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Extremely thick cranial vaults have been noted as a diagnostic characteristic of Homo erectus since the first fossil of the species was identified, but potential mechanisms underlying this seemingly unique trait have not been rigorously investigated. Cranial vault thickness (CVT) is not a monolithic trait, and the responsiveness of its

Extremely thick cranial vaults have been noted as a diagnostic characteristic of Homo erectus since the first fossil of the species was identified, but potential mechanisms underlying this seemingly unique trait have not been rigorously investigated. Cranial vault thickness (CVT) is not a monolithic trait, and the responsiveness of its layers to environmental stimuli is unknown. Identifying factors that affect CVT would be exceedingly valuable in teasing apart potential contributors to thick vaults in the Pleistocene. Four hypotheses were tested using CT scans of skulls of more than 1100 human and non-human primates. Data on total frontal, parietal, and occipital bone thickness and bone composition were collected to test the hypotheses: H1. CVT is an allometric consequence of brain or body size. H2. Thick cranial vaults are a response to long, low cranial vault shape. H3. High masticatory stress causes localized thickening of cranial vaults. H4. Activity-mediated systemic hormone levels affect CVT. Traditional comparative methods were used to identify features that covary with CVT across primates to establish behavior patterns that might correlate with thick cranial vaults. Secondly, novel experimental manipulation of a model organism, Mus musculus, was used to evaluate the relative plasticity of CVT. Finally, measures of CVT in fossil hominins were described and discussed in light of the extant comparative and experimental results. This dissertation reveals previously unknown variation among extant primates and humans and illustrates that Homo erectus is not entirely unique among primates in its CVT. The research suggests that it is very difficult to make a mouse grow a thick head, although it can be genetically programmed to have one. The project also identifies a possible hominin synapomorphy: high diploë ratios compared to non-human primates. It also found that extant humans differ from non-human primates in overall pattern of which cranial vault bones are thickest. What this project was unable to do was definitively provide an explanation for why and how Homo erectus grew thick skulls. Caution is required when using CVT as a diagnostic trait for Homo erectus, as the results presented here underscore the complexity inherent in its evolution and development.
ContributorsCopes, Lynn (Author) / Kimbel, William H. (Thesis advisor) / Schwartz, Gary T (Committee member) / Spencer, Mark A. (Committee member) / Ravosa, Matthew J. (Committee member) / Arizona State University (Publisher)
Created2012
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Description
Fundamental hypotheses about the life history, complex cognition and social dynamics of humans are rooted in feeding ecology - particularly in the experiences of young animals as they grow. However, the few existing primate developmental data are limited to only a handful of species of monkeys and apes. Without comparative

Fundamental hypotheses about the life history, complex cognition and social dynamics of humans are rooted in feeding ecology - particularly in the experiences of young animals as they grow. However, the few existing primate developmental data are limited to only a handful of species of monkeys and apes. Without comparative data from more basal primates, such as lemurs, we are limited in the scope of our understanding of how feeding has shaped the evolution of these extraordinary aspects of primate biology. I present a developmental view of feeding ecology in the ring-tailed lemur (Lemur catta) using a mixed longitudinal sample (infant through adult) collected at the Beza Mahafaly Special Reserve in southwestern Madagascar from May 2009 to March 2010. I document the development of feeding, including weaning, the transition to solid food, and how foods are included in infant diets. Early in juvenility ring-tailed lemurs efficiently process most foods, but that hard ripe fruits and insects require more time to master. Infants and juveniles do not use many of the social learning behaviors that are common in monkeys and apes, and instead likely rely both on their own trial and error and simple local enhancement to learn appropriate foods. Juvenile ring-tailed lemurs are competent and efficient foragers, and that mitigating ecological risks may not best predict the lemur juvenile period, and that increases in social complexity and brain size may be at the root of primate juvenility. Finally, from juvenility through adulthood, females have more diverse diets than males. The early emergence of sex differences in dietary diversity in juvenility that are maintained throughout adulthood indicate that, in addition to reproductive costs incurred by females, niche partitioning is an important aspect of sex differential feeding ecology, and that ontogenetic studies of feeding are particularly valuable to understanding how selection shapes adult, species-typical diets. Overall, lemur juvenility is a time to play, build social relationships, learn about food, and where the kernels of sex-typical feeding develop. This study of the ontogeny of feeding ecology contributes an important phylogenetic perspective on the relationship between juvenility and the emergent foraging behaviors of developing animals
ContributorsO'Mara, Michael Teague (Author) / Nash, Leanne T. (Thesis advisor) / Reed, Kaye E (Committee member) / Schwartz, Gary T (Committee member) / Sauther, Michelle L (Committee member) / Arizona State University (Publisher)
Created2012
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Description
The pattern and strength of genetic covariation is shaped by selection so that it is strong among functionally related characters and weak among functionally unrelated characters. Genetic covariation is expressed as phenotypic covariation within species and acts as a constraint on evolution by limiting the ability of linked characters to

