Functional adaptation of the infant craniofacial system to mechanical loadings arising from masticatory forces.

The morphology and biomechanics of infant crania undergo significant changes between the pre- and post-weaning phases due to increasing loading of the masticatory system. The aims of this study were to characterize the changes in muscle forces, bite forces and the pattern of mechanical strain and stress arising from the aforementioned forces across crania in the first 48 months of life using imaging and finite element methods. A total of 51 head computed tomography scans of normal individuals were collected and analysed from a larger database of 217 individuals. The estimated mean muscle forces of temporalis, masseter and medial pterygoid increase from 30.9 to 87.0 N, 25.6 to 69.6 N and 23.1 to 58.9 N, respectively (0-48 months). Maximum bite force increases from 90.5 to 184.2 N (3-48 months). There is a change in the pattern of strain and stress from the calvaria to the face during postnatal development. Overall, this study highlights the changes in the mechanics of the craniofacial system during normal development. It further raises questions as to how and what level of changes in the mechanical forces during the development can alter the morphology of the craniofacial system.

Icex: Advances in the automatic extraction and volume calculation of cranial cavities.

The use of non-destructive approaches for digital acquisition (e.g. computerised tomography-CT) allows detailed qualitative and quantitative study of internal structures of skeletal material. Here, we present a new R-based software tool, Icex, applicable to the study of the sizes and shapes of skeletal cavities and fossae in 3D digital images. Traditional methods of volume extraction involve the manual labelling (i.e. segmentation) of the areas of interest on each section of the image stack. This is time-consuming, error-prone and challenging to apply to complex cavities. Icex facilitates rapid quantification of such structures. We describe and detail its application to the isolation and calculation of volumes of various cranial cavities. The R tool is used here to automatically extract the orbital volumes, the paranasal sinuses, the nasal cavity and the upper oral volumes, based on the coordinates of 18 cranial anatomical points used to define their limits, from 3D cranial surface meshes obtained by segmenting CT scans. Icex includes an algorithm (Icv) for the calculation of volumes by defining a 3D convex hull of the extracted cavity. We demonstrate the use of Icex on an ontogenetic sample (0-19 years) of modern humans and on the fossil hominin crania Kabwe (Broken Hill) 1, Gibraltar (Forbes' Quarry) and Guattari 1. We also test the tool on three species of non-human primates. In the modern human subsample, Icex allowed us to perform a preliminary analysis on the absolute and relative expansion of cranial sinuses and pneumatisations during growth. The performance of Icex, applied to diverse crania, shows the potential for an extensive evaluation of the developmental and/or evolutionary significance of hollow cranial structures. Furthermore, being open source, Icex is a fully customisable tool, easily applicable to other taxa and skeletal regions.

Climate change research and action must look beyond 2100.

Anthropogenic activity is changing Earth's climate and ecosystems in ways that are potentially dangerous and disruptive to humans. Greenhouse gas concentrations in the atmosphere continue to rise, ensuring that these changes will be felt for centuries beyond 2100, the current benchmark for projection. Estimating the effects of past, current, and potential future emissions to only 2100 is therefore short-sighted. Critical problems for food production and climate-forced human migration are projected to arise well before 2100, raising questions regarding the habitability of some regions of the Earth after the turn of the century. To highlight the need for more distant horizon scanning, we model climate change to 2500 under a suite of emission scenarios and quantify associated projections of crop viability and heat stress. Together, our projections show global climate impacts increase significantly after 2100 without rapid mitigation. As a result, we argue that projections of climate and its effects on human well-being and associated governance and policy must be framed beyond 2100.

morphomap: An R package for long bone landmarking, cortical thickness, and cross-sectional geometry mapping.

OBJECTIVES: This study describes and demonstrates the functionalities and application of a new R package, morphomap, designed to extract shape information as semilandmarks in multiple sections, build cortical thickness maps, and calculate biomechanical parameters on long bones. METHODS: morphomap creates, from a single input (an oriented 3D mesh representing the long bone surface), multiple evenly spaced virtual sections. morphomap then directly and rapidly computes morphometric and biomechanical parameters on each of these sections. The R package comprises three modules: (a) to place semilandmarks on the inner and outer outlines of each section, (b) to extract cortical thicknesses for 2D and 3D morphometric mapping, and (c) to compute cross-sectional geometry. RESULTS: In this article, we apply morphomap to femora from Homo sapiens and Pan troglodytes to demonstrate its utility and show its typical outputs. morphomap greatly facilitates rapid analysis and functional interpretation of long bone form and should prove a valuable addition to the osteoarcheological analysis software toolkit. CONCLUSIONS: Long bone loading history is commonly retrodicted by calculating biomechanical parameters such as area moments of inertia, analyzing external shape and measuring cortical thickness. morphomap is a software written in the open source R environment, it integrates the main methodological approaches (geometric morphometrics, cortical morphometric maps, and cross-sectional geometry) used to parametrize long bones.

