Charles A. Lockwood

Quantifying temporal bone morphology of great apes and humans: an approach using geometric morphometrics

The hominid temporal bone offers a complex array of morphology that is linked to several different functional systems. Its frequent preservation in the fossil record gives the temporal bone added significance in the study of human evolution, but its morphology has proven difficult to quantify. In this study we use techniques of 3D geometric morphometrics to quantify differences among humans and great apes and discuss the results in a phylogenetic context. Twenty‐three landmarks on the ectocranial surface of the temporal bone provide a high level of anatomical detail. Generalized Procrustes analysis (GPA) is used to register (adjust for position, orientation and scale) landmark data from 405 adults representing Homo, Pan, Gorilla and Pongo. Principal components analysis of residuals from the GPA shows that the major source of variation is between humans and apes. Human characteristics such as a coronally orientated petrous axis, a deep mandibular fossa, a projecting mastoid process, and reduced lateral extension of the tympanic element strongly impact the analysis. In phenetic cluster analyses, gorillas and orangutans group together with respect to chimpanzees, and all apes group together with respect to humans. Thus, the analysis contradicts depictions of African apes as a single morphotype. Gorillas and orangutans lack the extensive preglenoid surface of chimpanzees, and their mastoid processes are less medially inflected. These and other characters shared by gorillas and orangutans are probably primitive for the African hominid clade.

The recognition and evaluation of homoplasy in primate and human evolution.

Homoplasy has been a prominent issue in primate systematics and phylogeny for as long as people have been studying human evolution. In the past, homoplasy, in the form of parallel evolution, was often considered the dominant theme in primate evolution. Today, it receives blame for difficulties in phylogenetic analysis, but is essential in the study of adaptation. This paper reviews the history of study of homoplasy, methods of defining homoplasy, and methodological and biological factors that generate homoplasy. A post hoc definition of homology and homoplasy, based on patterns of character distributions and their congruence or incongruence on a cladogram, is the most consistent method of recognizing these phenomena. Defined this way, homology and homoplasy are mutually exclusive. However, when different levels of analysis are examined, it is seen that homoplasy at one level, such as adult phenotype, often exists simultaneously with homology at a different level, such as developmental process. Thus, in some cases, patterns of homoplasy may point to underlying similarities that reflect the shared heritage of a particular clade. This is an old concept that is being renewed on the strength of recent trends in developmental biology. Factors that influence homoplasy include character definition and a host of biological factors, such as developmental constraints, allometry, and adaptation. These interact with one another to provide explanations of homoplastic patterns. Because of the repetition of events, explanations of homoplastic features are often more reliable than those for homologous features, and serve as effective tests for hypotheses of evolutionary process. In some cases, particular explanations of homoplasy lead to generalizations about the likelihood of homoplasy in a type of structure. The structure may be adaptive or highly epigenetic, or it may belong to an anatomical system considered to be more prone to homoplasy than others. However, our review shows that these generalizations are usually based on theory, and contradictory expectations can be developed under different theoretical models. More rigorous empirical studies are necessary to discover what, if any, generalizations can be made about the likelihood of homoplasy in different types of characters.

SEXUAL DIMORPHISM IN THE FACE OF AUSTRALOPITHECUS AFRICANUS

Recently discovered crania of Australopithecus africanus from Sterkfontein Member 4 and Makapansgat enlarge the size range of the species and encourage a reappraisal of both the degree and pattern of sexual dimorphism. Resampling methodology (bootstrapping) is used here to establish that A. africanus has a greater craniofacial size range than chimpanzees or modern humans, a range which is best attributed to a moderately high degree of sexual dimorphism. Compared to other fossil hominins, this variation is similar to that of Homo habilis (sensu lato) but less than that of A. boisei. The finding of moderately high dimorphism is corroborated by a CV-based estimate and ratios between those specimens considered to be male and those considered to be female. Inferences about the pattern of craniofacial dimorphism in the A. africanus face currently rely on the relationship of morphology and size. Larger specimens, particularly Stw 505, show prominent superciliary eminences and glabellar regions, but in features related in part to canine size, such as the curvature of the infraorbital surface, large and small specimens of A. africanus are similar. In this respect, the pattern resembles that of modern humans more so than chimpanzees or lowland gorillas. A. africanus may also show novel patterns of sexual dimorphism when compared to extant hominines, such as in the form of the anterior pillar. However, males of the species do not exhibit characteristics of more derived hominins, such as A. robustus.

