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Understanding why animals form social groups is a fundamental aim of sociobiology. To date, the field has been dominated by studies of kin groups, which have emphasized indirect fitness benefits as key drivers of grouping among relatives. Nevertheless, many animal groups are comprised of unrelated individuals. These cases provide unique

Understanding why animals form social groups is a fundamental aim of sociobiology. To date, the field has been dominated by studies of kin groups, which have emphasized indirect fitness benefits as key drivers of grouping among relatives. Nevertheless, many animal groups are comprised of unrelated individuals. These cases provide unique opportunities to illuminate drivers of social evolution beyond indirect fitness, especially ecological factors. This dissertation combines behavioral, physiological, and ecological approaches to explore the conditions that favor group formation among non-kin, using as a model the facultatively social carpenter bee, Xylocopa sonorina. Using behavioral and genetic techniques, I found that nestmates in this species are often unrelated, and that non-kin groups form following extensive inter-nest migration.Group living may arise as a strategy to mitigate constraints on available breeding space. To test the hypothesis that nest construction is prohibitively costly for carpenter bees, I measured metabolic rates of excavating bees and used imaging techniques to quantify nest volumes. From these measurements, I found that nest construction is highly energetically costly, and that bees who inherit nests through social queuing experience substantial energetic savings. These costs are exacerbated by limitations on the reuse of existing nests. Using repeated CT scans of nesting logs, I examined changes in nest architecture over time and found that repeatedly inherited tunnels become indefensible to intruders, and are subsequently abandoned. Together, these factors underlie intense competition over available breeding space. The imaging analysis of nesting logs additionally revealed strong seasonal effects on social strategy, with social nesting dominating during winter. To test the hypothesis that winter social nesting arises from intrinsic physiological advantages of grouping, I experimentally manipulated social strategy in overwintering bees. I found that social bees conserve heat and body mass better than solitary bees, suggesting fitness benefits to grouping in cold, resource-scarce conditions. Together, these results suggest that grouping in X. sonorina arises from dynamic strategies to maximize direct fitness in response to harsh and/or competitive conditions. These studies provide empirical insights into the ecological conditions that favor non-kin grouping, and emphasize the importance of ecology in shaping sociality at its evolutionary origins.
ContributorsOstwald, Madeleine (Author) / Fewell, Jennifer H (Thesis advisor) / Amdam, Gro (Committee member) / Harrison, Jon (Committee member) / Pratt, Stephen (Committee member) / Kapheim, Karen (Committee member) / Arizona State University (Publisher)
Created2022
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ABSTRACT Over the past several decades, the dilemma of free-roaming horses in the U.S. has proven to be one of the most divisive issues in management of public lands. According to federal land management agencies, without population regulation, horses can increase at the rate of 15-20% a year on arid

ABSTRACT Over the past several decades, the dilemma of free-roaming horses in the U.S. has proven to be one of the most divisive issues in management of public lands. According to federal land management agencies, without population regulation, horses can increase at the rate of 15-20% a year on arid rangelands with inadequate numbers of natural, large predators. Horses compete for valuable forage and water resources alongside cattle and native wildlife in delicate riparian areas highly susceptible to the negative ecological effects of soil compaction and overgrazing. Most U.S. management policies, therefore, call for increased removal of free-roaming horses as they are categorized as “un-authorized livestock” or "non-native" species. Wild horse advocates, however, continue to petition for improvement in animal welfare and expansion of the horses’ territory. With heightened social conflict spurred by animal rights and ecological concerns, not to mention the often-stark differences over what really “belongs” on the landscape, the success of appropriate management strategies hinges on managing agencies’ preparedness and ability to respond in a timely and inclusive manner. A critical element of the management context is the public’s views toward the wild horse and the science used to manage them. Synthesizing the vast literature in the history and philosophy of wildlife management in the American West, and utilizing an ethnographic and case study approach, my research examines the range of stakeholder concerns and analyzes the factors that have led to the disconnect between public values of wild horses and public policy for the management of the federally protected free-roaming horses in Arizona’s Apache-Sitgreaves National Forests.
ContributorsMurphree, Julie Joan (Author) / Minteer, Ben A. (Thesis advisor) / Schoon, Michael (Thesis advisor) / Bradshaw, Karen (Committee member) / Chew, Matthew (Committee member) / Arizona State University (Publisher)
Created2022
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Speciation, or the process by which one population diverges into multiple populations that can no longer interbreed with each other, has brought about the incredible diversity of life. Mechanisms underlying this process can be more visible in the early stages of the speciation process. The mechanisms that restrict gene flow