Jaw kinematics and mandibular morphology in humans.

Understanding the influence of feeding behavior on mandibular morphology is necessary for interpreting dietary change in fossil hominins. However, mandibular morphology is also likely to have an effect on feeding behavior, including jaw kinematics. Here we examine the relationships between mandibular morphology and jaw kinematics in humans using landmark-based morphometrics to quantify jaw movement. Three-dimensional movements of reflective markers coupled to the mandible and cranium were used to capture jaw movements while subjects chewed cubes of raw and cooked sweet potato. Geometric morphometric methods were adapted to quantify and analyze gape cycle motion paths. Gape cycles varied significantly across chewing sequences and between raw and cooked sweet potato. Variation in gape cycle size and shape is related to the width (intergonial distance) and length of the mandible. These results underline the fact that jaw kinematic variation within and between taxa is related to and may be influenced by mandibular morphology. Future studies examining kinematic variation should assess the influence of morphological differences on movement.

Assessing the reliability of virtual reconstruction of mandibles.

OBJECTIVES: Mandibular morphological variation is often used to examine various aspects of human palaeobiology. However, fossil and archeological skeletal remains are often fragmented/distorted and so are frequently excluded from studies. This leads to decreased sample sizes and, potentially, to biased results. Thus, it is of interest to restore the original anatomy of incomplete/distorted specimens. Thin plate splines (TPS), commonly used in Geometric Morphometrics (GM), offer the prospect of reconstruction of missing parts and particularly of interest here, missing landmarks. MATERIALS AND METHODS: Here, the reliability of TPS based mandibular reconstruction is tested. To that end missing landmarks were simulated in originally complete hemimandibles. TPS was then used to restore the location of simulated missing data and the predicted landmarks were compared to the original ones. RESULTS: Results show that error varies according to the number and location of estimated landmarks. Notwithstanding, estimation error is usually considerably smaller than the morphological differences between individuals from the same species. DISCUSSION: TPS based reconstruction allows fragmentary mandibles to be used in studies of whole mandibular variation, provided the above mentioned caveats are considered.

Three-dimensional analysis of sexual dimorphism in ribcage kinematics of modern humans.

OBJECTIVES: Sexual dimorphism is an important biological factor underlying morphological variation in the human skeleton. Previous research found sex-related differences in the static ribcage, with males having more horizontally oriented ribs and a wider lower ribcage than females. Furthermore, a recent study found sex-related differences in the kinematics of the human lungs, with cranio-caudal movements of the caudal part of the lungs accounting for most of the differences between sexes. However, these movements cannot be quantified in the skeletal ribcage, so we do not know if the differences observed in the lungs are also reflected in sex differences in the motion of the skeletal thorax. MATERIALS AND METHODS: To address this issue, we quantified the morphological variation of 42 contemporary human ribcages (sex-balanced) in both maximal inspiration and expiration using 526 landmarks and semilandmarks. Thoracic centroid size differences between sexes were assessed using a t test, and shape differences were assessed using Procrustes shape coordinates, through mean comparisons and dummy regressions of shape on kinematic status. A principal components analysis was used to explore the full range of morphological variation. RESULTS: Our results show significant size differences between males and females both in inspiration and expiration (p < .01) as well as significant shape differences, with males deforming more than females during inspiration, especially in the mediolateral dimension of the lower ribcage. Finally, dummy regressions of shape on kinematic status showed a small but statistically significant difference in vectors of breathing kinematics between males and females (14.78°; p < .01). DISCUSSION: We support that sex-related differences in skeletal ribcage kinematics are discernible, even when soft tissues are not analyzed. We hypothesize that this differential breathing pattern is primarily a result of more pronounced diaphragmatic breathing in males, which might relate to differences in body composition, metabolism, and ultimately greater oxygen demand in males compared to females. Future research should further explore the links between ribcage morphological variation and basal metabolic rate.

