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Over the past century in the southwestern United States human actions have altered hydrological processes that shape riparian ecosystems. One change, release of treated wastewater into waterways, has created perennial base flows and increased nutrient availability in ephemeral or intermittent channels. While there are benefits to utilizing treated wastewater for

Over the past century in the southwestern United States human actions have altered hydrological processes that shape riparian ecosystems. One change, release of treated wastewater into waterways, has created perennial base flows and increased nutrient availability in ephemeral or intermittent channels. While there are benefits to utilizing treated wastewater for environmental flows, there are numerous unresolved ecohydrological issues regarding the efficacy of effluent to sustain groundwater-dependent riparian ecosystems. This research examined how nutrient-rich effluent, released into waterways with varying depths to groundwater, influences riparian plant community development. Statewide analysis of spatial and temporal patterns of effluent generation and release revealed that hydrogeomorphic setting significantly influences downstream riparian response. Approximately 70% of effluent released is into deep groundwater systems, which produced the lowest riparian development. A greenhouse study assessed how varying concentrations of nitrogen and phosphorus, emulating levels in effluent, influenced plant community response. With increasing nitrogen concentrations, vegetation emerging from riparian seed banks had greater biomass, reduced species richness, and greater abundance of nitrophilic species. The effluent-dominated Santa Cruz River in southern Arizona, with a shallow groundwater upper reach and deep groundwater lower reach, served as a study river while the San Pedro River provided a control. Analysis revealed that woody species richness and composition were similar between the two systems. Hydric pioneers (Populus fremontii, Salix gooddingii) were dominant at perennial sites on both rivers. Nitrophilic species (Conium maculatum, Polygonum lapathifolium) dominated herbaceous plant communities and plant heights were greatest in effluent-dominated reaches. Riparian vegetation declined with increasing downstream distance in the upper Santa Cruz, while patterns in the lower Santa Cruz were confounded by additional downstream agricultural input and a channelized floodplain. There were distinct longitudinal and lateral shifts toward more xeric species with increasing downstream distance and increasing lateral distance from the low-flow channel. Patterns in the upper and lower Santa Cruz reaches indicate that water availability drives riparian vegetation outcomes below treatment facilities. Ultimately, this research informs decision processes and increases adaptive capacity for water resources policy and management through the integration of ecological data in decision frameworks regarding the release of effluent for environmental flows.
ContributorsWhite, Margaret Susan (Author) / Stromberg, Juliet C. (Thesis advisor) / Fisher, Stuart G. (Committee member) / White, Dave (Committee member) / Holway, James (Committee member) / Wu, Jianguo (Committee member) / Arizona State University (Publisher)
Created2011
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Description
The origin and function of color in animals has been a subject of great interest for taxonomists and ecologists in recent years. Coloration in animals is useful for many important functions like species identification, camouflage and understanding evolutionary relationships. Quantitative measurements of color signal and patch size in mammals, birds

The origin and function of color in animals has been a subject of great interest for taxonomists and ecologists in recent years. Coloration in animals is useful for many important functions like species identification, camouflage and understanding evolutionary relationships. Quantitative measurements of color signal and patch size in mammals, birds and reptiles, to name a few are strong indicators of sexual selection cues and individual health. These measurements provide valuable insights into the impact of environmental conditions on habitat and breeding of mammals, birds and reptiles. Recent advances in the area of digital cameras and sensors have led to a significant increase in the use of digital photography as a means of color quantification in animals. Although a significant amount of research has been conducted on ways to standardize image acquisition conditions and calibrate cameras for use in animal color quantification, almost no work has been done on designing automated methods for animal color quantification. This thesis presents a novel perceptual"–"based framework for the automated extraction and quantification of animal coloration from digital images with slowly varying (almost homogenous) background colors. This implemented framework uses a combination of several techniques including color space quantization using a few dominant colors, foreground"–"background identification, Bayesian classification and mixture Gaussian modelling of conditional densities, edge"–"enhanced model"–"based classification and Saturation"–"Brightness quantization to extract the colored patch. This approach assumes no prior information about the color of either the subject or the background and also the position of the subject in the image. The performance of the proposed method is evaluated for the plumage color of the wild house finches. Segmentation results obtained using the implemented framework are compared with manually scored results to illustrate the performance of this system. The segmentation results show a high correlation with manually scored images. This novel framework also eliminates common problems in manual scoring of digital images such as low repeatability and inter"–"observer error.
ContributorsBorkar, Tejas (Author) / Karam, Lina J (Thesis advisor) / Li, Baoxin (Committee member) / McGraw, Kevin J. (Committee member) / Arizona State University (Publisher)
Created2013
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Description
Colorful ornaments in animals often serve as sexually selected signals of quality. While pigment-based colors are well-studied in these regards, structural colors that result from the interaction of light with photonic nanostructures are comparatively understudied in terms of their consequences in social contexts, their costs of production, and even the

