Evaluating the muscle attachment hypothesis for sagittal cresting in Gorilla and Pongo.

Primate mandibular morphology is often associated with jaw functionality of the masticatory complex in the context of variation in diets. Recent research into the disparities between the diet and jaw functionality in male and female hominoids is inconclusive and suggests that sexual dimorphism in the mandible may be influenced by external factors such as temporalis and masseter muscle morphology, which in turn may be influenced by sexual selection. As the muscles associated with mastication (i.e., the type of chewing exhibited by primates and other mammals) encompass the mandible as well as the neurocranium, including the sagittal crest among some individuals, this study investigates sex-specific associations between regions of the mandibular ramus and neurocranium associated with mastication in a dentally mature sample of Gorilla and Pongo. A total of four cranial and mandibular variables were measured in two Gorilla taxa (Gorilla gorilla gorilla and Gorilla beringei graueri) and one Pongo taxon (Pongo pygmaeus pygmaeus) (n = 220). For all three taxa, we investigate (a) whether the degree of sexual dimorphism in cranial regions associated with sagittal cresting (sagittal crest size (SCS) and temporalis muscle attachment area (TMAA)) is proportional to the degree of mandibular ramus area (MRA) and coronoid process height (CPH) sexual dimorphism, (b) whether there are sex differences in scaling relationships between TMAA and MRA, and (c) whether there are sex differences in the strength of association between TMAA and CPH. We show that for G. g. gorilla, variables associated with sagittal cresting show higher sexual dimorphism values than our two mandibular ramus variables, which is not the case for G. b. graueri or for P. p. pygmaeus. All three taxa show similar sex-specific scaling relationships between TMAA and MRA, where for males this relationship does not diverge from isometry, and for females there is a negative allometric relationship. Our findings also show intraspecific sex differences in allometric slopes between MRA and TMAA for all three taxa. Only G. g. gorilla shows a significant association between TMAA and CPH, which is observed in both sexes. Although there are some statistical associations between the cranial and mandibular regions associated with mastication, our results show that among male gorillas and orangutans, patterns of variation in the sagittal crest, TMAA, mandibular ramus and the coronoid process cannot be explained by the muscle attachment hypothesis alone. These findings have implications surrounding the associations between social behaviour and the morphology of the craniofacial complex.

The Application of 3D Landmark-Based Geometric Morphometrics towards Refinement of the Piglet Grimace Scale.

Proper methods of assessment to objectively identify pain are essential for welfare improvements of piglets undergoing surgical castration on commercial farms. The Piglet Grimace Scale (PGS) is used to identify changes in facial expressions caused by acute pain in piglets undergoing tail docking and castration. However, subjective scoring methods are a concern for the validation of the PGS. The objectives of this study were to evaluate and refine the PGS through 3D landmark geometric morphometrics. Male piglets (n = 88) were randomly allocated to one of two treatments: castration and sham-castration. Piglet facial images were taken at four time points (before treatment, immediately post-treatment, 1 h post-treatment, and 4 h post-treatment) using a photogrammetry rig. Images were scored by four raters using five facial action units (FAUs): orbital tightening, ear position, temporal tension, lip contraction, and nose bulge/cheek tension. Three-dimensional facial models were generated and landmarked denoting 3 FAUs (orbital tightening, lip contraction, and nose bulge/cheek tension). Results suggest that orbital tightening and ear position may be reliable FAUs for the PGS. However, neither the PGS nor 3D landmark-based geometric morphometrics were able (both p > 0.10) to reliably identify facial indicators of pain in piglets undergoing castration.

Facial asymmetry tracks genetic diversity among Gorilla subspecies.

Mountain gorillas are particularly inbred compared to other gorillas and even the most inbred human populations. As mountain gorilla skeletal material accumulated during the 1970s, researchers noted their pronounced facial asymmetry and hypothesized that it reflects a population-wide chewing side preference. However, asymmetry has also been linked to environmental and genetic stress in experimental models. Here, we examine facial asymmetry in 114 crania from three Gorilla subspecies using 3D geometric morphometrics. We measure fluctuating asymmetry (FA), defined as random deviations from perfect symmetry, and population-specific patterns of directional asymmetry (DA). Mountain gorillas, with a current population size of about 1000 individuals, have the highest degree of facial FA (explaining 17% of total facial shape variation), followed by Grauer gorillas (9%) and western lowland gorillas (6%), despite the latter experiencing the greatest ecological and dietary variability. DA, while significant in all three taxa, explains relatively less shape variation than FA does. Facial asymmetry correlates neither with tooth wear asymmetry nor increases with age in a mountain gorilla subsample, undermining the hypothesis that facial asymmetry is driven by chewing side preference. An examination of temporal trends shows that stress-induced developmental instability has increased over the last 100 years in these endangered apes.

Temporomandibular joint shape in anthropoid primates varies widely and is patterned by size and phylogeny.

