Apes share a distinct set of morphological and anatomical characteristics that allow us to use our arms and hands in unique ways. Apes also have relatively larger brains with similar sulcal patterning indicating there is a distinctively hominoid brain structure. These features in great apes have consistently been linked with higher cognitive skills and are considered to be the precursors leading to the exceptional developments of humans over evolutionary time – establishing the physiological basis that allows us to make and use tools to modify our environments and build our unique cultures. This study examines the general model that orthogrady and suspensory postures were the antecedents for enhanced manipulative ability in apes, which consequently set the stage for enhanced cognitive abilities in early hominoids. The primary hypothesis is that if single-handed prehensility is enhanced by suspensory orthogrady, then we may predict ape feeding style will differ significantly from that of pronograde monkeys, allowing them to access foods in unique ways. Using sympatric white-handed gibbons, Hylobates lar, representative of orthograde apes, and pig-tailed macaques, Maccaca leonina, representative of pronograde monkeys, as models, the comparative feeding styles of primates were analyzed focusing on their positional behaviour and manual skill. Results support the hypothesis that gibbons exhibit a unique feeding style associated with their orthogrady/suspensory postures in comparison with pronograde macaques. This was demonstrated by their increased access to food in the trees with more stable postures, an expanded foraging radius, and more frequent use of the terminal branches, and was evident in their complex manipulative skills with larger manual repertoires, more variability in wrist use, and more sophisticated manual techniques. Moreover, significant differences in positional behaviour and manual skill demonstrated by gibbons and macaques were evident even when feeding on the same types of foods within their shared environment. This study proposes that the combined uniquely ape traits to forage in suspensory orthograde postures with precision dexterity have allowed apes to become highly selective feeders within their environments, leading to advance manual dexterity and cognitive prowess in apes.

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A comparative understanding of the patterns and processes of hominoid evolution is critical for determining the evolutionary trajectories of our own species. Several traits, including the development of relatively large-body size, a torso-orthograde (or upright; TO) -Bauplan, and suspensory adaptations are often considered key adaptations within the Hominoidea. Unfortunately, comparisons of the neontological and paleontological records have resulted in divergent theories regarding the origin of these traits, with one extreme advocating their homologous origins and the other for rampant homoplasy. It is argued here that a key factor that has continued to limit our understanding of hominoid and human evolution is the underutilization of hylobatids within comparative studies. The Hylobatidae are an extremely successful radiation of moderate-sized primates from the superfamily Hominoidea. The extant hylobatid family currently comprises four genera and 20 species spanning 11 countries in East, South, and Southeast Asia, with three known extinct genera from China and potentially one from India. Hylobatids are the smallest of the extant apes yet size-variable and are among the most orthograde/suspensory. Nevertheless, their natural variation is often condensed into a single observation point in comparative studies. As such, the goal of this dissertation is to document the variation in positional behavior and postcranial osteology among hylobatids, utilizing an ecological morphological framework, and to integrate this detailed evaluation with previous neontological and paleontological studies. Several interrelated studies are presented within this dissertation, including a new comparative look at hylobatid postcranial osteology and several new studies of hylobatid positional behavior from the wild. These investigations shed light on the adaptive niche of hylobatids and provide insights into the evolutionary processes that shaped this uniquely successful hominoid family. Moreover, the new results presented here allow for a more critical understanding of hominoid evolution and facilitate the synchronization of the neontological and paleontological records. In doing so, this study provides support to the theory that hylobatids and hominids have developed their body size regimes, TO-Bauplan, suspensory morphological adaptions, and accompanying TO-positional behavioral repertoire independently.

Positional behavior is the measurable and observable link between the biology and behavior of an animal in its environment. In this dissertation, I examine ontogenetic patterns of positional behavior in infant, juvenile, and adult white-faced capuchins (Cebus capucinus) and mantled howling monkeys (Alouatta palliata) inhabiting the same tropical forest in Costa Rica. During growth and development ontogenetic changes in body size, limb proportions, and motor skills are likely to influence locomotion and posture through the arboreal canopy. I collected data on positional behavior, activity, prehensile-tail use, branch size, branch angle, and crown location during a 12 month period at Estación Biológica La Suerte in northeastern Costa Rica. The data set is comprised of 401.3 hours of data on Cebus capucinus and 554.3 hours of data on Alouatta palliata totaling 955.7 hours of data or 57,344 individual activity records.Life history timing and differences in rates of growth did not predictably influence the development of adult-like positional behaviors in Cebus and Alouatta. In both species, infancy was characterized by high proportions of dorsal, ventral, and side riding on the mother with smaller proportions of independent positional modes observed during play, explore, active posture, and feed/forage. Young Cebus resembled the adult pattern of positional behavior by six months of age while howlers exhibited significant differences in several positional behavior categories through 24 months of age. The positional repertoire of both species revealed similarities in the types of modes used during feed/forage and travel in juveniles and adults. For example, in juvenile and adult age categories of Cebus, feeding and foraging included high proportions of quadrupedal walk, sit, and squat in conjunction with climbing, leaping, and suspensory behaviors. In howlers, the degree to which coordination and increases in body mass during ontogeny as limiting factors in the development of adult-like positional competence is unclear. Data presented here suggest that the environment exerts different pressures on growing Cebus and Alouatta that may relate to diet, energy expenditure, foraging skill, and/or social learning.

abstract: 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. / Dissertation/Thesis / Ph.D. Anthropology 2013

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