Colorful ornaments in animals often serve as sexually selected signals of quality. While pigment-based colors are well-studied in these regards, structural colors that result from the interaction of light with photonic nanostructures are comparatively understudied in terms of their consequences in social contexts, their costs of production, and even the best way to measure them. Iridescent colors are some of the most brilliant and conspicuous colors in nature, and I studied the measurement, condition-dependence, and signaling role of iridescence in Anna's hummingbirds (Calypte anna). While most animal colors are easily quantified using well-established spectrophotometric techniques, the unique characteristics of iridescent colors present challenges to measurement and opportunities to quantify novel color metrics. I designed and tested an apparatus for careful control and measurement of viewing geometry and highly repeatable measurements. These measurements could be used to accurately characterize individual variation in iridescent Anna's hummingbirds to examine their condition-dependence and signaling role. Next, I examined the literature published to date for evidence of condition-dependence of structural colors in birds. Using meta-analyses, I found that structural colors of all three types - white, ultra-violet/blue, and iridescence - are significantly condition-dependent, meaning that they can convey information about quality to conspecifics. I then investigated whether iridescent colors were condition-dependent in Anna's hummingbirds both in a field correlational study and in an experimental study. Throughout the year, I found that iridescent feathers in both male and female Anna's hummingbirds become less brilliant as they age. Color was not correlated with body condition in any age/sex group. However, iridescent coloration in male Anna's hummingbirds was significantly affected by experimental protein in the diet during feather growth, indicating that iridescent color may signal diet quality. Finally, I examined how iridescent colors were used to mediate social competitions in male and female Anna's hummingbirds. Surprisingly, males that were less colorful won significantly more contests than more colorful males, and colorful males received more aggression. Less colorful males may be attempting to drive away colorful neighbors that may be preferred mates. Female iridescent ornament size and color was highly variable, but did not influence contest outcomes or aggression.
ContributorsMeadows, Melissa (Author) / McGraw, Kevin J. (Thesis advisor) / Rutowski, Ronald L (Committee member) / Sabo, John L (Committee member) / Alcock, John (Committee member) / Deviche, Pierre (Committee member) / Arizona State University (Publisher)
Created2012
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Description
Differences between males and females can evolve through a variety of mechanisms, including sexual and ecological selection. Because coloration is evolutionarily labile, sexually dichromatic species are good models for understanding the evolution of sex differences. While many jumping spiders exhibit diverse and brilliant coloration, they have been notably absent from

