Primate dietary ecology in the context of food mechanical properties.

Substantial variation exists in the mechanical properties of foods consumed by primate species. This variation is known to influence food selection and ingestion among non-human primates, yet no large-scale comparative study has examined the relationships between food mechanical properties and feeding strategies. Here, we present comparative data on the Young's modulus and fracture toughness of natural foods in the diets of 31 primate species. We use these data to examine the relationships between food mechanical properties and dietary quality, body mass, and feeding time. We also examine the relationship between food mechanical properties and categorical concepts of diet that are often used to infer food mechanical properties. We found that traditional dietary categories, such as folivory and frugivory, did not faithfully track food mechanical properties. Additionally, our estimate of dietary quality was not significantly correlated with either toughness or Young's modulus. We found a complex relationship among food mechanical properties, body mass, and feeding time, with a potential interaction between median toughness and body mass. The relationship between mean toughness and feeding time is straightforward: feeding time increases as toughness increases. However, when considering median toughness, the relationship with feeding time may depend upon body mass, such that smaller primates increase their feeding time in response to an increase in median dietary toughness, whereas larger primates may feed for shorter periods of time as toughness increases. Our results emphasize the need for additional studies quantifying the mechanical and chemical properties of primate diets so that they may be meaningfully compared to research on feeding behavior and jaw morphology.

Dental microwear texture analysis of hominins recovered by the Olduvai Landscape Paleoanthropology Project, 1995-2007.

Dental microwear analysis has proven to be a valuable tool for the reconstruction of aspects of diet in early hominins. That said, sample sizes for some groups are small, decreasing our confidence that results are representative of a given taxon and making it difficult to assess within-species variation. Here we present microwear texture data for several new specimens of Homo habilis and Paranthropus boisei from Olduvai Gorge, bringing sample sizes for these species in line with those published for most other early hominins. These data are added to those published to date, and microwear textures of the enlarged sample of H. habilis (n = 10) and P. boisei (n = 9) are compared with one another and with those of other early hominins. New results confirm that P. boisei does not have microwear patterns expected of a hard-object specialist. Further, the separate texture complexity analyses of early Homo species suggest that Homo erectus ate a broader range of foods, at least in terms of hardness, than did H. habilis, P. boisei, or the "gracile" australopiths studied. Finally, differences in scale of maximum complexity and perhaps textural fill volume between H. habilis and H. erectus are noted, suggesting further possible differences between these species in diet.

Validation of bone surface modification models for inferring fossil hominin and carnivore feeding interactions, with reapplication to FLK 22, Olduvai Gorge, Tanzania.

Resolving the issue of how Early Stone Age hominins acquired large mammal carcasses requires information on their feeding interactions with large carnivores. This ecological information and its behavioral and evolutionary implications are revealed most directly from the tooth, cut, and percussion marks on bone surfaces generated by hominin and carnivore feeding activities. This paper employs a bootstrap method, a form of random resampling with replacement, to refine published neotaphonomic models that use the assemblage-wide proportions of long bones bearing feeding traces to infer the sequences in which Plio-Pleistocene hominins and carnivores accessed flesh, marrow, and/or grease from carcasses. Results validate the sensitivity of the models for inferring hominin feeding ecology, which have been questioned on grounds shown here to be unfounded. The bootstrapped feeding trace models are applied to the late Pliocene larger mammal fossil assemblage from FLK 22 (Zinjanthropus site), Olduvai Gorge, Tanzania. High frequencies of tooth and percussion marking on long bone midshaft fragments from FLK 22 are most consistent with those feeding trace models that simulate hominin scavenging from carcasses defleshed by carnivores, while cut mark data indicate that hominins more often had access to upper forelimb flesh than upper hind limb flesh. Together, the bone surface modification data indicate that hominins typically gained secondary access to partially defleshed carnivore kills, but they also allow for the possibility of some carcasses being processed only by carnivores and only by hominins.

Dental microwear texture and anthropoid diets.