Extended Male Growth in a Fossil Hominin Species

In primates that are highly sexually dimorphic, males often reach maturity later than females, and young adult males do not show the size, morphology, and coloration of mature males. Here we describe extended male development in a hominin species, Paranthropus robustus. Ranking a large sample of facial remains on the basis of dental wear stages reveals a difference in size and robusticity between young adult and old adult males. Combined with estimates of sexual dimorphism, this pattern suggests that male reproductive strategy focused on monopolizing groups of females, in a manner similar to that of silverback gorillas. However, males appear to have borne a substantial cost in the form of high rates of predation.

Homoplasy and adaptation in the atelid postcranium

Homoplasy is a ubiquitous phenomenon in phylogenetic investigations, but it is rarely investigated on its own. As a case study in the pattern and basis of homoplasy in primates, the atelid postcranium is discussed here. Characters available from Fords ([1986] in Erwin J, Swindler DR, eds: Comparative Primate Biology I: Systematics, Evolution, and Anatomy (New York: Alan R. Liss), p 73-135; [1994] in Fleagle JG, Kay RF, eds: Anthropoid Origins (New York: Plenum Press), p 595-674) analyses of New World monkeys are mapped onto alternative phylogenetic trees for the family Atelidae to contrast patterns of character evolution and to develop explanatory hypotheses for differences in the trees. In an unrooted phylogenetic network, pitheciines do not group together because those pitheciines that routinely adopt hind limb suspensory postures (Chiropotes, Cacajao) share traits with atelines. Fords (1986) work on phylogeny has shown that these traits are homoplastic and also identified potential synapomorphies of a clade comprised of modern pitheciins and atelines. However, following that work, congruence between studies of craniodental and molecular data suggested a still broader definition of atelids (including Callicebus and Cebupithecia), and in this case only one trait may define atelids, and several traits arise in parallel. The homoplastic characters in this phylogeny suggest that the phylogenetic signal in this set of postcranial data is overwhelmed by parallel adaptations to the use of climbing behaviors in all of Fords atelids and suspensory postures in a more restricted set of taxa. These parallelisms probably indicate a bias of selective pressures in the South American environment, especially given the frequent, independent evolution of suspensory mammals there. This highlights the fact that homoplasy can be a dominant source of similarity in data partitions strongly influenced by a particular behavioral regime, in this case positional behavior.

Adaptation and Behavior in the Primate Fossil Record

Few topics in functional morphology have been discussed as extensively with so little consensus as adaptation. Controversies swirl around both theoretical and methodological issues in the study of adaptation. Persistent questions concern definitions of adaptation, criteria for recognizing adaptations, and how to integrate phylogenetic and functional data. Here we review debates about how one identifies an adaptation, discuss recent developments in the study of adaptation, and focus on how these relate to our ability to reconstruct behavior in the fossil record. In keeping with the theme of this volume we will take our examples from the vast literature on primate functional anatomy, a resource that in many respects is more extensive and detailed than that for other orders of mammals. At the same time we will draw on the literature of other animals to introduce perspectives on the study of adaptation that have not been considered extensively by primatologists, but which hold considerable promise for future research.

Technical note: A new method for measuring long bone curvature using 3D landmarks and semi-landmarks.

