The energy frugality hypothesis states that in response to Madagascar’s unpredictable habitat, lemurs should adopt strategies of energy optimization. I have applied this hypothesis to lemur behavioural ecology in response to forest edges. I compared two groups of Propithecus coquereli living less than 1-km from a forest edge with two groups living greater than 1-km from the edge in Ampijoroa forest station, Ankarafantsika National Park, NW Madagascar. Edge effects in Ampijoroa penetrated up to 625-m into the forest. Propithecus coquereli were edge avoiders, with 94.54% of sightings of Propithecus coquereli found outside of the area of edge influence. There was no difference between group ranges for density of food trees, however tree diameter at breast height (dbh) and tree height did differ between groups. These habitat differences did not neatly divide edge versus interior groups, but appeared to be more nuanced. I found no differences between groups for activity budgets, food quality, or spatial patterns of plant species/ parts consumed. However, groups nearer to the edge had home ranges that were more than double in size to interior groups. Groups in the interior had higher group-specific densities and more frequent intergroup encounters which may have led groups to adopt smaller ranges to avoid expending energy in intergroup encounters. More evidence of human impact was found in the edge, therefore groups near the edge might also range further to avoid humans. Groups showed differences in the spatial pattern of behaviours, activity by age-sex category, substrate size and vertical location used while traveling, and dietary overlap. However, these differences may not be reflective of distance to the forest edge - all groups avoided the edge - but instead may reflect subtle differences in habitat structure between the four groups.

Research in hominoid cranial ontogeny has provided significant insight into the similarities and differences between apes and humans. Additionally, questions about within-species variability, allometry, and levels of sexual dimorphism in fossil hominoids are commonly addressed using the extant great apes as a comparative framework. However, this model is incomplete without the addition of the lesser apes, gibbons and siamangs. This analysis completes the hominoid record of cranial ontogeny with the addition of the Hylobatidae, and provides a full description of their cranial ontogeny and adult variation. Three-dimensional coordinates of 145 landmarks and 313 semilandmarks were measured on CT and surface scans from an ontogenetic sample of hominoid crania, comprising Pan, Gorilla, Pongo, Hylobates, and Symphalangus, with a cercopithecoid out-group, Colobus. After Procrustes superimposition, principal component analyses were computed in shape space. Regressions of shape coordinates on centroid size were used to assess within-group ontogenetic and static allometric trajectories. Results indicate that the shape changes during ontogeny in gibbons are similar to the shape changes previously reported for great apes. Genus-specific differences are already observable early in ontogeny, and the subsequent ontogenetic trajectories are almost parallel. Sexual dimorphism for both shape and size is found in Symphalangus adults, which was previously unobserved. Analysis of all taxa in the sample shows nearly parallel ontogenetic trajectories within the Hominoidea, which is consistent with previous studies. With the addition of hylobatids and Colobus, this analysis demonstrates that cranial ontogeny is highly conserved in the Catarrhini. Given the existence of this basic catarrhine growth trajectory, it should be possible in the future to predict fossil taxa morphologies at any stage of growth.

Research in hominoid cranial ontogeny has provided significant insight into the similarities and differences between apes and humans. Additionally, questions about within-species variability, allometry, and levels of sexual dimorphism in fossil hominoids are commonly addressed using the extant great apes as a comparative framework. However, this model is incomplete without the addition of the lesser apes, gibbons and siamangs. This analysis completes the hominoid record of cranial ontogeny with the addition of the Hylobatidae, and provides a full description of their cranial ontogeny and adult variation. Three-dimensional coordinates of 145 landmarks and 313 semilandmarks were measured on CT and surface scans from an ontogenetic sample of hominoid crania, comprising Pan, Gorilla, Pongo, Hylobates, and Symphalangus, with a cercopithecoid out-group, Colobus. After Procrustes superimposition, principal component analyses were computed in shape space. Regressions of shape coordinates on centroid size were used to assess within-group ontogenetic and static allometric trajectories. Results indicate that the shape changes during ontogeny in gibbons are similar to the shape changes previously reported for great apes. Genus-specific differences are already observable early in ontogeny, and the subsequent ontogenetic trajectories are almost parallel. Sexual dimorphism for both shape and size is found in Symphalangus adults, which was previously unobserved. Analysis of all taxa in the sample shows nearly parallel ontogenetic trajectories within the Hominoidea, which is consistent with previous studies. With the addition of hylobatids and Colobus, this analysis demonstrates that cranial ontogeny is highly conserved in the Catarrhini. Given the existence of this basic catarrhine growth trajectory, it should be possible in the future to predict fossil taxa morphologies at any stage of growth.

