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Speciation is the fundamental process that has generated the vast diversity of life on earth. The hallmark of speciation is the evolution of barriers to gene flow. These barriers may reduce gene flow either by keeping incipient species from hybridizing at all (pre-zygotic), or by reducing the fitness of hybrids

Speciation is the fundamental process that has generated the vast diversity of life on earth. The hallmark of speciation is the evolution of barriers to gene flow. These barriers may reduce gene flow either by keeping incipient species from hybridizing at all (pre-zygotic), or by reducing the fitness of hybrids (post-zygotic). To understand the genetic architecture of these barriers and how they evolve, I studied a genus of wasps that exhibits barriers to gene flow that act both pre- and post-zygotically. Nasonia is a genus of four species of parasitoid wasps that can be hybridized in the laboratory. When two of these species, N. vitripennis and N. giraulti are mated, their offspring suffer, depending on the generation and cross examined, up to 80% mortality during larval development due to incompatible genic interactions between their nuclear and mitochondrial genomes. These species also exhibit pre-zygotic isolation, meaning they are more likely to mate with their own species when given the choice. I examined these two species and their hybrids to determine the genetic and physiological bases of both speciation mechanisms and to understand the evolutionary forces leading to them. I present results that indicate that the oxidative phosphorylation (OXPHOS) pathway, an essential pathway that is responsible for mitochondrial energy generation, is impaired in hybrids of these two species. These results indicate that this impairment is due to the unique evolutionary dynamics of the combined nuclear and mitochondrial origin of this pathway. I also present results showing that, as larvae, these hybrids experience retarded growth linked to the previously observed mortality and I explore possible physiological mechanisms for this. Finally, I show that the pre-mating isolation is due to a change in a single pheromone component in N. vitripennis males, that this change is under simple genetic control, and that it evolved neutrally before being co-opted as a species recognition signal. These results are an important addition to our overall understanding of the mechanisms of speciation and showcase Nasonia as an emerging model for the study of the genetics of speciation.
ContributorsGibson, Joshua D (Author) / Gadau, Jürgen (Thesis advisor) / Harrison, Jon (Committee member) / Pratt, Stephen (Committee member) / Verrelli, Brian (Committee member) / Willis, Wayne (Committee member) / Arizona State University (Publisher)
Created2013
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Description
To address the need of scientists and engineers in the United States workforce and ensure that students in higher education become scientifically literate, research and policy has called for improvements in undergraduate education in the sciences. One particular pathway for improving undergraduate education in the science fields is to reform

To address the need of scientists and engineers in the United States workforce and ensure that students in higher education become scientifically literate, research and policy has called for improvements in undergraduate education in the sciences. One particular pathway for improving undergraduate education in the science fields is to reform undergraduate teaching. Only a limited number of studies have explored the pedagogical content knowledge of postsecondary level teachers. This study was conducted to characterize the PCK of biology faculty and explore the factors influencing their PCK. Data included semi-structured interviews, classroom observations, documents, and instructional artifacts. A qualitative inquiry was designed to conduct an in-depth investigation focusing on the PCK of six biology instructors, particularly the types of knowledge they used for teaching biology, their perceptions of teaching, and the social interactions and experiences that influenced their PCK. The findings of this study reveal that the PCK of the biology faculty included eight domains of knowledge: (1) content, (2) context, (3) learners and learning, (4) curriculum, (5) instructional strategies, (6) representations of biology, (7) assessment, and (8) building rapport with students. Three categories of faculty PCK emerged: (1) PCK as an expert explainer, (2) PCK as an instructional architect, and (3) a transitional PCK, which fell between the two prior categories. Based on the interpretations of the data, four social interactions and experiences were found to influence biology faculty PCK: (1) teaching experience, (2) models and mentors, (3) collaborations about teaching, and (4) science education research. The varying teaching perspectives of the faculty also influenced their PCK. This study shows that the PCK of biology faculty for teaching large introductory courses at large research institutions is heavily influenced by factors beyond simply years of teaching experience and expert content knowledge. Social interactions and experiences created by the institution play a significant role in developing the PCK of biology faculty.
ContributorsHill, Kathleen M. (Author) / Luft, Julie A. (Thesis advisor) / Baker, Dale (Committee member) / Orchinik, Miles (Committee member) / Arizona State University (Publisher)
Created2013
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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
The repression of reproductive competition and the enforcement of altruism are key components to the success of animal societies. Eusocial insects are defined by having a reproductive division of labor, in which reproduction is relegated to one or few individuals while the rest of the group members maintain the colony