Differences between males and females can evolve through a variety of mechanisms, including sexual and ecological selection. Because coloration is evolutionarily labile, sexually dichromatic species are good models for understanding the evolution of sex differences. While many jumping spiders exhibit diverse and brilliant coloration, they have been notably absent from such studies. In the genus Habronattus, females are drab and cryptic while males are brilliantly colored, displaying some of these colors to females during elaborate courtship dances. Here I test multiple hypotheses for the control and function of male color. In the field, I found that Habronattus males indiscriminately court any female they encounter (including other species), so I first examined the role that colors play in species recognition. I manipulated male colors in H. pyrrithrix and found that while they are not required for species recognition, the presence of red facial coloration improves courtship success, but only if males are courting in the sun. Because light environment affects transmission of color signals, the multi-colored displays of males may facilitate communication in variable and unpredictable environments. Because these colors can be costly to produce and maintain, they also have the potential to signal reliable information about male quality to potential female mates. I found that both red facial and green leg coloration is condition dependent in H. pyrrithrix and thus has the potential to signal quality. Yet, surprisingly, this variation in male color does not appear to be important to females. Males of many Habronattus species also exhibit conspicuous markings on the dorsal surface of their abdomens that are not present in females and are oriented away from females during courtship. In the field, I found that these markings are paired with increased leg-waving behavior in a way that resembles the pattern and behavior of wasps; this may provide protection by exploiting the aversions of predators. My data also suggest that different activity levels between the sexes have placed different selection pressures on their dorsal color patterns. Overall, these findings challenge some of the traditional ways that we think about color signaling and provide novel insights into the evolution of animal coloration.
ContributorsTaylor, Lisa Anne (Author) / McGraw, Kevin J. (Thesis advisor) / Clark, David L. (Committee member) / Johnson, James C. (Committee member) / Alcock, John (Committee member) / Rutowski, Ronald L (Committee member) / Arizona State University (Publisher)
Created2012
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Description

River and riparian areas are important foraging habitat for insectivorous bats. Numerous studies have shown that aquatic insects provide an important trophic resource to terrestrial consumers, including bats, and are key in regulating population size and species interactions in terrestrial food webs. Yet these studies have generally ignored how structural

River and riparian areas are important foraging habitat for insectivorous bats. Numerous studies have shown that aquatic insects provide an important trophic resource to terrestrial consumers, including bats, and are key in regulating population size and species interactions in terrestrial food webs. Yet these studies have generally ignored how structural characteristics of the riverine landscape influence trophic resource availability or how terrestrial consumers respond to ensuing spatial and temporal patterns of trophic resources. Moreover, few studies have examined linkages between a stream's hydrologic regime and the timing and magnitude of aquatic insect availability. The main objective of my dissertation is to understand the causes of bat distributions in space and time. Specifically, I examine how trophic resource availability, structural components of riverine landscapes (channel confinement and riparian vegetation structure), and hydrologic regimes (flow permanence and timing of floods) mediate spatial and temporal patterns in bat activity. First, I show that river channel confinement determines bat activity along a river's longitudinal axis (directly above the river), while trophic resources appear to have stronger effects across a river's lateral (with distance from the river) axis. Second, I show that flow intermittency affects bat foraging activity indirectly via its effects on trophic resource availability. Seasonal river drying appears to have complex effects on bat foraging activity, initially causing imperfect tracking by consumers of localized concentrations of resources but later resulting in disappearance of both insects and bats after complete river drying. Third, I show that resource tracking by bats varies among streams with contrasting patterns of trophic resource availability and this variation appears to be in response to differences in the timing of aquatic insect emergence, duration and magnitude of emergence, and adult body size of emergent aquatic insects. Finally, I show that aquatic insects directly influence bat activity along a desert stream and that riparian vegetation composition affects bat activity, but only indirectly, via effects on aquatic insect availability. Overall, my results show river channel confinement, riparian vegetation structure, flow permanence, and the timing of floods influence spatial and temporal patterns in bat distributions; but these effects are indirect by influencing the ability of bats to track trophic resources in space and time.