Dental microwear has long been used as evidence concerning the diets of extinct species. Here, we present a comparative baseline series of dental microwear textures for a sample of 21 anthropoid primate species displaying interspecific and intraspecific dietary variability. Four dental microwear texture variables (complexity, anisotropy, textural fill volume, and heterogeneity) were computed based on scale-sensitive fractal analysis and high-resolution three-dimensional renderings of microwear surfaces collected using a white-light confocal profiler. The purpose of this analysis was to assess the extent to which these variables reflect variation in diet. Significant contrasts between species with diets known to include foods with differing material properties are clearly evident for all four microwear texture variables. In particular, species that consume more tough foods, such as leaves, tended to have high levels of anisotropy and low texture complexity. The converse was true for species including hard and brittle items in their diets either as staples or as fallback foods. These results reaffirm the utility of dental microwear texture analysis as an important tool in making dietary inferences based on fossil primate samples.

Dental microwear texture analysis shows within-species diet variability in fossil hominins.

Reconstructing the diets of extinct hominins is essential to understanding the paleobiology and evolutionary history of our lineage. Dental microwear, the study of microscopic tooth-wear resulting from use, provides direct evidence of what an individual ate in the past. Unfortunately, established methods of studying microwear are plagued with low repeatability and high observer error. Here we apply an objective, repeatable approach for studying three-dimensional microwear surface texture to extinct South African hominins. Scanning confocal microscopy together with scale-sensitive fractal analysis are used to characterize the complexity and anisotropy of microwear. Results for living primates show that this approach can distinguish among diets characterized by different fracture properties. When applied to hominins, microwear texture analysis indicates that Australopithecus africanus microwear is more anisotropic, but also more variable in anisotropy than Paranthropus robustus. This latter species has more complex microwear textures, but is also more variable in complexity than A. africanus. This suggests that A. africanus ate more tough foods and P. robustus consumed more hard and brittle items, but that both had variable and overlapping diets.

Craniofacial biomechanics and functional and dietary inferences in hominin paleontology.

Finite element analysis (FEA) is a potentially powerful tool by which the mechanical behaviors of different skeletal and dental designs can be investigated, and, as such, has become increasingly popular for biomechanical modeling and inferring the behavior of extinct organisms. However, the use of FEA to extrapolate from characterization of the mechanical environment to questions of trophic or ecological adaptation in a fossil taxon is both challenging and perilous. Here, we consider the problems and prospects of FEA applications in paleoanthropology, and provide a critical examination of one such study of the trophic adaptations of Australopithecus africanus. This particular FEA is evaluated with regard to 1) the nature of the A. africanus cranial composite, 2) model validation, 3) decisions made with respect to model parameters, 4) adequacy of data presentation, and 5) interpretation of the results. Each suggests that the results reflect methodological decisions as much as any underlying biological significance. Notwithstanding these issues, this model yields predictions that follow from the posited emphasis on premolar use by A. africanus. These predictions are tested with data from the paleontological record, including a phylogenetically-informed consideration of relative premolar size, and postcanine microwear fabrics and antemortem enamel chipping. In each instance, the data fail to conform to predictions from the model. This model thus serves to emphasize the need for caution in the application of FEA in paleoanthropological enquiry. Theoretical models can be instrumental in the construction of testable hypotheses; but ultimately, the studies that serve to test these hypotheses - rather than data from the models - should remain the source of information pertaining to hominin paleobiology and evolution.

Ingestive behaviors in bearded capuchins (Sapajus libidinosus).

The biomechanical and adaptive significance of variation in craniodental and mandibular morphology in fossil hominins is not always clear, at least in part because of a poor understanding of how different feeding behaviors impact feeding system design (form-function relationships). While laboratory studies suggest that ingestive behaviors produce variable loading, stress, and strain regimes in the cranium and mandible, understanding the relative importance of these behaviors for feeding system design requires data on their use in wild populations. Here we assess the frequencies and durations of manual, ingestive, and masticatory behaviors from more than 1400 observations of feeding behaviors video-recorded in a wild population of bearded capuchins (Sapajus libidinosus) at Fazenda Boa Vista in Piauí, Brazil. Our results suggest that ingestive behaviors in wild Sapajus libidinosus were used for a range of food material properties and typically performed using the anterior dentition. Coupled with previous laboratory work indicating that ingestive behaviors are associated with higher mandibular strain magnitudes than mastication, these results suggest that ingestive behaviors may play an important role in craniodental and mandibular design in capuchins and may be reflected in robust adaptations in fossil hominins.

Folivory or fruit/seed predation for Mesopithecus, an earliest colobine from the late Miocene of Eurasia?