The repression of reproductive competition and the enforcement of altruism are key components to the success of animal societies. Eusocial insects are defined by having a reproductive division of labor, in which reproduction is relegated to one or few individuals while the rest of the group members maintain the colony and help raise offspring. However, workers have retained the ability to reproduce in most insect societies. In the social Hymenoptera, due to haplodiploidy, workers can lay unfertilized male destined eggs without mating. Potential conflict between workers and queens can arise over male production, and policing behaviors performed by nestmate workers and queens are a means of repressing worker reproduction. This work describes the means and results of the regulation of worker reproduction in the ant species Aphaenogaster cockerelli. Through manipulative laboratory studies on mature colonies, the lack of egg policing and the presence of physical policing by both workers and queens of this species are described. Through chemical analysis and artificial chemical treatments, the role of cuticular hydrocarbons as indicators of fertility status and the informational basis of policing in this species is demonstrated. An additional queen-specific chemical signal in the Dufour's gland is discovered to be used to direct nestmate aggression towards reproductive competitors. Finally, the level of actual worker-derived males in field colonies is measured. Together, these studies demonstrate the effectiveness of policing behaviors on the suppression of worker reproduction in a social insect species, and provide an example of how punishment and the threat of punishment is a powerful force in maintaining cooperative societies.
ContributorsSmith, Adrian A. (Author) / Liebig, Juergen (Thesis advisor) / Hoelldobler, Bert (Thesis advisor) / Gadau, Juergen (Committee member) / Johnson, Robert A. (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2011
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Description
Gene-centric theories of evolution by natural selection have been popularized and remain generally accepted in both scientific and public paradigms. While gene-centrism is certainly parsimonious, its explanations fall short of describing two patterns of evolutionary and social phenomena: the evolution of sex and the evolution of social altruism. I review

Gene-centric theories of evolution by natural selection have been popularized and remain generally accepted in both scientific and public paradigms. While gene-centrism is certainly parsimonious, its explanations fall short of describing two patterns of evolutionary and social phenomena: the evolution of sex and the evolution of social altruism. I review and analyze current theories on the evolution of sex. I then introduce the conflict presented to gene-centric evolution by social phenomena such as altruism and caste sterility in eusocial insects. I review gene-centric models of inclusive fitness and kin selection proposed by Hamilton and Maynard Smith. Based their assumptions, that relatedness should be equal between sterile workers and reproductives, I present several empirical examples that conflict with their models. Following that, I introduce a unique system of genetic caste determination (GCD) observed in hybrid populations of two sister-species of seed harvester ants, Pogonomyrmex rugosus and Pogonomyrmex barbatus. I review the evidence for GCD in those species, followed by a critique of the current gene-centric models used to explain it. In chapter two I present my own theoretical model that is both simple and extricable in nature to explain the origin, evolution, and maintenance of GCD in Pogonomyrmex. Furthermore, I use that model to fill in the gaps left behind by the contributing authors of the other GCD models. As both populations in my study system formed from inter-specific hybridization, I review modern discussions of heterosis (also called hybrid vigor) and use those to help explain the ecological competitiveness of GCD. I empirically address the inbreeding depression the lineages of GCD must overcome in order to remain ecologically stable, demonstrating that as a result of their unique system of caste determination, GCD lineages have elevated recombination frequencies. I summarize and conclude with an argument for why GCD evolved under selective mechanisms which cannot be considered gene-centric, providing evidence that natural selection can effectively operate on non-heritable genotypes appearing in groups and other social contexts.
ContributorsJacobson, Neal (Author) / Gadau, Juergen (Thesis advisor) / Laubichler, Manfred (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2012
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Description
Though for most of the twentieth century, dogma held that the adult brain was post-mitotic, it is now known that adult neurogenesis is widespread among vertebrates, from fish, amphibians, reptiles and birds to mammals including humans. Seasonal changes in adult neurogenesis are well characterized in the song control system of