ContributorsHagen, Elizabeth M (Author) / Sabo, John L (Thesis advisor) / Fisher, Stuart G. (Committee member) / Grimm, Nancy (Committee member) / Schmeeckle, Mark W (Committee member) / Stromberg, Juliet C. (Committee member) / Arizona State University (Publisher)
Created2010
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Description

More than half of all accessible freshwater has been appropriated for human use, and a substantial portion of terrestrial ecosystems have been transformed by human action. These impacts are heaviest in urban ecosystems, where impervious surfaces increase runoff, water delivery and stormflows are managed heavily, and there are substantial anthropogenic

More than half of all accessible freshwater has been appropriated for human use, and a substantial portion of terrestrial ecosystems have been transformed by human action. These impacts are heaviest in urban ecosystems, where impervious surfaces increase runoff, water delivery and stormflows are managed heavily, and there are substantial anthropogenic sources of nitrogen (N). Urbanization also frequently results in creation of intentional novel ecosystems. These "designed" ecosystems are fashioned to fulfill particular needs of the residents, or ecosystem services. In the Phoenix, Arizona area, the augmentation and redistribution of water has resulted in numerous component ecosystems that are atypical for a desert environment. Because these systems combine N loading with the presence of water, they may be hot spots of biogeochemical activity. The research presented here illustrates the types of hydrological modifications typical of desert cities and documents the extent and distribution of common designed aquatic ecosystems in the Phoenix metropolitan area: artificial lakes and stormwater retention basins. While both ecosystems were designed for other purposes (recreation/aesthetics and flood abatement, respectively), they have the potential to provide the added ecosystem service of N removal via denitrification. However, denitrification in urban lakes is likely to be limited by the rate of diffusion of nitrate into the sediment. Retention basins export some nitrate to groundwater, but grassy basins have higher denitrification rates than xeriscaped ones, due to higher soil moisture and organic matter content. An economic valuation of environmental amenities demonstrates the importance of abundant vegetation, proximity to water, and lower summer temperatures throughout the region. These amenities all may be provided by designed, water-intensive ecosystems. Some ecosystems are specifically designed for multiple uses, but maximizing one ecosystem service often entails trade-offs with other services. Further investigation into the distribution, bundling, and tradeoffs among water-related ecosystem services shows that some types of services are constrained by the hydrogeomorphology of the area, while for others human engineering and the creation of designed ecosystems has enabled the delivery of hydrologic ecosystem services independent of natural constraints.

ContributorsLarson, Elisabeth Knight (Author) / Grimm, Nancy (Thesis advisor) / Hartnett, Hilairy E (Committee member) / Fisher, Stuart G. (Committee member) / Anderies, John M (Committee member) / Lohse, Kathleen A (Committee member) / Arizona State University (Publisher)
Created2010
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Description
Animals have evolved a diversity of signaling traits, and in some species, they co-occur and are used simultaneously to communicate. Although much work has been done to understand why animals possess multiple signals, studies do not typically address the role of inter-signal interactions, which may vary intra- and inter-specifically and

Animals have evolved a diversity of signaling traits, and in some species, they co-occur and are used simultaneously to communicate. Although much work has been done to understand why animals possess multiple signals, studies do not typically address the role of inter-signal interactions, which may vary intra- and inter-specifically and help drive the evolutionary diversity in signals. For my dissertation, I tested how angle-dependent structural coloration, courtship displays, and the display environment interact and co-evolved in hummingbird species from the “bee” tribe (Mellisugini). Most “bee” hummingbird species possess an angle-dependent structurally colored throat patch and stereotyped courtship (shuttle) display. For 6 U.S. “bee” hummingbird species, I filmed male shuttle displays and mapped out the orientation- and-position-specific movements during the displays. With such display paths, I was able to then recreate each shuttle display in the field by moving plucked feathers from each male in space and time, as if they were naturally displaying, in order to measure each male’s color appearance during their display (i.e. the interactions between male hummingbird plumage, shuttle displays, and environment) from full-spectrum photographs. I tested how these interactions varied intra- and inter-specifically, and which of these originating traits might explain that variation. I first found that the solar-positional environment played a significant role in explaining variation in male color appearance within two species (Selasphorus platycercus and Calypte costae), and that different combinations of color-behavior-environment interactions made some males (in both species) appear bright, colorful, and flashy (i.e. their color appearance changes throughout a display), while other males maintained a consistent (non-flashing) color display. Among species, I found that plumage flashiness positively co-varied with male display behaviors, while another measure of male color appearance (average brightness/colorfulness) co-varied with the feather reflectance characteristics themselves. Additionally, species that had more exaggerated plumage features had less exaggerated shuttle displays. Altogether, my dissertation work illustrates the complexity of multiple signal evolution and how color-behavior-environment interactions are vital to understanding the evolution of colorful and behavioral display traits in animals.
ContributorsSimpson, Richard Kendall (Author) / McGraw, Kevin J. (Thesis advisor) / Rutowski, Ronald L (Committee member) / Pratt, Stephen C (Committee member) / Clark, Christopher J (Committee member) / McGuire, Jimmy A. (Committee member) / Arizona State University (Publisher)
Created2018
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Description
Sexual and social signals have long been thought to play an important role in speciation and diversity; hence, investigations of intraspecific communication may lead to important insights regarding key processes of evolution. Though we have learned much about the control, function, and evolution of animal communication by studying several very

