., 2009; Idrisova, 2018; Sanger et al., 2021) and strongly modulate morphological, physiological and performance traits in lizards, including Tropidurus (De-Lima et al., 2022; Rossigalli-Costa and Kohlsdorf, 2022).

It is important to note that our experimental design included both constant and fluctuating thermal regimes, reflecting the range of incubation conditions experienced by eggs in natural nests. While approaches such as constant temperature equivalents and development-rate weighted metrics have been proposed to account for the nonlinear effects of temperature on embryonic development (Georges, 1989; Georges et al., 1994), subsequent modeling work has demonstrated that thermal variability itself can alter developmental trajectories independently of mean temperature (Georges et al., 2005). Accordingly, our primary objective was to quantify biologically realistic developmental outcomes under distinct thermal scenarios rather than to standardize developmental time across treatments. Empirical studies further indicate that thermal variance can impose developmental stress and elevate mortality independently of mean temperature (Shine and Harlow, 1993; Telemeco et al., 2009; Du and Shine, 2015).

5 Conclusion

Our study expands previous evidence of temperature-dependent developmental sensitivity in the species (De-Lima et al., 2022) and shows that embryonic development in Tropidurus torquatus is highly vulnerable to rising temperatures, highlighting a critical pathway through which climate warming may affect Neotropical lizards. As suitable, vegetated, and thermally buffered nesting microhabitats decline due to habitat loss in the Cerrado (Rodrigues et al., 2022), females may become increasingly constrained in their ability to avoid extreme nest temperatures, a pattern also documented in lizard species inhabiting open dry savanna formations (Angilletta et al., 2009; Ortega et al., 2019; Dayananda et al., 2021). Such limitations are likely to elevate the frequency of lethal and sublethal developmental outcomes, ultimately reducing effective recruitment. Considering that Tropidurus species already shows climate-associated shifts in growth rate, morphology, physiology, performance, and reproductive timing (Piantoni et al., 2019; De-Lima et al., 2022; Rossigalli-Costa and Kohlsdorf, 2022), integrating embryonic responses into models of population persistence is essential for predicting how Neotropical lizards will cope with accelerating thermal stress.

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Footnotes

Edited by: Luisa Maria Diele Viegas, Federal University of Bahia (UFBA), Brazil

Reviewed by: Thea Gessler, Independent researcher, Ames, IA, United States

Miguel Domínguez Godoy, National Autonomous University of Mexico, Mexico