Applying Geometric Morphometrics to Digital Reconstruction and Anatomical Investigation.

Virtual imaging, image manipulation and morphometric methods are increasingly used in medicine and the natural sciences. Virtual imaging hardware and image manipulation software allows us to readily visualise, explore, alter, repair and study digital objects. This suite of equipment and tools combined with statistical tools for the study of form variation and covariation using Procrustes based analyses of landmark coordinates, geometric morphometrics, makes possible a wide range of studies of human variation pertinent to biomedicine. These tools for imaging, quantifying and analysing form have already led to new insights into organismal growth, development and evolution and offer exciting prospects in future biomedical applications. This chapter presents a review of commonly used methods for digital acquisition, extraction and landmarking of anatomical structures and of the common geometric morphometric statistical methods applied to investigate them: generalised Procrustes analysis to derive shape variables, principal component analysis to examine patterns of variation, multivariate regression to examine how form is influenced by meaningful factors and partial least squares analysis to examine associations among structures or between these and other interesting variables. An example study of human facial and maxillary sinus ontogeny illustrates these approaches.

The biomechanical significance of the frontal sinus in Kabwe 1 (Homo heidelbergensis).

Paranasal sinuses are highly variable among living and fossil hominins and their function(s) are poorly understood. It has been argued they serve no particular function and are biological 'spandrels' arising as a structural consequence of changes in associated bones and/or soft tissue structures. In contrast, others have suggested that sinuses have one or more functions, in olfaction, respiration, thermoregulation, nitric oxide production, voice resonance, reduction of skull weight, and craniofacial biomechanics. Here we assess the extent to which the very large frontal sinus of Kabwe 1 impacts on the mechanical performance of the craniofacial skeleton during biting. It may be that the browridge is large and the sinus has large trabecular struts traversing it to compensate for the effect of a large sinus on the ability of the face to resist forces arising from biting. Alternatively, the large sinus may have no impact and be sited where strains that arise from biting would be very low. If the former is true, then infilling of the sinus would be expected to increase the ability of the skeleton to resist biting loads, while removing the struts might have the opposite effect. To these ends, finite element models with hollowed and infilled variants of the original sinus were created and loaded to simulate different bites. The deformations arising due to loading were then compared among different models and bites by contrasting the strain vectors arising during identical biting tasks. It was found that the frontal bone experiences very low strains and that infilling or hollowing of the sinus has little effect on strains over the cranial surface, with small effects over the frontal bone. The material used to infill the sinus experienced very low strains. This is consistent with the idea that frontal sinus morphogenesis is influenced by the strain field experienced by this region such that it comes to lie entirely within a region of the cranium that would otherwise experience low strains. This has implications for understanding why sinuses vary among hominin fossils.

The biting performance of Homo sapiens and Homo heidelbergensis.

Modern humans have smaller faces relative to Middle and Late Pleistocene members of the genus Homo. While facial reduction and differences in shape have been shown to increase biting efficiency in Homo sapiens relative to these hominins, facial size reduction has also been said to decrease our ability to resist masticatory loads. This study compares crania of Homo heidelbergensis and H. sapiens with respect to mechanical advantages of masticatory muscles, force production efficiency, strains experienced by the cranium and modes of deformation during simulated biting. Analyses utilize X-ray computed tomography (CT) scan-based 3D models of a recent modern human and two H. heidelbergensis. While having muscles of similar cross-sectional area to H. heidelbergensis, our results confirm that the modern human masticatory system is more efficient at converting muscle forces into bite forces. Thus, it can produce higher bite forces than Broken Hill for equal muscle input forces. This difference is the result of alterations in relative in and out-lever arm lengths associated with well-known differences in midfacial prognathism. Apparently at odds with this increased efficiency is the finding that the modern human cranium deforms more, resulting in greater strain magnitudes than Broken Hill when biting at the equivalent tooth. Hence, the facial reduction that characterizes modern humans may not have evolved as a result of selection for force production efficiency. These findings provide further evidence for a degree of uncoupling between form and function in the masticatory system of modern humans. This may reflect the impact of food preparation technologies. These data also support previous suggestions that differences in bite force production efficiency can be considered a spandrel, primarily driven by the midfacial reduction in H. sapiens that occurred for other reasons. Midfacial reduction plausibly resulted in a number of other significant changes in morphology, such as the development of a chin, which has itself been the subject of debate as to whether or not it represents a mechanical adaptation or a spandrel.

