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.




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.




Hormones, Brain, and Behavior


Book Description

Volume 18.




Natural Hybridization and Evolution


Book Description

Michael L. Arnold offers an exploration of the evolutionary process of natural hybridisation, and presents data from various sources that support the paradigm of natural hybridisation as an important evolutionary process.




The Biology and Identification of the Coccidia (Apicomplexa) of Turtles of the World


Book Description

The Biology and Identification of the Coccidia (Apicomplexa) of Turtles of the World is an invaluable resource for researchers in protozoology, coccidia, and parasitology, veterinary sciences, animal sciences, zoology, and biology. This first-of-its-kind work offers a taxonomic guide to apicomplexan parasites of turtles that enables easy parasite identification, with a summary of virtually everything known about the biology of each known parasite species. It is an important documentation of this specific area, useful to a broad base of readers, including researchers in biology, parasitology, animal husbandry, diseases of wild and domestic animals, veterinary medicine, and faculty members in universities with graduate programs in these areas. There are about 330 turtle species on Earth; many are endangered, a growing number of species are kept as pets, and some are still used as food by humans. Turtles, like other vertebrate animals have many different kinds of parasites (viruses, bacteria, protozoa, worms, arthropods, and others). Coccidiosis in turtles has prevented large-scale turtle breeding, and represents a serious problem in need of control. This succinct and highly focused book will aid in that effort. Offers line drawings and photomicrographs of each parasite from each hosts species Provides methods of identification and treatment Presents a complete historical rendition of all known publications on coccidia (and their closest relatives) from all turtle species on Earth, and evaluates the scientific and scholarly merit of each Provides a complete species analysis of the known biology of every coccidian described from turtles Reviews the most current taxonomy of turtles and their phylogenetic relationships needed to help assess host-specificity and evaluate what little cross-transmission work is available




Molecular Markers, Natural History and Evolution


Book Description

Molecular approaches have opened new windows on a host of ecological and evolutionary disciplines, ranging from population genetics and behavioral ecology to conservation biology and systematics. Molecular Markers, Natural History and Evolution summarizes the multi-faceted discoveries about organisms in nature that have stemmed from analyses of genetic markers provided by polymorphic proteins and DNAs. The first part of the book introduces rationales for the use of molecular markers, provides a history of molecular phylogenetics, and describes a wide variety of laboratory methods and interpretative tools in the field. The second and major portion of the book provides a cornucopia of biological applications for molecular markers, organized along a scale from micro-evolutionary topics (such as forensics, parentage, kinship, population structure, and intra-specific phylogeny) to macro-evolutionary themes (including species relationships and the deeper phylogenetic structure in the tree of life). Unlike most prior books in molecular evolution, the focus is on organismal natural history and evolution, with the macromolecules being the means rather than the ends of scientific inquiry. Written as an intellectual stimulus for the advanced undergraduate, graduate student, or the practicing biologist desiring a wellspring of research ideas at the interface of molecular and organismal biology, this book presents material in a manner that is both technically straightforward, yet rich with concepts and with empirical examples from the world of nature.




Pheromones and Reproduction


Book Description