Dissertations / Theses on the topic 'Paleoanthropology'

Today’s Chinese ethno-nationalism exploits nativist ancestral claims back to antiquity to legitimize its geo-political occupation of the entire territory of modern China, which includes areas where many non-Han people live. It also insists on the inseparability of the non-Han nationalities as an integrated part of Zhonghua minzu. This dissertation traces the origin of this nationalism to the two major waves of scientific investigation in the fields of paleoanthropology and anthropology in the Chinese frontier during the first half of the twentieth century. Prevailing theories and discoveries in the two scientific disciplines inspired the ways in which the Chinese intellectuals constructed their national identity. The first wave concerns the international quest for human ancestors in North China and the northwestern frontier in the 1920s and 1930s. Foreign scientists, such as Pierre Teilhard de Chardin, Amadeus Grabau, Roy Chapman Andrews, and Davidson Black, came to China to search for the first human fossils. With the discovery of Peking Man, they made Beijing one of the most prestigious places for the study of human paleontology and popularized the evolutionary Asiacentric theory that designated Chinese Central Asia and Mongolia as the cradle of humans. Inspired by the theory and the study of the Peking Man fossils, Chinese intellectuals turned Peking Man into the first Chinese and a common ancestor of all humans. In the second wave, from the late 1930s to the early 1950s, Chinese anthropologists like Rui Yifu, Cen Jiawu, Fei Xiaotong, and Li Anzhai made enormous efforts to inscribe the non-Han people of the southwestern frontier into the genealogy of the Chinese nation (Zhonghua minzu). Their interpretations of the relationship between the Han and the non-Han and between the frontier and the center were influenced by various Western anthropological theories. However, their intensive studies of the southwestern non-Han societies advocated the ethnic integration and nationalization of China’s southwestern frontier. By linking the two waves of scientific endeavor, this dissertation asserts that the Chinese intellectual construction of modern Chinese ethnogenesis and nationalism was not a parochial and reactionary nationalist “invention” but a series of indigenizing attempts to appropriate and interpret scientific theories and discoveries.
History

This thesis presents the results of geoarchaeological investigations conducted at two new test pits, M11 and M12, at the paleoanthropological site of Dmanisi during the 2012 field season. This research is important for understanding the site formation processes occurring along the north-south axis of the Dmanisi site and how that affects the chronostratigraphic sequence and interpretation of archaeological materials here. With these excavations we can build a stronger interpretation for how broader areas of this site formed and changed both geologically and archaeologically. The geologic results of this study indicate that changes in sediment deposition and development episodes can affect interpretations of how long these sediments accumulated, how likely bones are to preserve, as well as how secondary gravel deposition can influence several archaeological interpretations. The archaeological results suggest that there could have been changes in occupation intensity between the stratum A and B phases although different rates of sediment deposition and surface stability could affect such artifact accumulations. In addition, during the stratum B phase there appears to be little change in artifact procurement behaviors and reduction characteristics by these hominins. The overall results of this research indicate that geologic factors should be addressed and cautions should be taken prior to making interpretations about archaeological assemblages.

The early human paleoanthropological site at Ngandong, Central Java, Indonesia has significant impact on the models for human migration and evolution out of the African continent. Located on an abandoned stream bank above the Solo River, Ngandong archaeological digs have uncovered fourteen Homo erectus fossils that, based on their unique shape, are believed to have lived more recently than any other known examples of Homo erectus. However, this hypothesis has not been substantiated by previous studies at Ngandong due to a general lack of understanding about the formation of the site as a whole. This study seeks to overcome the limits of these previous studies by thoroughly examining the grain size, grain shape, mineralogy, geochemistry, and stratigraphy of the site to understand how it formed, and in turn, provide a necessary geological context to the Ngandong Homo erectus fossils. The results outlined in this dissertation suggest the fossil-bearing layers were deposited at the site (at the time a channel bottom) over a short period of time and were sourced from the volcanic arc that forms the southern portion of Java island.

Els canals diploicos es col·loquen en l'os esponjós entre les capes corticals de la volta (externa i interna). Les venes diploicas són grans, de parets primes i sense vàlvules que passen per una xarxa de canals microscòpics. Les venes diploicas semblen estar més desenvolupades en humans que en primats no humans, el que suggereix un paper clau en l'evolució humana. Es descriuen les xarxes vasculars diplóicas, proporcionant comparacions quantitatives i correlacions entre el gruix de l'os esponjós i compacte, la longitud de la branca del vas i la mida del lumen, les asimetries vasculars i la distribució volumètrica en els ossos frontal, parietal i occipital. Es van notar les comunicacions entre la xarxa diploica amb l'artèria meníngia a la fossa temporal, amb les venes emisarias en l'os occipital i amb els sins venosos a la confluència del si. En vista de les probables diferències metabòliques i les limitacions associades amb la mida i la forma del cervell humà modern, s'ha d'avaluar si aquestes diferències vasculars poden ser deguts a la termoregulació endocranial. Per tant, considerant la possible contribució dels gots diploicos en la termoregulació del cervell i el maneig de la calor, l'anàlisi de la xarxa diploica pot ser rellevant en antropologia, medicina i paleontologia.
Los canales diploicos se colocan en el hueso esponjoso entre las capas corticales de la bóveda (externa e interna). Las venas diploicas son grandes, de paredes delgadas y sin válvulas que pasan por una red de canales microscópicos. Las venas diploicas parecen estar más desarrolladas en humanos que en primates no humanos, lo que sugiere un papel clave en la evolución humana. Se describen las redes vasculares diplóicas, proporcionando comparaciones cuantitativas y correlaciones entre el grosor del hueso esponjoso y compacto, la longitud de la rama del vaso y el tamaño del lumen, las asimetrías vasculares y la distribución volumétrica en los huesos frontal, parietal y occipital. Se notaron las comunicaciones entre la red diploica con la arteria meníngea en la fosa temporal, con las venas emisarias en el hueso occipital y con los senos venosos en la confluencia del seno. En vista de las probables diferencias metabólicas y las limitaciones asociadas con el tamaño y la forma del cerebro humano moderno, se debe evaluar si estas diferencias vasculares pueden deberse a la termorregulación endocraneal. Por lo tanto, considerando la posible contribución de los vasos diploicos en la termorregulación del cerebro y el manejo del calor, el análisis de la red diploica puede ser relevante en antropología, medicina y paleontología.
The diploic channels are placed in the cancellous bone between the vault cortical layers (external and internal). The diploic veins are large, thin-walled and valveless running through a network of microscopic channels. Diploic veins appear to be more developed in humans than in non-human primates, suggesting a key role in human evolution. We describe their vascular networks, providing quantitative comparisons and correlations between compact and cancellous bone thickness, vessels branch length and lumen size, vascular asymmetries and volumetric distribution in frontal, parietal and occipital bones. Communications between the diploic network with the meningeal artery at the temporal fossa, with the emissary veins at the occipital bone, and with the venous sinuses at the confluence of sinus were noted. In view of probable metabolic differences and constraints associated with modern human brain size and shape, it should be evaluated whether these vascular differences can be due to endocranial thermoregulation. Therefore, considering the possible contribution of diploic vessels in brain thermoregulation and heat management, the analysis of the diploic network may be relevant in anthropology, medicine, and paleontology.

Thesis (Ph. D.)--University of California, San Diego, 2009.
Title from first page of PDF file (viewed January 19, 2010). Available via ProQuest Digital Dissertations. Vita. Includes bibliographical references (p. 283-295).

Thesis (Ph. D.)--University of Oregon, 2006.
Typescript. Includes vita and abstract. Includes bibliographical references (leaves 729-756). Also available for download via the World Wide Web; free to University of Oregon users.

2006 Dozier Award Winner
Agent-based models can provide paleoanthropologists with a view of behavioral dynamics and site formation processes as they unfold in digital caricatures of past societies and paleoenvironments. This paper argues that the agent-based methodology has the most to offer when used to conduct controlled, repeatable experiments within the context of behavioral laboratories. To illustrate the potential of this decidedly heuristic approach, I provide a case study of a simple agent-based model currently being used to investigate the evolution of Plio-Pleistocene hominin food sharing in East Africa. The results of this null model demonstrate that certain levels of ecological patchiness can facilitate the evolution of even simple food sharing strategies among equally simple hominin foragers. More generally, they demonstrate the potential that agent-based models possess for helping historical scientists act as their own informants as to what could have happened in the past.

Dental morphology and tooth shape have been used to recreate the dietary adaptations for extinct species, and thus dental variation can provide information on the relationship between fossil species and their paleoenvironments. Variation in living species with known behaviors can provide a baseline for interpreting morphology, and behavior, in the fossil record. Tooth occlusal surface outlines in hominins and non-hominin primates, and other mammals, have been used for assessments of taxonomic significance, with variability often considered as being primarily phylogenetic. Few studies have attempted to assess how diet might influence the pattern of variability in closely related species. Here the occlusal surface shape variability in anterior and postcanine maxillary dentition in primates is measured to assess whether the relationship between diet and variability is consistent. Data were collected from five non-hominin primates in a range of dietary categories, as well as two hominin species, including the derived Paranthropus robustus and a gracile australopith. Mapping a series of 50 sliding semilandmarks based on 2-D photographs using tpsDig software, occlusal surfaces were outlined. Thereafter, outline shapes were quantified using Elliptical Fourier Functional Analysis, and principle components and multivariate analyses were preformed to explore the pattern of intra and interspecific variability in occlusal outlines.These results suggest that there is not a clear relationship between dietary feeding adaptations for all categories examined and selection for larger premolars and molars, as well as smaller incisors, led to less variation in both anterior and post-canine teeth of the fossil hominin Paranthropus robustus.