Open data and digital morphology.

Over the past two decades, the development of methods for visualizing and analysing specimens digitally, in three and even four dimensions, has transformed the study of living and fossil organisms. However, the initial promise that the widespread application of such methods would facilitate access to the underlying digital data has not been fully achieved. The underlying datasets for many published studies are not readily or freely available, introducing a barrier to verification and reproducibility, and the reuse of data. There is no current agreement or policy on the amount and type of data that should be made available alongside studies that use, and in some cases are wholly reliant on, digital morphology. Here, we propose a set of recommendations for minimum standards and additional best practice for three-dimensional digital data publication, and review the issues around data storage, management and accessibility.

The earliest evidence for anatomically modern humans in northwestern Europe.

The earliest anatomically modern humans in Europe are thought to have appeared around 43,000-42,000 calendar years before present (43-42 kyr cal BP), by association with Aurignacian sites and lithic assemblages assumed to have been made by modern humans rather than by Neanderthals. However, the actual physical evidence for modern humans is extremely rare, and direct dates reach no farther back than about 41-39 kyr cal BP, leaving a gap. Here we show, using stratigraphic, chronological and archaeological data, that a fragment of human maxilla from the Kent's Cavern site, UK, dates to the earlier period. The maxilla (KC4), which was excavated in 1927, was initially diagnosed as Upper Palaeolithic modern human. In 1989, it was directly radiocarbon dated by accelerator mass spectrometry to 36.4-34.7 kyr cal BP. Using a Bayesian analysis of new ultrafiltered bone collagen dates in an ordered stratigraphic sequence at the site, we show that this date is a considerable underestimate. Instead, KC4 dates to 44.2-41.5 kyr cal BP. This makes it older than any other equivalently dated modern human specimen and directly contemporary with the latest European Neanderthals, thus making its taxonomic attribution crucial. We also show that in 13 dental traits KC4 possesses modern human rather than Neanderthal characteristics; three other traits show Neanderthal affinities and a further seven are ambiguous. KC4 therefore represents the oldest known anatomically modern human fossil in northwestern Europe, fills a key gap between the earliest dated Aurignacian remains and the earliest human skeletal remains, and demonstrates the wide and rapid dispersal of early modern humans across Europe more than 40 kyr ago.

Validity and sensitivity of a human cranial finite element model: implications for comparative studies of biting performance.

Finite element analysis (FEA) is a modelling technique increasingly used in anatomical studies investigating skeletal form and function. In the case of the cranium this approach has been applied to both living and fossil taxa to (for example) investigate how form relates to function or infer diet or behaviour. However, FE models of complex musculoskeletal structures always rely on simplified representations because it is impossible completely to image and represent every detail of skeletal morphology, variations in material properties and the complexities of loading at all spatial and temporal scales. The effects of necessary simplifications merit investigation. To this end, this study focuses on one aspect, model geometry, which is particularly pertinent to fossil material where taphonomic processes often destroy the finer details of anatomy or in models built from clinical CTs where the resolution is limited and anatomical details are lost. We manipulated the details of a finite element (FE) model of an adult human male cranium and examined the impact on model performance. First, using digital speckle interferometry, we directly measured strains from the infraorbital region and frontal process of the maxilla of the physical cranium under simplified loading conditions, simulating incisor biting. These measured strains were then compared with predicted values from FE models with simplified geometries that included modifications to model resolution, and how cancellous bone and the thin bones of the circum-nasal and maxillary regions were represented. Distributions of regions of relatively high and low principal strains and principal strain vector magnitudes and directions, predicted by the most detailed FE model, are generally similar to those achieved in vitro. Representing cancellous bone as solid cortical bone lowers strain magnitudes substantially but the mode of deformation of the FE model is relatively constant. In contrast, omitting thin plates of bone in the circum-nasal region affects both mode and magnitude of deformation. Our findings provide a useful frame of reference with regard to the effects of simplifications on the performance of FE models of the cranium and call for caution in the interpretation and comparison of FEA results.

Extreme climate, rather than population history, explains mid-facial morphology of Northern Asians.