Though for most of the twentieth century, dogma held that the adult brain was post-mitotic, it is now known that adult neurogenesis is widespread among vertebrates, from fish, amphibians, reptiles and birds to mammals including humans. Seasonal changes in adult neurogenesis are well characterized in the song control system of song birds, and have been found in seasonally breeding mammals as well. In contrast to more derived vertebrates, such as mammals, where adult neurogenesis is restricted primarily to the olfactory bulb and the dentate gyrus of the hippocampus, neurogenesis is widespread along the ventricles of adult amphibians. I hypothesized that seasonal changes in adult amphibian brain cell proliferation and survival are a potential regulator of reproductive neuroendocrine function. Adult, male American bullfrogs (Rana catesbeiana; aka Lithobates catesbeianus), were maintained in captivity for up to a year under season-appropriate photoperiod. Analysis of hormone levels indicated seasonal changes in plasma testosterone concentration consistent with field studies. Using the thymidine analogue 5-bromo-2-deoxyuridine (BrdU) as a marker for newly generated cells, two differentially regulated aspects of brain cell neogenesis were tracked; that is, proliferation and survival. Seasonal differences were found in BrdU labeling in several brain areas, including the olfactory bulb, medial pallium, nucleus accumbens and the infundibular hypothalamus. Clear seasonal differences were also found in the pars distalis region of the pituitary gland, an important component of neuroendocrine pathways. BrdU labeling was also examined in relation to two neuropeptides important for amphibian reproduction: arginine vasotocin and gonadotropin releasing hormone. No cells co-localized with BrdU and either neuropeptide, but new born cells were found in close proximity to neuropeptide-containing neurons. These data suggest that seasonal differences in brain and pituitary gland cell neogenesis are a potential neuroendocrine regulatory mechanism.
ContributorsMumaw, Luke (Author) / Orchinik, Miles (Thesis advisor) / Deviche, Pierre (Committee member) / Chandler, Douglas (Committee member) / Arizona State University (Publisher)
Created2012
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Description
Guided by Tinto’s Theory of College Student Departure, I conducted a set of five studies to identify factors that influence students’ social integration in college science active learning classes. These studies were conducted in large-enrollment college science courses and some were specifically conducted in undergraduate active learning biology courses.

Guided by Tinto’s Theory of College Student Departure, I conducted a set of five studies to identify factors that influence students’ social integration in college science active learning classes. These studies were conducted in large-enrollment college science courses and some were specifically conducted in undergraduate active learning biology courses. Using qualitative and quantitative methodologies, I identified how students’ identities, such as their gender and LGBTQIA identity, and students’ perceptions of their own intelligence influence their experience in active learning science classes and consequently their social integration in college. I also determined factors of active learning classrooms and instructor behaviors that can affect whether students experience positive or negative social integration in the context of active learning. I found that students’ hidden identities, such as the LGBTQIA identity, are more relevant in active learning classes where students work together and that the increased relevance of one’s identity can have a positive and negative impact on their social integration. I also found that students’ identities can predict their academic self-concept, or their perception of their intelligence as it compares to others’ intelligence in biology, which in turn predicts their participation in small group-discussion. While many students express a fear of negative evaluation, or dread being evaluated negatively by others when speaking out in active learning classes, I identified that how instructors structure group work can cause students to feel more or less integrated into the college science classroom. Lastly, I identified tools that instructors can use, such as name tents and humor, which can positive affect students’ social integration into the college science classroom. In sum, I highlight inequities in students’ experiences in active learning science classrooms and the mechanisms that underlie some of these inequities. I hope this work can be used to create more inclusive undergraduate active learning science courses.
ContributorsCooper, Katelyn M (Author) / Brownell, Sara E (Thesis advisor) / Stout, Valerie (Committee member) / Collins, James (Committee member) / Orchinik, Miles (Committee member) / Zheng, Yi (Committee member) / Arizona State University (Publisher)
Created2018
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Description
Persistent cooperation between unrelated conspecifics rarely occurs in mature eusocial insect societies. In this dissertation, I present evidence of non-kin cooperation in the Nearctic honey ant Myrmecocystus mendax. Using microsatellite markers, I show that mature colonies in the Sierra Ancha Mountain of central Arizona contain multiple unrelated matrilines, an observation

