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Why do many animals possess multiple classes of photoreceptors that vary in the wavelengths of light to which they are sensitive? Multiple spectral photoreceptor classes are a requirement for true color vision. However, animals may have unconventional vision, in which multiple spectral channels broaden the range of wavelengths that can

Why do many animals possess multiple classes of photoreceptors that vary in the wavelengths of light to which they are sensitive? Multiple spectral photoreceptor classes are a requirement for true color vision. However, animals may have unconventional vision, in which multiple spectral channels broaden the range of wavelengths that can be detected, or in which they use only a subset of receptors for specific behaviors. Branchiopod crustaceans are of interest for the study of unconventional color vision because they express multiple visual pigments in their compound eyes, have a simple repertoire of visually guided behavior, inhabit unique and highly variable light environments, and possess secondary neural simplifications. I first tested the behavioral responses of two representative species of branchiopods from separate orders, Streptocephalus mackini Anostracans (fairy shrimp), and Triops longicaudatus Notostracans (tadpole shrimp). I found that they maintain vertical position in the water column over a broad range of intensities and wavelengths, and respond behaviorally even at intensities below those of starlight. Accordingly, light intensities of their habitats at shallow depths tend to be dimmer than terrestrial habitats under starlight. Using models of how their compound eyes and the first neuropil of their optic lobe process visual cues, I infer that both orders of branchiopods use spatial summation from multiple compound eye ommatidia to respond at low intensities. Then, to understand if branchiopods use unconventional vision to guide these behaviors, I took electroretinographic recordings (ERGs) from their compound eyes and used models of spectral absorptance for a multimodel selection approach to make inferences about the number of photoreceptor classes in their eyes. I infer that both species have four spectral classes of photoreceptors that contribute to their ERGs, suggesting unconventional vision guides the described behavior. I extended the same modeling approach to other organisms, finding that the model inferences align with the empirically determined number of photoreceptor classes for this diverse set of organisms. This dissertation expands the conceptual framework of color vision research, indicating unconventional vision is more widespread than previously considered, and explains why some organisms have more spectral classes than would be expected from their behavioral repertoire.
ContributorsLessios, Nicolas (Author) / Rutowski, Ronald L (Thesis advisor) / Cohen, Jonathan H (Thesis advisor) / Harrison, John (Committee member) / Neuer, Susanne (Committee member) / McGraw, Kevin (Committee member) / Arizona State University (Publisher)
Created2016
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Description
Foraging has complex effects on whole-organism homeostasis, and there is considerable evidence that foraging behavior is influenced by both environmental factors (e.g., food availability, predation risk) and the physiological condition of an organism. The optimization of foraging behavior to balance costs and benefits is termed state-dependent foraging (SDF) while behavior

Foraging has complex effects on whole-organism homeostasis, and there is considerable evidence that foraging behavior is influenced by both environmental factors (e.g., food availability, predation risk) and the physiological condition of an organism. The optimization of foraging behavior to balance costs and benefits is termed state-dependent foraging (SDF) while behavior that seeks to protect assets of fitness is termed the asset protection principle (APP). A majority of studies examining SDF have focused on the role that energy balance has on the foraging of organisms with high metabolism and high energy demands ("high-energy systems" such as endotherms). In contrast, limited work has examined whether species with low energy use ("low-energy systems" such as vertebrate ectotherms) use an SDF strategy. Additionally, there is a paucity of evidence demonstrating how physiological and environmental factors other than energy balance influence foraging behavior (e.g. hydration state and free-standing water availability). Given these gaps in our understanding of SDF behavior and the APP, I examined the state-dependency and consequences of foraging in a low-energy system occupying a resource-limited environment - the Gila monster (Heloderma suspectum, Cope 1869). In contrast to what has been observed in a wide variety of taxa, I found that Gila monsters do not use a SDF strategy to manage their energy reserves and that Gila monsters do not defend their energetic assets. However, hydration state and free-standing water availability do affect foraging behavior of Gila monsters. Additionally, as Gila monsters become increasingly dehydrated, they reduce activity to defend hydration state. The SDF behavior of Gila monsters appears to be largely driven by the fact that Gila monsters must separately satisfy energy and water demands with food and free-standing water, respectively, in conjunction with the timescale within which Gila monsters balance their energy and water budgets (supra-annually versus annually, respectively). Given these findings, the impact of anticipated changes in temperature and rainfall patterns in the Sonoran Desert are most likely going to pose their greatest risks to Gila monsters through the direct and indirect effects on water balance.
ContributorsWright, Christian (Author) / Denardo, Dale F. (Thesis advisor) / Harrison, Jon (Committee member) / McGraw, Kevin (Committee member) / Sullivan, Brian (Committee member) / Wolf, Blair (Committee member) / Arizona State University (Publisher)
Created2014
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Description
Warning coloration deters predators from attacking prey that are defended, usually by being distasteful, toxic, or otherwise costly for predators to pursue and consume. Predators may have an innate response to warning colors or learn to associate them with a defense through trial and error. In general, predators should select

