An inclusive anatomical network analysis of human craniocerebral topology.

The human brain's complex morphology is spatially constrained by numerous intrinsic and extrinsic physical interactions. Spatial constraints help to identify the source of morphological variability and can be investigated by employing anatomical network analysis. Here, a model of human craniocerebral topology is presented, based on the bony elements of the skull at birth and a previously designed model of the brain. The goal was to investigate the topological components fundamental to the craniocerebral geometric balance, to identify underlying phenotypic patterns of spatial arrangement, and to understand how these patterns might have influenced the evolution of human brain morphology. Analysis of the craniocerebral network model revealed that the combined structure of the body and lesser wings of the sphenoid bone, the parahippocampal gyrus, and the parietal and ethmoid bones are susceptible to sustain and apply major spatial constraints that are likely to limit or channel their morphological evolution. The results also showcase a high level of global integration and efficient diffusion of biomechanical forces across the craniocerebral system, a fundamental aspect of morphological variability in terms of plasticity. Finally, community detection in the craniocerebral system highlights the concurrence of a longitudinal and a vertical modular partition. The former reflects the distinct morphogenetic environments of the three endocranial fossae, while the latter corresponds to those of the basicranium and calvaria.

What do brain endocasts tell us? A comparative analysis of the accuracy of sulcal identification by experts and perspectives in palaeoanthropology.

Palaeoneurology is a complex field as the object of study, the brain, does not fossilize. Studies rely therefore on the (brain) endocranial cast (often named endocast), the only available and reliable proxy for brain shape, size and details of surface. However, researchers debate whether or not specific marks found on endocasts correspond reliably to particular sulci and/or gyri of the brain that were imprinted in the braincase. The aim of this study is to measure the accuracy of sulcal identification through an experiment that reproduces the conditions that palaeoneurologists face when working with hominin endocasts. We asked 14 experts to manually identify well-known foldings in a proxy endocast that was obtained from an MRI of an actual in vivo Homo sapiens head. We observe clear differences in the results when comparing the non-corrected labels (the original labels proposed by each expert) with the corrected labels. This result illustrates that trying to reconstruct a sulcus following the very general known shape/position in the literature or from a mean specimen may induce a bias when looking at an endocast and trying to follow the marks observed there. We also observe that the identification of sulci appears to be better in the lower part of the endocast compared to the upper part. The results concerning specific anatomical traits have implications for highly debated topics in palaeoanthropology. Endocranial description of fossil specimens should in the future consider the variation in position and shape of sulci in addition to using models of mean brain shape. Moreover, it is clear from this study that researchers can perceive sulcal imprints with reasonably high accuracy, but their correct identification and labelling remains a challenge, particularly when dealing with extinct species for which we lack direct knowledge of the brain.

A comprehensive anatomical network analysis of human brain topology.

A network approach to the macroscopic anatomy of the human brain can be used to model physical interactions among regions in order to study their topological properties, as well as the topological properties of the overall system. Here, a comprehensive model of human brain topology is presented, based on traditional macroanatomical divisions of the whole brain, which includes its subcortical regions. The aim was to localise anatomical elements that are essential for the geometric balance of the brain, as to identify underlying phenotypic patterns of spatial arrangement and understand how these patterns may influence brain morphology in ontogeny and phylogeny. The model revealed that the parahippocampal gyrus, the anterior lobe of the cerebellum and the ventral portion of the midbrain are subjected to major topological constraints that are likely to limit or channel their morphological evolution. The present model suggests that the brain can be divided into a superior and an inferior morphological block, linked with extrinsic topological constraints imposed by the surrounding braincase. This information should be considered duly both in ontogenetic and phylogenetic studies of primate neuroanatomy.

The brain of Homo habilis: Three decades of paleoneurology.

