Browsing by Subject "geometric morphometrics"

Fossil taxa are crucial to studies of brain evolution, as they allow us to identify evolutionary trends in relative brain size and brain shape that may not otherwise be identifiable in comparative studies using only extant taxa, owing to multiple events of parallel encephalization among primate clades. This thesis combines indirect and direct approaches to understanding primate evolution, by evaluating variation in the endocranial morphology of extant primates and their fossil representatives. I use a comparative approach to examine the relationships between interspecific adult endocranial volume and shape, and brain evolution and cranial form among extant primate clades and their fossil representatives. The associations are evaluated via phylogenetically informed statistics perfomed on volumetric measurements and three-dimensional geometric morphometric analyses of virtual endocasts constructed from micro-CT scans of primate crania. Fossil taxa included in these analyses are: 1) anthropoids Parapithecus, Aegyptopithecus (Early Oligocene, Egypt), Homunculus and Tremacebus (Early Miocene, Argentina), and 2) Eocene euprimates Adapis and Leptadapis (Eocene adapoids, France), and the Rooneyia (Eocene omomyoid, Texas).

The first part of this work (Chapter 2) explores variation in residual mass of brain components (taken from the literature) among primates, and evaluates the correlated evolution of encephalization and brain proportions with endocast shape, quantified via three-dimensional geometric morphometric techniques. Analyses reveal a broad range of variation in endocast shape among primates. Endocast shape is influenced by a complex array of factors, including phylogeny, body size, encephalization, and brain proportions (residual mass of brain components). The analysis supports previous research, which concludes that anthropoids and tarsiers (Haplorhini) share the enlargement of several key brain regions including the neocortex and visual systems, and a reduction of the olfactory system. Anthropoids further differ from strepsirrhines in endocranial features associated with encephalization--a more flexed brain base, an inferiorly deflected olfactory fossa--and those associated with brain proportions--a small olfactory fossa, and a more caudally extended cerebrum that extends posteriorly past the cerebellar poles. Tarsiers are unique in having a mediolaterally broad and rostro-caudally short endocast with an attenuated anterior and middle cranial fossae. This morphology is likely related to the extreme orbital enlargement in this taxon, which limits anterior expansion of the endocranium. Finally, despite the correlation between residual endocranial volume and endocast shape among modern primates, early anthropoid fossils demonstrate a disconnect between these factors in sharing key features of endocast shape with extant anthropoids at a relatively small brain size.

The second part of this thesis (Chapter 3) explores the relationship between craniofacial organization--cranial base angle, facial size, facial hafting--and encephalization via the lens of the Spatial Constraints and Facial Packing Hypotheses. These hypotheses predict that interspecific adult variation in encephalization correlates with endocranial shape such that a larger brain for a given body size will be more "globular" or spherical in shape. These hypotheses futher predict that basicranial angle covaries with encephalization and that the relative size of the endocranium and facial skeleton will have an antagonistic effect on basicranial angle and facial hafting. Results show that various measures of globularity have inconsistent and weak relationships to phylogeny, encephalization, and basicranial flexion, owing to a diversity of clade-specific scaling patterns between the maximum length, breadth, and width of the endocast. Among extant primates, there is weak but significant evidence to suggest that both facial size and encephalization influence variation in basicranial flexion. Considering the fossil specimens in isolation, their relative ranks in encephalization, basicranial flexion, and midline facial size and shape follow the pattern expected from the Spatial and Facial Packing Hypotheses outlined above; however, relative to modern species, the early fossil anthropoids have more flexed cranial bases and shorter facial skeletons at much smaller level of encephalization than seen in modern anthropoids.

Together, the extant data suggest a moderately conserved pattern of correlated evolution among endocranial size, endocranial shape, brain proportions, and craniofacial organization, which may explain differences in endocranial and facial shape between extant strepsirrhine and anthropoid primates; however, the fossil record for early anthropoid evolution demonstrates that a shift towards key anthropoid-like traits of the endocranium, basicranium, and facial skeleton were initiated early in anthropoid evolution, with subsequent encephalization occurring within and among members of this clade. Thus, these anthropoid cranial traits evolved in tandem with changes in the relative size of brain components, rather than absolute or relative brain size alone. Basicranial flexion, facial length and orientation are influenced by both: 1) shifts in endocranial shape associated with changes in brain proportion--accounting for the initiation of the anthropoid-like craniofacial plan early in the evolution of the clade--and 2) encephalization, which influenced subsequent morphological divergence among extant anthropoid groups.