Warning coloration deters predators from attacking prey that are defended, usually by being distasteful, toxic, or otherwise costly for predators to pursue and consume. Predators may have an innate response to warning colors or learn to associate them with a defense through trial and error. In general, predators should select for warning signals that are easy to learn and recognize. Previous research demonstrates long-wavelength colors (e.g. red and yellow) are effective because they are readily detected and learned. However, a number of defended animals display short-wavelength coloration (e.g. blue and violet), such as the pipevine swallowtail butterfly (Battus philenor). The role of blue coloration in warning signals had not previously been explicitly tested. My research showed in laboratory experiments that curve-billed thrashers (Toxostoma curvirostre) and Gambel's quail (Callipepla gambelii) can learn and recognize the iridescent blue of B. philenor as a warning signal and that it is innately avoided. I tested the attack rates of these colors in the field and blue was not as effective as orange. I concluded that blue colors may function as warning signals, but the effectiveness is likely dependent on the context and predator.

Blue colors are often iridescent in nature and the effect of iridescence on warning signal function was unknown. I reared B. philenor larvae under varied food deprivation treatments. Iridescent colors did not have more variation than pigment-based colors under these conditions; variation which could affect predator learning. Learning could also be affected by changes in appearance, as iridescent colors change in both hue and brightness as the angle of illuminating light and viewer change in relation to the color surface. Iridescent colors can also be much brighter than pigment-based colors and iridescent animals can statically display different hues. I tested these potential effects on warning signal learning by domestic chickens (Gallus gallus domesticus) and found that variation due to the directionality of iridescence and a brighter warning signal did not influence learning. However, blue-violet was learned more readily than blue-green. These experiments revealed that the directionality of iridescent coloration does not likely negatively affect its potential effectiveness as a warning signal.
ContributorsPegram, Kimberly Vann (Author) / Rutowski, Ronald L (Thesis advisor) / Hoelldobler, Berthold (Committee member) / Liebig, Juergen (Committee member) / McGraw, Kevin (Committee member) / Smith, Brian H. (Committee member) / Arizona State University (Publisher)
Created2015
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Description
The Great Bustard (Otis tarda) is an iconic species of the temperate grasslands of Europe and Asia, a habitat that is among the least protected ecosystems in the world. A distinct subspecies, the Asian Great Bustard (O. t. dybowskii), is poorly understood due to its wary nature and remote range

The Great Bustard (Otis tarda) is an iconic species of the temperate grasslands of Europe and Asia, a habitat that is among the least protected ecosystems in the world. A distinct subspecies, the Asian Great Bustard (O. t. dybowskii), is poorly understood due to its wary nature and remote range in Siberia, Mongolia, and northern China. This subspecies is now endangered by rapid development.