In 1987, Phillip Tobias published a comprehensive anatomical analysis of the endocasts attributed to Homo habilis, discussing issues dealing with brain size, sulcal patterns, and vascular traces. He suggested that the neuroanatomy of this species evidenced a clear change toward many cerebral traits associated with our genus, mostly when concerning the morphology of the frontal and parietal cortex. After more than 30 years, the fossil record associated with this taxon has not grown that much, but we have much more information on cranial and brain biology, and we are using a larger array of digital methods to investigate the paleoneurological variation observed in the human genus. Brain volume, the size of the frontal lobe, or the gross hemispheric asymmetries are still relevant issues, but they are considered to be less central than before. More attention is instead being paid to the cortical organization, the relationships with the cranial architecture, and the influence of molecular or ecological factors. Although the field of paleoneurology can currently count on a larger range of tools and principles, there is still a general lack of anatomical information on many endocranial traits. This aspect is probably crucial for the agenda of paleoneurology. More importantly, the whole science is undergoing a delicate change, because of the growing influence of the social environment. In this sense, the disciplines working with fossils (and, in particular, with brain evolution) should take particular care to maintain a healthy professional situation, avoiding an excess of speculation and overstatement.

Craniofacial orientation and parietal bone morphology in adult modern humans.

In adult humans, the orbits vary mostly in their orientation in relation to the frontal bone profile, while the orientation of the cranial base and face are associated with the anteroposterior dimensions of the parietal bone. Here we investigate the effect of parietal bone length on the orientation of the orbits, addressing craniofacial integration and head orientation. We applied shape analysis to a sample of computed tomography scans from 30 adult modern humans, capturing the outlines of the parietal and frontal bones, the orbits, and the lateral and midline cranial base, to investigate shape variation, covariation, and modularity. Results show that the orientation of the orbits varies in accordance with the anterior cranial base, and in association with changes in parietal bone longitudinal extension. Flatter, elongated parietal bones are associated with downwardly oriented orbits and cranial bases. Modularity analysis points to a significant integration among the orbits, anterior cranial base, and the frontal profile. While the orbits are morphologically integrated with the adjacent structures in terms of shape, the association with parietal bone size depends on the spatial relationship between the two blocks. Complementary changes in orbit and parietal bone might play a role in accommodating craniofacial variability and may contribute to maintain the functional axis of the head. To better understand how skull morphology and head posture relate, future studies should account for the spatial relationship between the head and the neck.

Reconstruction and analysis of the DAN5/P1 and BSN12/P1 Gona Early Pleistocene Homo fossils.

Two Early Pleistocene fossils from Gona, Ethiopia, were originally assigned to Homo erectus, and their differences in size and robusticity were attributed to either sexual dimorphism or anagenetic evolution. In the current study, we both revisit the taxonomic affinities of these fossils and assess whether morphological differences between them reflect temporal evolution or sexual variation. We generated virtual reconstructions of the mostly complete ∼1.55 Ma DAN5/P1 calvaria and the less complete 1.26 Ma BSN12/P1 fossil, allowing us to directly compare their anterior vault shapes using landmark-based shape analysis. The two fossils are similar in calvaria shape to H. erectus and also to other Early Pleistocene Homo species based on a geometric morphometric analysis of calvaria landmarks and semilandmarks. The DAN5/P1 fossil bears a particularly close affinity to the Georgian H. erectus fossils and to KNM-ER 1813 (H. habilis), probably reflecting allometric influences on vault shape. Combined with species-specific traits of the neurocranium (e.g., midline keeling, angular torus), we confirm that these fossils are likely early African H. erectus. We calculated regression-based estimates of endocranial volume for BSN12/P1 of 882-910 cm3 based on three virtual reconstructions. Although BSN12/P1 is markedly larger than DAN5/P1 (598 cm3), both fossils represent the smallest adult H. erectus known from their respective time periods in Africa. Some of the difference in endocranial volume between the two Gona fossils reflects broader species-level brain expansion from 1.77 to 0.01 Ma, confirmed here using a large sample (n = 38) of H. erectus. However, shape differences between these fossils did not reflect species-level changes to calvaria shape. Moreover, the analysis failed to recover a clear pattern of sexually patterned size or shape differences within H. erectus based on our current assessments of sex for individual fossils.

Not a matter of shape: The influence of tool characteristics on electrodermal activity in response to haptic exploration of Lower Palaeolithic tools.