La paléoanthropologie utilise la bipédie comme critère d'interprétation des vestiges fossiles permettant d'établir leur appartenance à la lignée humaine. Ainsi, la bipédie devient une caractéristique propre à la lignée humaine et qui en marque l'origine. Nous identifions ici un « cercle herméneutique » de la paléoanthropologie, puisque l'humain y est défini par la bipédie et, réciproquement, tout bipédie est interprétée comme étant nécessairement humaine. Du fait de cette circularité, les traits associés à la bipédie sont surinterprétés dans la description des vestiges fossiles, qui se voient alors conférer une signification fonctionnelle et phylogénétique univoque. L'unicité de la bipédie humaine constitue un principe d'interprétation resté ininterrogé en paléoanthropologie. Ce point révèle l'attachement de cette discipline scientifique à une conception philosophique du propre de l'homme qui semble pourtant difficilement compatible avec l'approche évolutionniste. Au contraire, une véritable épistémologie de la paléoanthropologie doit mettre en perspective la signification accordée à cette caractéristique anthropologique : nous nous y employons dans cette thèse, en montrant que le concept métaphysique de « station droite » trouve son écho dans le concept naturaliste de « bipédie ». Cette continuité ou cette capillarité de la philosophie à la science est mise en évidence à travers les usages du critère de la bipédie dans la description de plusieurs espèces fossiles. De Pithecanthropus erectus à Ardipithecus ramidus, nous analysons différents modes de l'équivalence entre « bipède » et « humain ». Cette équivalence, souvent implicite, représente une source de confusion majeure pour la paléoanthropologie, impliquant un concept d’ « humain » aux contours flous. Notre épistémologie de la paléoanthropologie dégage donc différents problèmes que cette discipline se doit d'affronter pour que le débat sur l'évolution de la bipédie dans la lignée humaine puisse avancer
Bipedalism represents for paleoanthropology a criterion to decipher fossil records in order to establish their belonging to the human lineage. As such, bipedalism is considered as a unique characteristic of the human lineage that marks its origin. In this thesis, we identify a "hermeneutic circle" of paleoanthropology, since the human is defined by its bipedal characteristics and, conversely, any bipedalism is interpreted as necessarily human. Because of this circularity, traits associated with bipedalism are overstated in describing the fossil record, hence conferring an unambiguous functional and phylogenetic significance. The uniqueness of human bipedalism is a principle of interpretation remained unquestioned in paleoanthropology. This shows the commitment of this scientific discipline to a philosophical conception of the human uniqueness who hardly seems consistent with the evolutionary approach. On the contrary, a real epistemology of paleoanthropology has put into perspective the meaning given to this anthropological characteristic. We address this latter in showing that the metaphysical concept of "upright station" is echoed in the naturalist concept of « bipedalism ». This continuity or capillary from philosophy to science is emphasized through the use of the criterion of bipedalism in the description of several fossil species. From Pithecanthropus erectus to Ardipithecus ramidus, we analyze various modes of equivalence between "biped" and "human". This equivalence, albeit implicit, represents a major source of confusion for paleoanthropology, implying an unclear concept of « human ». Our epistemology of paleoanthropology put thus upfront several philosophical and epistemological problems that this discipline has to challenge in order to the debate on the evolution of bipedalism in the human lineage can move forward

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Anthropology

Els estudis en Evolució Humana han conclòs en general que els trets derivats en humans van facilitar comportaments relacionats amb l'ús d'eines i ens van fer primats especialment destres. La manera com els humans adquirim aquesta configuració única de caràcters musclo-esquelètics té profundes implicacions en el nostre enteniment de l'Evolució Humana en general, ja que la tecnologia lítica és un element clau per definir cultures. El propòsit d'aquesta Tesi és avaluar si la mà humana va evolucionar per selecció natural en adaptació a l'ús d'eines lítiques, la qual és la hipòtesi més estesa per a l'evolució de la mà. Amb aquest propòsit, una mostra de mans de primats actuals, principalment humans, ximpanzés i goril·les, ha sigut analitzada amb diferents mètodes que permeten relacionar forma i funció. El primer objectiu ha sigut avaluar el comportament del polze sota escenaris simulats d'ús d'un percutor en hominoïdeus, a través de l'anàlisi d'elements finits. Els resultats indiquen que la distribució de l'estrès als ossos probablement no va ser una pressió selectiva prou forta com per modelar la forma dels ossos de la mà en humans. El segon objectiu ha sigut avaluar si els humans mostren una estructura de covariació en els carps que és diferent a la de ximpanzés i goril·les. Els resultats indiquen que diferents patrons de covariació poden haver modelat l'evolució del canell en aquests primats, podent estar relacionat amb demandes funcionals. L'últim objectiu ha estat provar si la mida de les èntesis de lligaments i músculs són proxis fiables per inferir patrons d'activitat. Els resultats indiquen que l'ús d'eines lítiques pot ser inferit en hominins analitzant regions de la mà on s'uneixen tendons (amb èntesis fibro-cartilaginoses) i lligaments. Això pot contribuir a la discussió de quan els hominins començaren a utilitzar eines lítiques i amb quina freqüència.
Los estudios en evolución humana han concluido en general que los rasgos derivados en humanos facilitaron comportamientos relacionados con el uso de herramientas y nos hicieron primates especialmente diestros. Cómo los humanos adquirimos esta configuración única de caracteres musculoesqueletales tiene profundas implicancias en nuestro entendimiento de la evolución humana en general, ya que la tecnología lítica es un elemento clave para definir culturas humanas. El propósito de esta tesis es evaluar si la mano humana evolucionó por selección natural en adaptación al uso de herramientas líticas, la que es la hipótesis más extendida para la evolución de la mano. Con este propósito, una muestra de manos de primates actuales, principalmente humanos, chimpancés y gorillas, fue analizada con distintos métodos que permiten relacionar forma y función biológica. El primer objetivo fue evaluar el comportamiento del pulgar bajo escenarios simulados de uso de un percutor en hominoideos, a través del análisis de elementos finitos. Los resultados indican que la distribución del estrés en los huesos probablemente no fue una presión selectiva suficientemente fuerte como para modelar la forma de los huesos de la mano en humanos. El segundo objetivo fue evaluar si los humanos muestran una estructura de covariación en los carpos que es diferente a la de chimpancés y gorilas. Los resultados indican que patrones de covariación distintos pueden haber modelado la evolución de la muñeca en estos primates, lo que puede estar relacionado con demandas funcionales distintas de locomoción y manipulación. El último objetivo fue probar si el tamaño de las entesis de ligamentos y músculos son proxis confiables para inferir patrones de actividad. Los resultados indican que el uso de herramientas líticas puede ser inferido en homininos analizando regiones de la mano donde se unen tendones (con entesis fibrocartilaginosas) y ligamentos. Esto puede servir para discutir desde cuándo y con qué frecuencia los homininos utilizaron herramientas líticas.
Studies in hand morphology have generally concluded that derived traits in humans facilitated stone tool-related behaviors and made us a highly dexterous primate. How humans acquired this unique configuration of musculoskeletal traits has profound implications for our understanding of human evolution overall, considering that stone technology is a key element defining culture in our species. The aim of this thesis was to evaluate whether the human hand evolved by natural selection in adaptation to stone tool use, which is the most widespread hypothesis for the evolution of the human hand. For this purpose, a comparative sample of hands of extant primates, mainly humans, chimpanzees and gorillas, were analyzed under different methods that relate biological form and function. The first objective of this thesis was to evaluate the performance of the thumb during simulated hammerstone use through a finite element analysis. The results that stress distribution in hand bones was unlikely to be a selective pressure strong enough to shape the hand of humans. The second objective was to evaluate whether humans have a different covariation structure at the wrist than that of chimpanzees and gorillas. Results indicate that different covariation patterns may be shaping the evolution of the wrist in these primates, which might be related to different functional demands of locomotion and manipulation. The last objective was to test whether entheseal size of ligaments and muscles are reliable for inferring activity patterns in the hand. The results indicate that stone tool use may be inferred in hominins by analyzing hand regions where tendons (with fibrocartilaginous entheses) and ligaments attach. This serves to discuss since when and to which extent hominins were using stone tools.

