Keeping it cool to take the heat: tropical lizards have greater thermal tolerance in less disturbed habitats.

Global climate change has profound effects on species, especially those in habitats already altered by humans. Tropical ectotherms are predicted to be at high risk from global temperature increases, particularly those adapted to cooler temperatures at higher altitudes. We investigated how one such species, the water anole (Anolis aquaticus), is affected by temperature stress similar to that of a warming climate across a gradient of human-altered habitats at high elevation sites. We conducted a field survey on thermal traits and measured lizard critical thermal maxima across the sites. From the field survey, we found that (1) lizards from the least disturbed site and (2) operative temperature models of lizards placed in the least disturbed site had lower temperatures than those from sites with histories of human disturbance. Individuals from the least disturbed site also demonstrated greater tolerance to high temperatures than those from the more disturbed sites, in both their critical thermal maxima and the time spent at high temperatures prior to reaching critical thermal maxima. Our results demonstrate within-species variability in responses to high temperatures, depending on habitat type, and provide insight into how tropical reptiles may fare in a warming world.

The power of community science to quantify ecological interactions in cities.

Studying animals in urban environments is especially challenging because much of the area is private property not easily accessible to professional scientists. In addition, collecting data on animals that are cryptic, secretive, or rare is also challenging due to the time and resources needed to amass an adequate dataset. Here, we show that community science can be a powerful tool to overcome these challenges. We used observations submitted to the community science platform iNaturalist to assess predation and parasitism across urbanization gradients in a secretive, 'hard-to-study' species, the Southern Alligator Lizard (Elgaria multicarinata). From photographs, we quantified predation risk by assessing tail injuries and quantified parasitism by counting tick loads on lizards. We found that tail injuries increased with age and with urbanization, suggesting that urban areas are risky habitats. Conversely, parasitism decreased with urbanization likely due to a loss of hosts and anti-tick medications used on human companion animals. This community science approach generated a large dataset on a secretive species rapidly and at an immense spatial scale that facilitated quantitative measures of urbanization (e.g. percent impervious surface cover) as opposed to qualitative measures (e.g. urban vs. rural). We therefore demonstrate that community science can help resolve ecological questions that otherwise would be difficult to address.

Physiological Stress Integrates Resistance to Rattlesnake Venom and the Onset of Risky Foraging in California Ground Squirrels.

Using venom for predation often leads to the evolution of resistance in prey. Understanding individual variation in venom resistance is key to unlocking basic mechanisms by which antagonistic coevolution can sustain variation in traits under selection. For prey, the opposing challenges of predator avoidance and resource acquisition often lead to correlated levels of risk and reward, which in turn can favor suites of integrated morphological, physiological and behavioral traits. We investigate the relationship between risk-sensitive behaviors, physiological resistance to rattlesnake venom, and stress in a population of California ground squirrels. For the same individuals, we quantified foraging decisions in the presence of snake predators, fecal corticosterone metabolites (a measure of "stress"), and blood serum inhibition of venom enzymatic activity (a measure of venom resistance). Individual responses to snakes were repeatable for three measures of risk-sensitive behavior, indicating that some individuals were consistently risk-averse whereas others were risk tolerant. Venom resistance was lower in squirrels with higher glucocorticoid levels and poorer body condition. Whereas resistance failed to predict proximity to and interactions with snake predators, individuals with higher glucocorticoid levels and in lower body condition waited the longest to feed when near a snake. We compared alternative structural equation models to evaluate alternative hypotheses for the relationships among stress, venom resistance, and behavior. We found support for stress as a shared physiological correlate that independently lowers venom resistance and leads to squirrels that wait longer to feed in the presence of a snake, whereas we did not find evidence that resistance directly facilitates latency to forage. Our findings suggest that stress may help less-resistant squirrels avoid a deadly snakebite, but also reduces feeding opportunities. The combined lethal and non-lethal effects of stressors in predator-prey interactions simultaneously impact multiple key traits in this system, making environmental stress a potential contributor to geographic variation in trait expression of toxic predators and resistant prey.

Evolutionary dynamics in the Anthropocene: Life history and intensity of human contact shape antipredator responses.