Here we describe and evaluate a new method for quantifying long bone curvature using geometric morphometric and semi-landmark analysis of the human femur. The technique is compared with traditional ways of measuring subtense and point of maximum curvature using either coordinate calipers or projection onto graph paper. Of the traditional methods the graph paper method is more reliable than using coordinate calipers. Measurement error is consistently lower for measuring point of maximum curvature than for measuring subtense. The results warrant caution when comparing data collected by the different traditional methods. Landmark data collection proves reliable and has a low measurement error. However, measurement error increases with the number of semi-landmarks included in the analysis of curvature. Measurements of subtense can be estimated more reliably using 3D landmarks along the curve than using traditional techniques. We use equidistant semi-landmarks to quantify the curve because sliding the semi-landmarks masks the curvature signal. Principal components analysis of these equidistant semi-landmarks provides the added benefit of describing the shape of the curve. These results are promising for functional and forensic analysis of long bone curvature in modern human populations and in the fossil record.

Understanding paleoclimate and human evolution through the hominin sites and paleolakes drilling project

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Size variation in early human mandibles and molars from Klasies River, South Africa: Comparison with other middle and late Pleistocene assemblages and with modern humans†

Previous studies of the Middle Stone Age human remains from Klasies River have concluded that they exhibited more sexual dimorphism than extant populations, but these claims have not been assessed statistically. We evaluate these claims by comparing size variation in the best-represented elements at the site, namely the mandibular corpora and M(2)s, to that in samples from three recent human populations using resampling methods. We also examine size variation in these same elements from seven additional middle and late Pleistocene sites: Skhūl, Dolní Vestonice, Sima de los Huesos, Arago, Krapina, Shanidar, and Vindija. Our results demonstrate that size variation in the Klasies assemblage was greater than in recent humans, consistent with arguments that the Klasies people were more dimorphic than living humans. Variation in the Skhūl, Dolní Vestonice, and Sima de los Huesos mandibular samples is also higher than in the recent human samples, indicating that the Klasies sample was not unusual among middle and late Pleistocene hominins. In contrast, the Neandertal samples (Krapina, Shanidar, and Vindija) do not evince relatively high mandibular and molar variation, which may indicate that the level of dimorphism in Neandertals was similar to that observed in extant humans. These results suggest that the reduced levels of dimorphism in Neandertals and living humans may have developed independently, though larger fossil samples are needed to test this hypothesis.

Sexual Dimorphism and Facial Growth Beyond Dental Maturity in Great Apes and Gibbons

The great apes and gibbons are characterized by extensive variation in degree of body size and cranial dimorphism, but although some studies have investigated how sexual dimorphism in body mass is attained in these species, for the majority of taxa concerned, no corresponding work has explored the full extent of how sexual dimorphism is attained in the facial skeleton. In addition, most studies of sexual dimorphism combine dentally mature individuals into a single “adult” category, thereby assuming that no substantial changes in size or dimorphism take place after dental maturity. We investigated degree and pattern of male and female facial growth in Pan troglodytes troglodytes, Pan paniscus, Gorilla gorilla gorilla, Pongo pygmaeus, and Hylobates lar after dental maturity through cross-sectional analyses of linear measurements and geometric mean values of the facial skeleton and age-ranking of individuals based on molar occlusal wear. Results show that overall facial size continues to increase after dental maturity is reached in males and females of Gorilla gorilla gorilla and Pongo pygmaeus, as well as in the females of Hylobates lar. In male Pongo pygmaeus, adult growth patterns imply the presence of a secondary growth spurt in craniofacial dimensions. There is suggestive evidence of growth beyond dental maturity in the females of Pan troglodytes troglodytes and Pan paniscus, but not in the males of those species. The results show the presence of statistically significant facial size dimorphism in young adults of Pan paniscus and Hylobates lar, and of near statistical significance in Pan troglodytes troglodytes, but not in older adults of those species; adults of Gorilla gorilla gorilla and Pongo pygmaeus are sexually dimorphic at all ages after dental maturity. The presence of sex-specific growth patterns in these hominoid taxa indicates a complex relationship between socioecological selective pressures and growth of the facial skeleton.