The Feeding Biomechanics and Dietary Ecology of Paranthropus boisei

The African Plio‐Pleistocene hominins known as australopiths evolved derived craniodental features frequently interpreted as adaptations for feeding on either hard, or compliant/tough foods. Among australopiths, Paranthropus boisei is the most robust form, exhibiting traits traditionally hypothesized to produce high bite forces efficiently and strengthen the face against feeding stresses. However, recent mechanical analyses imply that P. boisei may not have been an efficient producer of bite force and that robust morphology in primates is not necessarily strong. Here we use an engineering method, finite element analysis, to show that the facial skeleton of P. boisei is structurally strong, exhibits a strain pattern different from that in chimpanzees (Pan troglodytes) and Australopithecus africanus, and efficiently produces high bite force. It has been suggested that P. boisei consumed a diet of compliant/tough foods like grass blades and sedge pith. However, the blunt occlusal topography of this and other species suggests that australopiths are adapted to consume hard foods, perhaps including grass and sedge seeds. A consideration of evolutionary trends in morphology relating to feeding mechanics suggests that food processing behaviors in gracile australopiths evidently were disrupted by environmental change, perhaps contributing to the eventual evolution of Homo and Paranthropus This is the peer reviewed version of the following article: Smith, A. L., Benazzi, S. , Ledogar, J. A., Tamvada, K. , Pryor Smith, L. C., Weber, G. W., Spencer, M. A., Lucas, P. W., Michael, S. , Shekeban, A. , Al‐Fadhalah, K. , Almusallam, A. S., Dechow, P. C., Grosse, I. R., Ross, C. F., Madden, R. H., Richmond, B. G., Wright, B. W., Wang, Q. , Byron, C. , Slice, D. E., Wood, S. , Dzialo, C. , Berthaume, M. A., van, Casteren, A. and Strait, D. S. (2015), The Feeding Biomechanics and Dietary Ecology of Paranthropus boisei, which has been published in final form at https://doi.org/10.1002/ar.23073. This article may be used for non-commercial purposes in accordance with Wiley Terms and Conditions for Use of Self-Archived Version

Ambient occlusion and PCV (portion de ciel visible): A new dental topographic metric and proxy of morphological wear resistance

Recently, ambient occlusion, quantified through portion de ciel visible (PCV) was introduced as a method for quantifying dental morphological wear resistance and reconstructing diet in mammals. Despite being used to reconstruct diet and investigate the relationship between dental form and function, no rigorous analysis has investigated the correlation between PCV and diet. Using a sample of platyrrhine and prosimians M2s, we show average PCV was significantly different between most dietary groups. In prosimian, insectivores had the lowest PCV, followed by folivores, omnivores, frugivores, and finally hard-object feeders. In platyrrhines, omnivores had the lowest average PCV, followed by folivores, frugivores, and finally hard-object feeders. PCV was correlated to two topographic variables (Dirichlet normal energy, DNE, and relief index, RFI) but uncorrelated to three others (orientation patch count rotated, OPCR, tooth surface area, and tooth size). The OPCR values here differed greatly from previously published values using the same sample, showing how differences in data acquisition (i.e., using 2.5D vs. 3D surfaces) can lead to drastic differences in results. Compared to other popular topographic variables, PCV performed as well or better at predicting diet in these groups, and when combined with a metric for size, the percent of successful dietary classifications reached 90%. Further, using an ontogenetic series of hominin (Paranthropus robustus) M2s, we show that PCV correlates well with probability of wear, with PCV values being higher on the portions of the occlusal surface that experience more wear (e.g., cusps and crest tips, wear facets) than the portions of the tooth that experience less. This relationship is strongest once wear facets have begun to form on the occlusal surface. These results highlight the usefulness of PCV in quantifying morphological wear resistance and predicting diet in mammals

Effects of cropping, smoothing, triangle count, and mesh resolution on 6 dental topographic metrics

Dental topography is a widely used method for quantifying dental morphology and inferring dietary ecology in animals. Differences in methodology have brought into question the comparability of different studies. Using primate mandibular second molars, we investigated the effects of mesh preparation parameters smoothing, cropping, and triangle count/mesh resolution (herein, resolution) on six topographic variables (Dirichlet normal energy, DNE; orientation patch count rotated, OPCR; relief index, RFI; ambient occlusion, portion de ciel visible, PCV; enamel surface area, SA; tooth size) to determine the effects of smoothing, cropping, and triangle count/resolution on topographic values and the relationship between these values and diet. All topographic metrics are sensitive to smoothing, cropping method, and triangle count/resolution. In general, smoothing decreased DNE, OPCR, RFI, and SA, increased PCV, and had no predictable effect on tooth size. Relative to the basin cut off (BCO) cropping method, the entire enamel cap (EEC) method increased RFI, SA, and size, and had no predictable effect on DNE and OPCR. Smoothing and cropping affected DNE/OPCR and surfaces with low triangle counts more than other metrics and surfaces with high triangle counts. There was a positive correlation between DNE/OPCR and triangle count/resolution, and the rate of increase was weakly correlated to diet. PCV tended to converge or decrease with increases in triangle count/resolution, and RFI, SA, and size converged. Finally, there appears to be no optimal triangle count or resolution for predicting diet from this sample, and constant triangle count appeared to perform better than constant resolution for predicting diet