The energy frugality hypothesis states that in response to Madagascar’s unpredictable habitat, lemurs should adopt strategies of energy optimization. I have applied this hypothesis to lemur behavioural ecology in response to forest edges. I compared two groups of Propithecus coquereli living less than 1-km from a forest edge with two groups living greater than 1-km from the edge in Ampijoroa forest station, Ankarafantsika National Park, NW Madagascar. Edge effects in Ampijoroa penetrated up to 625-m into the forest. Propithecus coquereli were edge avoiders, with 94.54% of sightings of Propithecus coquereli found outside of the area of edge influence. There was no difference between group ranges for density of food trees, however tree diameter at breast height (dbh) and tree height did differ between groups. These habitat differences did not neatly divide edge versus interior groups, but appeared to be more nuanced. I found no differences between groups for activity budgets, food quality, or spatial patterns of plant species/ parts consumed. However, groups nearer to the edge had home ranges that were more than double in size to interior groups. Groups in the interior had higher group-specific densities and more frequent intergroup encounters which may have led groups to adopt smaller ranges to avoid expending energy in intergroup encounters. More evidence of human impact was found in the edge, therefore groups near the edge might also range further to avoid humans. Groups showed differences in the spatial pattern of behaviours, activity by age-sex category, substrate size and vertical location used while traveling, and dietary overlap. However, these differences may not be reflective of distance to the forest edge - all groups avoided the edge - but instead may reflect subtle differences in habitat structure between the four groups.

The origin and diversification of great apes and humans is among the most researched and debated series of events in the evolutionary history of the Primates. A fundamental part of understanding these events involves reconstructing paleoenvironmental and paleogeographic patterns in the Eurasian Miocene; a time period and geographic expanse rich in evidence of lineage origins and dispersals of numerous mammalian lineages, including apes. Traditionally, the geographic origin of the African ape and human lineage is considered to have occurred in Africa, however, an alternative hypothesis favouring a Eurasian origin has been proposed. This hypothesis suggests that that after an initial dispersal from Africa to Eurasia at ~17Ma and subsequent radiation from Spain to China, fossil apes disperse back to Africa at least once and found the African ape and human lineage in the late Miocene. The purpose of this study is to test the Eurasian origin hypothesis through the analysis of spatial and temporal patterns of distribution, in situ evolution, interprovincial and intercontinental dispersals of Eurasian terrestrial mammals in response to environmental factors. Using the NOW and Paleobiology databases, together with data collected through survey and excavation of middle and late Miocene vertebrate localities in Hungary and Romania, taphonomic bias and sampling completeness of Eurasian faunas are assessed. Previous bioprovincial zonations of Europe and Western Asia are evaluated and modified based on statistical analysis of Eurasian faunas and consideration of geophysical, climatic and eustatic events. Within these bioprovinces, occurrences of in situ evolution and directionality of dispersals of land mammals are used as a framework to address and evaluate these same processes in Eurasian apes. The results of this analysis support previous hypotheses regarding first occurrences and phyletic relations among Eurasian apes and propose new ideas regarding the relations of these taxa to previously known and newly discovered late Miocene African apes. Together with analysis of environmental data, Eurasian mammals support the hypothesis that the descendant of a Eurasian ape dispersed to Africa in the early late Miocene (top of MN7/8 or base of MN9), however the question of whether this taxon founded the African ape and human lineage remains equivocal.