Els primats catarrins s’originaren a l’Àfrica durant l’Oigocè. A més dels cercopithecoïdeus i els hominoïdeus, inclouen diversos llinatges del Miocè de l’Àfrica i Euràsia. Les seves relacions filogenètiques són controversides a causa del registre fòssil incomplet, l’abundant homoplàsia, i l’evolució en mosaic. La identifiació d’àrees anatòmiques no gaire proclius a l’homoplàsia, com els canals semicirculars (CSs) de l’orella interna, és crucial per inferir la filogènia dels catarrins extints. Aquesta tesi explora la utilitat dels CSs per inferir la filogènia dels antropoïdeus, i especialment dels catarrins miocens. Els objectius específics inclouen: (1) testar la idoneïtat de diferents aproximacions de morfometria geomètrica tridimensional (3DGM); (2) quantificar el senyal filogenètic inclòs en la morfologia dels CSs i identificar-hi caràcters susceptibles de ser analitzats cladísticament; i (3) testar hipòtesis filogenètiques alternatives per als catarrins miocens d’Europa (pliopitecoïdeus i hominoïdeus). Els mètodes es fonamenten en escanejos de microtomografia computada i 3DGM estàndard basada en landmarks, així com una innovadora aproximació alternativa basada en deformació (sense landmarks). Les dades de forma s’analitzaren mitjançant regressions (al·lomètriques) bivariants i anàlisis multivariants (anàlisi de components principals entre groups), juntament amb mètodes informats filogenèticament (filomorfoespai amb reconstrucció de nodes ancestrals, índexs cladístics), mesures de senyal filogenètic (λ de Pagel i K de Blomberg), i anàlisi de clústers. Els resultats es proporcionen en tres articles publicats, que posteriorment són discutits amb relació a la bibliografia i les perspectives de futur. El primer article analitza els patrons de variació en la forma dels CSs en els catarrins actuals i en testa la utilitat per fer inferències filogenètiques en hominoïdeus extints. Els altres dos articles testen hipòtesis filogenètiques alternatives per al pliopitecoïdeu Epipliopithecus i els driopitecins Hispanopithecus i Rudapithecus. S’arriba a les següents conclusions: (1) La 3DGM basada en deformació captura adequadament la variació en la forma dels CSs i elimina els biaixos de la 3DGM estàndard, que depèn del disseny subjectiu del protocol de landmarks. (2) La forma dels CSs inclou un fort senyal filogenètic i discrimina entre els principals clades d’antropoïdeus i, fins i tot, entre els gèneres d’hominoïdeus, i és per tant potencialment útil per testar hipòtesis filogenètiques per als tàxons extints. (3) La reconstrucció dels morfotips ancestrals permet la definició de diversos caràcters dels CSs potencialment sinapomòrfics per als catarrins corona, els hominoïdeus corona, i els homínids corona. Aquesta darrers són els més distintius, deut a les seves proporcions volumètriques robustes. (4) La morfologia dels CSs del pliopitecoïdeu Epipliopithecus confirma l’opinió àmpliament sostinguda que aquest tàxon s’interpreta més parsimoniosament com un catarrí basal més derivat que els propliopitecoïdeus que no pas com un hominoïdeu (com suggerien alguns estudis recents). (5) L’enigmàtic antropomorf Oreopithecus presenta un mosaic de característiques dels CSs primitives i derivades, que suggereixen un estatus d’hominoïdeu basal juntament amb l’adquisició independent d’algunes sinapomorfies dels homínids. (6) La morfologia dels CSs dels driopitecins Hispanopithecus i Rudapithecus confirma que són gèneres de grans antropomorfs diferents, però no afavoreix de manera conclusiva un estatus d’homínid o d’hominí basals, perquè els antropomorfs africans són en gran mesura plesiomòrfics amb relació als orangutans, més derivats. (7) Aquesta tesi és el primer estudi de la morfologia dels CSs en els antropoïdeus mitjançant 3DGM basada en deformació. Confirma el potencial dels CSs per fer inferències filogenètiques en catarrins miocens. Aquestes mètodes s’haurien d’estendre a la còclea i els ossicles de l’orella mitjana per contribuir encara més a les anàlisis cladístiques dels catarrins actuals i fòssils.
“Los primates catarrinos se originaron en África durante el Oligoceno. Además de los cercopitecoideos y los hominoideos, incluyen varios linajes del Mioceno de África y Eurasia. Sus relaciones filogenéticas son controvertidas a causa del registro fósil incompleto, la abundante homoplasia, y la evolución en mosaico. La identificación de áreas anatómicas no muy proclives a la homoplasia, como los canales semicirculares (CSs) del oído interno, es crucial para inferir la filogenia de los catarrinos extintos. Esta tesis explora la utilidad de los CSs para inferir la filogenia de los antropoides, y especialmente los catarrinos miocénicos. Los objetivos específicos incluyen: (1) testar la idoneidad de diferentes aproximaciones de morfometría geométrica tridimensional (3DGM); (2) cuantificar la señal filogenética presente en la morfología de los CSs e identificar caracteres susceptibles de ser analizados cladísticamente; y (3) testar hipótesis filogenéticas alternativas para los catarrinos miocénicos de Europa (pliopitecoideos y hominoideos). Los métodos se basan en escaneos de microtomografía computarizada y 3DGM estándar basada en landmarks, así como una innovadora aproximación alternativa basada en deformación (sin landmarks). Los datos de forma se analizaron mediantes regresiones alométricas bivariantes y análisis multivariantes (análisis de componentes principales entre grupos), junto con métodos informados filogenéticamente (filomorfoespacio y reconstrucción de nodos ancestrales, índices cladísticos), medidas de señal filogenética (λ de Pagel y K de Blomberg), y análisis de clúster. Los resultados se proporcionan en tres artículos publicados, que posteriormente son discutidos en relación con la bibliografía y las perspectivas de futuro. El primer artículo analiza los patrones de variación en la forma de los CSs en los catarrinos actuales, y testa su utilidad para realizar inferencias filogenéticas en hominoideos extintos. Los otros dos artículos testan hipótesis filogenéticas alternativas para el pliopitecoideo Epipliopithecus y los driopitecinos Hispanopithecus y Rudapithecus. Se llega a las siguientes conclusiones: (1) La 3DGM basada en deformación captura adecuadamente la variación en la en la forma de los CSs y elimina los sesgos de la 3DGM estándar, que dependen del diseño subjetivo del protocolo de landmarks. (2) La forma de los CSs incluye una fuerte señal filogenética y discrimina entre los principales clados de antropoides e, incluso, entre los géneros de hominoideos, y es por lo tanto potencialmente útil para testar hipótesis filogenéticas para los taxones extintos. (3) La reconstrucción de los morfotipos ancestrales permite la definición de varios caracteres de los CSs potencialmente sinapomórficos para los catarrinos corona, los hominoideos corona, y los homínidos corona. Estos últimos son los más distintivos debido a sus proporciones volumétricas robustas. (4) La morfología de los CSs del pliopitecoideo Epipliopithecus confirma la opinión ampliamente sostenida que este taxón se interpreta más parsimoniosamente como un catarrino basal más derivado que los propliopitecoideos que como un hominoideo (como sugerían algunos estudios recientes). (5) El enigmático antropomorfo Oreopithecus presenta un mosaico de características de los CSs primitivas y derivadas, que sugieren un estatus de hominodeo basal junto con la adquisición independiente de algunas sinapomorfías de los homínidos. (6) La morfología de los CSs de los driopitecinos Hispanopithecus y Rudapithecus confirma que son géneros distintos de grandes antropomorfos, pero no favorece conclusivamente un estatus de homínido o de hominino basales, porque los grandes antropomorfos africanos son en gran medida plesiomórficos con relación a los orangutanes, más derivados. (7) Esta tesis es el primer estudio de la morfología de los CSs en los antropoides mediante 3DGM basada en deformación. Confirma el potencial de los CSs para realizar inferencias filogenéticas en catarrinos miocénicos. Estos métodos deberían extenderse a la cóclea y los huesecillos del oído medio para contribuir aún más a los análisis cladísticos en catarrinos actuales y fósiles.
Catarrhine primates originated in Africa by the Oligocene. Besides cercopithecoids and hominoids they include several lineages from the Miocene of Africa and Eurasia. Their phylogenetic relationships are controversial due to their incomplete fossil record, abundant homoplasy, and mosaic evolution. The identification of anatomical areas not very prone to homoplasy, such as the inner ear semicircular canals (SCs), is crucial for inferring the phylogeny of extinct catarrhines. The present dissertation explores the utility of SCs for phylogenetic inference among anthropoids, and particularly Miocene catarrhines. Specific aims include: (1) testing the suitability of different three-dimensional geometric morphometric (3DGM) approaches; (2) quantifying the phylogenetic signal embedded in SC morphology and identifying characters amenable to cladistic analysis; and (3) testing competing phylogenetic hypotheses for European Miocene catarrhines (pliopithecoids and hominids). The methods rely on microcomputed tomography scans and landmark-based standard 3DGM, as well as an innovative deformation-based (landmark-free) alternative approach. Shape data were analyzed by means of bivariate (allometric) regressions and multivariate analyses (between-group principal component analysis), coupled with phylogenetically-informed methods (phylomorphospace with ancestral node reconstruction, cladistic indices), metrics of phylogenetic signal (Pagel’s λ and Blomberg’s K), and cluster analysis. The results are provided in three published articles, which are subsequently discussed in relation to the literature and prospects for the future. The first article analyzes patterns of SC shape variation in extant catarrhines and tests its utility for making phylogenetic inferences in extinct hominoids. The remaining two articles test alternative phylogenetic hypotheses in the pliopithecoid Epipliopithecus and the dryopithecines Hispanopithecus and Rudapithecus. The following conclusions are reached: (1) Deformation-based 3DGM adequately captures SC shape variation and eliminates biases of standard 3DGM that depend on the subjective design of the landmarking protocol. (2) SC shape embeds strong phylogenetic signal and discriminates among major anthropoid clades and even extant hominoid genera, thus being potentially useful to test phylogenetic hypotheses for extinct taxa. (3) The reconstruction of ancestral morphotypes enables the definition of several characters of the SCs potentially synapomorphic for crown catarrhines, crown hominoids, and crown hominids. The latter are most distinctive due to their stout volumetric proportions. (4) The SC morphology of the pliopithecoid Epipliopithecus confirms the widely held view that this taxon is most parsimoniously interpreted as a stem catarrhine more derived than propliopithecoids, rather than a hominoid (as suggested by some recent studies). (5) The enigmatic ape Oreopithecus displays a mosaic of primitive and derived SC features suggestive of a stem hominoid status coupled with the independent acquisition of some hominid synapomorphies. (6) The SC morphology of the dryopithecines Hispanopithecus and Rudapithecus confirms they are distinct great ape genera, but does not conclusively favor a stem hominid vs. hominine status, because African apes are largely plesiomorphic as compared with the more derived orangutans. (7) This dissertation is the first study devoted to the SC morphology among anthropoids by means of deformation-based 3DGM. It confirms the potential of SCs for phylogenetic inference in Miocene catarrhines. These methods should be extended to the cochlea and middle ear ossicles to further contribute to cladistic analyses of extant and fossil catarrhines.
Universitat Autònoma de Barcelona. Programa de Doctorat en Biodiversitat

Au Levant, le Paléolithique moyen (environ 200-50 Ka B.P.) correspond à une période cruciale pour la compréhension des mécanismes évolutifs et la dispersion des Homo sapiens hors d’Afrique. Le site de Qafzeh (Basse Galilée), au croisement des trois grandes aires géographiques de l’ancien monde, comprend des niveaux archéologiques chronologiquement situés au cœur de cette période (92 ± 5 ka BP). Il est exceptionnel en raison à la fois d’une riche documentation anthropologique comprenant des individus de toutes les classes d’âges (du périnatal à l’adulte) et de la présence de pratiques funéraires uniques (sépulture double, dépôt funéraire). Si de nombreuses études ont déjà été publiées sur cette documentation fossile, les progrès technologiques récents de la Paléoimagerie permettent d’extraire des données inédites. Ainsi grâce à des outils 2D (radiographies, coupes CT) et 3D (reconstructions virtuelles à partir de données photogrammétriques ou de [micro]CT-scan) des analyses sont réalisées sur la tête osseuse de trois individus (Qafzeh 6, 9 et 25). Deux types d’analyses sont menés : une étude morphométrique de structures internes (i.e. labyrinthe osseux) et une étude paléopathologique. L’examen de l’oreille interne dévoile ainsi une diversité inattendue pour la population présente à Qafzeh ; par ailleurs des rapprochements au sein des échantillons de comparaison, entre quelques individus classiquement considérés comme archaïques et d’autres modernes, se dégagent. L’étude paléopathologique quant à elle révèle sur Qafzeh 9 la présence de troubles de la croissance osseuse et dentaire, qui font écho aux anomalies du développement précédemment décrites sur des fossiles immatures du site. Ces données nous permettent de mieux appréhender la variabilité individuelle normale et pathologique documentée à Qafzeh et de replacer ces chasseurs-cueilleurs nomades dans le contexte plus large des groupes locaux et eurasiatiques associés au contexte archéologique Moustérien
In the Levant, the Middle Paleolithic (about 200-50 Ka B.P.) corresponds to a period of time that is crucial for understanding evolutionary mechanisms and the spread of Homo sapiens out of Africa. The Qafzeh site (Lower Galilee) located at the crossroads of the three major geographical areas of the Old World is rich of archaeological deposits dated to the middle of this time period (92 ± 5 ka BP). The site is exceptional both for its important anthropological documentation of all age groups (from perinatal to adults) and for unique funeral practices (double burial, funeral repository). While many studies have already been published on the human remains, recent technological advances in Paleoimaging permit new investigation and data improvement. Using 2D (radiographs, CT sections) and 3D (virtual reconstructions from photogrammetric data or [micro]CT-scan) tools, two types of analyses are conducted on the skull and mandible of three individuals (Qafzeh 6, 9 and 25): a morphometric study of internal structures (i.e. bony labyrinth) and a palaeopathological investigation. Examination of the inner ear of these Mousterian individuals brings evidence of an unexpected diversity within the overall Qafzeh sample; furthermore, among hominins of the comparative sample, similarities between few individuals classically considered as archaic and modern specimens are recognized. Bone and dental growth disorders are described on Qafzeh 9 that are consistent with developmental abnormalities previously identified on immature individuals from this site. The data collected illustrate the normal and pathological variability documented by the nomadic hunter-gatherers from Qafzeh, and in addition, to compare them with local and Eurasian groups associated with a Mousterian archaeological context

