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Despite the critical role that the vertebral column plays in postural and locomotor behaviors, the functional morphology of the cervical region (i.e., the bony neck) remains poorly understood, particularly in comparison to that of the thoracic and lumbar sections. This dissertation tests the hypothesis that morphological variation in cervical vertebrae

Despite the critical role that the vertebral column plays in postural and locomotor behaviors, the functional morphology of the cervical region (i.e., the bony neck) remains poorly understood, particularly in comparison to that of the thoracic and lumbar sections. This dissertation tests the hypothesis that morphological variation in cervical vertebrae reflects differences in positional behavior (i.e., suspensory vs. nonsuspensory and orthograde vs. pronograde locomotion and postures). Specifically, this project addresses two broad research questions: (1) how does the morphology of cervical vertebrae vary with positional behavior and cranial morphology among primates and (2) where does fossil hominoid morphology fall within the context of the extant primates. Three biomechanical models were developed for the primate cervical spine and their predictions were tested by conducting a comparative analysis using a taxonomically and behaviorally diverse sample of primates. The results of these analyses were used to evaluate fossil hominoid morphology. The two biomechanical models relating vertebral shape to positional behaviors are not supported. However, a number of features distinguish behavioral groups. For example, the angle of the transverse process in relation to the cranial surface of the vertebral body--a trait hypothesized to reflect the deep spinal muscles' ability to extend and stabilize the neck--tends to be greater in pronograde species; this difference is in the opposite of the direction predicted by the biomechanical models. Other traits distinguish behavioral groups (e.g., spinous process length and cross-sectional area), but only in certain parts of the cervical column. The correlation of several vertebral features, especially transverse process length and pedicle cross-sectional area, with anterior cranial length supports the predictions made by the third model that links cervical morphology with head stabilization (i.e., head balancing). Fossil hominoid cervical remains indicate that the morphological pattern that characterizes modern humans was not present in Homo erectus or earlier hominins. These hominins are generally similar to apes in having larger neural arch cross-sectional areas and longer spinous processes than modern humans, likely indicating the presence of comparatively large nuchal muscles. The functional significance of this morphology remains unclear.
ContributorsNalley, Thierra Kénnec (Author) / Kimbel, William H. (Thesis advisor) / Reed, Kaye (Committee member) / Shapiro, Liza (Committee member) / Arizona State University (Publisher)
Created2013
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Description
Arguments of human uniqueness emphasize our complex sociality, unusual cognitive capacities, and language skills, but the timing of the origin of these abilities and their evolutionary causes remain unsolved. Though not unique to primates, kin-biased sociality was key to the success of the primate order. In contrast to ancestral solitary

Arguments of human uniqueness emphasize our complex sociality, unusual cognitive capacities, and language skills, but the timing of the origin of these abilities and their evolutionary causes remain unsolved. Though not unique to primates, kin-biased sociality was key to the success of the primate order. In contrast to ancestral solitary mammals, the earliest primates are thought to have maintained dispersed (non-group living) social networks, communicating over distances via vocalizations and scent marks. If such ancestral primates recognized kin, those networks may have facilitated the evolution of kin-biased sociality in the primate order and created selection for increased cognitive and communicative abilities. I used the gray mouse lemur (Microcebus murinus) to model whether vocalizations could have facilitated matrilineal and patrilineal kin recognition in ancestral primates. Much like mouse lemurs today, ancestral primates are thought to have been small-bodied, nocturnal creatures that captured insects and foraged for fruit in the thin, terminal ends of tree branches. Thus, the mouse lemur is an excellent model species because its ecological niche is likely to be similar to that of ancestral primates 55-90 million years ago. I conducted playback experiments in Ankarafantsika National Park, Madagascar testing whether mouse lemur agonistic calls contain matrilineal kin signatures and whether the lemurs recognize matrilineal kin. In contrast to large-brained, socially complex monkeys with frequent coalitionary behavior, mouse lemurs did not react differently to the agonistic calls of matrilineal kin and nonkin, though moderate signatures were present in the calls. I tested for patrilineal signatures and patrilineal kin recognition via mating and alarm calls in a colony with known pedigree relationships. The results are the first to demonstrate that a nocturnal, solitary foraging mammal gives mating calls with patrilineal signatures and recognizes patrilineal kin. Interestingly, alarm calls did not have signatures and did not facilitate kin recognition, suggesting that selection for kin recognition is stronger in some call types than others. As this dissertation is the first investigation of vocal kin recognition in a dispersed-living, nocturnal strepsirrhine primate, it greatly advances our knowledge of the role of vocal communication in the evolution of primate social complexity.
ContributorsKessler, Sharon E (Author) / Nash, Leanne (Thesis advisor) / Reed, Kaye (Thesis advisor) / Radespiel, Ute (Committee member) / Zimmermann, Elke (Committee member) / Arizona State University (Publisher)
Created2014
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This dissertation research describes the hunting behavior of early modern humans through the analysis of vertebrate faunal remains from Contrebandiers Cave, Morocco. Contrebandiers Cave is located in the town of Témara and is roughly 250 meters from the current shoreline of the Atlantic Ocean. The cave was excavated in the