During the last six million years, humans shifted from a primarily arboreal lifestyle to a habitually bipedal, terrestrial lifestyle. Australopithecus had a significant bipedal component to its locomotion; whether suspensory and climbing behavior were also important has remained unclear. Morphological features of the forelimb have been linked to locomotor differences among primates, but the interpretation of human fossils has remained problematic.

This dissertation examined the total morphological pattern of the forelimb, specifically the functional integration of the musculature and joint systems. This approach employed both geometric morphometrics and a biomechanical modeling approach to studying how and how well the forelimb morphology of living suspensory and quadrupedal primates, as well as humans and fossil hominins, accommodate climbing and suspensory locomotion. Data collected with a microscribe 3-D digitizer on the scapula, humerus, radius, and ulna of Australopithecus sediba, Australopithecus afarensis, and Homo erectus were compared to a sample of Homo sapiens, Pan troglodytes, Pan paniscus, Gorilla gorilla, Pongo pygmaeus, Hylobates lar, and Macaca fuscicularis.

The hominin upper limb is a rich mosaic of primitive and derived traits. The blade of Australopithecus sediba, although appearing most similar to extant orangutans, is in fact functionally most similar to chimpanzees. The overall morphology of the australopith elbow joint appears most similar to Pan, as does the elbow joint of Homo erectus, suggesting that the modern human configuration happened more recently than 1.5 million years ago.

Au. afarensis and Au. sediba share important similarities, but are clearly distinct species. While their overall elbow joint shape is strikingly similar, the articular surface is not identical. Au. afarensis is more similar in this respect to Pan and Homo, while Au. sediba is more similar to extant taxa that spend substantially more time engaging in vertical climbing and suspensory behavior.

The results from this study support previous interpretations that not all australopiths across time were employing the same locomotor repertoire. While this study does not present unambiguous conclusions regarding early hominin arboreal locomotion, this study suggests that the morphology of the upper limb is varied, and caution must be taken when interpreting single skeletal elements in the hominin fossil record.

Primates span incredible behavioral and ecological diversity, and this diversity is reflected in the shape of the skull. This dissertation asks two questions surrounding the evolution of primate skull shape: 1) what are the macroevolutionary correlates of primate skull shape? And 2) what is the pattern of phenotypic integration in the primate skull at a macroevolutionary scale? To address these questions, I compiled a broad comparative dataset of anatomical landmarks identified from 3D scans of primate skulls and analyzed this data using statistical methods that combine geometric morphometrics and phylogenetic comparative methods. To investigate the macroevolutionary correlates of skull shape, I used multivariate phylogenetic generalized linear models to test for relationships between skull shape and several variables that are predicted to be correlated with skull shape: allometry, typical diet, tree gouging behavior, activity pattern, and sexual dimorphism. I found strong phylogenetic signal for primate skull shape, confirming the need for phylogenetic comparative methods. Allometry was a significant predictor of skull shape, with larger primates having relatively small, convergent orbits, and anteroposteriorly short skulls compared to small primates. Sexual dimorphism was associated with a dramatically lengthened rostrum, probably to facilitate a large gape in aggressive displays. Folivory was associated with deeper mandibles, which may improve mechanical advantage and strain resistance. To investigate patterns of phenotypic integration in the skull, I performed hierarchical clustering analyses on phylogenetically corrected inter-landmark covariance matrices. In contrast to previous research, I did not find evidence for distinct phenotypic modules in the primate skull, and I argue that this discrepancy is due to methodological shortcomings of past research that biased results towards identifying different anatomical regions as discrete modules. This dissertation represents one of the first investigations of primate skull shape at a macroevolutionary scale, and demonstrates that the combination of geometric morphometrics and phylogenetic comparative methods can yield novel insights into evolutionary morphology.