Le gisement moustérien des Pradelles (Marillac-le-Franc, Charente, France) a été occupé saisonnièrement par les Néandertaliens en tant que halte de chasse, durant le stade isotopique 4. Il a livré près d’une centaine de restes humains (crâniens, dentaires et des membres) appartenant à au moins sept individus (enfants, adolescents et adultes). Ils présentent des traits morphologiques et des dimensions métriques intégrant la variabilité néandertalienne. Ils nous fournissent ainsi des données supplémentaires quant à la connaissance morphométrique des individus de ce taxon. En outre, certains vestiges présentent à leur surface des modifications anthropiques que l’on retrouve sur les restes de faune du site : impacts de fracturation, stries de découpe et de raclage ... L’étude taphonomique réalisée révèle un transport sélectif des parties du squelette des Néandertaliens sur le gisement ainsi que leur exploitation compatible avec une visée nutritive
The Mousterian site of Les Pradelles (Marillac-le-Franc, Charente, France) was occupied seasonally by the Neanderthal as hunting camp, during the isotopic stage 4. About a hundred human remains were unearthed (cranial, dental and limbs fragments) belonging to at least seven individuals (children, teenagers and adults). They present morphological features and metric dimensions integrating the Neanderthal variability. Thus, they provide additional data to our morphometric knowledge of the individuals of this taxon. Furthermore, some of these remains present anthropological modifications on their surface, the same that on the faunal remains from the site: percussion impacts, cutmarks, scraping marks ... The taphonomical study realized reveals a selective transport of the Neandertal skeletal parts to the site and their exploitation compatible with nutritive aim
El yacimiento musteriense de Les Pradelles (Marillac-le-Franc, Charente, Francia) ha sido ocupado temporadamente por los Neandertales como alto de caza, durante el estadio isotópico 4. Entregó un centenar restos humanos (craneanos, dentarios y de los miembros) perteneciente a un mínimo de siete individuos (niños, adolescentes y adultos). Presentan rasgos morfológicos y dimensiones métricas que integran la variabilidad de Neandertal. Así nos proporcionan datos suplementarios en cuanto al conocimiento morfometrico de este taxón. Además, un cierto número de estos vestigios presenta en su superficie modificaciones antrópicas que se encuentran en los restos de fauna del sitio: impactos de fracturación, estrías de recorte y de raspado... El estudio tafonómico realizado revela un transporte selectivo de las partes del esqueleto de los Neandertales en el yacimiento así como su explotación compatible con un objetivo nutritivo

L'évolution biologique des organismes peut être étudiée comme une succession de transformations morphologiques qui sont caractérisées par le changement de leur géométrie tridimensionnelle globale et locale. Dans ce contexte, il est nécessaire de développer des outils mathématiques et informatiques comparatifs de formes tridimensionnelles afin d'étudier ces transformations et de pouvoir les comparer avec les variabilités inter- et intra-espèces. Dans la chaîne de traitement des données tridimensionnelles (images 3D ou maillages 3D) employée en " paléoanthropologie virtuelle ", la méthode la plus souvent utilisée en analyse comparative est basée sur des points de repère (en général, anatomiques) dont les coordonnées sont analysées à l'aide d'outils mathématiques tels que la " morphométrie géométrique ". Plus récemment, une autre classe de méthodes a été proposée. Elle permet des comparaisons globales entre les surfaces complètes de structures anatomiques sans avoir besoin de définir des points de repère. On obtient ainsi une analyse statistique de la forme moyenne et de sa variabilité en tout point. Dans cette thèse, nous proposons d'étudier la chaîne d'analyse morphométrique des données 3D utilisées en paléoanthropologie, de la numérisation à l'exploitation des données par les chercheurs. Cette thèse présente des méthodes analytiques pour le traitement des données issues de la paléoanthropologie, depuis la numérisation des sites de fouilles jusqu'à l'acquisition et l'analyse des spécimens. Nous établissons des modèles numériques de terrain (analyses multidimensionnelles de données issues de différentes modalités d'acquisition telles que les scans laser et la photogrammétrie) qui permettent d'appréhender les vestiges dans leur contexte et nous proposons des analyses qui répondent à des problématiques qui sont spécifiques aux études en biologie. En particulier, nous apportons des outils d'analyse et de visualisation (cartographies 3D et analyses statistiques) pour des problématiques de déformation basées sur des recalages surfaciques. Nous proposons également une méthode d'analyse sur des données partielles afin de pouvoir exploiter l'ensemble des données disponibles dans les registres fossiles et modernes. Nos résultats mettent en évidence que les méthodes par recalage surfacique augmentent non seulement les possibilités de capter les formes et leurs variations, mais permettent également de travailler sur des formes globales et non uniquement sur certains points. Nous montrons notamment que ces méthodes permettent le développement d'outils qui sont bien adaptés pour les études en paléoanthropologie
The biological evolution of organisms can be studied as a set of morphological transformations which are characterized by the modification of their global three- dimensional geometry and by some discrete traits. In this context, it is necessary to develop comparative mathematical and computational tools for the study of the inter- and intraspecific variation. Within the three-dimensional data processing workflow (3D images or 3D meshes) employed in " virtual paleoanthropology ", the method that is most commonly used in comparative analysis is based on landmarks (most often anatomical landmarks) from which coordinates are analyzed by using mathematical tools such as " geometric morphometrics ". More recently, other methods allowing global comparisons between three-dimensional reconstructions without landmarks have been proposed. They allow for example the statistical analysis of a global shape and its variability. We suggest to study the process for morphometric analysis of 3D data commonly used in paleoanthropology, from the digitization to the exploration of 3D data. This dissertation introduces analytical methods for the processing of data provided by paleoanthropological studies, from the digitization of the excavation sites to the acquisition and the analysis of specimens. We established digital ground models (multidimensional analyses of data from various modalities of acquisition such as laser scanner and photogrammetry) that contribute to a comprehensive understanding of fossil remains in their context and we proposed relevant analyses for resolving specific problems inherent to biological studies. In particular, we developed appropriate tools for analyses and viewing (3D mappings and statistical analyses) dedicated specifically to problems of deformation-based registrations. Additionally, we introduced a method for the analysis of partial data in order to use all the specimens available in the fossil and modern records. Besides opening up new possibilities of capturing shape variation, our results highlight that techniques based on surface registration provide a reliable methodological framework for working on global shapes without focusing on specific points. We reported in particular that these methods allow the development of tools which are particularly suitable for the paleoanthropological studies

Ce travail de thèse porte sur l'intégration des structures craniofaciales au sein de la famille des hominidés. Au cours de l'évolution, une réduction du prognathisme et une diminution de la longueur de la face sont observées chez les taxons appartenant au rameau humaine. Cette réduction des structures faciales est associée à une base du crâne plus fléchie et à une mandibule plus gracile. L'objectif de ce travail est de définir le rôle joué par les changements basicrâniens et mandibulaires dans la mise en place de la face courte et droite des humains modernes. Dans ce contexte, les schémas d'intégration liant la face et les autres structures crâniennes (basicrâne et mandibule) ont été décrits et quantifiés dans ce mémoire. Ce travail a été effectué sur la base d'un échantillon de crânes incluant l'ensemble des genres d'hominidés actuels : les humains modernes, les chimpanzés, les gorilles et les orangs-outans. Les crânes ont été préalablement numérisés à l'aide d'un scanner médical. Les schémas d'intégration craniofaciaux ont alors été étudiés à l'aide d'outils statistiques et de méthodes d'analyses en morphométrie géométrique. Ce travail a permis de mettre en avant plusieurs mécanismes d'intégration craniofaciale, propres aux humains modernes. Ces schémas d'intégration spécifiques permettent d'expliquer en grande partie la mise en place de la face réduite des humains modernes. Ces résultats permettent donc d'éclairer les mécanismes d'évolution et de mise en place des structures faciales chez les hominidés et dans le rameau humain
This thesis dissertation is dedicated to the study of craniofacial structures within the hominid family. Throughout evolution, a reduction of facial prognathism and a diminution of the facial length are observed in the taxa which belong to the human lineage. This reduction of facial structures is associated to a more flexed cranial base and to a shorter mandible. The aim of this work is to define the role played by the basicranial and mandibular changes in the set up of the short and straight face of modern humans. In this context, the patterns of integration linking the face and the other cranial structures (basicranium and mandible) are decrypted and quantified in this thesis dissertation. This work has been done with a sample including all the extant hominid genera: modern humans, chimpanzees, gorillas, orang-utans. The skulls were first scanned using a medical scanner. Patterns of craniofacial integration were then studied using statistical tools and geometric morphometric analysis methods. This work underlined several mechanisms of craniofacial integration, unique to modern humans. These specific patterns of integration can explain an important part of the set up of modern humans reduced face. Thus, these results enlighten the evolution mechanisms and the set up of facial structures in hominids and in the human lineage

There is a universally recognized need for a more refined chronological understanding of the Early Holocene in the Southeast. The Paleoindian/Early Archaic period deposits at the Hester site provide an opportunity to refine the local chronology for northeast Mississippi. Three-dimensional spatial analysis of these deposits revealed the presence of at least four distinct occupations. Attribute-level morphological analysis of projectile points excavated from these occupations indicated that base width, shoulder width, and basal indentation ratio, changed significantly through time. A paradigmatic classification system employing these attributes was used to classify the projectile points from Hester. These classes were successfully seriated indicating that they are true historical classes. The range of historical classes indicates that this alternative to the traditional point typology may provide greater chronological precision for the Early Holocene in northeast Mississippi.