Speciation, or the process by which one population diverges into multiple populations that can no longer interbreed with each other, has brought about the incredible diversity of life. Mechanisms underlying this process can be more visible in the early stages of the speciation process. The mechanisms that restrict gene flow in highly mobile species with no absolute barriers to dispersal, especially marine species, are understudied. Similarly, human impacts are reshaping ecosystems globally, and we are only just beginning to understand the implications of these rapid changes on evolutionary processes. In this dissertation, I investigate patterns of speciation and evolution in two avian clades: a genus of widespread tropical seabirds (boobies, genus Sula), and two congeneric passerine species in an urban environment (cardinals, genus Cardinalis). First, I explore the prevalence of gene flow across land barriers within species and between sympatric species in boobies. I found widespread evidence of gene flow over all land barriers and between 3 species pairs. Next, I compared the effects of urbanization on the spatial distributions of two cardinal species, pyrrhuloxia (Cardinalis sinuatus) and northern cardinals (Cardinalis cardinalis), in Tucson, Arizona. I found that urbanization has different effects on the spatial distributions of two closely related species that share a similar environmental niche, and I identified environmental variables that might be driving this difference. Then I tested for effects of urbanization on color and size traits of these two cardinal species. In both of these species, urbanization has altered traits involved in signaling, heat tolerance, foraging, and maneuverability. Finally, I tested for evidence of selection on the urban populations of both cardinal species and found evidence of both parallel selection and introgression between the species, as well as selection on different genes in each species. The functions of the genes that experienced positive selection suggest that light at night, energetics, and air pollution may have acted as strong selective pressures on these species in the past. Overall, my dissertation emphasizes the role of introgression in the speciation process, identifies environmental stressors faced by wildlife in urban environments, and characterizes their evolutionary responses to those stressors.
ContributorsJackson, Daniel Nelson (Author) / McGraw, Kevin J (Thesis advisor) / Amdam, Gro (Committee member) / Sweazea, Karen (Committee member) / Taylor, Scott (Committee member) / Arizona State University (Publisher)
Created2023
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Understanding the dynamic interactions between humans and wildlife is essential to establishing sustainable wildlife-based ecotourism (WBE). Animal behavior exists within a complex feedback loop that affects overall ecosystem function, tourist satisfaction, and socioeconomics of local communities. However, the specific value that animal behavior plays in provisioning ecosystem services has not