The pattern and strength of genetic covariation is shaped by selection so that it is strong among functionally related characters and weak among functionally unrelated characters. Genetic covariation is expressed as phenotypic covariation within species and acts as a constraint on evolution by limiting the ability of linked characters to evolve independently of one another. Such linked characters are "constrained" and are expected to express covariation both within and among species. In this study, the pattern and magnitude of covariation among aspects of dental size and shape are investigated in anthropoid primates. Pleiotropy has been hypothesized to play a significant role in derivation of derived hominin morphologies. This study tests a series of hypotheses; including 1) that negative within- and among-species covariation exists between the anterior (incisors and canines) and postcanine teeth, 2) that covariation is strong and positive between the canines and incisors, 3) that there is a dimorphic pattern of within-species covariation and coevolution for characters of the canine honing complex, 4) that patterns of covariation are stable among anthropoids, and 5) that genetic constraints have been a strong bias on the diversification of anthropoid dental morphology. The study finds that patterns of variance-covariance are conserved among species. Despite these shared patterns of variance-covariance, dental diversification has frequently occurred along dimensions not aligned with the vector of genetic constraint. As regards the canine honing complex, there is no evidence for a difference in the pleiotropic organization or the coevolution of characters of the complex in males and females, which undermines arguments that the complex is selectively important only in males. Finally, there is no evidence for strong or negative pleiotropy between any dental characters, which falsifies hypotheses that predict such relationships between incisors and postcanine teeth or between the canines and the postcanine teeth.
ContributorsDelezene, Lucas (Author) / Kimbel, William H. (Thesis advisor) / Schwartz, Gary T (Committee member) / Spencer, Mark (Committee member) / Verrelli, Brian C (Committee member) / Arizona State University (Publisher)
Created2011
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Description
Evolution is the foundation of biology, yet it remains controversial even among college biology students. Acceptance of evolution is important for students if we want them to incorporate evolution into their scientific thinking. However, students’ religious beliefs are a consistent barrier to their acceptance of evolution due to a perceived

Evolution is the foundation of biology, yet it remains controversial even among college biology students. Acceptance of evolution is important for students if we want them to incorporate evolution into their scientific thinking. However, students’ religious beliefs are a consistent barrier to their acceptance of evolution due to a perceived conflict between religion and evolution. Using pre-post instructional surveys of students in introductory college biology, Study 1 establishes instructional strategies that can be effective for reducing students' perceived conflict between religion and evolution. Through interviews and qualitative analyses, Study 2 documents how instructors teaching evolution at public universities may be resistant towards implementing strategies that can reduce students' perceived conflict, perhaps because of their own lack of religious beliefs and lack of training and awareness about students' conflict with evolution. Interviews with religious students in Study 3 reveals that religious college biology students can perceive their instructors as unfriendly towards religion which can negatively impact these students' perceived conflict between religion and evolution. Study 4 explores how instructors at Christian universities, who share the same Christian backgrounds as their students, do not struggle with implementing strategies that reduce students' perceived conflict between religion and evolution. Cumulatively, these studies reveal a need for a new instructional framework for evolution education that takes into account the religious cultural difference between instructors who are teaching evolution and students who are learning evolution. As such, a new instructional framework is then described, Religious Cultural Competence in Evolution Education (ReCCEE), that can help instructors teach evolution in a way that can reduce students' perceived conflict between religion and evolution, increase student acceptance of evolution, and create more inclusive college biology classrooms for religious students.
ContributorsBarnes, Maryann Elizabeth (Author) / Brownell, Sara (Thesis advisor) / Nesse, Randolph (Committee member) / Collins, James (Committee member) / Husman, Jenefer (Committee member) / Maienschein, Jane (Committee member) / Arizona State University (Publisher)
Created2018
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Description
How fast is evolution? In this dissertation I document a profound change that occurred around the middle of the 20th century in the way that ecologists conceptualized the temporal and spatial scales of adaptive evolution, through the lens of British plant ecologist Anthony David Bradshaw (1926–2008). In the early 1960s,

How fast is evolution? In this dissertation I document a profound change that occurred around the middle of the 20th century in the way that ecologists conceptualized the temporal and spatial scales of adaptive evolution, through the lens of British plant ecologist Anthony David Bradshaw (1926–2008). In the early 1960s, one prominent ecologist distinguished what he called “ecological time”—around ten generations—from “evolutionary time”— around half of a million years. For most ecologists working in the first half of the 20th century, evolution by natural selection was indeed a slow and plodding process, tangible in its products but not in its processes, and inconsequential for explaining most ecological phenomena. During the 1960s, however, many ecologists began to see evolution as potentially rapid and observable. Natural selection moved from the distant past—a remote explanans for both extant biological diversity and paleontological phenomena—to a measurable, quantifiable mechanism molding populations in real time.