Previous studies have examined mid-facial cold adaptation among either widely dispersed and genetically very diverse groups of humans isolated for tens of thousands of years, or among very closely related groups spread over climatically different regions. Here we present a study of one East Asian and seven North Asian populations in which we examine the evidence for convergent adaptations of the mid-face to a very cold climate. Our findings indicate that mid-facial morphology is strongly associated with climatic variables that contrast the temperate climate of East Asians and the very cold and dry climate of North Asians. This is also the case when either maxillary or nasal cavity measurements are considered alone. The association remains significant when mtDNA distances among populations are taken into account. The morphological contrasts between populations are consistent with physiological predictions and prior studies of mid-facial cold adaptation in more temperate regions, but among North Asians there appear to be some previously undescribed morphological features that might be considered as adaptive to extreme cold. To investigate this further, analyses of the seven North Asian populations alone suggest that mid-facial morphology remains strongly associated with climate, particularly winter precipitation, contrasting coastal Arctic and continental climates. However, the residual covariation among North Asian mid-facial morphology and climate when genetic distances are considered, is not significant. These findings point to modern adaptations to extreme climate that might be relevant to our understanding of the mid-facial morphology of fossil hominins that lived during glaciations.

Exploring motion using geometric morphometrics in microscopic aquatic invertebrates: 'modes' and movement patterns during feeding in a bdelloid rotifer model species.

BACKGROUND: Movement is a defining aspect of animals, but it is rarely studied using quantitative methods in microscopic invertebrates. Bdelloid rotifers are a cosmopolitan class of aquatic invertebrates of great scientific interest because of their ability to survive in very harsh environment and also because they represent a rare example of an ancient lineage that only includes asexually reproducing species. In this class, Adineta ricciae has become a model species as it is unusually easy to culture. Yet, relatively little is known of its ethology and almost nothing on how it behaves during feeding. METHODS: To explore feeding behaviour in A. ricciae, as well as to provide an example of application of computational ethology in a microscopic invertebrate, we apply Procrustes motion analysis in combination with ordination and clustering methods to a laboratory bred sample of individuals recorded during feeding. RESULTS: We demonstrate that movement during feeding can be accurately described in a simple two-dimensional shape space with three main 'modes' of motion. Foot telescoping, with the body kept straight, is the most frequent 'mode', but it is accompanied by periodic rotations of the foot together with bending while the foot is mostly retracted. CONCLUSIONS: Procrustes motion analysis is a relatively simple but effective tool for describing motion during feeding in A. ricciae. The application of this method generates quantitative data that could be analysed in relation to genetic and ecological differences in a variety of experimental settings. The study provides an example that is easy to replicate in other invertebrates, including other microscopic animals whose behavioural ecology is often poorly known.

Divergent evolutionary processes associated with colonization of offshore islands.

Oceanic islands have been a test ground for evolutionary theory, but here, we focus on the possibilities for evolutionary study created by offshore islands. These can be colonized through various means and by a wide range of species, including those with low dispersal capabilities. We use morphology, modern and ancient sequences of cytochrome b (cytb) and microsatellite genotypes to examine colonization history and evolutionary change associated with occupation of the Orkney archipelago by the common vole (Microtus arvalis), a species found in continental Europe but not in Britain. Among possible colonization scenarios, our results are most consistent with human introduction at least 5100 bp (confirmed by radiocarbon dating). We used approximate Bayesian computation of population history to infer the coast of Belgium as the possible source and estimated the evolutionary timescale using a Bayesian coalescent approach. We showed substantial morphological divergence of the island populations, including a size increase presumably driven by selection and reduced microsatellite variation likely reflecting founder events and genetic drift. More surprisingly, our results suggest that a recent and widespread cytb replacement event in the continental source area purged cytb variation there, whereas the ancestral diversity is largely retained in the colonized islands as a genetic 'ark'. The replacement event in the continental M. arvalis was probably triggered by anthropogenic causes (land-use change). Our studies illustrate that small offshore islands can act as field laboratories for studying various evolutionary processes over relatively short timescales, informing about the mainland source area as well as the island.

Concordance of traditional osteometric and volume-rendered MSCT interlandmark cranial measurements.