Understanding the dynamic interactions between humans and wildlife is essential to establishing sustainable wildlife-based ecotourism (WBE). Animal behavior exists within a complex feedback loop that affects overall ecosystem function, tourist satisfaction, and socioeconomics of local communities. However, the specific value that animal behavior plays in provisioning ecosystem services has not been thoroughly evaluated. People enjoy activities that facilitate intimate contact with animals, and there are many perceived benefits associated with these experiences, such as encouraging pro-environmental attitudes that can lead to greater motivation for conservation. There is extensive research on the effects that unregulated tourism activity can have on wildlife behavior, which include implications for population health and survival. Prior to COVID-19, WBE was developing rapidly on a global scale, and the pause in activity caused by the pandemic gave natural systems the chance to recover from environmental damage from over-tourism and provided insights into how tourism could be less impactful in the future. Until now it has been undetermined how changes in animal behavior can alter the relationships and socioeconomics of this multidimensional system. This dissertation provides a thorough exploration of the behavioral, ecological, and economic parameters required to model biosocial interactions and feedbacks within the whale watching system in Las Perlas Archipelago, Panama. Through observational data collected in the field, this project assessed how unmanaged whale watching activity is affecting the behavior of Humpback whales in the area as well as the socioeconomic and conservation contributions of the industry. Additionally, it is necessary to consider what a sustainable form of wildlife tourism might be, and whether the incorporation of technology will help enhance visitor experience while reducing negative impacts on wildlife. To better ascertain whether this concept of this integration would be favorably viewed, a sample of individuals was surveyed about their experiences about using technology to enhance their interactions with nature. This research highlights the need for more deliberate identification and incorporation of the perceptions of all stakeholders (wildlife included) to develop a less-impactful WBE industry that provides people with opportunities to establish meaningful relationships with nature that motivate them to help meet the conservation challenges of today.
ContributorsSurrey, Katie (Author) / Gerber, Leah (Thesis advisor) / Guzman, Hector (Committee member) / Minteer, Ben (Committee member) / Schoon, Michael (Committee member) / Arizona State University (Publisher)
Created2023
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Heavy metals and persistent organic pollutants contribute to human health risks worldwide. Among the most common routes of exposure to pollutants for humans are through the consumption of contaminated water and food, with fish being among the greatest vectors for ingestion of heavy metals in humans, particularly mercury.This dissertation consists

Heavy metals and persistent organic pollutants contribute to human health risks worldwide. Among the most common routes of exposure to pollutants for humans are through the consumption of contaminated water and food, with fish being among the greatest vectors for ingestion of heavy metals in humans, particularly mercury.This dissertation consists of three chapters with a central theme of investigating heavy metal and persistent organic pollutant concentrations in fish and corned beef, which are two commonly consumed food items in American Samoa. A literature review illustrated that historically the primary pollutants of concern in fish muscle tissue from American Samoa have been mercury, arsenic, and polycyclic aromatic hydrocarbon mixtures. To better understand the changes in heavy metals and persistent organic pollutants in fish, this study reports an updated data set, comparing concentrations in pollutants as they have changed over time. To further investigate pollutants in fish tissue, 77 locally caught and commonly consumed fish were analyzed for heavy metals and persistent organic pollutants, and baseline human health risk assessments were calculated for contaminants that had available oral reference doses. While in American Samoa collecting fish for contaminant analyses, it was realized that canned corned beef appeared to be more commonly consumed than fresh fish. An IRB approved consumption survey revealed that 89% of American Samoan adults regularly consume fish, which is the same percentage of people that reported eating canned corned beef, indicating a dramatic increase in this food item to their diet since its introduction in the 20th century. Results of this study indicate that fish muscle tissue generally has higher heavy metal concentrations than canned corned beef, and that mercury continues to be a main contaminant of concern when consuming fresh and canned fish in American Samoa. While none of the heavy metal concentrations in corned beef exceeded calculated action levels, these foods might contribute to negative health outcomes in other ways. One of the main findings of this study is that either the presence or the ability to detect persistent organic pollutant concentrations are increasing in fish tissue and should be periodically monitored to adequately reflect current conditions.
ContributorsLewis, Tiffany Beth (Author) / Polidoro, Beth (Thesis advisor) / Neuer, Susanne (Thesis advisor) / Halden, Rolf (Committee member) / Schoon, Michael (Committee member) / Arizona State University (Publisher)
Created2023
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Effective collaboration and cooperation across difference are at the heart of present and future sustainability challenges and solutions. Collaboration among social groups (intragenerational), across time (intergenerational), and across species (interspecies) is each central to achieving sustainability transitions in the 21st century. In practice, there are three types of

