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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
The nests of the Curve-billed Thrasher (Toxostoma curvirostre) were studied across the greater Phoenix area from 2020-2022 in order to assess any significant relationships between their composition and the composition of their environment. Nests were collected and measured, and the vegetation was surveyed to 100 m for potential nest material

The nests of the Curve-billed Thrasher (Toxostoma curvirostre) were studied across the greater Phoenix area from 2020-2022 in order to assess any significant relationships between their composition and the composition of their environment. Nests were collected and measured, and the vegetation was surveyed to 100 m for potential nest material type. In the lab, nests were separated by material type and tallied. The dense cores of the nests received a 100-piece sampling, with the first hundred pieces plucked from the structure, sorted by type, and massed. Ordinary least squares (OLS) and binomial regression analyses were performed on the body tallies and their corresponding site tallies. Core material weights and their corresponding site tallies only received OLS regression analyses. Beta regression analyses were also performed on the mass proportions of core samples and their corresponding environmental tallies. OLS regression yielded a significant relationship between the spiny body material tally and its site tallies at 25 and 100 m. While failing the assumption of normality, the tally of barrel cactus in a nest body yielded significant p-values in OLS and binomial regression, as well as the Spearman’s correlation test, supporting a strong correlation with the 100m site tally. The tally of anthropogenic materials and the distance to the nearest man-made structure failed the test of normality, but yielded significant p-values in binomial regression and the Spearman’s correlation test. OLS regression of log anthropogenic tally and log distance to nearest structure failed normality but yielded a significant p-value as well. In beta regression analyses, only the spiny core mass proportion yielded a significant relationship at the 100 m site tally.
ContributorsMotta, Anthony Joseph (Author) / Taylor, Jay (Thesis advisor) / Makings, Elizabeth (Committee member) / Puente, Raul (Committee member) / Arizona State University (Publisher)
Created2022
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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
Understanding the effects of fire on the Sonoran Desert is of critical importance as rising temperatures and changing weather patterns increase the frequency and size of wildfires. Seed banks are an important component in post-fire landscape recovery as the seeds that remain in the soil are an indicator of a

Understanding the effects of fire on the Sonoran Desert is of critical importance as rising temperatures and changing weather patterns increase the frequency and size of wildfires. Seed banks are an important component in post-fire landscape recovery as the seeds that remain in the soil are an indicator of a landscape’s future trajectory. The purpose of this study is to determine the lasting impacts of fire on the soil seed bank of the Sonoran Desert and to identify potential concerns affecting post-fire recovery and restoration. The study site was located in the Arizona Upland division of the Sonoran Desert, Arizona, United States. Soil samples were collected from five burned sites with increasing time since fire, and five nearby unburned sites used as a control. A seedling emergence test was conducted to investigate the density and richness of the seed bank of burned and unburned sites. Seed densities and species richness for sites were calculated using germination results. Findings were analyzed using non-parametric analyses comparing changes in burned and unburned sites over time. Results found that burn status and time since fire had no significant impact on seed density. Graminoid and forb densities were statistically consistent across burn status and time since fire. While species richness was consistent across both plot types, burned samples typically had fewer species than unburned samples. Burned and unburned plots revealed a dominance of annual species with limited presence of woody perennials. While seed densities and species richness are relatively unchanged across burned and unburned sites over time, the lack of woody perennials in the seed bank raises concerns about landscape recovery trajectories in burned sites. These results suggest that restoration efforts focused on maintaining the presence of woody trees and shrubs in the landscape may have the most impact.
ContributorsCountryman, Kristen (Author) / Makings, Elizabeth (Thesis advisor) / Pigg, Kathleen (Thesis advisor) / Lata, Mary (Committee member) / Arizona State University (Publisher)
Created2023
Description
Urban wetland ecosystems provide myriad ecosystem services and are shaped by diverse social and ecological factors. In rapidly urbanizing parts of the desert Southwest, wetlands are especially vital. Across less than 60 km as it enters the Phoenix area, the Salt River is dammed, diverted, re-filled, clear-cut, restored, and ignored.