The statistical quantification of error and uncertainty is inherently intertwined with ascertaining the admissibility of forensic evidence in a court of law. In the forensic anthropological discipline, the robustness of any given standard should not only be evaluated according to its stated error but by the accuracy and precision of the raw data (measurements) from which they are derived. In the absence of Australian contemporary documented skeletal collections, medical scans (e.g. multislice computed tomography-MSCT) offer a source of contemporary population-specific data for the formulation of skeletal standards. As the acquisition of morphometric data from clinical MSCT scans is still relatively novel, the purpose of this study is to assess validity of the raw data that is being used to formulate Australian forensic standards. Six human crania were subjected to clinical MSCT at a slice thickness of 0.9 mm. Each cranium and its corresponding volume-rendered three-dimensional MSCT image were measured multiple times. Whether differences between MSCT and dry bone interlandmark measurements are negligible is statistically quantified; intra- and inter-observer measurement error is also assessed. We found that traditional bone measurements are more precise than their MSCT counterparts, although overall differences between the two data acquisition methods are negligible compared to sample variance. Cranial variation accounted on average for more than 20× the variance explained by MSCT vs. bone measurements. Similarly, although differences between operators were sometimes significant compared to intra-operator variance, they were negligible when compared to sample variance, which was on average 12× larger than that due to inter-operator differences.

A Comparison of Semilandmarking Approaches in the Visualisation of Shape Differences.

In landmark-based analyses of size and shape variation and covariation among biological structures, regions lacking clearly identifiable homologous landmarks are commonly described by semilandmarks. Different algorithms may be used to apply semilandmarks, but little is known about the consequences of analytical results. Here, we assess how different approaches and semilandmarking densities affect the estimates and visualisations of mean and allometrically scaled surfaces. The performance of three landmark-driven semilandmarking approaches is assessed using two different surface mesh datasets with different degrees of variation and complexity: adult human head and ape cranial surfaces. Surfaces fitted to estimates of the mean and allometrically scaled landmark and semilandmark configurations arising from geometric morphometric analyses of these datasets are compared between semilandmarking approaches and different densities, as well as with those from warping to landmarks alone. We find that estimates of surface mesh shape (i.e., after re-semilandmarking and then re-warping) made with varying numbers of semilandmarks are generally consistent, while the warping of surfaces using landmarks alone yields surfaces that can be quite different to those based on semilandmarks, depending on landmark coverage and choice of template surface for warping. The extent to which these differences are important depends on the particular study context and aims.

A Comparison of Semilandmarking Approaches in the Analysis of Size and Shape.

Often, few landmarks can be reliably identified in analyses of form variation and covariation. Thus, 'semilandmarking' algorithms have increasingly been applied to surfaces and curves. However, the locations of semilandmarks depend on the investigator's choice of algorithm and their density. In consequence, to the extent that different semilandmarking approaches and densities result in different locations of semilandmarks, they can be expected to yield different results concerning patterns of variation and co-variation. The extent of such differences due to methodology is, as yet, unclear and often ignored. In this study, the performance of three landmark-driven semilandmarking approaches is assessed, using two different surface mesh datasets (ape crania and human heads) with different degrees of variation and complexity, by comparing the results of morphometric analyses. These approaches produce different semilandmark locations, which, in turn, lead to differences in statistical results, although the non-rigid semilandmarking approaches are consistent. Morphometric analyses using semilandmarks must be interpreted with due caution, recognising that error is inevitable and that results are approximations. Further work is needed to investigate the effects of using different landmark and semilandmark templates and to understand the limitations and advantages of different semilandmarking approaches.

Normal human craniofacial growth and development from 0 to 4 years.

Knowledge of human craniofacial growth (increase in size) and development (change in shape) is important in the clinical treatment of a range of conditions that affects it. This study uses an extensive collection of clinical CT scans to investigate craniofacial growth and development over the first 48 months of life, detail how the cranium changes in form (size and shape) in each sex and how these changes are associated with the growth and development of various soft tissues such as the brain, eyes and tongue and the expansion of the nasal cavity. This is achieved through multivariate analyses of cranial form based on 3D landmarks and semi-landmarks and by analyses of linear dimensions, and cranial volumes. The results highlight accelerations and decelerations in cranial form changes throughout early childhood. They show that from 0 to 12 months, the cranium undergoes greater changes in form than from 12 to 48 months. However, in terms of the development of overall cranial shape, there is no significant sexual dimorphism in the age range considered in this study. In consequence a single model of human craniofacial growth and development is presented for future studies to examine the physio-mechanical interactions of the craniofacial growth.