Using satellite telemetry and remote sensing, I investigated three aspects of the Asian Great Bustard’s ecology critical to its conservation: migratory routes, migratory cues, and habitat use patterns. I found that Asian Great Bustards spent one-third of the year on a 2000 km migratory pathway, a distance twice as far as has previously been recorded for the species. Tracked individuals moved nomadically over large winter territories and did not repeat migratory stopovers, complicating conservation planning. Migratory timing was variable and migratory movements were significantly correlated with weather cues. Specifically, bustards migrated on days when wind support was favorable and temperature presaged warmer temperatures on the breeding grounds (spring) or advancing winter weather (fall). On the breeding grounds, Asian Great Bustards used both steppe and wheat agriculture habitat. All recorded reproductive attempts failed, regardless of habitat in which the nest was placed. Agricultural practices are likely to intensify in the coming decade, which would present further challenges to reproduction. The distinct migratory behavior and habitat use patterns of the Asian Great Bustard are likely adaptations to the climate and ecology of Inner Asia and underscore the importance of conserving these unique populations.

My research indicates that conservation of the Asian Great Bustard will require a landscape-level approach. This approach should incorporate measures at the breeding grounds to raise reproductive success, alongside actions on the migratory pathway to ensure appropriate habitat and reduce adult mortality. To secure international cooperation, I proposed that an increased level of protection should be directed toward the Great Bustard under the Convention on Migratory Species (CMS). That proposal, accepted by the Eleventh Conference of Parties to CMS, provides recommendations for conservation action and illustrates the transdisciplinary approach I have taken in this research.
ContributorsKessler, Aimee (Author) / Smith, Andrew T. (Thesis advisor) / Brown, David (Committee member) / Franklin, Janet (Committee member) / McGraw, Kevin (Committee member) / Wu, Jianguo (Committee member) / Arizona State University (Publisher)
Created2015
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Description
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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Description
For animals that experience annual cycles of gonad development, the seasonal timing (phenology) of gonad growth is a major adaptation to local environmental conditions. To optimally time seasonal gonad growth, animals use environmental cues that forecast future conditions. The availability of food is one such environmental cue. Although the importance

For animals that experience annual cycles of gonad development, the seasonal timing (phenology) of gonad growth is a major adaptation to local environmental conditions. To optimally time seasonal gonad growth, animals use environmental cues that forecast future conditions. The availability of food is one such environmental cue. Although the importance of food availability has been appreciated for decades, the physiological mechanisms underlying the modulation of seasonal gonad growth by this environmental factor remain poorly understood.

Urbanization is characterized by profound environmental changes, and urban animals must adjust to an environment vastly different from that of their non-urban conspecifics. Evidence suggests that birds adjust to urban areas by advancing the timing of seasonal breeding and gonad development, compared to their non-urban conspecifics. A leading hypothesis to account for this phenomenon is that food availability is elevated in urban areas, which improves the energetic status of urban birds and enables them to initiate gonad development earlier than their non-urban conspecifics. However, this hypothesis remains largely untested.

My dissertation dovetailed comparative studies and experimental approaches conducted in field and captive settings to examine the physiological mechanisms by which food availability modulates gonad growth and to investigate whether elevated food availability in urban areas advances the phenology of gonad growth in urban birds. My captive study demonstrated that energetic status modulates reproductive hormone secretion, but not gonad growth. By contrast, free-ranging urban and non-urban birds did not differ in energetic status or plasma levels of reproductive hormones either in years in which urban birds had advanced phenology of gonad growth or in a year that had no habitat-related disparity in seasonal gonad growth. Therefore, my dissertation provides no support for the hypothesis that urban birds begin seasonal gonad growth because they are in better energetic status and increase the secretion of reproductive hormones earlier than non-urban birds. My studies do suggest, however, that the phenology of key food items and the endocrine responsiveness of the reproductive system may contribute to habitat-related disparities in the phenology of gonad growth.
ContributorsDavies, Scott (Author) / Deviche, Pierre (Thesis advisor) / Sweazea, Karen (Committee member) / McGraw, Kevin (Committee member) / Orchinik, Miles (Committee member) / Warren, Paige (Committee member) / Arizona State University (Publisher)
Created2014
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Description
Many animals thermoregulate to maximize performance. However, interactions with other animals, such as competitors or predators, limit access to preferred microclimates. For instance, an animal may thermoregulate poorly when fighting rivals or avoiding predators. However, the distribution of thermal resources should influence how animals perceive and respond to risk. When