Effective collaboration and cooperation across difference are at the heart of present and future sustainability challenges and solutions. Collaboration among social groups (intragenerational), across time (intergenerational), and across species (interspecies) is each central to achieving sustainability transitions in the 21st century. In practice, there are three types of differences that limit collaboration and cooperation toward sustainability outcomes: differences among social groups, differences across time, and differences across species. Each of these differences have corresponding cognitive biases that challenge collaboration. Social cognitive biases challenge collaboration among social groups; temporal cognitive biases challenge collaboration across time; and anthropocentric cognitive biases challenge collaboration across species. In this work, I present three correctives to collaboration challenges spanning the social, temporal, and species cognitive biases through intervention-specific methods that build beyond traditional framings of empathy, toward social, futures, and ecological empathy. By re-theorizing empathy across these domains, I seek to construct a multidimensional theory of empathy for sustainability, and suggest methods to build it, to bridge differences among people, time horizons, and species for sustainability practice.
ContributorsLambert, Lauren Marie-Jasmine (Author) / Selin, Cynthia (Thesis advisor) / Schoon, Michael (Thesis advisor) / Tomblin, David (Committee member) / Berbés-Blázquez, Marta (Committee member) / Arizona State University (Publisher)
Created2023
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Though controversial in its utility to the scientific study of nonhuman animals, anthropomorphism, or the attribution of human characteristics to a nonhuman being, is omnipresent in our interactions with other animals. Anthropomorphism is undeniably a fixture of modern zoos, but how anthropomorphism relates to zoos’ contributions to conservation is unclear.

Though controversial in its utility to the scientific study of nonhuman animals, anthropomorphism, or the attribution of human characteristics to a nonhuman being, is omnipresent in our interactions with other animals. Anthropomorphism is undeniably a fixture of modern zoos, but how anthropomorphism relates to zoos’ contributions to conservation is unclear. In this dissertation, I investigate these potentially dueling, potentially overlapping, messages within great ape exhibits in accredited zoos. Given the complexity of both anthropomorphism and conservation, this dissertation reveals some nuances of how both play out in zoological spaces. Human psychology literature on anthropomorphism indicates that there are a variety of uses for this lens that benefit humans; from feeling we can understand a confusing animal action, to feeling social connection. Whereas the comparative psychology literature highlights a contested utility of anthropomorphism in studies of nonhuman animals. The main findings from this study are four-fold. Firstly, surveys conducted with zoo visitors show that many bring anthropomorphic beliefs with them on their trek through the zoo. Visitors are prone to viewing great apes as strikingly like humans in terms of emotions, emotional expression, and understanding of the world. Secondly, surveyed zoo visitors who agreed more with anthropomorphic statements also agreed more with statements about feeling interconnected with nature. Thirdly, there is no uniform understanding within the zoo community about how zoo exhibits do or should contribute to conservation efforts given that exhibits have multiple goals, one being the safety and wellness of their animal residents. Fourthly, interviews of zoo staff show that they mediate a variety of messages for zoo visitors and walk a sometimes-divisive line between when it’s acceptable to use anthropomorphic framing to discuss zoo animals and when it’s inaccurate. By leveraging a better understanding of these attitudes and relationships, zoos can further empower their staff to navigate these complex issues and improve their mission-based goal of promoting conservation outcomes by acknowledging the human practices embedded in our perceptions of and interactions with zoo animals. This work speaks to the importance of carefully considering the ways we understand animals in zoos, in the wild, and all the places in-between.
ContributorsLyon, Cassandra (Author) / Minteer, Ben A. (Thesis advisor) / Wynne, Clive D.L. (Committee member) / Maynard, Lily (Committee member) / Schoon, Michael (Committee member) / Arizona State University (Publisher)
Created2024
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With global environmental systems under increasing Anthropogenic influence, conservationists and environmental managers are under immense pressure to protect and recover the world’s imperiled species and ecosystems. This effort is often motivated by a sense of moral responsibility, either to nature itself, or to the end of promoting human wellbeing over