Humans profoundly impact landscapes, ecosystems, and animal behavior. In many cases, animals living near humans become tolerant of them and reduce antipredator responses. Yet, we still lack an understanding of the underlying evolutionary dynamics behind these shifts in traits that affect animal survival. Here, we used a phylogenetic meta-analysis to determine how the mean and variability in antipredator responses change as a function of the number of generations spent in contact with humans under 3 different contexts: urbanization, captivity, and domestication. We found that any contact with humans leads to a rapid reduction in mean antipredator responses as expected. Notably, the variance among individuals over time observed a short-term increase followed by a gradual decrease, significant for domesticated animals. This implies that intense human contact immediately releases animals from predation pressure and then imposes strong anthropogenic selection on traits. In addition, our results reveal that the loss of antipredator traits due to urbanization is similar to that of domestication but occurs 3 times more slowly. Furthermore, the rapid disappearance of antipredator traits was associated with 2 main life-history traits: foraging guild and whether the species was solitary or gregarious (i.e., group-living). For domesticated animals, this decrease in antipredator behavior was stronger for herbivores than for omnivores or carnivores and for solitary than for gregarious species. By contrast, the decrease in antipredator traits was stronger for gregarious, urbanized species, although this result is based mostly on birds. Our study offers 2 major insights on evolution in the Anthropocene: (1) changes in traits occur rapidly even under unintentional human "interventions" (i.e., urbanization) and (2) there are similarities between the selection pressures exerted by domestication and by urbanization. In all, such changes could affect animal survival in a predator-rich world, but through understanding evolutionary dynamics, we can better predict when and how exposure to humans modify these fitness-related traits.

What can we learn from wildlife sightings during the COVID-19 global shutdown?

During the worldwide shutdown in response to the COVID-19 pandemic, many reports emerged of urban wildlife sightings. While these images garnered public interest and declarations of wildlife reclaiming cities, it is unclear whether wildlife truly reoccupied urban areas or whether there were simply increased detections of urban wildlife during this time. Here, we detail key questions and needs for monitoring wildlife during the COVID-19 shutdown and then link these with future needs and actions with the intent of improving conservation within urban ecosystems. We discuss the tools ecologists and conservation scientists can use to safely and effectively study urban wildlife during the shutdown. With a coordinated, multicity effort, researchers and community scientists can rigorously investigate the responses of wildlife to changes in human activities, which can help us address long-standing questions in urban ecology, inspire conservation of wildlife, and inform the design of sustainable cities.

Urban invaders are not bold risk-takers: a study of 3 invasive lizards in Southern California.

Biological invasions threaten biodiversity worldwide, and therefore, understanding the traits of successful invaders could mitigate their spread. Many commonly invasive species do well in disturbed habitats, such as urban environments, and their abilities to effectively respond to disturbances could contribute to their invasiveness. Yet, there are noninvasive species that also do well in disturbed habitats. The question remains whether urban invaders behave differently in urban environments than noninvaders, which could suggest an "urban-exploiting" phenotype. In Southern California, the co-occurrence of invasive Italian wall lizards Podarcis siculus, brown anoles Anolis sagrei, and green anoles A. carolinensis, and native western fence lizards Sceloporus occidentalis offers an opportunity to test whether invasives exhibit consistent differences in risk-taking within human-altered habitats compared with a native species. We predicted that invasive lizards would exhibit more bold behavior by having shorter flight-initiation distances (FIDs) and by being found farther from a refuge (behaviors that would presumably maximize foraging in low-risk environments). Invasive populations had similar or longer FIDs, but were consistently found at distances closer to a refuge. Collectively, invasive lizards in urban habitats were not bolder than a native species. Reliance on nearby refuges might help species successfully invade urban habitats, and if a general pattern, may pose an added challenge in detecting or eliminating them.

Dewlap size in male water anoles associates with consistent inter-individual variation in boldness.

Male sexually selected signals can indicate competitive ability by honestly signaling fitness-relevant traits such as condition or performance. However, behavior can also influence contest outcomes; in particular, boldness often predicts dominance rank and mating success. Here, we sought to determine whether male ornament size is associated with consistent individual differences in boldness in water anoles Anolis aquaticus. We measured the relative size of the dewlap, a flap of skin under the chin that is a sexually selected ornament in Anolis lizards, and tested for associations with responses to a novel and potentially risky environment: time to emerge from a refuge into an arena and number of head scans post-emergence. We found that individuals consistently differed in both time to emerge and head scanning (i.e., individual responses were repeatable), and that dewlap size was negatively related to number of head scans. This suggests that ornament size could indicate male boldness if scanning represents antipredator vigilance. We found that males that had larger relative dewlaps were also in better body condition, but boldness (i.e., head scanning) was not related to condition. Lastly, we found consistent differences in behavior between trials, showing that anoles were becoming habituated or sensitized to the testing arena. Overall, our study shows that in addition to indicating condition and performance, dewlap size could also honestly indicate male boldness in Anolis lizards.

Downsizing for downtown: Limb lengths, toe lengths, and scale counts decrease with urbanization in western fence lizards (Sceloporus occidentalis).