This dissertation research describes the hunting behavior of early modern humans through the analysis of vertebrate faunal remains from Contrebandiers Cave, Morocco. Contrebandiers Cave is located in the town of Témara and is roughly 250 meters from the current shoreline of the Atlantic Ocean. The cave was excavated in the 1950s and 1970s by l’Abbé Roche, and again starting in 2007 by Dibble and El Hajraoui with total station plotting of finds. Contrebandiers Cave contains Middle Stone Age (MSA) deposits dated to Marine Isotope Stages (MIS) 5e, 5d and 5c, ~120,000 to ~96,000 years ago. The Later Stone Age (LSA) deposits are dated to MIS 2, ~20,000 years ago. The entirety of the ~12,000 vertebrate faunal remains from Dibble and El Hajraoui’s excavation were analyzed for taxonomic and taphonomic identification.

A total of 67 vertebrate taxa were identified and include ungulates, carnivores, lagomorphs, birds, tortoises, snakes and fish. The faunal remains from Contrebandiers Cave preserve surface modification that indicates both humans and carnivores acted as agents of prey accumulation. Skeletal element representation and surface modification of ungulate remains suggest that humans had primary access to small, medium and large-bodied prey. In the MSA levels, carnivore skeletal remains preserve surface modification that is interpreted as being indicative of behavior associated with skinning for fur removal.

The vertebrate faunal remains from MIS 5e and 5d indicate that humans were hunting grazers and mixed feeders from open habitats and suids from mixed habitats. The faunal remains from MIS 5c indicate that humans focused less on suids and more on mixed feeders from open habitats. The vertebrate faunal remains from MIS 2 reveal humans hunting grazers from dry, open habitats. This research provides a description of human hunting behavior in North Africa, and contributes to our understanding of early modern human behavior prior to dispersal out of Africa.
ContributorsHallett, Emily Yuko (Author) / Marean, Curtis W (Thesis advisor) / Reed, Kaye E (Committee member) / Dibble, Harold L. (Committee member) / Arizona State University (Publisher)
Created2018
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This study explores how early modern humans used stone tool technology to adapt to changing climates and coastlines in the Middle Stone Age of South Africa. The MSA is associated with the earliest fossil evidence for modern humans and complex cultural behaviors during a time period of dramatic climate change.