With global environmental systems under increasing Anthropogenic influence, conservationists and environmental managers are under immense pressure to protect and recover the world’s imperiled species and ecosystems. This effort is often motivated by a sense of moral responsibility, either to nature itself, or to the end of promoting human wellbeing over the long run. In other words, it is the purview of environmental ethics, a branch of applied philosophy that emerged in the 1970s and that for decades has been devoted to understanding and defending an attitude of respect for nature, usually for its own sake. Yet from the very start, environmental ethics has promoted itself as contributing to the resolution of real-world management and policy problems. By most accounts, however, the field has historically failed to deliver on this original promise, and environmental ethicists continue to miss opportunities to make intellectual inroads with key environmental decisionmakers. Inspired by classical and contemporary American philosophers such as Charles Sanders Peirce, William James, John Dewey, and Richard Rorty, I defend in this dissertation the virtues of a more explicitly pragmatic approach to environmental ethics. Specifically, I argue that environmental pragmatism is not only commensurate with pro-environmental attitudes but that it is more likely to lead to viable and sustainable outcomes, particularly in the context of eco-social resilience-building activities (e.g., local experimentation, adaptation, cooperation). In doing so, I call for a recasting of environmental ethics, a project that entails: 1) a conceptual reorientation involving the application of pragmatism applied to environmental problems; 2) a methodological approach linking a pragmatist environmentalism to the tradition and process of adaptive co-management; and 3) an empirical study of stakeholder values and perspectives in conservation collaboratives in Arizona. I conclude that a more pragmatic environmental ethics has the potential to bring a powerful set of ethical and methodological tools to bear in real-world management contexts and, where appropriate, can ground and justify coordinated conservation efforts. Finally, this research responds to critics who suggest that, because it strays too far from the ideological purity of traditional environmental ethics, the pragmatic decision-making process will, in the long run, weaken rather than bolster our commitment to conservation and environmental protection.
ContributorsRojas, Christopher A (Author) / Minteer, Ben A (Thesis advisor) / Carr Kelman, Candice (Committee member) / Kinzig, Ann (Committee member) / Schoon, Michael (Committee member) / Arizona State University (Publisher)
Created2019
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This dissertation consists of three substantive chapters. The first substantive chapter investigates the premature harvesting problem in fisheries. Traditionally, yield-per-recruit analysis has been used to both assess and address the premature harvesting of fish stocks. However, the fact that fish size often affects the unit price suggests that this approach

This dissertation consists of three substantive chapters. The first substantive chapter investigates the premature harvesting problem in fisheries. Traditionally, yield-per-recruit analysis has been used to both assess and address the premature harvesting of fish stocks. However, the fact that fish size often affects the unit price suggests that this approach may be inadequate. In this chapter, I first synthesize the conventional yield-per-recruit analysis, and then extend this conventional approach by incorporating a size-price function for a revenue-per-recruit analysis. An optimal control approach is then used to derive a general bioeconomic solution for the optimal harvesting of a short-lived single cohort. This approach prevents economically premature harvesting and provides an "optimal economic yield". By comparing the yield- and revenue-per-recruit management strategies with the bioeconomic management strategy, I am able to test the economic efficiency of the conventional yield-per-recruit approach. This is illustrated with a numerical study. It shows that a bioeconomic strategy can significantly improve economic welfare compared with the yield-per-recruit strategy, particularly in the face of high natural mortality. Nevertheless, I find that harvesting on a revenue-per-recruit basis improves management policy and can generate a rent that is close to that from bioeconomic analysis, in particular when the natural mortality is relatively low.

The second substantive chapter explores the conservation potential of a whale permit market under bounded economic uncertainty. Pro- and anti-whaling stakeholders are concerned about a recently proposed, "cap and trade" system for managing the global harvest of whales. Supporters argue that such an approach represents a novel solution to the current gridlock in international whale management. In addition to ethical objections, opponents worry that uncertainty about demand for whale-based products and the environmental benefits of conservation may make it difficult to predict the outcome of a whale share market. In this study, I use population and economic data for minke whales to examine the potential ecological consequences of the establishment of a whale permit market in Norway under bounded but significant economic uncertainty. A bioeconomic model is developed to evaluate the influence of economic uncertainties associated with pro- and anti- whaling demands on long-run steady state whale population size, harvest, and potential allocation. The results indicate that these economic uncertainties, in particular on the conservation demand side, play an important role in determining the steady state ecological outcome of a whale share market. A key finding is that while a whale share market has the potential to yield a wide range of allocations between conservation and whaling interests - outcomes in which conservationists effectively "buy out" the whaling industry seem most likely.