Many animals thermoregulate to maximize performance. However, interactions with other animals, such as competitors or predators, limit access to preferred microclimates. For instance, an animal may thermoregulate poorly when fighting rivals or avoiding predators. However, the distribution of thermal resources should influence how animals perceive and respond to risk. When thermal resources are concentrated in space, individuals compete for access, which presumably reduces the thermoregulatory performance while making their location more predictable to predators. Conversely, when thermal resources are dispersed, several individuals can thermoregulate effectively without occupying the same area. Nevertheless, interactions with competitors or predators impose a potent stress, often resulting in both behavioral and physiological changes that influence thermoregulation. To assess the costs of intraspecific competition and predation risk during thermoregulation, I measured thermoregulation, movement, and hormones of male lizards (Sceloporus jarrovi) in experiment landscapes, with clumped to patchy distributions of microclimates. I found lizards aggressively competed for access to microclimates, with larger males gaining priority access when thermal resources were aggregated. Competition reduced thermoregulatory performance, increased movements, and elevated plasma corticosterone in large and small males. However, the magnitude of these responses decreased as the patchiness of the thermal environment increased. Similarly, under simulated predation risk, lizards reduced thermoregulatory performance, decreased movements, and elevated plasma corticosterone. Again, with the magnitude of these responses decreased with increasing thermal patchiness. Interestingly, even without competitors or predators, lizards in clumped arenas moved greater distances and circulated more corticosterone than did lizards in patchy arenas, indicating the thermal quality of the thermal landscape affected the energetic demands on lizards. Thus, biologists should consider species interactions and spatial structure when modeling impacts of climate change on thermoregulation.
ContributorsRusch, Travis W (Author) / Angilletta, Michael (Thesis advisor) / Sears, Mike (Committee member) / DeNardo, Dale (Committee member) / Deviche, Pierre (Committee member) / McGraw, Kevin (Committee member) / Arizona State University (Publisher)
Created2017
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Description
The molt from pupae to adult stage, called eclosion, occurs at specific times of the day in many holometabolous insects. These events are not well studied within Lepidopteran species. It was hypothesized that the eclosion timing in a species may be shaped by strong selective pressures, such as sexual selection

The molt from pupae to adult stage, called eclosion, occurs at specific times of the day in many holometabolous insects. These events are not well studied within Lepidopteran species. It was hypothesized that the eclosion timing in a species may be shaped by strong selective pressures, such as sexual selection in the context of male-male competition. The daily timing of eclosion was measured for six species of nymphalid butterflies. This was done by rearing individuals to pupation, placing the pupa in a greenhouse, and video recording eclosion to obtain the time of day at which it occurred. Four species exhibited clustered eclosion distributions that were concentrated to within 201 minutes after sunrise and were significantly different from one another. The other two species exhibited eclosion times that were non-clustered. There were no differences between sexes within species. The data support a relationship between the timing of eclosion each day and the timing of mating activities, but other as of yet undetermined selective pressures may also influence eclosion timing.
ContributorsSencio, Kaylon (Author) / Rutowski, Ron (Thesis advisor) / McGraw, Kevin (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2017
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Description
In wild birds, the stress response can inhibit the activity of the innate immune system, which serves as the first line of defense against pathogens. By elucidating the mechanisms which regulate the interaction between stress and innate immunity, researchers may be able to predict when birds experience increased susceptibility to

