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Climate change has the potential to affect vegetation via changes in temperature and precipitation. In the semi-arid southwestern United States, heightened temperatures will likely lead to accelerated groundwater pumping to meet human needs, and altered storm patterns may lead to changes in flood regimes. All of these hydrologic changes have

Climate change has the potential to affect vegetation via changes in temperature and precipitation. In the semi-arid southwestern United States, heightened temperatures will likely lead to accelerated groundwater pumping to meet human needs, and altered storm patterns may lead to changes in flood regimes. All of these hydrologic changes have the potential to alter riparian vegetation. This research, consisting of two papers, examines relationships between hydrology and riparian vegetation along the Verde River in central Arizona, from applied and theoretical perspectives. One paper investigates how dominance of tree and shrub species and cover of certain functional groups change along hydrologic gradients. The other paper uses the Verde River flora along with that river's flood and moisture gradients to answer the question of whether functional groups can be defined universally. Drying of the Verde River would lead to a shift from cottonwood-willow streamside forest to more drought adapted desert willow or saltcedar, a decline in streamside marsh species, and decreased species richness. Effects drying will have on one dominant forest tree, velvet ash, is unclear. Increase in the frequency of large floods would potentially increase forest density and decrease average tree age and diameter. Correlations between functional traits of Verde River plants and hydrologic gradients are consistent with "leaf economics," or the axis of resource capture, use, and release, as the primary strategic trade-off for plants. This corresponds to the competitor-stress tolerator gradient in Grime's life history strategy theory. Plant height was also a strong indicator of hydrologic condition, though it is not clear from the literature if plant height is independent enough of leaf characteristics on a global scale to be considered a second axis. Though the ecohydrologic relationships are approached from different perspectives, the results of the two papers are consistent if interpreted together. The species that are currently dominant in the near-channel Verde River floodplain are tall, broad-leaf trees, and the species that are predicted to become more dominant in the case of the river drying are shorter trees or shrubs with smaller leaves. These results have implications for river and water management, as well as theoretical ecology.
ContributorsHazelton, Andrea Florence (Author) / Stromberg, Juliet C. (Thesis advisor) / Schmeeckle, Mark W (Committee member) / Franklin, Janet (Committee member) / Arizona State University (Publisher)
Created2011
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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
Climate and land use change are projected to threaten biodiversity over the coming century. However, the combined effects of these threats on biodiversity and the capacity of current conservation networks to protect species' habitat are not well understood. The goals of this study were to evaluate the effect of climate

Climate and land use change are projected to threaten biodiversity over the coming century. However, the combined effects of these threats on biodiversity and the capacity of current conservation networks to protect species' habitat are not well understood. The goals of this study were to evaluate the effect of climate change and urban development on vegetation distribution in a Mediterranean-type ecosystem; to identify the primary source of uncertainty in suitable habitat predictions; and to evaluate how well conservation areas protect future habitat in the Southwest ecoregion of the California Floristic Province. I used a consensus-based modeling approach combining three different species distribution models to predict current and future suitable habitat for 19 plant species representing different plant functional types (PFT) defined by fire-response (obligate seeders, resprouting shrubs), and life forms (herbs, subshurbs). I also examined the response of species grouped by range sizes (large, small). I used two climate models, two emission scenarios, two thresholds, and high-resolution (90m resolution) environmental data to create a range of potential scenarios. I evaluated the effectiveness of an existing conservation network to protect suitable habitat for rare species in light of climate and land use change. The results indicate that the area of suitable habitat for each species varied depending on the climate model, emission scenario, and threshold combination. The suitable habitat for up to four species could disappear from the ecoregion, while suitable habitat for up to 15 other species could decrease under climate change conditions. The centroid of the species' suitable environmental conditions could shift up to 440 km. Large net gains in suitable habitat were predicted for a few species. The suitable habitat area for herbs has a small response to climate change, while obligate seeders could be the most affected PFT. The results indicate that the other two PFTs gain a considerable amount of suitable habitat area. Several rare species could lose suitable habitat area inside designated conservation areas while gaining suitable habitat area outside. Climate change is predicted to be more important than urban development as a driver of habitat loss for vegetation in this region in the coming century. These results indicate that regional analyses of this type are useful and necessary to understand the dynamics of drivers of change at the regional scale and to inform decision making at this scale.
ContributorsBeltrán Villarreal, Bray de Jesús (Author) / Franklin, Janet (Thesis advisor) / Fenichel, Eli P (Committee member) / Kinzig, Ann P (Committee member) / Collins, James P. (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
In many social groups, reproduction is shared between group members, whocompete for position in the social hierarchy for reproductive dominance. This reproductive conflict can lead to different means of enforcing reproductive differences, such as dominance displays or limited control of social hierarchy through antagonistic encounters. In eusocial insects, archetypal colonies contain a single,