The temporomandibular joint is the direct interface between the mandible and the cranium and is critical for transmitting joint reaction forces and determining mandibular range of motion. As a consequence, understanding variation in the morphology of this joint and how it relates to other aspects of craniofacial form is important for better understanding masticatory function. Here, we present a detailed three-dimensional (3D) geometric morphometric analysis of the cranial component of this joint, the glenoid fossa, across a sample of 17 anthropoid primates, and we evaluate covariation between the glenoid and the cranium and mandible. We find high levels of intraspecific variation in glenoid shape that is likely linked to sexual dimorphism and joint remodeling, and we identify differences in mean glenoid shape across taxonomic groups and in relation to size. Analyses of covariation reveal strong relationships between glenoid shape and a variety of aspects of cranial and mandibular form. Our findings suggest that intraspecific variation in glenoid shape in primates could further be reflective of high levels of functional flexibility in the masticatory apparatus, as has also been suggested for primate jaw kinematics and muscle activation patterns. Conversely, interspecific differences likely reflect larger scale differences between species in body size and/or masticatory function. Results of the covariation analyses dovetail with those examining covariation in the cranium of canids and may be indicative of larger patterns across mammals.

How does scanner choice and 3D model resolution affect data accuracy?

Researchers using digital methods often collect data from 3D models at different resolutions, obtained using different scanning techniques. Although previous research has sought to understand whether scanning method and model resolution affect data accuracy, no study has systematically evaluated the sources of error associated with scanning method, data acquisition method and model resolution with the aim of providing practical recommendations about the model resolution required to yield sufficiently accurate data for specimens of given sizes. In this study, using data taken from primate specimens of three broad size categories, we test whether 3D models obtained using five different scanners (Breuckmann SmartSCAN, DAVID/HP 3D Pro S3, NextEngine 2020i, Creaform Go!Scan 20 and microCT/clinicalCT) yield accurate measurements. We assess whether caliper measurements can be used alongside measurements collected from 3D surface models, whether scanning resolution affects measurement accuracy, and how scan resolution, estimated using each scanner's proprietary software, compares to model resolution measured in a standardized way. Each scanner produces 3D models that yield accurate measurements for each size category, however, combining caliper data with those taken from digital models can be problematic. Our results indicate that the accuracy of measurements taken from 3D models depends on both object size and model resolution. Based on our findings, we recommend that small specimens should be scanned at <0.3 mm, medium specimens at 0.3-0.7 mm, and large specimens at 0.3-0.5 mm resolutions if data taken from 3D surface models are to be combined with caliper datasets. We further show, for the first time, that discrepancies in estimated final model resolution are frequently observed across software packages. We therefore recommend that researchers ensure that final model resolutions are adequate based on specimen size and are independently verified using a software package other than the scanner's proprietary software. Finally, we consider the implications of the findings that measurements obtained from surface models are variably consistent with those obtained using calipers.

Mandibular corpus shape is a taxonomic indicator in extant hominids.

OBJECTIVES: The aim of this study is to understand whether the shape of three sub-regions of the mandibular corpus (the alveolar arch, corpus at M1 and posterior symphysis) are useful for making taxonomic assessments at the genus and species levels in extant hominids. MATERIALS AND METHODS: We use data taken from 3D surface scans of the mandibular corpus of seven extant hominid taxa: Gorilla gorilla gorilla, Gorilla beringei graueri, Homo sapiens, Pan paniscus, Pan troglodytes schweinfurthii, Pongo abelii, and Pongo pygmaeus pygmaeus to generate four shape variables: alveolar arch shape (AAS), corpus shape at M1 (CSM1 ), posterior symphysis shape at the midline (PSSM), and posterior symphysis shape (PSS). To ascertain how reliable each mandibular shape variable is for assessing taxonomy, we ran canonical discriminant and discriminant function analysis, reporting cross-validated results. RESULTS: Using a combination of three mandibular corpus shape variables, 99% of specimens were classified correctly for genus-level analyses. A maximum of 100% of Pan specimens, 94% of Gorilla specimens and 96% of Pongo specimens were classified correctly at the species level when up to three mandibular shape variables were included in the analyses. When mandibular corpus variables were considered in isolation, posterior symphysis shape yielded the highest overall correct classification results. DISCUSSION: The high taxonomic classification rates at both the genus and species level, using 3D surface data and advanced quantification techniques, show that the shape of the alveolar arch, corpus at M1 and symphysis can distinguish extant hominid taxa. These findings have implications for assessing the taxonomy of extinct hominid specimens which preserve these mandibular sub-regions.

Early brain growth cessation in wild Virunga mountain gorillas (Gorilla beringei beringei).

Understanding the life history correlates of ontogenetic differences in hominoid brain growth requires information from multiple species. At present, however, data on how brain size changes over the course of development are only available from chimpanzees and modern humans. In this study, we examined brain growth in wild Virunga mountain gorillas using data derived from necropsy reports (N = 34) and endocranial volume (EV) measurements (N = 86). The youngest individual in our sample was a 10-day-old neonatal male with a brain mass of 208 g, representing 42% of the adult male average. Our results demonstrate that Virunga mountain gorillas reach maximum adult-like brain mass by 3-4 years of age; adult-sized EV is reached by the time the first permanent molars emerge. This is in contrast to the pattern observed in chimpanzees, which despite their smaller absolute brain size, reportedly attain adult brain mass approximately 1 year later than Virunga mountain gorillas. Our findings demonstrate that brain growth is completed early in Virunga mountain gorillas compared to other great apes studied thus far, in a manner that appears to be linked with other life history characteristics of this population.