OBJECTIVES: Haptics involves somatosensory perception through the skin surface and dynamic touch based on the proprioceptive response of the whole body. Handling Palaeolithic stone tools influences the arousal and attentional engagement, which can be detected and measured through electrodermal activity. Although tool shape has generally been studied to consider tool functions or tool making, it is also a major factor in tool sensing and haptic perception. The purpose of this survey is to analyze whether the electrodermal reactions are influenced by stone tool morphology. METHODS: We first quantify the morphological variability of 72 stone tools through geometric morphometrics. Then, 12 stone tools from the previous sample were randomly selected to perform the electrodermal analysis in a sample of 46 right-handed adults. RESULTS: Elongation is the main factor involved in Lower Palaeolithic shape variation, followed by the position of the maximum thickness. Attention and manipulation time are mainly influenced by tool size, while arousal mostly correlates with tool weight. Electrodermal activity is apparently not influenced by the overall tool shape. Tool size, weight, and base morphology are the variables that mainly trigger an electrodermal reaction. CONCLUSIONS: Electrophysiological reaction is more sensitive to specific physical features of the tool than to its general outline. These features are not particularly different in worked pebbles and handaxes in terms of grasping, but underwent remarkable changes in other technological traditions. That changes associated with behavioral performances can be employed in cognitive archaeology to investigate the relationships between tool sensing and tool use.

Visual Attention and Cognitive Archaeology: An Eye-Tracking Study of Palaeolithic Stone Tools.

The study of lithic technology can provide information on human cultural evolution. This article aims to analyse visual behaviour associated with the exploration of ancient stone artefacts and how this relates to perceptual mechanisms in humans. In Experiment 1, we used eye tracking to record patterns of eye fixations while participants viewed images of stone tools, including examples of worked pebbles and handaxes. The results showed that the focus of gaze was directed more towards the upper regions of worked pebbles and on the basal areas for handaxes. Knapped surfaces also attracted more fixation than natural cortex for both tool types. Fixation distribution was different to that predicted by models that calculate visual salience. Experiment 2 was an online study using a mouse-click attention tracking technique and included images of unworked pebbles and 'mixed' images combining the handaxe's outline with the pebble's unworked texture. The pattern of clicks corresponded to that revealed using eye tracking and there were differences between tools and other images. Overall, the findings suggest that visual exploration is directed towards functional aspects of tools. Studies of visual attention and exploration can supply useful information to inform understanding of human cognitive evolution and tool use.

Craniovascular traits and braincase morphology in craniosynostotic human skulls.

Middle meningeal vessels, dural venous sinuses, and emissary veins leave imprints and canals in the endocranium, and thus provide evidence of vascular patterns in osteological samples. This paper investigates whether craniovascular morphology undergoes changes in craniosynostotic human skulls, and if specific alterations may reflect structural and functional relationships in the cranium. The analyzed osteological sample consists of adult individuals with craniosynostoses generally associated with dolichocephalic or brachycephalic proportions, and a control sample of anatomically normal adult skulls. The pattern and dominance of the middle meningeal artery, the morphology of the confluence of the sinuses, and the size and number of the emissary foramina were evaluated. Craniovascular morphology was more diverse in craniosynostotic skulls than in anatomically normal skulls. The craniosynostotic skulls often displayed enlarged occipito-marginal sinuses and more numerous emissary foramina. The craniosynostotic skulls associated with more brachycephalic morphology often presented enlarged emissary foramina, while the craniosynostotic skulls associated with dolichocephalic effects frequently displayed more developed posterior branches of the middle meningeal artery. The course and morphology of the middle meningeal vessels, dural venous sinuses, and emissary veins in craniosynostotic skulls can be related to the redistribution of growth forces, higher intracranial pressure, venous hypertension, or thermal constraints. These functional and structural changes are of interest in both anthropology and medicine, involving epigenetic traits that concern the functional and ontogenetic balance between soft and hard tissues.

A morphometric comparison of the parietal lobe in modern humans and Neanderthals.