Durante el Pleistoceno Medio y Superior ocuparon Europa al menos tres poblaciones: los heidelbergensis, los neandertales y los humanos modernos. La economía de estas sociedades habría dependido en última instancia de los recursos vegetales y animales disponibles, que habrían variado según las oscilaciones climáticas a lo largo de los 600 Ka en que el continente fue ocupado de manera continua. La variación en la dieta implicaría un cambio en las estrategias de subsistencia que podrían haber tenido repercusiones en las diferentes adaptaciones (culturales y biológicas) y en la estructura de las sociedades. Los estudios de microestriación ofrecen la posibilidad de analizar qué tipo de alimentación habrían tenido las poblaciones estudiadas diferenciando entre dietas más blandas o más abrasivas según la cantidad de estrías que se encuentren en el esmalte dental y los índices derivados de la orientación preferente, discriminan la tendencia de los grupos distinguiendo entre dietas vegetarianas, mixtas y carnívoras. Los resultados obtenidos apuntan a una diferencia clara entre los heidelbergensis, con una dieta más abrasiva y altamente dependiente del clima y del medio, y las poblaciones paleolíticas más modernas. La dieta de los H. heidelbergensis habría sido menos refinada, incorporando partículas muy abrasivas, como por ejemplo, los vegetales. El Pleistoceno Superior, por el contrario, habría tenido un patrón más similar aunque los humanos modernos (AMH) y los neandertales son claramente diferentes, incluso cuando se considera el mismo periodo (periodo isotópico 3) e incluso en un ambiente similar consiguiendo porcentajes de clasificación correcta de más del 80%. El clima también influyó a estas poblaciones, pero en menor medida, ya que, ante un cambio de ambiente, se observa desde el Pleistoceno Medio (heidelbergensis) hasta el Paleolítico Superior Final (LUP AMH) una disminución de la distancia entre los patrones ante la variación climática de forma que cada vez ésta influyó menos en las estrategias de las poblaciones del Paleolítico. Este patrón se observa también en la variación en el tamaño dental por lo que, la respuesta al cambio ambiental en los humanos más antiguos habría sido eminentemente biológica mientras que en los más recientes, la cultura habría jugado un papel preponderante, disminuyendo la dependencia (aunque no independencia) del medio en el LUP-AMH con una importancia muy relevante de la cultura a partir del Paleolítico Superior. Estos datos concuerdan con los resultados de otras metodologías como el tamaño dental y las tasas de desgaste oclusal. Igualmente, los datos de la respuesta de cada sexo al cambio ambiental indican una variación que implicaría cambios en la estructura demográfica de las poblaciones con una importancia en las diferencias inter-sexuales diferente en cada población. La dieta de los humanos del Pleistoceno Medio y Superior habría sido por lo tanto omnívora y muy diversa pero la comparación con grupos de cazadores-recoletores actuales con dietas conocidas de diferentes medios y ambientes, indica que habría habido una tendencia hacia el aumento de la ingesta de carne y de una dieta menos abrasiva siendo los heidelbergensis los más vegetarianos y los humanos modernos los más carnívoros.
During the Middle and Upper Pleistocene at least three populations occupied Europe: heidelbergensis, Neanderthals and modern humans. The economy of these societies would have ultimately depended on plant and animal resources available, which would have varied according to climatic oscillations along the 600 Ka that the continent was continuously occupied. Microwear studies offer the possibility to analyze what kind of food the populations would have consumed differentiating between soft or more abrasive diets by the number and orientation of grooves found in the enamel The results indicate a clear difference between heidelbergensis, with a more abrasive diet and highly dependent on climate, and modern Paleolithic populations. The diet of H. heidelbergensis would have been less refined, incorporating very abrasive items, such as vegetables. The Upper Pleistocene, however, would have had a closer pattern although modern humans (AMH) and Neanderthals are clearly different, even when considering the same period (isotopic stage 3) and even in a similar environment with correct classification rates higher than 80%. The climatic variation would have also affected Upper Pleistocene groups but to a lesser extent: from the Middle Pleistocene (heidelbergensis) to the Final to the Final Upper Paleolithic (LUP AMH) a decrease of the distance between microwear patterns when climate changes is observed highlighting the importance of the culture in the more recent populations. The Middle and Upper Pleistocene human’s diet would have been diverse, mainly omnivorous but, if compared with hunter-gatherers from known diets from different environments, the results indicate a trend towards an increase in the meat percentage together with a decrease in the abrassivity being heidelbergensis the most vegetarian group and modern humans the most carnivorous.

Amb el títol L’Esquerda. Arqueologia funerària i estudi antropològic d’una població medieval (segles VIII-XIV) el present treball presenta un cas d’estudi sobre les dues àrees funeràries–extramurs i eclesial- del poblat medieval de l’Esquerda. La recerca s’obre en el front arqueotanatològic i l’arqueoantropològic a partir de les intervencions arqueològiques realitzades entre 2008 i 2012 sota una única mirada, que s’obre en panoràmica per obtenir una visió documentada i el més àmplia possible sobre els diferents punts d’observació i valorar-los en el seu context geogràfic, històric i social. L’anàlisi de les pràctiques funeràries i la seva evolució en els diferents espais i estrats descriu i s’aproxima tant a l’aplicació com a la transgressió de les normatives litúrgiques en matèria sepulcral, a la persistència de tradicions antigues, a l’acomodació de la societat a les exigències socials i espacials que afecten l’àrea funerària i a l’adaptació enfront de situacions d’emergència per conflictes epidèmics o de caràcter bèl·lic. La caracterització antropològica de les mostres esquelètiques de 181 individus obté els corresponents perfils poblacionals per a cada fase i per a cada una de les necròpolis, mitjançant la seva descripció morfològica. L’estudi aprofundeix en aquesta línia de recerca amb la contrastació estadística dels resultats i la seva valoració argumentada a nivell evolutiu i en relació a les diferents classes d’edat i per ambdós sexes. En l’anàlisi dels diferents aspectes patològics l’estudi registra tot el ventall d’anomalies degeneratives articulars i estructurals que és habitual en mostres antropològiques medievals, altres afectacions menys freqüents i un gran nombre de traumatismes, accidentals i per causes violentes, que es descriuen de manera individual i es valoren en conjunt. El resultat és un treball exhaustiu sobre un jaciment funerari paradigmàtic, el de l’Esquerda, que pretén ésser un referent per a posteriors recerques, no només per a Osona, no només per als jaciments funeraris, sinó per als diferents àmbits de coneixement del període medieval de Catalunya.
With the title L’Esquerda. Funerary Archeology and Anthropological Study of a Medieval Population (8th-14th centuries), this work presents a case of study on the two funerary areas (extramuros and ecclesiastical) of the medieval village of Esquerda. The research opens up on the archaeothanatological and archaeoanthropological front from the archaeological interventions carried out on the site between 2008 and 2012 under a single glance, which opens up in panoramic views to obtain a documented and a wide view as possible on the different points of observation, in order to value them in their geographical, historical and social context. The analysis of funerary practices and their evolution in different spaces describes and approaches both the application and the transgression of liturgical regulations in the burial subject. Also, the persistence of ancient traditions, the accommodation from society to the social and spatial demands that affect the funeral area, and the adaptation to emergency situations due to epidemic or warlike conflicts. The anthropological characterization of the skeletal samples obtains the corresponding population profiles for each phase and for each one of the necropolis, by means of their morphological description. The study explores this line of research with the statistical comparison of the results and their evaluation argued at an evolutionary level and in relation to different age classes and for both sexes. In the analysis of the different pathological aspects, the study records the whole range of joint and structural degenerative anomalies that are common in anthropological medial samples, other less frequent affections and a large number of traumatisms, accidental and violent causes, which are described individually and they are valued as a whole. The result is an exhaustive work on a paradigmatic funerary deposit which seeks to be a reference for further research, not only for Osona, not only for funerary sites, but for the different fields of knowledge of the Medieval period of Catalonia.

The Doctoral Thesis entitled "the glenohumeral joint of hominoid primates: locomotor Correlates, anatomical variation and evolution" is about the anatomical adaptations in the shoulder joint (glenohumeral joint) of hominoid primates. The action of the forces exerted during locomotion model the shape of the joint determining the range of motion animals can achieve. The hominoid primates stand out as having very mobile joints, with the ability to raise the arm above the shoulder enabling the use of suspensory locomotion, such as brachiation. Here, the morphological characters related to the types of locomotion of extant hominoid primates in the proximal humerus and the glenoid cavity of the scapula were identified. These morphological characters were used to identify locomotor capacities in extinct hominoid taxa and basal Miocene catarrhines, as well as hominins (Plio-Pleistocene human ancestors). Several taxa of basal Miocene catarrhines might have acquired suspensor locomotion without first acquiring an upright body plan (orthogrady), and Plio-Pleistocene hominins showed affinities with extant hominoids but also generalized taxa such as the woolly monkeys (New World primate). Thus hominins Plio-Plesitocene mezcaldas presented (mosaic) morphological affinities, thus supporting the notion that the common ancestor between humans and chimpanzees, as well as many of the ancestral nodes in the hominoid lineage, presented fairly generalized arboreal features, instead of a derived morphology as found in African great apes, in turn related to the use of knuckle-walking locomotion. Thus, knuckle-walking may be a result of convergent evolution gorillas and chimpanzees. Importantly, the PhD provided evidence of decoupling between orthograde characters (upright trunk) and suspensory locomotion. The consequences of this finding regarding the evolutionary history of hominoids is that orthogrady and suspension have been independently acquired by a number hominoid taxa, and is at least result of evolutionary convergence between hylobatids and great apes.
La Tesis Doctoral con título "the glenohumeral joint of hominoid primates: locomotor correlates, anatomical variation and evolution" trata sobre las adaptaciones anatómicas en la articulación del hombro (articulación glenohumeral) de los primates hominoideos. La acción de las fuerzas ejercidas durante la locomoción modelan la forma de la articulación, determinado el rango de movimientos que los animales pueden alcanzar. Los primates hominoideos destacan por tener articulaciones muy móviles, con la capacidad de elevar el brazo por encima del hombro para desplazarse con usando tipos de locomoción suspensora extrema como a braquiación. En esta tesis, se identificaron los caracteres morfológicos relacionados con los tipos de locomoción de los primates hominoideos actuales en el húmero proximal y la cavidad glenoides de la escápula. Estos caracteres morfológicos se usaron para identificar capacidades locomotoras en taxones hominoideos i catarrinos basales extintos del Mioceno así como homininos (ancestros de los humanos del Plio-Pleistoceno). Varios taxones de catarrinos basales del Mioceno podrían haber adquirido locomoción suspensoria sin adquirir primero un plano corporal erecto (ortogradía), y los homininos del Plio-Pleistoceno mostraron afinidades con los gropus de hominoideos actuales pero también con taxones generalizados (morfológicamente y de uso de locomoción) como la mona lanuda platirrina (primate del nuevo mundo). Así, los homininos del Plio-Plesitoceno presentaron afinidades morfológicas mezcaldas (mosaico), respaldando así la noción de que el ancestro común entre humanos y chimpanzés, así como muchos de los nodos ancestrales en el linage hominoideo, pudo presentar unas caracteristicas arborícolas bastante generalizadas, en vez de una morfología derivada como la que se encuentra en los grandes simios africanos, a su vez relacionada con el uso del tipo de locomoción knuckle-walking. Así, este tipo de locomoción puede ser resultado de evolución convergente en estos dos taxones (gorilas y chimpancés). Otro resultado destacable es la evidencia de de disociación entre caracteres ortógrados (tronco erecto) y el tipo de locomoción suspensora, ya que en esta tesis se presentan evidencias de primates no ortógrados actuales (mona lanuda) y catarrinos no hominoideos extintos que presentan adaptaciones a la suspensión sin presentar un cambio de patrón corporal de (pronógrado a ortógrado). Las consecuencias de este hallazgo respecto a la historia evolutiva del los hominoideos es que, junto con las evidencias de la tendéncia contrária (presencia de caracteres ortógrados sin adaptaciones a la suspensión) implican que la ortogradia y la suspensión pudieron haber sido adquiridas independientemente por un número de taxones hominoideos, siendo al menos resultado de convergencia evolutiva entre hilobátidos y grandes simios. A su vez, el hecho de que taxones de tamaño pequeño como los pliopitécidos o la mona lanuda puedan hacer uso de la locomoción suspensora sin exhibir a la vez caracteres ortógrados da pie a pensar que los ancestros de los hilobátidos puedieron haber sido primates de pronógrados de tamaño pequeño, no siendo el tamaño corporal pequeño de los hilóbatidos resultado de enanismo des de un ancestro común ortógrado y de tamaño grande.