Sexual and social signals have long been thought to play an important role in speciation and diversity; hence, investigations of intraspecific communication may lead to important insights regarding key processes of evolution. Though we have learned much about the control, function, and evolution of animal communication by studying several very common signal types, investigating rare classes of signals may provide new information about how and why animals communicate. My dissertation research focused on rapid physiological color change, a rare signal-type used by relatively few taxa. To answer longstanding questions about this rare class of signals, I employed novel methods to measure rapid color change signals of male veiled chameleons Chamaeleo calyptratus in real-time as seen by the intended conspecific receivers, as well as the associated behaviors of signalers and receivers. In the context of agonistic male-male interactions, I found that the brightness achieved by individual males and the speed of color change were the best predictors of aggression and fighting ability. Conversely, I found that rapid skin darkening serves as a signal of submission for male chameleons, reducing aggression from winners when displayed by losers. Additionally, my research revealed that the timing of maximum skin brightness and speed of brightening were the best predictors of maximum bite force and circulating testosterone levels, respectively. Together, these results indicated that different aspects of color change can communicate information about contest strategy, physiology, and performance ability. Lastly, when I experimentally manipulated the external appearance of chameleons, I found that "dishonestly" signaling individuals (i.e. those whose behavior did not match their manipulated color) received higher aggression from unpainted opponents. The increased aggression received by dishonest signalers suggests that social costs play an important role in maintaining the honesty of rapid color change signals in veiled chameleons. Though the color change abilities of chameleons have interested humans since the time of Aristotle, little was previously known about the signal content of such changes. Documenting the behavioral contexts and information content of these signals has provided an important first step in understanding the current function, underlying control mechanisms, and evolutionary origins of this rare signal type.
ContributorsLigon, Russell (Author) / McGraw, Kevin J. (Committee member) / DeNardo, Dale F (Committee member) / Karsten, Kristopher B (Committee member) / Rutowski, Ronald L (Committee member) / Deviche, Pierre (Committee member) / Arizona State University (Publisher)
Created2015
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Description
Although mimetic animal coloration has been studied since Darwin's time, many questions on the efficacy, evolution, and function of mimicry remain unanswered. Müller (1879) hypothesized that unpalatable individuals converge on the same conspicuous coloration to reduce predation. However, there are many cases where closely related, unpalatable species have diverged from

