The Causes and Consequences of Adaptation to a Novel Invader in the Eastern Fence Lizard (Sceloporus Undulatus).


Book Description

Many environments are changing rapidly due to human impacts including habitat alteration, climate change, and the introduction of invasive species. This global environmental change has numerous direct and indirect effects on species that can exert novel pressures. In some cases organisms must adapt to these changes or face extirpation or extinction. Understanding the diversity and extent of these effects of global change, and the processes by which species adapt to it, is critical to future conservation and can provide important insights into ecological and evolutionary processes. My dissertation addresses questions about the effects imposed by invasive species on native species and how native species adapt to associated novel pressures by using a system of native fence lizards (Sceloporus undulatus) subject to differing levels of exposure to an invasive predatory fire ant (Solenopsis invicta). In the southeastern United States, fence lizard populations have adapted to this novel predator via an increase in anti-ant behaviors, including twitching and fleeing which promote escape from and removal of attacking ants; longer hind limbs which support this behavior; and a heightened stress response. Novel pressures induced by invasive species may include lethal (e.g. predation, injury) and sublethal (e.g. altered energy or habitat availability) effects. The nature of these effects may vary with characteristics of an organism (e.g., sex, age, size) and history of exposure to the invader. I explored how the effects of fire ants on fence lizards varied with life stage of the lizard and their evolutionary exposure (adaptation) to the ants. I first investigated the effects of fire ants on the vulnerable life history stage of fence lizard eggs by creating artificial nests in the field and monitoring them daily. I found that fire ants do prey on fence lizard eggs in the field and may predate up to 60% of fence lizard nests in fire ant-invaded areas. Eggs from lizards from fire ant-invaded and -uninvaded sites were equally susceptible to predation, suggesting a lack of adaptation to this threat. I next quantified the effects of fire ants on juvenile and adult lizards from fire ant-invaded and -uninvaded sites by placing them in large field enclosures with natural and reduced densities of fire ants for two weeks. Fire ants reduced survival of adult but not juvenile lizards, regardless of their evolutionary history with fire ants. Juvenile lizards grew less in enclosures with fire ants than in fire ant-free enclosures, but there was no effect of fire ant density on adult growth. This research shows that fire ants have both lethal and sublethal effects on lizards, and that these vary with lizard life stage. I conducted field surveys and manipulations to examine the consequences of behavioral adaptations of fence lizards to fire ants. I exposed lizards from fire ant-invaded and -uninvaded sites to encounters with predatory fire ants and non-threatening native ants, and to simulated attacks by a native predator (a taxidermied American Kestrel). While lizards responded similarly to the native predator, lizards from fire ant-invaded sites showed a generalized increase in responsiveness to all ants. While this behavior is adaptive against fire ants, it may also attract attention from native predators as evidenced by my findings of higher injury rates of lizards at fire ant-invaded sites. At a geographic scale, I found that the presence of fire ants reverses natural latitudinal gradients in behavior, stress responsiveness, and morphology of fence lizards. This implies that fire ants are driving these traits away from historical values that evolved under fire ant-free conditions and suggests potential costs to adapting to fire ants. As a whole, this work shows that invasive species can have broad impacts on native taxa, and that these impacts may vary with the life stage of affected native species and the degree to which they have adapted to the invader. Adaptive responses by native species may be strong enough to reverse pre-existing latitudinal clines in relevant traits over large portions of an affected species' range. However, while native species can adapt to novel invaders, these adaptations may incur also costs that should be considered when assessing population persistence and management in the face of these increasing perturbations.










Advances in the Study of Behavior


Book Description

Advances in the Study of Behavior, Volume 49 provides users with the latest insights in this ever-evolving field. Users will find new information on a variety of species, including social behaviors in reptiles, the behavioral evidence of felt emotions, a section on developmental plasticity, a chapter on covetable corpses and plastic beetles and the socioecological behavior of burying beetles, and a section on the mechanisms of communication and cognition in chickadees. This volume makes another important contribution to the development of the field by presenting theoretical ideas and research findings to professionals studying animal behavior and related fields. Researchers in a variety of behavioral fields will find this longstanding series, initiated over 40 years ago, to be a go-to resource for the study of animal behavior. Initiated over 40 years ago to serve the increasing number of scientists engaged in the study of animal behavior Makes another important contribution to the development of the field Presents theoretical ideas and research to those studying animal behavior and related fields







Geographic Variation in Life History Tactics, Adaptive Growth Rates, and Habitat-specific Adaptations in Phylogenetically Similar Species