Persistent cooperation between unrelated conspecifics rarely occurs in mature eusocial insect societies. In this dissertation, I present evidence of non-kin cooperation in the Nearctic honey ant Myrmecocystus mendax. Using microsatellite markers, I show that mature colonies in the Sierra Ancha Mountain of central Arizona contain multiple unrelated matrilines, an observation that is consistent with primary polygyny. In contrast, similar analyses suggest that colonies in the Chiricahua Mountains of southeastern Arizona are primarily monogynous. These interpretations are consistent with field and laboratory observations. Whereas cooperative colony founding was observed frequently among groups of Sierra Ancha foundresses, founding in the Chiricahua population was restricted to individual foundresses. Furthermore, Sierra Ancha foundresses successfully established incipient laboratory colonies without undergoing queen culling following emergence of the first workers. Multi-queen laboratory Sierra Ancha colonies also produced more workers and repletes than haplometrotic colonies, and when brood raiding was induced between colonies, queens of those with more workers had a higher survival probability.

Microsatellite analyses of additional locations within the M. mendax range suggest that polygyny is also present in some other populations, especially in central-northern Arizona, albeit at lower frequencies than that in the Sierra Anchas. In addition, analyses of multiple types of genetic data, including microsatellites, the mitochondrial barcoding region, and over 2000 nuclear ultra-conserved elements indicate that M. mendax populations within the southwestern U.S. and northwestern Mexico are geographically structured, with strong support for the existence of two or more divergent clades as well as isolation-by-distance within clades. This structure is further shown to correlate with variation in queen number and hair length, a diagnostic taxonomic feature used to distinguish honey ant species.

Together, these findings suggest that regional ecological pressures (e.g. colony density , climate) may have acted on colony founding and social strategy to select for increasing workforce size and, along with genetic drift, have driven geographically isolated M. mendax populations to differentiate genetically and morphologically. The presence of colony fusion in the laboratory and life history traits in honey ant that are influenced by colony size, including repletism, brood raiding, and tournament, support this evolutionary scenario.
ContributorsEriksson, Ti (Author) / Gadau, Jürgen (Thesis advisor) / Taylor, Jay (Thesis advisor) / Fewell, Jennifer (Committee member) / Hӧlldobler, Bert (Committee member) / Johnson, Robert (Committee member) / Pratt, Stephen (Committee member) / Arizona State University (Publisher)
Created2018
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Description
Human-inhabited or -disturbed areas pose many unique challenges for wildlife, including increased human exposure, novel challenges, such as finding food or nesting sites in novel structures, anthropogenic noises, and novel predators. Animals inhabiting these environments must adapt to such changes by learning to exploit new resources and avoid danger. To

Human-inhabited or -disturbed areas pose many unique challenges for wildlife, including increased human exposure, novel challenges, such as finding food or nesting sites in novel structures, anthropogenic noises, and novel predators. Animals inhabiting these environments must adapt to such changes by learning to exploit new resources and avoid danger. To my knowledge no study has comprehensively assessed behavioral reactions of urban and rural populations to numerous novel environmental stimuli. I tested behavioral responses of urban, suburban, and rural house finches (Haemorhous mexicanus) to novel stimuli (e.g. objects, noises, food), to presentation of a native predator model (Accipiter striatus) and a human, and to two problem-solving challenges (escaping confinement and food-finding). Although I found few population-level differences in behavioral responses to novel objects, environment, and food, I found compelling differences in how finches from different sites responded to novel noise. When played a novel sound (whale call or ship horn), urban and suburban house finches approached their food source more quickly and spent more time on it than rural birds, and urban and suburban birds were more active during the whale-noise presentation. In addition, while there were no differences in response to the native predator, rural birds showed higher levels of stress behaviors when presented with a human. When I replicated this study in juveniles, I found that exposure to humans during development more accurately predicted behavioral differences than capture site. Finally, I found that urban birds were better at solving an escape problem, whereas rural birds were better at solving a food-finding challenge. These results indicate that not all anthropogenic changes affect animal populations equally and that determining the aversive natural-history conditions and challenges of taxa may help urban ecologists better understand the direction and degree to which animals respond to human-induced rapid environmental alterations.
ContributorsWeaver, Melinda (Author) / McGraw, Kevin J. (Thesis advisor) / Rutowski, Ronald (Committee member) / Pratt, Stephen (Committee member) / Bateman, Heather (Committee member) / Deviche, Pierre (Committee member) / Arizona State University (Publisher)
Created2018
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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