The modern human brain and braincase have a characteristic globular shape including parietal and cerebellar bulging. In contrast, Neanderthals, although having similar endocranial volume, displayed more elongated endocrania with flatter parietal and cerebellar regions. Based on endocranial imprints, we compare the parietal lobe morphology of modern humans and Neanderthals, as this brain region is central to several cognitive functions including tool use and visual imaging. In paleoneurology, shape analyses of endocasts are based either on anatomical landmarks that represent endocranial surface features homologous to cortical convolutions (impressions of brain gyri and sulci) or on dense meshes of semilandmarks that capture overall endocranial shape. Previous analyses using the former suggested that modern humans have relatively longer and taller parietal lobes than extinct human species, while the latter emphasized parietal bulging without a significant size difference of parietal regions. In the present study, we combine both anatomical landmarks and surface semilandmarks to investigate the morphological differences of the parietal lobes between modern humans and Neanderthals. Despite limitations by landmark uncertainty, our analyses were able to detect and confirm average different parietal shapes, with modern humans displaying taller and anteroposteriorly extended parietal lobes. We also show mean size differences, with modern humans displaying slightly larger surface areas on the dorsal posterior parietal region, and on a lateral region comprising the supramarginal gyrus, angular gyrus, and intraparietal sulcus. While we observed average differences in the parietal form between the two species, their ranges of distribution overlap, indicating the differences could be a matter of degree. Thus, further analyses on intraspecific variation in parietal lobe morphology within modern human brains should help understand the differences between globular and elongated endocrania. This is crucial because changes to the parietal cortex might affect associative and integrative functions between somatic and visual primary inputs.

Is the middle cranial fossa a reliable predictor of temporal lobe volume in extant and fossil anthropoids?

OBJECTIVES: We investigate the suitability of middle cranial fossa (MCF) size as a proxy for temporal lobe volume (TLV), examining the strength of the association between TLV and MCF metrics and assess the reliability predicting TLV in fossil anthropoids. The temporal lobe of the primate brain is a multimodal association cortex involved in long-term memory, auditory, and visual processing with unique specializations in modern humans for language comprehension. The MCF is the bony counterpart for the temporal lobe providing inferences for fossil hominin temporal lobe evolution. We now investigate whether the MCF is a suitable proxy for the temporal lobe. METHODS: A sample of 23 anthropoid species (n = 232, including 13 fossil species) from computed tomography (CT) scans of ex vivo crania and magnetic resonance imaging (MRI) of the in vivo brain were generated into three-dimensional (3D) virtual models. Seven linear metrics were digitally measured on the right MCF with right TLV calculated from in vivo MRI. RESULTS: Regression analyses produced statistically significant correlations between TLV and all MCF metrics (r ≥ 0.85; p ≤ 0.0009) with TLV predictions within ±1 standard error and three MCF metrics (posterior-width, mid-length, and mid-width) the most reliable predictors of TLV with only one metric weakly associated with TLV. DISCUSSION: These findings indicate a strong association between the MCF and TLV, provide reliable predictors of fossil TLV that were previously unattainable, allow the inclusion of fragmentary fossil material, and enable inferences into the emergence of modern human temporal lobe morphology.

Ontogenetic changes of diploic channels in modern humans.

OBJECTIVES: The diploic channels are bony passages of veins, running within frontal, parietal, and occipital bones. In this study, we investigate ontogenetic changes of these channels in a sample of nonadult and adult modern humans. MATERIALS AND METHODS: Using computed tomography scans of dried crania, we provide quantitative comparisons of lumen size, branch length, volume, and vascular asymmetries, and correlations with age, cranial size, and bone thickness. RESULTS: The vascular system displays progressive but nonlinear changes throughout ontogeny, becoming even more complex with adulthood. Vascular variables are significantly different in frontal, parietal, and occipital bones for most of the postnatal ontogeny. Diploic channels of the left and right sides are developed similarly. Vascular variables display a nonlinear association with age and cranial size in modern humans. Cranial bone thickness is shown to be a major determinant of lumen size, branch length, and volume. CONCLUSIONS: A previous radiographic survey suggested that diploic channels are more developed in adult modern humans than in nonadults. Recent advances in digital anatomy have been used in this study to investigate this craniovascular structure. The complexity of the channels increases during development, with a noticeable boost in adults. Taking into account the potential metabolic differences and constraints associated with modern human brain size and shape, the vascular differences found might be related to endocranial thermoregulation.