Here we compare dental microwear textures from specimens of the fossil genus Mesopithecus (Cercopithecidae, Colobinae) from the late Miocene of Eastern Europe with dental microwear textures from four extant primate species with known dietary differences. Results indicate that the dental microwear textures of Mesopithecus differ from those of extant leaf eaters Alouatta palliata and Trachypithecus cristatus and instead resemble more closely those of the occasional hard-object feeders Cebus apella and Lophocebus albigena. Microwear texture data presented here in combination with results from previous analyses suggest that Mesopithecus was a widespread, opportunistic feeder that often consumed hard seeds. These data are consistent with the hypothesis that early colobines may have preferred hard seeds to leaves.

Trabecular Anisotropy in the Primate Mandibular Condyle Is Associated with Dietary Toughness.

Past attempts to establish a relationship between mandibular morphology and different dietary categories (e.g., frugivore, folivore, insectivore) have had mixed results, possibly because descriptive dietary categories are too broad and obscure variation within primate diets. Another potential reason is that not all aspects of skeletal architecture, especially trabecular anisotropy, have factored into functional assessments of dietary inputs into jaw form. Recent emphasis on quantifying food mechanical properties (FMPs) has provided an alternative to reliance on dietary categories. We used data on FMPs to test for correlations among dietary toughness and Young's modulus and the trabecular structure of the mandibular condyle, which is loaded during feeding and should reflect differences in masticatory stresses associated with different dietary FMPs. Adult primate mandibles from 11 species were imaged using high-resolution X-ray computed tomography, and trabecular structure was analyzed with BoneJ and Quant3D to assess common three-dimensional trabecular parameters. Results of phylogenetic generalized least squares analysis suggested a positive correlation between the degree of anisotropy (DA) and toughness, and weaker correlations between FMPs and various other trabecular variables. Because the DA contributes to the mechanical properties of bone, these results suggest a functional relationship between dietary toughness and trabecular anisotropy in the mandibular condyle. Such a perspective underscores the need to consider all aspects of skeletal morphology in evaluating the links between diet and jaw biomechanics. Anat Rec, 2018. © 2018 Wiley Periodicals, Inc.

Molar microwear textures and the diets of Australopithecus anamensis and Australopithecus afarensis.

Many researchers have suggested that Australopithecus anamensis and Australopithecus afarensis were among the earliest hominins to have diets that included hard, brittle items. Here we examine dental microwear textures of these hominins for evidence of this. The molars of three Au. anamensis and 19 Au. afarensis specimens examined preserve unobscured antemortem microwear. Microwear textures of these individuals closely resemble those of Paranthropus boisei, having lower complexity values than Australopithecus africanus and especially Paranthropus robustus. The microwear texture complexity values for Au. anamensis and Au. afarensis are similar to those of the grass-eating Theropithecus gelada and folivorous Alouatta palliata and Trachypithecus cristatus. This implies that these Au. anamensis and Au. afarensis individuals did not have diets dominated by hard, brittle foods shortly before their deaths. On the other hand, microwear texture anisotropy values for these taxa are lower on average than those of Theropithecus, Alouatta or Trachypithecus. This suggests that the fossil taxa did not have diets dominated by tough foods either, or if they did that directions of tooth-tooth movement were less constrained than in higher cusped and sharper crested extant primate grass eaters and folivores.

Dental microwear texture analysis: technical considerations.

Dental microwear analysis is commonly used to infer aspects of diet in extinct primates. Conventional methods of microwear analysis have usually been limited to two-dimensional imaging studies using a scanning electron microscope and the identification of apparent individual features. These methods have proved time-consuming and prone to subjectivity and observer error. Here we describe a new methodological approach to microwear: dental microwear texture analysis, based on three-dimensional surface measurements taken using white-light confocal microscopy and scale-sensitive fractal analysis. Surface parameters for complexity, scale of maximum complexity, anisotropy, heterogeneity, and textural fill volume offer repeatable, quantitative characterizations of three-dimensional surfaces, free of observer measurement error. Some results are presented to illustrate how these parameters distinguish extant primates with different diets. In this case, microwear surfaces of Cebus apella and Lophocebus albigena, which consume some harder food items, have higher average values for complexity than do folivores or soft fruit eaters.