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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
In social insect colonies, as with individual animals, the rates of biological processes scale with body size. The remarkable explanatory power of metabolic allometry in ecology and evolutionary biology derives from the great diversity of life exhibiting a nonlinear scaling pattern in which metabolic rates are not proportional to mass,

In social insect colonies, as with individual animals, the rates of biological processes scale with body size. The remarkable explanatory power of metabolic allometry in ecology and evolutionary biology derives from the great diversity of life exhibiting a nonlinear scaling pattern in which metabolic rates are not proportional to mass, but rather exhibit a hypometric relationship with body size. While one theory suggests that the supply of energy is a major physiological constraint, an alternative theory is that the demand for energy is regulated by behavior. The central hypothesis of this dissertation research is that increases in colony size reduce the proportion of individuals actively engaged in colony labor with consequences for energetic scaling at the whole-colony level of biological organization. A combination of methods from comparative physiology and animal behavior were developed to investigate scaling relationships in laboratory-reared colonies of the seed-harvester ant, Pogonomyrmex californicus. To determine metabolic rates, flow-through respirometry made it possible to directly measure the carbon dioxide production and oxygen consumption of whole colonies. By recording video of colony behavior, for which ants were individually paint-marked for identification, it was possible to reconstruct the communication networks through which information is transmitted throughout the colony. Whole colonies of P. californicus were found to exhibit a robust hypometric allometry in which mass-specific metabolic rates decrease with increasing colony size. The distribution of walking speeds also scaled with colony size so that larger colonies were composed of relatively more inactive ants than smaller colonies. If colonies were broken into random collections of workers, metabolic rates scaled isometrically, but when entire colonies were reduced in size while retaining functionality (queens, juveniles, workers), they continued to exhibit a metabolic hypometry. The communication networks in P. californicus colonies contain a high frequency of feed-forward interaction patterns consistent with those of complex regulatory systems. Furthermore, the scaling of these communication pathways with size is a plausible mechanism for the regulation of whole-colony metabolic scaling. The continued development of a network theory approach to integrating behavior and metabolism will reveal insights into the evolution of collective animal behavior, ecological dynamics, and social cohesion.
ContributorsWaters, James S., 1983- (Author) / Harrison, Jon F. (Thesis advisor) / Quinlan, Michael C. (Committee member) / Pratt, Stephen C. (Committee member) / Fewell, Jennifer H. (Committee member) / Gadau, Juergen (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
Body size plays a pervasive role in determining physiological and behavioral performance across animals. It is generally thought that smaller animals are limited in performance measures compared to larger animals; yet, the vast majority of animals on earth are small and evolutionary trends like miniaturization occur in every animal clade.

Body size plays a pervasive role in determining physiological and behavioral performance across animals. It is generally thought that smaller animals are limited in performance measures compared to larger animals; yet, the vast majority of animals on earth are small and evolutionary trends like miniaturization occur in every animal clade. Therefore, there must be some evolutionary advantages to being small and/or compensatory mechanisms that allow small animals to compete with larger species. In this dissertation I specifically explore the scaling of flight performance (flight metabolic rate, wing beat frequency, load-carrying capacity) and learning behaviors (visual differentiation visual Y-maze learning) across stingless bee species that vary by three orders of magnitude in body size. I also test whether eye morphology and calculated visual acuity match visual differentiation and learning abilities using honeybees and stingless bees. In order to determine what morphological and physiological factors contribute to scaling of these performance parameters I measure the scaling of head, thorax, and abdomen mass, wing size, brain size, and eye size. I find that small stingless bee species are not limited in visual learning compared to larger species, and even have some energetic advantages in flight. These insights are essential to understanding how small size evolved repeatedly in all animal clades and why it persists. Finally, I test flight performance across stingless bee species while varying temperature in accordance with thermal changes that are predicted with climate change. I find that thermal performance curves varied greatly among species, that smaller species conform closely to air temperature, and that larger bees may be better equipped to cope with rising temperatures due to more frequent exposure to high temperatures. This information may help us predict whether small or large species might fare better in future thermal climate conditions, and which body-size related traits might be expected to evolve.
ContributorsDuell, Meghan (Author) / Harrison, Jon F. (Thesis advisor) / Smith, Brian H. (Thesis advisor) / Rutowski, Ronald (Committee member) / Wcislo, William (Committee member) / Conrad, Cheryl (Committee member) / Arizona State University (Publisher)
Created2018
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Description
Early hominins present an unusual pattern of sexual dimorphism. On one hand, the canine teeth of these species are weakly size-dimorphic, vertically short, and nonhoning, suggesting a social system characterized by infrequent, low-intensity intermale competition and monogamous pair-bonding. On the other hand, marked size variation in skeletal remains attributed to

Early hominins present an unusual pattern of sexual dimorphism. On one hand, the canine teeth of these species are weakly size-dimorphic, vertically short, and nonhoning, suggesting a social system characterized by infrequent, low-intensity intermale competition and monogamous pair-bonding. On the other hand, marked size variation in skeletal remains attributed to species of Australopithecus is thought to reflect strong body-mass dimorphism, which is more consistent with intense intermale competition. Reconciling these conflicting signals and understanding their adaptive significance is a major goal of paleoanthropology. This dissertation research contributes to this objective by investigating factors that may constrain or reduce canine height in extant anthropoid primates. Two hypotheses regarding the relationship between canine height and other elements of the masticatory system were tested using phylogenetic comparative methods. According to the first hypothesis, canine reduction is a pleiotropic by-product of changes in the sizes of other components of the dentition. With respect to canine height, the results of this study fail to support this idea. There is limited evidence for a relationship between basal canine crown dimensions and incisor and postcanine size, but significant interspecific correlations between these variables are not strong and are restricted primarily to the female maxillary dentition. These results indicate that if pleiotropy influences canine size, then its effects are weak. The second hypothesis proposes that canine reduction is a consequence of selection for increased jaw-muscle leverage. This hypothesis receives some support: there is a clear inverse relationship between canine height and the leverage of the masseter muscle in male anthropoids. Females do not exhibit this association due to the fact that dimorphism in muscle leverage is weak or absent in most anthropoid species; in other words, female muscle leverage tracks male muscle leverage, which is linked to canine height. Leverage of the temporalis muscle is not correlated with canine height in either sex. Two specimens of the 3.0-3.7-million-year-old hominin Australopithecus afarensis fall at or beyond the upper end of the great ape range of variation in masseter leverage, which is consistent with the idea that hominin canine evolution was influenced by selection for increased jaw-muscle leverage.
ContributorsScott, Jeremiah Ezekiel (Author) / Kimbel, William H. (Thesis advisor) / Schwartz, Gary T. (Committee member) / Spencer, Mark A. (Committee member) / Arizona State University (Publisher)
Created2010