Urban wetland ecosystems provide myriad ecosystem services and are shaped by diverse social and ecological factors. In rapidly urbanizing parts of the desert Southwest, wetlands are especially vital. Across less than 60 km as it enters the Phoenix area, the Salt River is dammed, diverted, re-filled, clear-cut, restored, and ignored. This study documents how animal and plant communities in three perennially inundated reaches of the river changed over a decade under different social-ecological pressures. One wetland in the urban core is restored, another formed accidentally by human infrastructure, and the last is managed on the urban periphery. Surveys conducted since 2012 used point-count surveys to assess bird communities and visual encounter surveys to assess reptiles and amphibians. Plant communities were surveyed in 2012 and 2022 using cover classes. Between 2012 and 2022, accidental and restored wetlands close to the urban core displayed an increase in plant abundance, largely consisting of introduced species. While all sites saw an increase in plant species considered invasive by land management groups, both urban wetlands saw an increase in regionally native species, including plants that are culturally significant to local Indigenous groups. Reptile communities declined in richness and abundance in both urban sites, but birds grew in abundance and richness at the urban restored site while not changing at the urban accidental wetland. The non-urban site saw stable populations of both birds and herpetofauna. These trends in biotic communities reveal ecological tradeoffs under different management strategies for urban wetlands. These findings also create a portrait of wetland communities along a rapidly urbanizing arid river. As the Salt River watershed becomes more urbanized, it is important to establish a more empathetic and informed relationship between its plant and animal—including human—residents. To this end, these data were incorporated in a series of handmade paper artworks, crafted from the most abundant wetland plant species found at the study sites, harvested alongside local land management efforts. These artworks examine the potential of four common cosmopolitan wetland plants for papermaking, revealing the potential to align ecosystem management efforts with both materials production and fine arts. By using relief printmaking to visualize long-term ecological data, I explored an alternative, more creative and embodied way to engage with and visualize urban wetland communities. This alternate mode of engagement can complement ecological management and research to diversify disciplines and participants engaged with understanding and living alongside urban wetlands.
ContributorsRamsey-Wiegmann, Luke Dawson (Author) / Childers, Daniel L (Thesis advisor) / Makings, Elizabeth (Committee member) / Bateman, Heather (Committee member) / Arizona State University (Publisher)
Created2023
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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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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
Stream flow permanence plays a critical role in determining floristic composition, abundance, and diversity in the Sonoran Desert, but questions remain about the effects of stream flow permanence on butterfly composition, abundance, and diversity. Understanding the effects of flow permanence on butterflies and relevant subsets of butterflies (such as butterflies

Stream flow permanence plays a critical role in determining floristic composition, abundance, and diversity in the Sonoran Desert, but questions remain about the effects of stream flow permanence on butterfly composition, abundance, and diversity. Understanding the effects of flow permanence on butterflies and relevant subsets of butterflies (such as butterflies whose host plants are present) and comparing them to these same effects on plants and relevant subsets of plants (such as butterfly nectar plants and larval host plants) provided insight into pollinator and riparian conservation and restoration.

I surveyed four Sonoran desert stream sites, and found significant relationships between flow permanence and plant and butterfly species richness and abundance, as well as strong relationships between plant and butterfly abundance and between plant and butterfly species richness. Most notably, my results pointed to hosted butterflies as a break-out category of butterflies which may more clearly delineate ecological relationships between butterfly and plant abundance and diversity along Sonoran Desert streams; this can inform conservation decisions. Managing for hosted (resident) butterflies will necessarily entail managing for the presence of surface water, nectar forage, varying levels of canopy cover, and plant, nectar plant, and host plant diversity since the relationships between hosted butterfly species richness and/or abundance and all of these variables were significant, both statistically and ecologically.
ContributorsButler, Lane (Author) / Stromberg, Juliet C. (Thesis advisor) / Makings, Elizabeth (Committee member) / Pearson, David L (Committee member) / Boggess, May (Committee member) / Buchmann, Stephen (Committee member) / Arizona State University (Publisher)
Created2015