The third substantive chapter examines the sea lice externality between farmed fisheries and wild fisheries. A central issue in the debate over the effect of fish farming on the wild fisheries is the nature of sea lice population dynamics and the wild juvenile mortality rate induced by sea lice infection. This study develops a bioeconomic model that integrates sea lice population dynamics, fish population dynamics, aquaculture and wild capture salmon fisheries in an optimal control framework. It provides a tool to investigate sea lice control policy from the standpoint both of private aquaculture producers and wild fishery managers by considering the sea lice infection externality between farmed and wild fisheries. Numerical results suggest that the state trajectory paths may be quite different under different management regimes, but approach the same steady state. Although the difference in economic benefits is not significant in the particular case considered due to the low value of the wild fishery, I investigate the possibility of levying a tax on aquaculture production for correcting the sea lice externality generated by fish farms.
ContributorsHuang, Biao (Author) / Abbott, Joshua K (Thesis advisor) / Perrings, Charles (Thesis advisor) / Gerber, Leah R. (Committee member) / Muneepeerakul, Rachata (Committee member) / Schoon, Michael (Committee member) / Arizona State University (Publisher)
Created2014
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Neurotoxicology has historically focused on substances that directly damage nervous tissue. Behavioral assays that test sensory, cognitive, or motor function are used to identify neurotoxins. But, the outcomes of behavioral assays may also be influenced by the physiological status of non-neural organs. Therefore, toxin induced damage to non- neural organs

Neurotoxicology has historically focused on substances that directly damage nervous tissue. Behavioral assays that test sensory, cognitive, or motor function are used to identify neurotoxins. But, the outcomes of behavioral assays may also be influenced by the physiological status of non-neural organs. Therefore, toxin induced damage to non- neural organs may contribute to behavioral modifications. Heavy metals and metalloids are persistent environmental pollutants and induce neurological deficits in multiple organisms. However, in the honey bee, an important insect pollinator, little is known about the sublethal effects of heavy metal and metalloid toxicity though they are exposed to these toxins chronically in some environments. In this thesis I investigate the sublethal effects of copper, cadmium, lead, and selenium on honey bee behavior and identify potential mechanisms mediating the behavioral modifications. I explore the honey bees’ ability to detect these toxins, their sensory perception of sucrose following toxin exposure, and the effects of toxin ingestion on performance during learning and memory tasks. The effects depend on the specific metal. Honey bees detect and reject copper containing solutions, but readily consume those contaminated with cadmium and lead. And, exposure to lead may alter the sensory perception of sucrose. I also demonstrate that acute selenium exposure impairs learning and long-term memory formation or recall. Localizing selenium accumulation following chronic exposure reveals that damage to non-neural organs and peripheral sensory structures is more likely than direct neurotoxicity. Probable mechanisms include gut microbiome alterations, gut lining

damage, immune system activation, impaired protein function, or aberrant DNA methylation. In the case of DNA methylation, I demonstrate that inhibiting DNA methylation dynamics can impair long-term memory formation, while the nurse-to- forager transition is not altered. These experiments could serve as the bases for and reference groups of studies testing the effects of metal or metalloid toxicity on DNA methylation. Each potential mechanism provides an avenue for investigating how neural function is influenced by the physiological status of non-neural organs. And from an ecological perspective, my results highlight the need for environmental policy to consider sublethal effects in determining safe environmental toxin loads for honey bees and other insect pollinators.
ContributorsBurden, Christina Marie (Author) / Amdam, Gro (Thesis advisor) / Smith, Brian H. (Thesis advisor) / Gallitano-Mendel, Amelia (Committee member) / Harrison, Jon (Committee member) / Vu, Eric (Committee member) / Arizona State University (Publisher)
Created2016