This study explores how early modern humans used stone tool technology to adapt to changing climates and coastlines in the Middle Stone Age of South Africa. The MSA is associated with the earliest fossil evidence for modern humans and complex cultural behaviors during a time period of dramatic climate change. Human culture allows for the creation, use, and transmission of technological knowledge that can evolve with changing environmental conditions. Understanding the interactions between technology and the environment is essential to illuminating the role of culture during the origin of our species. This study is focused on understanding ancient tool use from the study of lithic edge damage patterns at archaeological assemblages in southern Africa by using image-based quantitative methods for analyzing stone tools. An extensive experimental program using replicated stone tools provides the comparative linkages between the archaeological artifacts and the tasks for which they were used. MSA foragers structured their tool use and discard behaviors on the landscape in several ways – by using and discarding hunting tools more frequently in the field rather than in caves/rockshelters, but similarly in coastal and interior contexts. This study provides evidence that during a significant microlithic technological shift seen in southern Africa at ~75,000 years ago, new technologies were developed alongside rather than replacing existing technologies. These results are compared with aspects of the European archaeological record at this time to identify features of early human technological behavior that may be unique to the evolutionary history of our species.
ContributorsSchoville, Benjamin J (Author) / Marean, Curtis W (Thesis advisor) / Barton, Michael (Committee member) / Hill, Kim (Committee member) / Arizona State University (Publisher)
Created2016
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Description
Early weaning, slow somatic and dental growth, and late age at reproduction are all part of a suite of energetic trade-offs that have shaped human evolution. A similar suite of energetic trade-offs has shaped the evolution of the indriid-palaeopropithecid clade, though members of this clade exhibit extremely fast dental development

Early weaning, slow somatic and dental growth, and late age at reproduction are all part of a suite of energetic trade-offs that have shaped human evolution. A similar suite of energetic trade-offs has shaped the evolution of the indriid-palaeopropithecid clade, though members of this clade exhibit extremely fast dental development and nearly vestigial deciduous teeth. The development and functional occlusion of the primary postcanine dentition (i.e., deciduous premolars and molars) coincides with several life history parameters in great apes and indriids. This dissertation explored great ape dental macrowear, molar development in indriids, and molar size in lemurs with a broader goal of improving reconstructions of life history profiles in extinct primates. To this aim, macrowear and dental development were analyzed in apes and lemurs, respectively. Occlusal casts (six great ape species; N=278) were scanned to track mandibular fourth deciduous premolar (dp4) macrowear. Utilizing dental topographic analyses, changes in occlusal gradient and terrain were quantified. A subset of the great ape data (four species; n=199) was analyzed to test if differences in dp4 wear correlate with age at weaning. Using dental histology, molar development was reconstructed for Indri indri (n=1) and Avahi laniger (n=1). Life history and molar size data were collected from the literature. The results of this dissertation demonstrate that most great apes exhibited evidence of topographic maintenance, suggesting dp4s wear in a manner that maintain functional efficiency during growth and development; however, the manner in which maintenance is achieved (e.g., preservation of relief or complexity) is species specific. Dp4 macrowear is not correlated with age at weaning in great apes and is probably unreliable to reconstruct age at weaning in hominins. The pace of molar development in members of the indriid- palaeopropithecid clade did not correlate with body or brain size, an association present in several other primates. Associations of molar size with age at weaning suggest that expanding other developmental models (e.g., the inhibitory cascade) to life history is worth consideration. The broad variation in macrowear, dental development, and size highlights how the primary dentition may correlate with different life history parameters depending on the species and ecological setting, an important consideration when using teeth to reconstruct life history profiles.
ContributorsCatlett, Kierstin Kay (Author) / Schwartz, Gary (Thesis advisor) / Barton, Michael (Committee member) / Godfrey, Laurie (Committee member) / Reed, Kaye (Committee member) / Arizona State University (Publisher)
Created2016
Description
Craniofacial morphology in primates can vary on the basis of their diet because foods are often disparate in the amount and duration of force required to break them down. Therefore diet has the potential to exercise considerable selective pressure on the morphology of the masticatory system. The zygomatic arch

Craniofacial morphology in primates can vary on the basis of their diet because foods are often disparate in the amount and duration of force required to break them down. Therefore diet has the potential to exercise considerable selective pressure on the morphology of the masticatory system. The zygomatic arch is a known site of relatively high masticatory strain and yet the relationship between arch form and load type is relatively unknown in primates. While the relative position and robusticity of the arch is considered a key indicator of craniofacial adaptations to a mechanically challenging diet, and central to efforts to infer diet in past species, the relationships between morphology and diet type in this feature are not well established.