In many social groups, reproduction is shared between group members, whocompete for position in the social hierarchy for reproductive dominance. This reproductive conflict can lead to different means of enforcing reproductive differences, such as dominance displays or limited control of social hierarchy through antagonistic encounters. In eusocial insects, archetypal colonies contain a single, singly-mated fertile queen, such that no reproductive conflict exists within a colony. However, many eusocial insects deviate from this archetype and have multiply-mated queens (polyandry), multiple queens in a single colony (polygyny), or both. In these cases, reproductive conflict exists between the matrilines and patrilines represented in a colony, specifically over the production of sexual offspring. A possible outcome of reproductive conflict may be the emergence of cheating lineages, which favor the production of sexual offspring, taking advantage of the worker force produced by nestmate queens and/or patrilines. In extreme examples, inquiline social parasites may be an evolutionary consequence of reproductive conflict between nestmate queens. Inquiline social parasitism is a type of social parasitism that is usually defined by a partial or total loss of the worker caste, and the “infiltration” of host colonies to take advantage of the host worker force for reproduction. It has been hypothesized that these inquiline social parasites evolve through the speciation of cheating queen lineages from within their incipient host species. This “intra- specific” origin model involves a foundational hypothesis that the common ancestor of host and parasite (and thus, putatively, the host at the time of speciation) should be functionally polygynous, and that parasitism evolves as a “resolution” of reproductive conflict in colonies. In this dissertation, I investigate the hypothesized role of polygyny in the evolution of inquiline social parasites. I use molecular ecology and statistical approaches to validate the role of polygyny in the evolution of some inquiline social parasites. I further discuss potential mechanisms for the evolution and speciation of social parasites, and discuss future directions to elucidate these mechanisms.
ContributorsDahan, Romain Arvid (Author) / Rabeling, Christian (Thesis advisor) / Amdam, Gro V (Committee member) / Fewell, Jennifer H (Committee member) / Pratt, Stephen C (Committee member) / Rüppell, Olav (Committee member) / Arizona State University (Publisher)
Created2021
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Description
Climate and environmental forcing are widely accepted to be important drivers of evolutionary and ecological change in mammal communities over geologic time scales. This paradigm has been particularly influential in studies of the eastern African late Cenozoic fossil record, in which aridification, increasing seasonality, and C4 grassland expansion are seen

Climate and environmental forcing are widely accepted to be important drivers of evolutionary and ecological change in mammal communities over geologic time scales. This paradigm has been particularly influential in studies of the eastern African late Cenozoic fossil record, in which aridification, increasing seasonality, and C4 grassland expansion are seen as having shaped the major patterns of human and faunal evolution. Despite the ubiquity of studies linking climate and environmental forcing to evolutionary and ecological shifts in the mammalian fossil record, many central components of this paradigm remain untested or poorly developed. To fill this gap, this dissertation employs biogeographical and macroecological analyses of present-day African mammal communities as a lens for understanding how abiotic change may have shaped community turnover and structure in the eastern African Plio-Pleistocene. Three dissertation papers address: 1) the role of ecological niche breadth in shaping divergent patterns of macroevolutionary turnover across clades; 2) the effect of climatic and environmental gradients on community assembly; 3) the relative influence of paleo- versus present-day climates in structuring contemporary patterns of community diversity. Results of these papers call into question many tenets of current theory, particularly: 1) that niche breadth differences (and, by extension, their influence on allopatric speciation) are important drivers of macroevolution, 2) that climate is more important than biotic interactions in community assembly, and 3) that communities today are in equilibrium with present-day climates. These findings highlight the need to critically reevaluate the role and scale-dependence of climate in mammal evolution and community ecology and to carefully consider potential time lags and disequilibrium dynamics in the fossil record.
ContributorsRowan, John (Author) / Reed, Kaye E (Thesis advisor) / Campisano, Christopher J (Committee member) / Franklin, Janet (Committee member) / Marean, Curtis W (Committee member) / Arizona State University (Publisher)
Created2018
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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
Urban riparian corridors have the capacity to maintain high levels of abundance and biodiversity. Additionally, urban rivers also offer environmental amenities and can be catalysts for social and economic revitalization in human communities. Despite its importance for both humans and wildlife, blue space in cities used by waterbirds has received

Urban riparian corridors have the capacity to maintain high levels of abundance and biodiversity. Additionally, urban rivers also offer environmental amenities and can be catalysts for social and economic revitalization in human communities. Despite its importance for both humans and wildlife, blue space in cities used by waterbirds has received relatively little focus in urban bird studies. My principal objective was to determine how urbanization and water availability affect waterbird biodiversity in an arid city. I surveyed 36 transects stratified across a gradient of urbanization and water availability along the Salt River, a LTER long-term study system located in Phoenix, Arizona. Water physiognomy (shape and size) was the largest factor in shaping the bird community. Connectivity was an important element for waterbird diversity, but not abundance. Urbanization had guild-specific effects on abundance but was not important for waterbird diversity. Habitat-level environmental characteristics were more important than land use on waterbird abundance, as well as diversity. Diving and fish-eating birds were positively associated with large open bodies of water, whereas dabbling ducks, wading birds, and marsh species favored areas with large amounts of shoreline and emergent vegetation. My study supports that Phoenix blue space offers an important subsidy to migrating waterbird communities; while alternative habitat is not a replacement, it is important to consider as part of the larger conservation picture as traditional wetlands decline. Additionally, arid cities have the potential to support high levels of waterbird biodiversity, heterogeneous land use matrix can be advantageous in supporting regional diversity, and waterbirds are tolerant of urbanization if proper resources are provided via the habitat. The implications of this study are particularly relevant to urban planning in arid cities; Phoenix alone contains over 1,400 bodies of water, offering the opportunity to design and improve urban blue space to optimize potential habitat while providing public amenities.
ContributorsBurnette, Riley (Author) / Bateman, Heather (Thesis advisor) / Franklin, Janet (Committee member) / Allen, Daniel (Committee member) / Arizona State University (Publisher)
Created2016
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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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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