The origin and diversification of great apes and humans is among the most researched and debated series of events in the evolutionary history of the Primates. A fundamental part of understanding these events involves reconstructing paleoenvironmental and paleogeographic patterns in the Eurasian Miocene; a time period and geographic expanse rich in evidence of lineage origins and dispersals of numerous mammalian lineages, including apes. Traditionally, the geographic origin of the African ape and human lineage is considered to have occurred in Africa, however, an alternative hypothesis favouring a Eurasian origin has been proposed. This hypothesis suggests that that after an initial dispersal from Africa to Eurasia at ~17Ma and subsequent radiation from Spain to China, fossil apes disperse back to Africa at least once and found the African ape and human lineage in the late Miocene. The purpose of this study is to test the Eurasian origin hypothesis through the analysis of spatial and temporal patterns of distribution, in situ evolution, interprovincial and intercontinental dispersals of Eurasian terrestrial mammals in response to environmental factors. Using the NOW and Paleobiology databases, together with data collected through survey and excavation of middle and late Miocene vertebrate localities in Hungary and Romania, taphonomic bias and sampling completeness of Eurasian faunas are assessed. Previous bioprovincial zonations of Europe and Western Asia are evaluated and modified based on statistical analysis of Eurasian faunas and consideration of geophysical, climatic and eustatic events. Within these bioprovinces, occurrences of in situ evolution and directionality of dispersals of land mammals are used as a framework to address and evaluate these same processes in Eurasian apes. The results of this analysis support previous hypotheses regarding first occurrences and phyletic relations among Eurasian apes and propose new ideas regarding the relations of these taxa to previously known and newly discovered late Miocene African apes. Together with analysis of environmental data, Eurasian mammals support the hypothesis that the descendant of a Eurasian ape dispersed to Africa in the early late Miocene (top of MN7/8 or base of MN9), however the question of whether this taxon founded the African ape and human lineage remains equivocal.

When human skeletal remains are found in prehistoric, historic, or forensic contexts, establishing age at death is an important step in reconstructing life histories, building demographic profiles, and identifying victims. Reliability for adult skeletal age estimations is generally lower than ideal, especially for individuals over the age of 40 years. A factor rarely considered in age estimation is that of body size; namely, how individuals of varying body sizes experience skeletal aging. This thesis explores the variables of BMI, stature, and body mass to quantify the influence of error from body size on adult skeletal age estimates. Eight age estimation methods were tested on 764 adult skeletons from the Hamann-Todd and William Bass Collections. These individuals were documented to have ranged in stature from 1.30m to 1.93m and body mass from 24.0kg to 99.8kg. Each age estimation method was evaluated separately. Analyses show that underweight individuals have the most error associated with their age estimations for all methods. All methods under-age underweight individuals by 5 to 15 years compared to average and obese individuals. The Kunos et al. method is the most reliable rib method, and the Lovejoy et al. and Suchey-Brooks methods are the most reliable pelvic methods. The İşcan et al. fourth rib method under-ages individuals by the highest degree compared to all other methods. The DiGangi et al. method is the most unreliable method with low accuracy (over twenty years) and high bias scores (under-aging individuals by almost 20 years). Body mass has a stronger influence on age estimation than stature. Studies have shown that body mass has a strong impact on bone resorption and remodelling rates, regardless of nutrition, and that tissue type (i.e. fat vs. muscle), physical activity, and mechanical loading influence bone remodelling. Skeletal remains of underweight individuals have fewer age markers while obese individuals show an increase in surface texture degeneration and osteophytic lipping. This is the first study to show that body size influences skeletal age estimation, and that age estimations are significantly different between weight-bearing and non-weight-bearing joints.