The science of human origins, known formally as ‘paleoanthropology’, was effectively born in the fierce late nineteenth century debate as to the human status of Neanderthal. Critical social theory on ‘scientism’ has generated a wealth of research on the ways the various human sciences contribute to the structuring and organizing of social relations. This includes Foucault’s well-known genealogical studies of clinical medicine, which have provided sociologists with crucial insight into how classifying and ordering practices actually create ‘Man’ in the way they operate as a field – or “technology” – of power (Foucault 1970). However, as yet there has been very little produced by sociologists interested in the impacts of science on society with regards to paleoanthropology specifically. This is especially surprising considering that Neanderthal, the quintessential ‘missing link’ and the hub of paleoanthropology’s speculative and explanatory universe, clearly occupies a central place in the socio-historical emergence of ‘humanness’ as an ontological category. Moving forward from the basic observation that the original 1856 discovery of fossilized Neanderthal remains in a cave in Germany’s Neander Valley generally coincided with the end of the colonial period, my dissertation seeks to fill a void in sociology via a genealogical study of paleoanthropological science. Drawing largely upon the insight of Foucault but also that of Saïd, I undertake a discourse analysis of the early debates surrounding Neanderthal with an aim toward shedding light upon the ways in which Neanderthal propagated or concealed certain anxieties, particularly as they relate to biological kinships. This is then applied to an exploration of how the debates surrounding Neanderthal were in turn pivotal to the emergence of today’s prevailing paleoanthropological models of human origins. The profound ontological and epistemological tensions embodied by these models, I argue, wholly reflect the inherently ambiguous nature of the missing link as both concept and metaphor. The result is that missing links, because of the discursive field in which they function, are a powerful source of normativity and stratification.
Thesis (Ph.D, Sociology) -- Queen's University, 2014-02-04 12:53:52.631

This thesis applies a single test method to a high number of proposed explanations (models) of hominin bipedalism. This has not been undertaken previously, due largely to the very variable ways in which the models have been built, and the disparity of their core ideas. Taken as a whole, however, the modelling literature contains a gap, and the thesis' main claim is that it can be exploited to assess relative levels of tenability, using non-statistics-based hypothesis testing. Infants are defined here as individuals too young to survive without caregivers and older infants are defined as those who are not completely dependent on others for nutrition and transport. Most models have omitted infants, and others have included them only in minor or circumscribed ways. A species' survival, however, depends on sufficient numbers of young reaching breeding age, and species which provide postnatal care need to provide adequate care Any viable model of how hominin bipedalism evolved ought, therefore, to be able to accommodate appropriate levels of infant survival and adequate caregiving, and factoring infants and caregivers into the models can thus serve as a test method. The method cannot assess overall robusticity, because models which accommodate infants and caregivers might be untenable relative to other criteria, but it can narrow down the possible ways in which bipedalism might have evolved by identifying models which are untenable in one crucial way. Seventy-six models, written between 1871 and 2017, are identified. They are classified into 12 model types, three of which are treated as obsolete. Seventy-one models are tested, 76.2% of which omit infants or include them only trivially. The models are tested either in blocs, according to type or variant, or individually. Forty-three tests are conducted. In order to factor a proto-biped infant into a model, it is necessary to have a clear idea of its likely basic characters. After its Introduction, Methods chapter and Literature Review, this thesis is therefore divided into two distinct parts. The first part extensively reviews available literature bases for character reconstruction, and one of these bases, the literature on nonhuman anthropoid infancy, is supplemented by a short-term captive study of clinging behaviour in an orthograde ape, presented as an appendix. The likely size of older proto-biped infants, relative to conspecific adult females, is then estimated, based on data for extant anthropoids. All this information is then synthesized to reconstruct the infant and to guide the tests' expectations of infant/adult differences and adequate caregiving behaviours. The second part of the thesis contains the 43 tests. The results show that 40.8% of models are untenable, including two whole model types, 43.7% are highly questionable, and only 11.3% accommodate infant survival well (4.2% did not support the test method). These results indicate that the ways in which bipedalism might have evolved are heavily constrained by the needs of infants. They also suggest that future models which consider the full implications of their proposed behaviours for infants and caregivers will have greater explanatory value than the majority of existing models.

The years between the two world wars, which just preceded the emergence of the neo-Darwinian "new synthesis," were intellectually difficult ones for paleoanthropology in America. Patterns of thought deeply ingrained in biology and anthropology pushed writers on hominid evolution into interpretive "blind alleys." Most prominent among the patterns was what Ernst Mayr has called "typological thinking," which often mixed with a tendency to project "scientific" racism back into the hominid past. A "splitting" habit in taxonomy combined with these and with belief in "orthogenetic" change to make polyphyletism the norm. Hesitance to accept as human ancestors any Pleistocene forms exhibiting "primitive" characters led to phylogenies which put the known fossils on side-branches. Anatomically modern humans were thus left "ancestorless" by most writers, though nearly all continued to use existing fossils in their evolutionary scenarios by designating them as "structural ancestors." Research conducted in Europe before 1914 on the Neanderthal skeleton and on the interperetation of endocranial casts, along with the Piltdown fraud, did much to establish these phylogenies and scenarios. In tandem with these general themes came the ascendancy of several specific hypotheses that eventually clashed with accumulating evidence. That the brain had led the way in hominid evolution, that Neanderthals and other "low-brows" could be ruled out as ancestors, and that modern Homo sapiens had appeared early in the Pleistocene, became even harder to maintain. The close evolutionary bond between humans and great apes theorized in England by Sir Arthur Keith and elaborated in America by William King Gregory remained vigorous, however, despite challenge. The present study examines these issues through an analysis of the five Americans whose writings on hominid evolution were most extensive and varied--Henry Fairfield Osborn, George Grant MacCurdy, Ales Hrdlicka, Earnest A. Hooton and William K. Gregory. The writings of each are analyzed separately, so that both general themes and responses to the changing state of the discipline can be traced. This approach reveals that shared patterns of thought did not prevent considerable diversity on nearly every main issue, a fact which rendered the field fertile for rapid growth later.

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.

Recently, the number of hominin genera has increased dramatically. Prior to the announcement of Ardipithecus, only two genera were used by paleoanthropologists: Australopithecus and Homo. Presently, up to eight hominin genera are used: Sahelanthropus, Orrorin, Ardipithecus, Australopithecus, Praeanthropus, Kenyanthropus, Paranthropus and Homo. Unlike species concepts, the genus category has not received wide critical examination. To investigate the use of the genus category in paleoanthropology, a comparative framework drawing on morphometric data from a large number of catarrhines is developed. Cranial variables include 36 standard linear measurements from representatives of catarrhine genera across the major tribes/families. This study seeks to assess whether too few or too many hominin genera have been recognized compared with extant catarrhines. Moreover, two published hypotheses about the use of Homo are examined: 1) Wood & Collard's (1999) proposal to transfer Homo habilis/rudolfensis to Australopithecus; and 2) Goodman et al's (1998) classification of both humans and chimpanzees in Homo. To analyze these cranial variables and a number of shape indices calculated from them, as well as to assess competing hypotheses, univariate, bivariate and multivariate statistical approaches are used. The results allow the identification of a set of variables and shape indices which distinguish genera across the catarrhines. Importantly, body size seems to be the major separator of catarrhine genera, reinforcing the idea that they occupy discrete adaptive zones. Moreover, differences between these genera mostly represent contrasts in the size of the neuroversus the viscerocranium. When applied to hominins, a picture emerges which distinguishes them from extant catarrhines: cranial shape rather than size is the major component distinguishing them; this suggests that extinct hominins occupied similar habitats and adaptive zones; variability in size and shape within hominin genera is much lower than extant catarrhines; and the major differences seen in shape among hominins are the result of encephalization in Homo. It is concluded here that both Wood & Collard's (1999) and Goodman et al.'s (1998) proposals appear to be premature. Moreover, while the earliest hominins may be too finely split at the genus level, the evidence for distinction of Australopithecus and Paranthropus is solid.

abstract: This dissertation is an historical analysis of the science of human origins, paleoanthropology, examining the intersection of science and culture around fossil human ancestors (hominins) over the last century and a half. Focusing on fossils as scientific objects, this work examines three controversial fossils from the science’s history asking, how do fossils formulate, challenge, and reconfigure notions of what it means to be human? The introduction reviews the historiography of paleoanthropology and the gaps that exist in the literature. Chapter two examines the first case study, the type specimen of Homo neanderthalensis, known as the Feldhofer Neanderthal, providing a biography of the object from its discovery in Germany in 1856 until its species designation in 1864. Chapter three briefly links the Neanderthal’s story in time and space to the next fossil’s story. Chapter four picks up the story of paleoanthropology in 1924 in South Africa, with the discovery and initial analysis of a specimen nicknamed the Taungs Baby, which was labeled a new hominin species, Australopithecus africanus. Chapter five is another brief chapter connecting the Taungs Baby story in time and space to the final specimen examined in this work at the end of the century. Chapter six examines the final case study, a specimen discovered in 2003 in Indonesia, designated a new species named Homo floresiensis and nicknamed the Hobbit. Through comparing contrasting, and connecting the stories of these three specimens, three major conclusions emerge about the field. First, the fossils themselves play an important role in knowledge production about the hominin past. Second, scientific practice shaped both interpretations of fossils and larger questions of what it means to be human. Third, the scientific practice is itself shaped by local culture, which continually interacts with attempts to establish a global perspective about the human past. The perspective gleaned through the eyes of these three fossils therefore reveals the way shifting, rather than eternally true, claims are embedded in culture and intertwined with the perspectives of the humans conducting the science.
Dissertation/Thesis
Doctoral Dissertation Human and Social Dimensions of Science and Technology 2020