The idea that adaptive evolution could be rapid and highly localized was a significant enabling condition for the emergence of ecological genetics in the second half of the 20th century. Most of what historians know about that conceptual shift and the rise of ecological genetics centers on the work of Oxford zoologist E. B. Ford and his students on polymorphism in Lepidotera, especially industrial melanism in Biston betularia. I argue that ecological genetics in Britain was not the brainchild of an infamous patriarch (Ford), but rather the outgrowth of a long tradition of pastureland research at plant breeding stations in Scotland and Wales, part of a discipline known as “genecology” or “experimental taxonomy.” Bradshaw’s investigative activities between 1948 and 1968 were an outgrowth of the specific brand of plant genecology practiced at the Welsh and Scottish Plant Breeding stations. Bradshaw generated evidence that plant populations with negligible reproductive isolation—separated by just a few meters—could diverge and adapt to contrasting environmental conditions in just a few generations. In Bradshaw’s research one can observe the crystallization of a new concept of rapid adaptive evolution, and the methodological and conceptual transformation of genecology into ecological genetics.
ContributorsPeirson, Bruce Richard Erick (Author) / Laubichler, Manfred D (Thesis advisor) / Maienschein, Jane (Thesis advisor) / Creath, Richard (Committee member) / Collins, James (Committee member) / Arizona State University (Publisher)
Created2015
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Description
Across primates, molar-emergence age is strongly correlated to life-history variables, such as age-at-first-reproduction and longevity. This relationship allows for the reconstruction of life-history parameters in fossil primates. The mechanism responsible for modulating molar-emergence age is unknown, however. This dissertation uses a biomechanical model that accurately predicts the position of molars

Across primates, molar-emergence age is strongly correlated to life-history variables, such as age-at-first-reproduction and longevity. This relationship allows for the reconstruction of life-history parameters in fossil primates. The mechanism responsible for modulating molar-emergence age is unknown, however. This dissertation uses a biomechanical model that accurately predicts the position of molars in adults to determine whether molar emergence is constrained by chewing biomechanics throughout ontogeny. A key aspect of chewing system configuration in adults is the position of molars: the distal-most molar is constrained to avoid tensile forces at the temporomandibular joint (TMJ). Using three-dimensional data from growth samples of 1258 skulls, representing 21 primate species, this research tested the hypothesis that the location and timing of molar emergence is constrained to avoid high and potentially dangerous tensile forces at the TMJ throughout growth. Results indicate that molars emerge in a predictable position to safeguard the TMJ during chewing. Factors related to the size of the buffer zone, a safety feature that creates greater stability at the TMJ during biting, account for a large portion of both ontogenetic and interspecific variation in the position of emergence. Furthermore, the rate at which space is made available in the jaws and the duration of jaw growth both determine the timing of molar emergence. Overall, this dissertation provides a mechanical and developmental model for explaining temporal and spatial variation in molar emergence and a framework for understanding how variation in the timing of molar emergence has evolved among primates. The findings suggest that life history is related to ages at molar emergence through its influence on the rate and duration of jaw growth. This dissertation provides support for the functionally integrated nature of craniofacial growth and has implications for the study of primate life history evolution and masticatory morphology in the fossil record.
ContributorsGlowacka, Halszka (Author) / Schwartz, Gary T (Thesis advisor) / Kimbel, William H. (Committee member) / Reed, Kaye E (Committee member) / Wright, Barth W (Committee member) / Arizona State University (Publisher)
Created2017
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Description
Transgenic experiments in Drosophila have proven to be a useful tool aiding in the

determination of mammalian protein function. A CNS specific protein, dCORL is a

member of the Sno/Ski family. Sno acts as a switch between Dpp/dActivin signaling.

dCORL is involved in Dpp and dActivin signaling, but the two homologous mCORL

protein functions

Transgenic experiments in Drosophila have proven to be a useful tool aiding in the

determination of mammalian protein function. A CNS specific protein, dCORL is a

member of the Sno/Ski family. Sno acts as a switch between Dpp/dActivin signaling.

dCORL is involved in Dpp and dActivin signaling, but the two homologous mCORL

protein functions are unknown. Conducting transgenic experiments in the adult wings,

and third instar larval brains using mCORL1, mCORL2 and dCORL are used to provide

insight into the function of these proteins. These experiments show mCORL1 has a

different function from mCORL2 and dCORL when expressed in Drosophila. mCORL2

and dCORL have functional similarities that are likely conserved. Six amino acid

substitutions between mCORL1 and mCORL2/dCORL may be the reason for the

functional difference. The evolutionary implications of this research suggest the

conservation of a switch between Dpp/dActivin signaling that predates the divergence of

arthropods and vertebrates.
ContributorsStinchfield, Michael J (Author) / Newfeld, Stuart J (Thesis advisor) / Capco, David (Committee member) / Laubichler, Manfred (Committee member) / Arizona State University (Publisher)
Created2019
Description