Sex-specific calls are used in male-male agonistic encounters and male-female courtship in many animal species. The ring-tailed lemur (Lemur catta) is a gregarious Malagasy strepsirhine with twenty-two distinct vocalizations for adults, including two male-specific vocalizations and an additional vocalization with male-specific functions: the howl, the squeal, and the purr. Proposed intra-sexual agonistic functions for these three vocalizations have never been empirically tested. This study’s purpose was to investigate the functions of howling, squealing, and purring in the ring-tailed lemur, and to assess the relationships between the rates of these vocalizations and male dominance. From March to July 2010, I collected 600 hours of total data and 480 hours of focal data on male ring-tailed lemurs aged three and older at Beza Mahafaly Special Reserve, Madagascar. I observed each male continuously for 30 minutes at a time and noted behaviours including all vocalizations and all agonism using one–zero sampling at 2.5-min intervals. I calculated male dominance rank and vocalization rates from these data. My results indicated that male dominance rank is correlated with male purring rate and with squealing rate, but not with howling rate. Male purring rate increased during intra-sexual agonism and was associated with aggression in agonistic encounters. Squealing rate increased during male-male agonism and indicated both aggression and submission in male-male encounters. Howling rate increased during inter-group encounters and a greater number of males participated in multi-male howling choruses when non-group members were present. Purring and squealing are agonistic vocalizations and used in male-male agonism in the ring-tailed lemur, while howling is used in inter-group encounters.

Sex-specific calls are used in male-male agonistic encounters and male-female courtship in many animal species. The ring-tailed lemur (Lemur catta) is a gregarious Malagasy strepsirhine with twenty-two distinct vocalizations for adults, including two male-specific vocalizations and an additional vocalization with male-specific functions: the howl, the squeal, and the purr. Proposed intra-sexual agonistic functions for these three vocalizations have never been empirically tested. This study’s purpose was to investigate the functions of howling, squealing, and purring in the ring-tailed lemur, and to assess the relationships between the rates of these vocalizations and male dominance. From March to July 2010, I collected 600 hours of total data and 480 hours of focal data on male ring-tailed lemurs aged three and older at Beza Mahafaly Special Reserve, Madagascar. I observed each male continuously for 30 minutes at a time and noted behaviours including all vocalizations and all agonism using one–zero sampling at 2.5-min intervals. I calculated male dominance rank and vocalization rates from these data. My results indicated that male dominance rank is correlated with male purring rate and with squealing rate, but not with howling rate. Male purring rate increased during intra-sexual agonism and was associated with aggression in agonistic encounters. Squealing rate increased during male-male agonism and indicated both aggression and submission in male-male encounters. Howling rate increased during inter-group encounters and a greater number of males participated in multi-male howling choruses when non-group members were present. Purring and squealing are agonistic vocalizations and used in male-male agonism in the ring-tailed lemur, while howling is used in inter-group encounters.

The late Holocene marks a period of significant population movement and subsistence change throughout much of sub-Saharan Africa. Around 3500 BP it appears that foraging populations in southernmost South Africa began to experience stress related to an increasing population and changing climatic conditions. Approximately 1500 years later a new form of subsistence - sheep herding – emerged in areas previously occupied solely by foragers, but was not exclusively adopted. The mechanisms surrounding the introduction of this new subsistence strategy – an indigenous adoption via diffusion or a foreign migration - remain unresolved. This study takes a biological approach to this significant question in southern African prehistory by exploring a collection of Later Stone Age skeletal remains that predate and postdate the appearance of pastoralism in order to determine if any significant changes in skeletal morphology indicative of population discontinuity can be identified at 2000 BP. A collection of seventy-three Later Stone Age adult skeletons (31 M, 42 F) with newly generated radiocarbon dates ranging from 8000 BP to 300 BP (uncalibrated) from the Eastern Cape Province of South Africa forms the basis of this study. Questions surrounding population continuity or discontinuity associated with the advent of sheep herding are investigated by examining metric variables collected from the cranium, post cranial skeleton and dentition, in conjunction with cranial discrete traits. Some changes in skeletal morphology are observed, but the timing, pattern and magnitude of these changes are not consistent with a foreign migration. A reduction in overall skeletal size in the absence of changes in shape corresponds with the period of forager intensification. Body size rebounds at around 2000 BP when evidence for a new form subsistence, sheep herding, is first observed in this region, suggesting that some foragers may have adopted sheep and the herding way of life as a stress relieving mechanism, while others maintained the foraging lifestyle. The timing of the observed changes in skeletal size, the absence of shape changes and the homogeneity in cranial discrete trait frequencies through time argues against the idea that sheep herding was introduced to the Cape region by outsiders. Rather, the findings of the current study suggest sheep herding was an indigenous development among existing foragers.