One method of investigating the palaeoecology of a site is to analyse the associated faunal remains. In palaeoanthropology faunal analysis is not uncommon, but on most occasions only a fraction of the overall fauna is analysed, often only large mammals. This discrepancy is addressed in this thesis. Here, the molluscan fauna from a range of sites is analysed from an ecological perspective, and the results compared to the palaeoecological results based on other faunal data, including, when available, small mammals, birds, reptiles, fish and amphibians. The aim of this study is to investigate whether the molluscs add anything to the overall palaeoecological picture of a site. The data are analysed from a palaeoanthropological perspective, with different hominine groups of sites compared from a palaeoecological perspective, based on the molluscan data. At the same time the value of molluscs as palaeoecological indicators is measured against data from other sources, such as other faunal data. The results show that molluscs are capable of adding important new and corroborating data that could be used to test established palaeoecological reconstructions of the past. The data support the idea that open habitats may have played an important role in the evolution of the genus Homo, but they also strongly suggest the presence of permanent water, whether in open or closed habitats.

The subject of this doctoral dissertation concerns the usefulness of virtual analysis approach in studies of dental tissues and their defects. The real symmetry and perfect balance between opposite jaw halves and antagonistic teeth is not the reality in the masticatory system. The sample of Yuendumu Aboriginal people used in Paper I consists of complete maxillary and mandibular dental arch 3D models from 19 individuals (young and adult). The analysis was carried out on first molars from all quadrants and only individuals with similar levels of wear were selected (76 teeth in total). Virtual methods were applied in order to inspect the palatal arch asymmetries in relation to the alveolar bones inclination and consequently, the alteration of dental crown. Crown alteration (Enamel and Dentine) could be produced by several factors such as masticatory activity, pathological, anthropic and cultural alterations. However, anthropological studies are sometimes not able to distinguish differences among them without an interdisciplinary approach. Particularly, pathological alterations (pits and fissures) are extremely difficult to interpret without microscopy analysis. That is the reason why a virtual approach in dental studies is useful to understand and distinguish any natural or anthropic alteration. Two examples are shown in Paper II and Paper III where two dental treatments discovered in two Italian sites have been described: Villabruna specimen and Fredian shelters. The modern human specimen Villabruna (Paper II) from a burial in Northern Italy is the earliest evidence of dental therapeutic intervention on a Late Upper Paleolithic (ca. 14,000 yr BP). Using Scanning Electron Microscopy (SEM) we showed the presence of striations deriving from the manipulation of a large occlusal carious cavity of the lower right third molar. The striations have a “V’’-shaped transverse section and several parallel micro-scratches at their base, as typically displayed by cutmarks on teeth. Based on in vitro experimental replication and a complete functional reconstruction of the Villabruna dental arches, we confirmed that the identified striations and the associated extensive enamel chipping on the mesial wall of the cavity were produced ante-mortem by pointed flint tools during scratching and chiseling activities. Similar situation was identified at the Fredian shelter (Paper III). The two upper incisors display exposed pulp chambers with circumferential enlargement, chipped dentine on the cavity margins and striations on the cavity walls. Histochemical analysis of the material embedded at the bottom of the cavities revealed a conglomerate of vegetal fibers and probably hairs. Moreover, Fourier transform infrared spectroscopy (FTIR), energy dispersion X-ray spectroscopy (EDS), and Raman microscopy analysis of black residue adhering to the walls of both cavities is consistent with organic substances, specifically bitumen. A direct chronometric date for Fredian 5 confirms a Late Upper Paleolithic context (between 13,000-12,735 calendar years ago). Overall, our results are consistent with in vivo dental drilling to remove necrotic or infected pulp tissue (pulpitis) and the subsequent use of a composite, organic dental filling in the cavity. The conservation of human remains is the main interest of good anthropological research and any organic matter identified within a dental cavity needs to be conserved. This is the topic of paper IV: Letter to the Editor I had written in order to clarify the reason why we have decided to use three different methods (infrared spectroscopy (FTIR), energy dispersion X-ray spectroscopy (EDS), and Raman microscopy analysis) rather than gas-chromatography. Finally, when a tooth does not show any pathological alteration or exhibits a normal/slight wear pattern on the occlusal surface (wear stage 1-2, 3 Smith, 1974) it is possible to obtain information regarding taxonomy based on volume (3D) or areas (2D) of Enamel, Dentine (paper V, VI, VII. VIII) and morphometric analysis of teeth's root (paper IX). This thesis therefore strives to provide a contribution to understanding how virtual approach to dental studies can be used to increase the knowledge of dental tissues and their defects.

Sterkfontein West Pit, dated to 1.7-1.9 Ma, has yielded a number of fossils that are difficult to classify, such as StW 252, which comprises cranial bone fragments, and a full set of robust maxillary anterior and posterior teeth. The purpose of this study is to ascertain whether dental measurements of StW 252 more closely align with those of Australopithecus africanus, Australopithecus robustus, Homo sapiens or the African apes. For M1 and M2, StW 252 is distinct from the comparative samples, and is particularly large buccolingually for M1 and mesiodistally for M2, partly resembling the dimensions of A. africanus, whereas for M3, StW 252 is mesiodistally and buccolingually large. Canonical scores axes show StW 252 as extreme in terms of size and polarized from A. robustus in terms of shape. StW 252 is distinct from both A. africanus and A. robustus, indicating an additional hominin taxon may be represented at Sterkfontein West Pit.

abstract: This research uses Peircean Semiotics to model the evolution of symbolic behavior in the human lineage and the potential material correlates of this evolutionary process in the archaeological record. The semiotic model states the capacity for symbolic behavior developed in two distinct stages. Emergent capacities are characterized by the sporadic use of non-symbolic and symbolic material culture that affects information exchange between individuals. Symbolic exchange will be rare. Mobilized capacities are defined by the constant use of non-symbolic and symbolic objects that affect both interpersonal and group-level information exchange. Symbolic behavior will be obligatory and widespread. The model was tested against the published archaeological record dating from ~200,000 years ago to the Pleistocene/Holocene boundary in three sub-regions of Africa and Eurasia. A number of Exploratory and Confirmatory Data Analysis techniques were used to identify patterning in artifacts through time consistent with model predictions. The results indicate Emergent symboling capacities were expressed as early as ~100,000 years ago in Southern Africa and the Levant. However, capacities do not appear fully Mobilized in these regions until ~17,000 years ago. Emergent symboling is not evident in the European record until ~42,000 years ago, but develops rapidly. The results also indicate both Anatomically Modern Humans and Neanderthals had the capacity for symbolic behavior, but expressed those capacities differently. Moreover, interactions between the two populations did not select for symbolic expression, nor did periodic aggregation within groups. The analysis ultimately situates the capacity for symbolic behavior in increased engagement with materiality and the ability to recognize material objects can be made meaningful– an ability that must have been shared with Anatomically Modern Humans’ and Neanderthals’ most recent common ancestor. Consequently, the results have significant implications for notions of ‘modernity’ and human uniqueness that drive human origins research. This work pioneers deductive approaches to cognitive evolution, and both strengths and weaknesses are discussed. In offering notable results and best practices, it effectively operationalizes the semiotic model as a viable analytical method for human origins research.
Dissertation/Thesis
Appendices A-N: Spreadsheets
Doctoral Dissertation Anthropology 2016

There is variation in Neandertal permanent dentition in both the size and shape of first molars. Ecogeography as well as chronology would be expected to account for at least some of the variation observed. Occlusal outlines of maxillary and mandibular first molar casts from European Neandertals, (Spy 1, Scladina 4A-4, Engis 2, l’Hortus 2, 4, 5, and 8, La Quina H5, Malarnaud 1) were generated through photostereomicroscopy and non-landmark smooth tracing methods, and occlusal areas, buccolingual breadths and mesiolingual lengths were measured by calibrated Motic 3.0MP microscope cameras. Principal component (PC) scores of elliptical Fourier harmonic descriptors were calculated using SHAPE v1.3, yielding a total of 76 and 67 PC scores for mandibular and maxillary data respectively. Of the maxillary outlines analyzed, a strong correlation exists between PC1 (58.4% of variance) and occlusal area, explaining size influence. On PC1, the smallest (Engis 2), is followed by Scladina 4A-A, Hortus 8, La Quina 5 and Spy 1. On PC2 (23% of variance), Scladina 4A-A is an outlier. On PC3 (12.6%), Hortus 8 is separated from the others whereas Scladina 4A-A is difficult to classify. PC4 accounts for 5.8% of the variance and separates Spy 1 from La Quina 5. Spy 1 appears as distinct on PC1, PC3, and PC4 while Scladina 4A-A is relatively distinct on all axes. In a cluster analysis of PC scores Spy 1 and La Quina 5 are linked by the shortest distance and joined secondarily to Hortus 8, whereas Scladina 4A-A and Engis 2 are relatively distinct from the others. Mandibular results indicate that Malarnaud is distinct on PC1 (40% of variance) and again on PC2 (27%), while La Quina 5 appears as slightly distinct on PC3 (14%), though grouped with Engis 2, while Hortus 2 and 4 group together this axis as well as PC1. PC4 (12%) again separates Malarnaud, and presents Hortus 5 as an outlier. Overall, chronology correlates better than geography to the variance observed in occlusal first molar shape in these European Neandertals, with some inconsistencies most likely due to individual biological variability. This study demonstrates a new method to compute the elliptical Fourier descriptors of molar occlusal outlines, and applies these to explain variation in these Neandertals with respect to ecogeographic and chronological situation.