Urbanization induced habitat loss and alteration causes significant challenges for the survival of many species. Identifying how species respond to urbanization can yield insights for the conservation of wildlife, but research on reptiles has been narrowly-focused. We compared morphology among four populations of western fence lizards (Sceloporus occidentalis) to determine whether a common native species affected by urbanization exhibits morphological differences consistent with habitat use. We quantified habitat differences across four sites in Los Angeles County, California, USA that varied in level of urbanization, measured how lizards used microhabitats, and assessed variation in body size, limb length, toe length, and scalation of lizards collected from each site. Urban and suburban populations of fence lizards mostly used human-made substrates while lizards from more natural areas mostly used natural woody substrates. Lizards from the most urban site also exhibited the widest breadth of substrates used, indicating that urban sites might offer more variable microhabitats. Urban lizards had reduced limb lengths and toe lengths consistent with how they used microhabitats and other habitat characteristics (e.g., percent impervious surface cover). Urban lizards also had fewer dorsal scales, which might be associated with changes in ambient temperature (e.g., urban heat island effect), given that lizards with fewer and larger scales typically have reduced evaporative water loss. Our results uniquely differ from past studies on lizard responses to urbanization, indicating that work on diverse taxa is necessary to assess the potential varied pathways of morphological adaptations to urban environments.

Fear no colors? Observer clothing color influences lizard escape behavior.

Animals often view humans as predators, leading to alterations in their behavior. Even nuanced aspects of human activity like clothing color affect animal behavior, but we lack an understanding of when and where such effects will occur. The species confidence hypothesis posits that birds are attracted to colors found on their bodies and repelled by non-body colors. Here, we extend this hypothesis taxonomically and conceptually to test whether this pattern is applicable in a non-avian reptile and to suggest that species should respond less fearfully to their sexually-selected signaling color. Responses to clothing color could also be impacted by habituation to humans, so we examine whether behavior varied between areas with low and high human activity. We quantified the effects of four T-shirt colors on flight initiation distances (FID) and on the ease of capture in western fence lizards (Sceloporus occidentalis), and we accounted for detectability against the background environment. We found no differences in lizard behavior between sites. However, lizards tolerated the closest approaches and were most likely to be captured when approached with the T-shirt that resembled their sexually-selected signaling color. Because changes in individual behavior affect fitness, choice of clothing color by people, including tourists, hikers, and researchers, could impact wildlife populations and research outcomes.

Behavioral thermal tolerances of free-ranging rattlesnakes (Crotalus oreganus) during the summer foraging season.

Increasing temperature due to climate change is one of the greatest challenges for wildlife worldwide. Behavioral data on free-ranging individuals is necessary to determine at what temperatures animals modify activity as this would determine their capacity to continue to move, forage, and mate under altered thermal regimes. In particular, high temperatures could limit available surface activity time and time spent on fitness-related activities. Conversely, performance, such as feeding rate, can increase with temperature potentially having positive fitness effects. Here, we examine how the hunting behaviors of free-ranging Northern Pacific Rattlesnakes (Crotalus oreganus) associate with air temperature and body temperature. We continuously recorded snakes in the field using videography, capturing behaviors rarely considered in past studies such as movements in and out of refuge and strikes on prey. We found that as mean daily air temperature increased, hunting activity and the likelihood of hunting at night decreased, while the number of movements and distance moved per day increased. Snakes typically retreated to refuge before body temperatures reached 31°C. Body temperatures of snakes hunting on the surface were lower compared to temperatures of non-hunting snakes in refuge in the morning, while this relationship was inverted in the afternoon. Snake body size influenced the disparity of these temperatures. Finally, strike initiation and success occurred across a wide range of body temperatures, indicating hunting performance may not be strongly constrained by temperature. These results on the temperatures at which free-ranging rattlesnakes exhibit fitness-related behaviors could be valuable for understanding their vulnerabilities to future climates.

Activity cycles and foraging behaviors of free-ranging sidewinder rattlesnakes (Crotalus cerastes): the ontogeny of hunting in a precocial vertebrate.

Predators often employ a complex series of behaviors to overcome antipredator defenses and effectively capture prey. Although hunting behaviors can improve with age and experience, many precocial species are necessarily effective predators from birth. Additionally, many predators experience innate ontogenetic shifts in predatory strategies as they grow, allowing them to adapt to prey more appropriate for their increased size and energetic needs. Understanding how the relative roles of innate age-specific adaptation and learning have evolved requires information on how predation behavior develops in situ, in free-ranging predators. However, most of the research on the ontogeny of predation behavior is based on laboratory studies of captive animals, largely due to the difficulty of following newborn individuals in nature. Here, we take advantage of the unique tracks left by juveniles of a precocial viperid, the sidewinder rattlesnake (Crotalus cerastes), which we used to follow free-ranging snakes in the field. We recorded details of their ambush hunting behavior, and compared the behaviors of these juveniles to adult snakes that we monitored in the field via radio telemetry. Although juvenile and adult behaviors were similar in most respects, we did find that adults chose more effective ambush sites, which may be due to their increased experience. We also found that juveniles (but typically not adults) perform periodic tail undulations while in ambush, and that juveniles displayed slightly different activity cycles. Both of these latter differences are likely the result of age-specific adaptations for juveniles' greater reliance on lizards versus small mammals as prey. We also compared the general predatory behavior of sidewinders to that of other species in the genus Crotalus. These findings will provide important baseline field information for more detailed empirical research on the ontogeny of predation behavior in precocial vertebrates.