In wild birds, the stress response can inhibit the activity of the innate immune system, which serves as the first line of defense against pathogens. By elucidating the mechanisms which regulate the interaction between stress and innate immunity, researchers may be able to predict when birds experience increased susceptibility to infections and can target specific mediators to mitigate stress-induced suppression of innate immune activity. Such elucidation is especially important for urban birds, such as the House Sparrow (Passer domesticus), because these birds experience higher pathogen prevalence and transmission when compared to birds in rural regions. I investigated the role of corticosterone (CORT) in stress-induced suppression of two measures of innate immune activity (complement- and natural antibody-mediated activity) in male House Sparrows. Corticosterone, the primary avian glucocorticoid, is elevated during the stress response and high levels of this hormone induce effects through the activation of cytosolic and membrane-bound glucocorticoid receptors (GR). My results demonstrate that CORT is necessary and sufficient for stress-induced suppression of complement-mediated activity, and that this relationship is consistent between years. Corticosterone, however, does not inhibit complement-mediated activity through cytosolic GR, and additional research is needed to confirm the involvement of membrane-bound GR. The role of CORT in stress-induced inhibition of natural antibody-mediated activity, however, remains puzzling. Stress-induced elevation of CORT can suppress natural antibody-mediated activity through the activation of cytosolic GR, but the necessity of this mechanism varies inter-annually. In other words, both CORT-dependent and CORT-independent mechanisms may inhibit natural antibody-mediated activity during stress in certain years, but the causes of this inter-annual variation are not known. Previous studies have indicated that changes in the pathogen environment or food availability can alter regulation of innate immunity, but further research is needed to test these hypotheses. Overall, my dissertation demonstrates that stress inhibits innate immunity through several mechanisms, but environmental pressures may influence this inhibitory relationship.
ContributorsGao, Sisi (Author) / Deviche, Pierre (Thesis advisor) / DeNardo, Dale (Committee member) / McGraw, Kevin (Committee member) / Orchinik, Miles (Committee member) / Moore, Michael C. (Committee member) / Arizona State University (Publisher)
Created2017
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Prior to the COVID-19 global pandemic, ecotourism represented the tourism industry’s fastest growing segment with projections estimating that ecotourism would become the world’s largest tourism type by 2030. While the tourism industry will need several years to rebound, if historic trends tell us anything, it is that ecotourism will continue

Prior to the COVID-19 global pandemic, ecotourism represented the tourism industry’s fastest growing segment with projections estimating that ecotourism would become the world’s largest tourism type by 2030. While the tourism industry will need several years to rebound, if historic trends tell us anything, it is that ecotourism will continue to represent a large portion of the overall industry and will continue to grow at a rate that outpaces all other tourism types. In theory, ecotourism promotes sustainable socioeconomic development while also minimizing negative environmental impacts. Unfortunately, research suggests that this is not always true, and many examples exist of ecotourism causing more harm than good. In order to combat these potential negative impacts, the ecotourism industry has become increasingly reliant on ecotourism certification programs to act as an assessment tool that identifies ecotourism’s best practitioners while minimizing false advertising present within the industry. Despite these beliefs in the efficacy of certification, there is a lack of empirical research to actually support certification as an effective assessment tool. Furthermore, little research has been conducted that assesses the impacts that certification itself has on ecotourism businesses (both certified and uncertified) and the local communities dependent on ecotourism.
My dissertation employs a mixed methods design and combines qualitative and quantitative research methods spanning multiple geographic scales to develop an understanding of certification programs as they exist today and to discern the impacts that certification itself may cause for all those either directly or indirectly involved in ecotourism. My findings ultimately suggest that certification reform is needed if certification programs are expected to be the assessment tool ecotourism experts claim them to be. Specifically, as certification exists presently, there is: no universal guideline or standard for existing certification programs to follow, a disconnect between the advertised benefits certification offers and the actual benefits received, and a lack of market penetration both amongst ecotourists and ecotourism businesses. Each of these must be addressed before certification can live up to its full potential. Furthermore, I found that certification may impact community socioeconomic dynamics, particularly by creating or exacerbating community wealth distribution.
ContributorsDavila, Ryan (Author) / Kinzig, Ann (Thesis advisor) / Perrings, Charles (Committee member) / Collins, James (Committee member) / Schoon, Michael (Committee member) / Buzinde, Christine (Committee member) / Arizona State University (Publisher)
Created2020