Previous research on the biomechanics of tool use has focused heavily on traits correlated with locomotion, tool manufacturing, and habitual tool use. Features like the breadth of the metacarpals, relative length of the thumb, styloid process of the third metacarpal, and the breadth of the apical tufts are skeletal features associated with the use and development of stone tools. However, there are many traits of the distal forelimb that may also be correlated directly with the development and use of tools. The purpose of this research is to analyze morphological features of the hands and compare them to features of the arm in humans, fossil Homo and the great apes to understand how the hominin distal arm functions as a mosaic in response to the use of stone tools. The results indicate a separation between tool-users and non-tool users when all distal forelimb dimensions are examined. Omo 40-19 falls closer to non-tool users when univariate plots of ulna length and breadth are examined. Ratios of hand measurements to radius length are better at polarizing the tool-users from non-tool users than are hand dimensions to ulna length ratios. These results highlight the role of the radius in stabilizing the hand during stone tool production.

Humans are more carnivorous than other hominoids. It has been hypothesized that, during the evolution of this increased carnivory, hominins transitioned through a scavenging niche made viable by certain carnivoran taxa (especially sabertooths) that may have lacked the morphology necessary to fully utilize all parts of carcasses (e.g., marrow), therefore leaving an open niche in the form of high-quality scavengable remains available for hominins. In this dissertation, I examine the postcanine dentition of modern carnivorans, using quantifications of occlusal radii of curvature and intercuspid notches, and study the correlation of this morphology with carcass-processing behavior. I use these correlations to deduce the carcass-processing capabilities of the Plio-Pleistocene carnivores of South Africa (a guild for which we have a good appreciation of taxonomic diversity, and that existed at an important time during the evolution of our lineage - possibly the time that we transitioned into that guild), and compare these results with those of previous studies that relied on more conventional morphological measures.

Both radius of curvature and intercuspid notch data do a good job of separating taxa by dietary category, revealing subtle patterns including possible differences in the carcass-processing abilities of fossil and modern members of some extant species. Other strong trends confirm that the "hunting-hyena," Chasmaporthetes, was probably a hypercarnivore, and not a durophage like its modern confamilial taxa. Somewhat surprisingly, results do not support the hypothesis that sabertooth felids were more hypercarnivorous than modern felids. Furthermore, though the sympatric hypercarnivorous taxa were more numerous, so to were the durophageous taxa, with one taxon, Pachycrocuta, probably exceeding the durophageous capabilities of modern durophages.

As such, this dissertation shows no evidence that members of the paleo-carnivore guild were capable of producing higher quality scavengable carcasses than are modern carnivorans, and thus, based on these analyses of fossil carnivorans, it does not appear that high-quality scavengable remains were more available in the Plio-Pleistocene than there are today. Therefore, though there is clear evidence from other sources that hominins did scavenge at least occasionally, this dissertation does not support the hypothesis that there was an open niche consisting of high-quality scavengable remains.

Dissertation

Aim of this work is to show the evolutionary connection between the chewing movements and the development of species-specific traits useful for speech in Homo Sapiens. This study will demonstrate how motor control of masticatory system affects the whole body homeostasis and its interactions are reciprocal for the production of human articulated language. In this perspective we consider speech as the product of a slow evolution of structures previously used for primary purposes and then adapted to new purely human functions ( refunctionalization ).

Dissertação de Mestrado em Evolução e Biologia Humanas apresentada à Faculdade de Ciências e Tecnologia
Do antigo alto fortificado “O Caramito”, ressaltam várias necrópoles diacrónicas, espalhadas no cabeço conhecido hoje por Alto do Calvário (Miranda do Corvo).Em 2011/12 foram feitas duas sondagens de diagnóstico: SA – junto à Torre Sineira e SB – junto à Cisterna, as duas únicas estruturas visíveis do castelo. Na SA foi possível identificar um Ossário, bem como 22 sepulturas rupestres e 3 em covacho, da Idade Média. Na SB foram identificadas 7 sepulturas rupestres datadas da Época Moderna. Das sepulturas e sedimentos associados recuperaram-se inumações primárias, reduções, ossários e ossos dispersos que correspondem a um número mínimo de 58 indivíduos da SA e 11 da SB. O material ósseo encontra-se em bom estado de preservação, com a SA a apresentar um índice de conservação anatómico (ICA) de 48%, e a SB de 58%. 63,8% (37/58) da amostra da SA é constituída por não adultos, dos quais 80% (n=33) revelam uma idade à morte inferior a 12 anos. Todos os não adultos recuperados da SB são da fase fetal, representando 27,3% (3/11) da amostra. Estes dados, sugerem que particularmente a área da SA, pode ter sido originalmente um local dedicado à inumação de crianças. A amostra de adultos da SA é constituída por 9 indivíduos femininos e 9 masculinos, e 5 femininos e 3 masculinos na SB. No âmbito do estudo morfológico, as duas amostras revelam dados semelhantes, incluindo uma elevada frequência de ossículos cranianos supranumerários e baixo dimorfismo sexual da estatura. Considerando a análise paleopatológica, a frequência mais elevada e de lesões mais severas de marcadores de stress ocupacional foram registradas para a amostra recuperada da SA, e na SB, maior e mais severa frequência dos vários parâmetros de patologia oral investigados. Estas duas condições podem indicar status social diferente entre os indivíduos das duas sondagens. Outros tipos de patologia observados nas duas subamostras incluem infeciosa, traumática e congénita, sendo de salientar a elevada frequência de lesões remodeladas, refletindo a superação das maleitas. No estudo paleoparasitológico identificou-se ainda os parasitas Giardia lamblia e Cryptosporidium spp., geralmente presente em indivíduos imuno-suprimidos.
From the ancient fortified hill, the “Caramito”, we can distinguish several and diachronic necropolis, spread in the mount known today as “Alto do Calvário”.In 2011/12 we dug two diagnostic polls: SA – next to the Bell Tower; SB – next to the medieval cistern: the only two remaining structures of the ancient castle. In SA we identified an Ossuary, as well as 22 stone-graves and 3 in the soil, from the Middle Ages. In SB we identified 7 stone graves from the Modern Ages. From the graves and the associated soil, we recovered primary inhumations, reductions, ossuary, and random loose bones, corresponding to a minimum of 58 individuals from the SA and 11 from the SB. This material was found in a good state of preservation, with 48% of anatomic conservation index (ICA), from SA, and 58%, from SB. 63,8% (37/58) of the sample from SA were non-adult individuals, 80% (n=33) of them with an age at death under 12 years. All the non-adults recovered from SB were in the fetus phase and represent 27,3% (3/11) of this sample. This data may suggest that the area of SA was initially used as a dedicated place for the burial of children. The sample of adults of SA is composed of 9 females and 9 males, while SB is composed of 5 females and 3 males. The morphological study revealed similar data, for the two areas: similar frequency of discreet traits, including high supernumerary skull bones, and low sexual dimorphism on stature. In the paleopathology study SA had the highest and more severe frequency of occupational stress markers, and SB had the highest frequency from the several aspects of the studied oral pathologies. This data leads us to the conclusion that the two samples probably had different socioeconomic status. Other pathological studies in the two samples, with different etiology, include infectious, traumatic, and congenital disease, noting a high frequency of remodeled lesions, reflecting an overcoming of this disease. Finally, the paleoparasitological study was able to collect data about infections from Giardia lamblia e Cryptosporidium spp. parasites.

Dans le cadre de ce mémoire, les relations entre morphologie, locomotion et croissance chez les hominoïdes sont analysées sous l'angle des proportions métaphysaires et de leur acquisition. Plusieurs niveaux d'analyse — intermembre, supérieur et inférieur — sont abordés dans une perspective ontogénique. La masse corporelle et la direction des charges influencent la morphologie des surfaces articulaires et métaphysaires mais aussi leur développement. Les charges étant dépendantes du mode locomoteur et celui-ci se modifiant en fonction de l'âge, on tente de voir à quel(s) moment(s) les changements proportionnels ont lieu et pourquoi ils apparaissent. Des mesures linéaires ont été recueillies sur l'humérus, le radius, le fémur et le tibia sur un échantillon squelettique des espèces H. sapiens, P. troglodytes, G. gorilla et P. pygmaeus. À partir de ces mesures et du calcul de certains ratios, des comparaisons intra et interspécifiques ont été réalisées. Les différences les plus significatives entre les espèces se dévoilent au niveau intermembre et sont relatives aux différents pourcentages d'utilisation des membres supérieurs ou inférieurs. Au sein des espèces, les résultats révèlent une similarité dans les réactions des surfaces métaphysaires au niveau intermembre, supérieur et inférieur. Les changements proportionnels ont lieu entre les stades 0 et 1 pour H. sapiens (première marché indépendante), entre les stades 2 et 4 pour P. troglodytes (majorité du poids corporel soutenue par les membres inférieurs) et entre les stades 3 et 5 pour G. gorilla (taille adulte et quadrupédie très majoritaire). Pour P. pygmaeus aucun stade en particulier n'a été ciblé par les analyses et cela concorde avec l'homogénéité de ses modes de locomotion employés au cours de la vie. Les différences proportionnelles répondent à des changements locomoteurs majeurs. Australopithecus afarensis est intermédiaire entre H. sapiens et les grands singes pour de nombreuses comparaisons. Au niveau du genou, les plus jeunes individus A. afarensis ne montrent pas de morphologie bipède, similaire aux humains.
This thesis analyses the relationship between morphology, locomotion and growth in hominoids by studying metaphyseal proportions and development. Several levels of analysis — interlimb, upper and lower limbs — are discussed in an ontogenic perspective. Body mass and direction of loads affect the morphology of articular and metaphyseal surfaces but also their development. Taking into account the locomotion of a species and related loads during growth, we try to determine when proportions change, if at all, and why they appear. Australopithecus afarensis is one species for which the debate about its locomotion is still ongoing, study of the ontogeny of its proportions may shed light on the functions of its limbs during locomotion. Linear measurements were collected on the humerus, radius, ulna, femur and tibia of H. sapiens, P. troglodytes, G. gorilla and P. pygmaeus. From these measurements, ratios have been calculated to intra and inter limb proportions of hominoid appendicular skeleton for different age groups. Differences between species are most significant at the interlimb level and relative to the different relative percentage of upper and lower limbs use. Within species, results reveal a similarity for metaphyseal surfaces responses to loads at all levels of analyses. Proportional changes take place between dental stages 0 and 1 for H. sapiens (acquisition of bipedality), between stages 2 and 4 for P. troglodytes (majority of body weight supported by the lower limb) and between stages 3 and 5 for G. gorilla (knuckle-walking for 85 % of the time). For P. pygmaeus, no proportional change occur at any specific stage, which corresponds to the absence of changes in locomotor behavior from birth to adulthood in that species. From these data, it appears that proportional differences are responses to major changes in the mode of locomotion. Australopithecus afarensis is intermediate between H. sapiens and apes for many proportional comparisons while the knee joint, contrarily to expectation, is not like the bipedal humans.