Book Description

ABSTRACT: To understand the evolutionary and ecological significance of geographic variation in life history traits, we must understand whether the patterns are induced through plastic or adaptive responses. The Eastern Fence Lizard, Sceloporus undulatus, exhibits countergradient variation (larger body sizes, et cetera, in northern, cooler environments; presumed adaptive) in life history traits across its large geographic range. However, cogradient variation (the expected result from a plastic response, although not necessarily inconsistent with adaptation) has been suggested as a null hypothesis, especially on fine geographic scales because of relatively small environmental changes. Here we focus on life history variation on a fine geographic scale to test whether cogradient variation is exhibited even though countergradient variation is exhibited at larger scales, and if so, what mechanisms are involved in the switch. We examined north and south populations (~2° latitude between) of the S. undulatus, and the Florida Scrub Lizard, S. woodi, by measuring adult body sizes, reproduction, and hatchling body sizes over a two year period and conducting reciprocal transplants of juvenile lizards each year. Our results indicate cogradient variation (larger body size in the southern population experiencing a warmer environment) in life history traits of S. undulatus and countergradient variation, a lack of variation in adult body size, in S. woodi along the Florida peninsula. Thus, S. undulatus exhibits cogradient variation at fine geographic scales and countergradient variation at larger scales. Reciprocal transplants revealed that the larger adult body sizes in the southern population of S. undulatus could be explained by longer growth periods allowed by greater intrinsic survival. In S. woodi, the larger than expected adult body sizes in the north could be explained by faster intrinsic and extrinsic juvenile growth rates in the northern population. Because S. undulatus and S. woodi remain distinct species associated with distinct, though adjacent, habitats, we also looked for habitat-specific adaptations. The second reciprocal transplant (between species and habitats) revealed habitat-specific adaptations in juvenile growth rates, but not juvenile survival. Each native species grew faster and had a higher average probability of reaching size at maturity in their native environment than did the foreign species.







The Effect of Corticosterone on Behavior in Sceloporus Undulatus


Book Description

Levels of the stress hormone, corticosterone (CORT) in the blood can be a great indicator of stress levels in lizards. This approach reveals that invasive species impose novel pressure on natives, elevating stress levels. Native species often behaviorally adapt to these pressures in order to increase their chances of surviving exposure to these nonnative threats. We tested the hypothesis that eastern fence lizards (Sceloporus undulatus) that have been exposed to predatory invasive fire ants (Solenopsis invicta) for several generations exhibit different behavioral responses to experimentally elevated CORT than do lizards whose habitat has not yet been invaded. Lizards from the uninvaded site respond to elevated CORT levels by hiding more and moving and basking less, whereas those from the invaded site exhibit the opposite pattern; hiding less, moving more, and spend more time on the basking log when CORT levels are elevated. These differences in response may facilitate survival in these different environments. Within natural uninvaded populations, lizards would benefit from becoming less active and hiding in response to a stressful event, as this would reduce their vulnerability to predators and conspecifics. In contrast, within fire ant invaded sites, increased CORT levels occur following encounters with fire ants. Responding to elevated CORT levels by moving away and off the ground would increase the lizards' chances of surviving these encounters in the presence of this invader. The changes in basking behavior following elevated CORT levels appears to be due to changes in the propensity of lizards escaping up off the ground rather than having any thermoregulatory relevance. A second study revealed that lizards body temperatures were not affected by experimentally elevated CORT levels, and this response did not differ between fire ant invaded and uninvaded sites. Together, these reveal that the behavioral response to physiological stress can be altered by the introduction of non-native species, allowing native species to persist in the face of this novel threat.







Factors Affecting the Physiological Consequences of Stress in Eastern Fence Lizards (Sceloporus Undulatus).


Book Description

Understanding the consequences of stress is integral to predicting how organisms will respond to global environmental change. The stress response is generally adaptive, promoting physiological changes that allow an organism to deal with and recover from a threat. However, the stress response can lead to negative fitness outcomes because it diverts energy away from growth, reproduction, and immune function. My dissertation broadly investigates how traits of a stressor and of an organism affect the physiological outcomes of stress. To do so, I utilized populations of eastern fence lizards (Sceloporus undulatus) that co-evolved with different levels of stress associated with presence of invasive predatory fire ants (Solenopsis invicta). Stress is typically characterized by duration, with short-duration ("acute") stress assumed to be neutral or beneficial and long-duration ("chronic") stress assumed to have negative outcomes. However, the outcomes of stressors that result from repeated short-duration stress (e.g. frequent predator attacks) are not well understood. I first investigated the immune outcomes of these "repeated acute" stressors. Lizards from both high-and low-stress populations enhanced immune function following exposure to repeated acute elevations of exogenous corticosterone (CORT). This demonstrates that repeated acute stressors produce immune outcomes more typical of those expected from short-duration stress (i.e. immune enhancement). I then investigated the role of stressor duration, frequency, and intensity in determining the immune outcomes of stress. My results reveal that stressor intensity is a major driver of immune outcomes of stress in this system and suggest that the current duration-centric terminology is not adequate. Finally, I explored how exposure to stress within a lifetime and across generations affects stress physiology and immune function by raising offspring of lizards from high- and low-stress sites under high and low-stress conditions in the lab and measuring physiological outcomes in adulthood. Early life stress did not affect adult stress physiology, but offspring of lizards from high-stress sites had more robust stress responses than those from uninvaded sites. This suggests that, in this system, stress experienced by an individual's ancestors may be more important in shaping adult stress physiology than stress that an individual faces within its lifetime. By contrast, within- and across-generational factors interacted to affect adult immune function; CORT-elevation during early life suppressed adult immune function in lizards from low-stress sites but enhanced immune function in lizards from high-stress sites. Together, these results further our understanding of how traits of a stressor and those of an organism influence physiological outcomes of stress. This insight allows us to better predict how organisms will be affected by stress, including that imposed as a result of global change, and to better allocate resources to manage and mitigate these fitness-relevant effects.