Hand morphometrics, electrodermal activity, and stone tools haptic perception.

OBJECTIVES: Tool use requires integration among sensorial, biomechanical, and cognitive factors. Taking into account the importance of tool use in human evolution, changes associated with the genus Homo are to be expected in all these three aspects. Haptics is based on both tactile and proprioceptive feedbacks, and it is associated with emotional reactions. Previous analyses have suggested a difference between males and females, and during haptic exploration of different typologies of stone tools. Here, we analyze the correlation between electrodermal reactions during stone tool handling and hand morphology to provide evidence of possible allometric factors shared by males and females. METHODS: Electrodermal analysis was used to investigate some specific parameters involved in these reactions, such as changes in the level of attention and arousal. We analyzed the responses of 46 right-handed adults to 20 distinct stone tools while blindfolded. RESULTS: Females have smaller hands and a wider range of electrodermal reactions. Within males and females, hand diameters and general hand size do not correlate with the degree of electrodermal level and response. CONCLUSIONS: Sex differences in electrodemal reaction during stone tool handling are apparently not due to the effect of hand size or proportions. Differences between males and females are better interpreted as real sex differences, either due to a biological or cultural influences. Hand size does not influence the degree of arousal or attention during tool exploration, suggesting that other factors trigger individual reactions. These results add to a general cognitive approach on hand-tool evolution and tool sensing.

Normal craniovascular variation in two modern European adult populations.

The vascular networks running into the meningeal layers, between the brain and braincase, leave imprints on the endocranial surface. These traces are visible in osteological specimens and skeletal collections, providing indirect evidence of vascular patterns in those cases in which bone remains are the only source of anatomical information, such as in forensic science, bioarchaeology and paleontology. The main vascular elements are associated with the middle meningeal artery, the venous sinuses of the dura mater, and the emissary veins. Most of these vascular systems have been hypothesized to be involved in endocranial thermal regulation. Although these traits deal with macroanatomical features, much information on their variation is still lacking. In this survey, we analyze a set of craniovascular imprints in two European dry skull samples with different neurocranial proportions: a brachycephalic Czech sample (n = 103) and a mesocephalic Italian sample (n = 152). We analyzed variation and distribution, correlation with cranial metrics, and sex differences in the dominance of the branches of the middle meningeal artery, the patterns of confluence of the sinuses, and the size of the emissary foramina. The descriptive statistics provide a reference to compare specimens and samples from different case studies. When compared with the Italian skulls, the Czech skulls display a greater dominance of the anterior branch of the middle meningeal artery and more asymmetric right-dominance of the confluence of the venous sinuses. There is no sex difference in the middle meningeal vessels, but males show a greater prevalence of the occipito-marginal draining system. Differences in the middle meningeal vessels or venous sinuses are apparently not influenced by cranial dimensions or proportions. The mastoid foramina are larger in larger and more brachycephalic skulls, which increases the emissary potential flow in the Czech sample and males, when compared with the Italian samples and females, respectively. The number of mastoid foramina increases in wider skulls. This anatomic information is necessary to develop further morphological and functional inferences on the relationships between neurocranial bones and vessels at the genetic, ontogenetic, and phylogenetic levels.

The cerebellum in Alzheimer's disease: evaluating its role in cognitive decline.