Dental topography and the diet of Homo naledi

Though late Middle Pleistocene in age, Homo naledi is characterized by a mosaic of Australopithecus-like (e.g., curved fingers, small brains) and Homo-like (e.g., elongated lower limbs) traits, which may suggest it occupied a unique ecological niche. Ecological reconstructions inform on niche occupation, and are particularly successful when using dental material. Tooth shape (via dental topography) and size were quantified for four groups of South African Plio-Pleistocene hominins (specimens of Australopithecus africanus, Paranthropus robustus, H. naledi, and Homo sp.) on relatively unworn M2s to investigate possible ecological differentiation in H. naledi relative to taxa with similar known geographical ranges. H. naledi has smaller, but higher-crowned and more wear resistant teeth than Australopithecus and Paranthropus. These results are found in both lightly and moderately worn teeth. There are no differences in tooth sharpness or complexity. Combined with the high level of dental chipping in H. naledi, this suggests that, relative to Australopithecus and Paranthropus, H. naledi consumed foods with similar fracture mechanics properties but more abrasive particles (e.g., dust, grit), which could be due to a dietary and/or environmental shift(s). The same factors that differentiate H. naledi from Australopithecus and Paranthropus may also differentiate it from Homo sp., which geologically predates it, in the same way. Compared to the great apes, all hominins have sharper teeth, indicating they consumed foods requiring higher shear forces during mastication. Despite some anatomical similarities, H. naledi likely occupied a distinct ecological niche from the South African hominins that predate it

Dental macrowear reveals ecological diversity of Gorilla spp.

Size and shape variation of molar crowns in primates plays an important role in understanding how species adapted to their environment. Gorillas are commonly considered to be folivorous primates because they possess sharp cusped molars which are adapted to process fibrous leafy foods. However, the proportion of fruit in their diet can vary significantly depending on their habitats. While tooth morphology can tell us what a tooth is capable of processing, tooth wear can help us to understand how teeth have been used during mastication. The objective of this study is to explore if differences in diet at the subspecies level can be detected by the analysis of molar macrowear. We analysed a large sample of second lower molars of Grauer’s, mountain and western lowland gorilla by combining the Occlusal Fingerprint Analysis method with other dental measurements. We found that Grauer’s and western lowland gorillas are characterised by a macrowear pattern indicating a larger intake of fruit in their diet, while mountain gorilla’s macrowear is associated with the consumption of more folivorous foods. We also found that the consumption of herbaceous foods is generally associated with an increase in dentine and enamel wear, confirming the results of previous studies.This study was supported by the Australian Research Council (Grant Number: DP190100465), by the the French National Research Agency (Grant Number: ANR-17-CE02-0010-01), by the Primate Research Institute Cooperative Research Program (Grant Number: 2016-B-91), and by the Spanish Ministerio de Ciencia e Innovación (Grant Number: PID2020-114517GB-I00)

Functional and evolutionary consequences of cranial fenestration in birds

Ostrich-like birds (Palaeognathae) show very little taxonomic diversity while their sister taxon (Neognathae) contains roughly 10,000 species. The main anatomical differences between the two taxa are in the crania. Palaeognaths lack an element in the bill called the lateral bar that is present in both ancestral theropods and modern neognaths, and have thin zones in the bones of the bill, and robust bony elements on the ventral surface of their crania. Here we use a combination of modeling and developmental experiments to investigate the processes that might have led to these differences. Engineering-based finite element analyses indicate that removing the lateral bars from a neognath increases mechanical stress in the upper bill and the ventral elements of the skull, regions that are either more robust or more flexible in palaeognaths. Surgically removing the lateral bar from neognath hatchlings led to similar changes. These results indicate that the lateral bar is load-bearing and suggest that this function was transferred to other bony elements when it was lost in palaeognaths. It is possible that the loss of the load-bearing lateral bar might have constrained diversification of skull morphology in palaeognaths and thus limited taxonomic diversity within the group

Data from: Functional and evolutionary consequences of cranial fenestration in birds

Ostrich-like birds (Palaeognathae) show very little taxonomic diversity while their sister taxon (Neognathae) contains roughly 10000 species. The main anatomical differences between the two taxa are in the crania. Palaeognaths lack an element in the bill called the lateral bar that is present in both ancestral theropods and modern neognaths, have thin zones in the bones of the bill, and robust bony elements on the ventral surface of their crania. Here we use a combination of modelling and developmental experiments to investigate the processes that might have led to these differences. Engineering-based finite element analyses indicate that removing the lateral bars from a neognath increases mechanical stress in the upper bill and the ventral elements of the skull, regions that are either more robust or more flexible in palaeognaths. Surgically removing the lateral bar from neognath hatchlings led to similar changes. These results indicate that the lateral bar is load-bearing and suggest that this function was transferred to other bony elements when it was lost in palaeognaths. It is possible that the loss of the load-bearing lateral bar might have constrained diversification of skull morphology in palaeognaths and thus limited taxonomic diversity within the group

Gussekloo_etal_Fenestration_Evolution

Finite Element Models in ANSYS format and measurement data of the in vivo experiment. For further information, please see included README files