This study tested hypotheses using two diet categorizations: total consumption percent and food material properties (FMPs). The first hypothesis that cortical bone area (CA) and section moduli (bone strength) are positively correlated with masticatory loading tests whether CA and moduli measures were greatest anteriorly and decreased posteriorly along the arch. The results found these measures adhered to this predicted pattern in the majority of taxa. The second hypothesis examines sutural complexity in the zygomaticotemporal suture as a function of dietary loading differences by calculating fractal dimensions as indices of complexity. No predictable pattern was found linking sutural complexity and diet in this primate sample, though hard object consumers possessed the most complex sutures. Lastly, cross-sectional geometric properties were measured to investigate whether bending and torsional resistance and cross-sectional shape are related to differences in masticatory loading. The highest measures of mechanical resistance tracked with areas of greatest strain in the majority of taxa. Cross-sectional shape differences do appear to reflect dietary differences. FMPs were not correlated with cross-sectional variables, however pairwise comparisons suggest taxa that ingest foods of greater stiffness experience relatively larger measures of bending and torsional resistance. The current study reveals that internal and external morphological factors vary across the arch and in conjunction with diet in primates. These findings underscore the importance of incorporating these mechanical differences in models of zygomatic arch mechanical behavior and primate craniofacial biomechanics.
ContributorsEdmonds, Hallie Margaret (Author) / Reed, Kaye (Thesis advisor) / Schwartz, Gary (Committee member) / Vinyard, Chris (Committee member) / Arizona State University (Publisher)
Created2017
Description
Crown Cercopithecoidea (Old World monkeys) share bilophodont molars characterized by four cusps arranged into two transversely-aligned pairs connected by crests or “loph(id)s”. This derived dental configuration provides a flexible template that has been modified in different lineages of Old World monkeys to meet the mechanical demands of food-processing in species

Crown Cercopithecoidea (Old World monkeys) share bilophodont molars characterized by four cusps arranged into two transversely-aligned pairs connected by crests or “loph(id)s”. This derived dental configuration provides a flexible template that has been modified in different lineages of Old World monkeys to meet the mechanical demands of food-processing in species with diverse and varied diets. This molar Bauplan evolved in the early stages of Old World monkey evolution, and one consequence of these morphological changes in occlusal morphology relative to apes and more basal catarrhines is a set of distinct patterns of tooth wear. Adaptive explanations for the origins of bilophodonty have emphasized dietary reconstructions but have not explored the implications of molar crown reorganization on the interaction between tooth wear and tooth function. This study combines description of new fossil material of early Miocene stem cercopithecoids and 3D dental topographic analyses of cross-sectional M2 wear series of extant catarrhines (n=511, 24 species) and Miocene fossil catarhines (n=81, 7 genera) to explore how functional aspects of molar topography are altered by tooth wear, to test whether the acquisition of bilophodont molars resulted in distinct occlusal topographies and patterns of topographic change with wear among Old World monkeys, and to determine whether differences in patterns of topographic change with wear reflect differences in diet.Descriptions of new fossils of the early Miocene stem cercopithecoid Noropithecus bulukensis confirm its generic distinction from Victoriapithecus macinnesi and highlight the dental metric and morphological variation that complicates identification of isolated teeth. Results of dental topographic analyses show that wear-mediated patterns of change in functional topographic metrics do not reflect broad dietary differences in extant catarrhines. While topographic features of unworn molars exhibit a phylogenetic signal, the pattern of wear-mediated topographic change does not. Molar topography of victoriapithecids is similar to extant cercopithecids with frugivorous and hard-object feeding diets, supporting previous dietary reconstructions. Victoriapithecid molar occlusal surfaces exhibit less complexity, less curvature, and higher relief than proconsulids prior to heavy wear stages. They are not distinct from occlusal topographies of small-bodied non-cercopithecoid catarrhines at any wear stage. Overall, these results suggest that the acquisition of bilophodont molar morphology in early and middle Miocene stem cercopithecoids was not associated with a shift in occlusal topography relative to more basal catarrhines. Rather, it is among proconsulids that shifts toward more complex, higher curvature occlusal surfaces are found.
ContributorsLocke, Ellis (Author) / Reed, Kaye (Thesis advisor) / Schwartz, Gary (Committee member) / Kelley, Jay (Committee member) / Arizona State University (Publisher)
Created2021