The cerebellum has long been regarded as essential only for the coordination of voluntary motor activity and motor learning. Anatomical, clinical and neuroimaging studies have led to a paradigm shift in the understanding of the cerebellar role in nervous system function, demonstrating that the cerebellum appears integral also to the modulation of cognition and emotion. The search to understand the cerebellar contribution to cognitive processing has increased interest in exploring the role of the cerebellum in neurodegenerative and neuropsychiatric disorders. Principal among these is Alzheimer's disease. Here we review an already sizeable existing literature on the neuropathological, structural and functional neuroimaging studies of the cerebellum in Alzheimer's disease. We consider these observations in the light of the cognitive deficits that characterize Alzheimer's disease and in so doing we introduce a new perspective on its pathophysiology and manifestations. We propose an integrative hypothesis that there is a cerebellar contribution to the cognitive and neuropsychiatric deficits in Alzheimer's disease. We draw on the dysmetria of thought theory to suggest that this cerebellar component manifests as deficits in modulation of the neurobehavioural deficits. We provide suggestions for future studies to investigate this hypothesis and, ultimately, to establish a comprehensive, causal clinicopathological disease model.

Electrodermal activity during Lower Paleolithic stone tool handling.

OBJECTIVES: Hand coordination is a key feature in primate evolution at both behavioral and cognitive levels. Humans further improved their manual abilities, and their cognitive niche is deeply associated with hand-tool relationships and technological capacity. A main cognitive change is thought to be related to the transition from Oldowan to Acheulean stone tool technology around 1.7 million years ago. In this survey, we test whether distinct Lower Paleolithic tools induce different electrophysiological reactions during haptic exploration. METHODS: Electrodermal activity is a relatively quick way to measure emotional and attentional changes during specific tasks. We analyzed changes of the electrodermal response and electrodermal level during Oldowan and Acheulean stone tool handling in a sample of 46 right-handed adult subjects with no previous archeological knowledge. RESULTS: Electrodermal proxies for attention and emotion display a skewed distribution. Females present more variable reactions than males and more emotional engagement. Acheulean tools require longer manipulation time and exert less emotional response than Oldowan tools. Attention is influenced by tool length and weight, emotion is sensitive to tool thickness and weight, and manipulation time depends on tool length and width. CONCLUSIONS: This study suggests subtle but detectable perceptual differences when handling Oldowan and Acheulean stone tools. Such variations associated with hand-tool interaction can provide information on haptic and prosthetic capacities associated with our specialized technological resources. Perceptual changes in the archeological record can reveal evolutionary changes in the corresponding body-tool cognitive mechanisms.

Parietal lobe variation in cercopithecid endocasts.

In extant primates, the posterior parietal cortex is involved in visuospatial integration, attention, and eye-hand coordination, which are crucial functions for foraging and feeding behaviors. Paleoneurology studies brain evolution through the analysis of endocasts, that is molds of the inner surface of the braincase. These may preserve imprints of cortical structures, such as sulci, which might be of interest for locating the boundaries of major cortical regions. Old World monkeys (Cercopithecidae) represent an interesting zoological group for evolutionary studies, because of their diverse ecologies and locomotor behaviors. In this study, we quantify parietal lobe variation within the cercopithecid family, in a sample of 30 endocasts including 11 genera and 17 species, by combining landmark-based and landmark-free geometric morphometric analyses. More specifically, we quantitatively assess variation of the parietal proportions based on landmarks placed on reliable anatomical references and of parietal lobe surface morphology through deformation-based methods. The main feature associated with the cercopithecid endocranial variation regards the inverse proportions of parietal and occipital lobes, with colobines, Theropithecus, and Papio displaying relatively larger parietal lobes and smaller occipital lobes compared with cercopithecins. The parietal surface is anteroposteriorly longer and mediolaterally flatter in colobines, while longitudinally shorter but laterally bulging in baboons. Large parietal lobes in colobines and baboons are likely to be independent evolutionary traits, and not necessarily associated with analogous functions or morphogenetic mechanisms.

Human Paleoneurology and the Evolution of the Parietal Cortex.