Differences in the postcanine dentition of primates likely represent dietary adaptations given that the teeth interact directly with foods during mastication. Among early hominins, changes to both molar and premolar morphology are purported to indicate consumption of foods differing in their material properties. Some early hominins, such as the robust

Differences in the postcanine dentition of primates likely represent dietary adaptations given that the teeth interact directly with foods during mastication. Among early hominins, changes to both molar and premolar morphology are purported to indicate consumption of foods differing in their material properties. Some early hominins, such as the robust australopiths, possess premolars that resemble molars with enhancements to the distal part of the tooth (i.e., the talonid), including additional cusps and/or expanded basins. Such molarized premolars are thought to indicate that these hominins were processing mechanically challenging foods; that is, food items that were either hard or tough. Hypotheses tested in this study evaluated the link between the degree of premolar molarization and consumption of mechanically challenging foods in extant primates. Surface anatomy of the distal-most mandibular premolar (the P4) was quantified using a combination of 3D scans of postcanine dental casts and craniodental landmark data collected from 541 individuals, representing 22 extant primate taxa with well-studied diets and known food material properties. Taxa with more mechanically challenging diets were expected to have premolars with expanded talonids and enlarged P4s (and/or molar rows) relative to several mechanically-relevant size proxies. Taxa consuming high proportions of structural carbohydrates were also expected to have postcanine teeth with high occlusal relief (RFI), sharpness (DNE), and complexity (OPCR). Taxa consuming harder food items were expected to have lower relief and higher complexity, with sharpness determined by the proportion of structural carbohydrates included in their diet. The work presented in this dissertation supports most of these expectations, though talonid expansion per se was not clearly linked to the consumption of any particular diet. Overall, taxa with more mechanically challenging diets generally had relatively enlarged premolars when compared to taxa with softer diets and also differed predictably in their occlusal topography. The results of this dissertation support the functional significance of P4 crown size and measures of dental topography as they relate to diet and have implications for improving dietary inferences from the fossil record.

ContributorsDaly, Elizabeth Susanne (Author) / Schwartz, Gary T (Thesis advisor) / Delezene, Lucas K (Committee member) / Kimbel, William H (Committee member) / Arizona State University (Publisher)
Created2021
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Description
Insecticide resistance is a continuing issue that negatively affects both public health and agriculture and allows vector-borne diseases to spread throughout the globe. To improve resistance management strategies (RMS), robust susceptibility bioassays need to be performed in order to fill the gap of the relationship between resistant and susceptible genotype

Insecticide resistance is a continuing issue that negatively affects both public health and agriculture and allows vector-borne diseases to spread throughout the globe. To improve resistance management strategies (RMS), robust susceptibility bioassays need to be performed in order to fill the gap of the relationship between resistant and susceptible genotype and phenotype, and a deeper knowledge of how bioassay data relates to vector control success or failure is imperative. A bioassay method that is infrequently used but yields robust results is the topical application bioassay, where the insect is directly treated with a constant volume and concentration of an insecticide via a syringe. To bring more attention to this method, my colleagues and I published a paper in the Journal of Visualized Experiments where the optimized protocol of the topical application bioassay for mosquitoes and fruit flies is described, and the strengths and limitations to the method are explained. To further investigate insecticide susceptibility tests, I set up my individual project where I used Aedes aegypti mosquitoes to compare the topical application bioassay to the commonly used Centers for Disease Control and Prevention (CDC) bottle bioassay and World Health Organization (WHO) tube test. The objective of this study was to test which method exhibited the most variability in mortality results, which would guide the choice of assay to determine the link between resistant and susceptible genotype and phenotype. The results showed that the topical application method did indeed exhibit the least amount of variation, followed by the CDC bottle bioassay (WHO data is currently being collected). This suggests that the topical application bioassay could be a useful tool in insecticide resistance surveillance studies, and, depending on the goal, may be better than the CDC and WHO tube tests for assessing resistance levels at a given site. This study challenges the value of the widely used CDC and WHO assays and provides a discussion on the importance of technical and practical resistance assays. This will help vector control specialists to collect accurate surveillance data that will inform effective RMS.
ContributorsAlthoff, Rachel (Author) / Huijben, Silvie (Thesis advisor) / Harris, Robin (Committee member) / Collins, James (Committee member) / Arizona State University (Publisher)
Created2022