Although mimetic animal coloration has been studied since Darwin's time, many questions on the efficacy, evolution, and function of mimicry remain unanswered. Müller (1879) hypothesized that unpalatable individuals converge on the same conspicuous coloration to reduce predation. However, there are many cases where closely related, unpalatable species have diverged from a shared conspicuous pattern. What selection pressures have led to divergence in warning colors? Environmental factors such as ambient light have been hypothesized to affect signal transmission and efficacy in animals. Using two mimetic pairs of Heliconius butterflies, Postman and Blue-white, I tested the hypothesis that animals with divergent mimetic colors segregate by light environment to maximize conspicuousness of the aposematic warning signal under their particular environmental conditions. Each mimetic pair was found in a light environment that differed in brightness and spectral composition, which affected visual conspicuousness differently depending on mimetic color patch. I then used plasticine models in the field to test the hypothesis that mimics had higher survival in the habitat where they occurred. Although predation rates differed between the two habitats, there was no interactive effect of species by habitat type. Through choice experiments, I demonstrated that mimetic individuals preferred to spend time in the light environment where they were most often found and that their absolute visual sensitivity corresponds to the ambient lighting of their respective environment. Eye morphology was then studied to determine if differences in total corneal surface area and/or facet diameters explained the differences in visual sensitivities, but the differences found in Heliconius eye morphology did not match predictions based upon visual sensitivity. To further understand how eye morphology varies with light environments, I studied many tropical butterflies from open and closed habitats to reveal that forest understory butterflies have larger facets compared to butterflies occupying open habitats. Lastly, I tested avian perception of mimicry in a putative Heliconius mimetic assemblage and show that the perceived mimetic resemblance depends upon visual system. This dissertation reveals the importance of light environments on mimicry, coloration, behavior and visual systems of tropical butterflies.
ContributorsSeymoure, Brett M (Author) / Rutowski, Ronald L (Thesis advisor) / McGraw, Kevin J. (Thesis advisor) / McMillan, W. Owen (Committee member) / Pratt, Stephen (Committee member) / Gadau, Jürgen (Committee member) / Arizona State University (Publisher)
Created2016
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Description
Urbanization provides an excellent opportunity to examine the effects of human-induced rapid environmental change (HIREC) on natural ecosystems. Certain species can dominate in urban habitats at the expense of biodiversity. Phenotypic plasticity may be the mechanism by which these 'urban exploiters' flourish in urban areas. Color displays and condition-dependent phenotypes

Urbanization provides an excellent opportunity to examine the effects of human-induced rapid environmental change (HIREC) on natural ecosystems. Certain species can dominate in urban habitats at the expense of biodiversity. Phenotypic plasticity may be the mechanism by which these 'urban exploiters' flourish in urban areas. Color displays and condition-dependent phenotypes are known to be highly plastic. However, conspicuous color displays are perplexing in that they can be costly to produce and may increase detection by enemies. The Western black widow spider () is a superabundant pest species that forms dense aggregations throughout metropolitan Phoenix, Arizona, USA. Adult female display a red hourglass on their abdomen, which is speculated to function as a conspicuous warning signal to enemies. Here, I performed field studies to identify how widow morphology and hourglass color differ between urban and desert subpopulations. I also conducted laboratory experiments to examine the dietary sensitivity of hourglass coloration and to identify its functional role in the contexts of agonism, mating, and predator defense. My field data reveal significant spatial variation across urban and desert subpopulations in ecology and color. Furthermore, hourglass coloration was significantly influenced by environmental factors unique to urban habitats. Desert spiders were found to be smaller and less colorful than urban spiders. Throughout, I observed a positive correlation between body condition and hourglass size. Laboratory diet manipulations empirically confirm the condition-dependence of hourglass size. Additionally, widows with extreme body conditions exhibited condition-dependent coloration. However, hourglass obstruction and enlargement did not produce any effects on the outcome of agonistic encounters, male courtship, or predator deterrence. This work offers important insights into the effects of urbanization on the ecology and coloration of a superabundant pest species. While the function of the hourglass remains undetermined, my findings characterize the black widow's hourglass as extremely plastic. Plastic responses to novel environmental conditions can modify the targets of natural selection and subsequently influence evolutionary outcomes. Therefore, assuming a heritable component to this plasticity, the response of hourglass plasticity to the abrupt environmental changes in urban habitats may result in the rapid evolution of this phenotype.
ContributorsGburek, Theresa (Author) / Johnson, James C. (Thesis advisor) / McGraw, Kevin J. (Committee member) / Rutowski, Ronald L (Committee member) / Arizona State University (Publisher)
Created2014