Paleoneurology deals with the study of brain anatomy in fossil species, as inferred from the morphology of their endocranial features. When compared with other living and extinct hominids, Homo sapiens is characterized by larger parietal bones and, according to the paleoneurological evidence, also by larger parietal lobes. The dorsal elements of the posterior parietal cortex (superior parietal lobules, precuneus, and intraparietal sulcus) may be involved in these morphological changes. This parietal expansion was also associated with an increase in the corresponding vascular networks, and possibly with increased heat loads. Only H. sapiens has a specific early ontogenetic stage in which brain form achieves such globular appearance. In adult modern humans, the precuneus displays remarkable variation, being largely responsible for the longitudinal parietal size. The precuneus is also much more expanded in modern humans than in chimpanzees. Parietal expansion is not influenced by brain size in fossil hominids or living primates. Therefore, our larger parietal cortex must be interpreted as a derived feature. Spatial models suggest that the dorsal and anterior areas of the precuneus might be involved in these derived morphological variations. These areas are crucial for visuospatial integration, and are sensitive to both genetic and environmental influences. This article reviews almost 20 years of my collaborations on human parietal lobe evolution, integrating functional craniology, paleoneurology, and evolutionary neuroanatomy.

Cranial shape variation in adult howler monkeys (Alouatta seniculus).

Howler monkeys (genus Alouatta) display a distinctive cranial architecture characterized by airorhynchy (or retroflexion of the facial skeleton on the cranial base), a small braincase, and a posteriorly oriented foramen magnum. This configuration has been associated with distinct factors including a high folivory diet, locomotion, and the presence of a specialized vocal tract characterized by large hyoid bone. However, the morphological relationships between the facial and neurocranial blocks in Alouatta have been scarcely investigated. In this study we quantitatively analyzed the cranial shape variation in Alouatta seniculus, to evaluate possible influences and constraints in face and braincase associated with airorhynchy. We also considered the structural role of the pteric area within the cranial functional matrix. We applied landmark-based analysis and multivariate statistics to 31 adult crania, computing shape analyses based on 3D coordinates registration as well as the analysis of the Euclidean distance matrix to investigate patterns of intraspecific morphological variability. Our results suggest that allometry is the main source of variation involved in shaping cranial morphology in howlers, influencing the degree of facial proportions and braincase flattening, and generating the main sexual differences. Larger individuals are characterized by a higher degree of airorhynchy, neurocranial flattening, and expansion of the zygomatic arch. Allometric variations influence the skull as a whole, without distinct patterns for face and braincase, which behave as an integrated morphological unit. A preliminary survey on the pteric pattern suggests that the morphology of this area may be the result of variations in the vertical growth rates between face and braincase. Future studies should be dedicated to the ontogenetic series and focus on airorhynchy in terms of differential growth among distinct cranial districts.

Shape analysis of spatial relationships between orbito-ocular and endocranial structures in modern humans and fossil hominids.

The orbits and eyes of modern humans are situated directly below the frontal lobes and anterior to the temporal lobes. Contiguity between these orbital and cerebral elements could generate spatial constraints, and potentially lead to deformation of the eye and reduced visual acuity during development. In this shape analysis we evaluate whether and to what extent covariation exists between ocular morphology and the size and spatial position of the frontal and temporal areas in adult modern humans. Magnetic resonance imaging (MRI) was used to investigate patterns of variation among the brain and eyes, while computed tomography (CT) was used to compare cranial morphology in this anatomical region among modern humans, extinct hominids and chimpanzees. Seventeen landmarks and semi-landmarks that capture the outline of the eye, frontal lobe, anterior fossa/orbital roof and the position of the temporal tips were sampled using lateral scout views in two dimensions, after projection of the average grayscale values of each hemisphere, with midsagittal and parasagittal elements overlapped onto the same plane. MRI results demonstrated that eye position in adult humans varies most with regard to its horizontal distance from the temporal lobes and, secondly, in its vertical distance from the frontal lobes. Size was mainly found to covary with the distance between the eye and temporal lobes. Proximity to these cerebral lobes may generate spatial constraints, as some ocular deformation was observed. Considering the CT analysis, modern humans vary most with regard to the orientation of the orbits, while interspecific variation is mainly associated with separation between the orbits and endocranial elements. These findings suggest that size and position of the frontal and temporal lobes can affect eye and orbit morphology, though potential effects on eye shape require further study. In particular, possible effects of these spatial and allometric relationships on the eye and vision should be examined using ontogenetic samples, vision parameters such as refractive error in diopters, and three-dimensional approaches that include measures of extraocular soft tissues within the orbit.