Hard-object Feeding Research Articles

Tooth chipping patterns and dental caries suggest a soft fruit diet in early anthropoids.

Fossils from the Fayum Depression, Egypt, are crucial for understanding anthropoid evolution due to the abundance of taxa and the time interval they represent (late Eocene to early Oligocene). Dietary and foraging behavioral interpretations suggest fruits were their dominant food source, although hard foods (e.g., seeds and nuts) and leaves could have been important dietary components for particular groups. In this study, we compare dental chipping patterns in five Fayum primate genera with chipping data for extant primates, to assess potential hard object feeding in early anthropoids. Original specimens were studied (Aegyptopithecus: n = 100 teeth; Parapithecus: n = 72, Propliopithecus: n = 99, Apidium: n = 82; Catopithecus: n = 68); with the number, severity, and position of chips recorded. Dental caries was also recorded, due to its association with soft fruit consumption in extant primates. Tooth chipping was low across all five genera studied, with a pooled chipping prevalence of 5% (21/421). When split into the three anthropoid families represented, chipping prevalence ranged from 2.6% (4/154) in Parapithecidae, 6% (12/199) in Propliopithecidae, and 7.4% (5/68) in Oligopithecidae. Three carious lesions were identified in Propliopithecidae. The chipping prevalence is low when compared to extant anthropoids (range from 4% to 40%) and is consistent with a predominantly soft fruit diet, but not with habitual hard food mastication. The presence of caries supports consumption of soft, sugary fruits, at least in Propliopithecidae. Our results add support for low dietary diversity in early anthropoids, with soft fruits as likely dominant food sources.

How mangabey molar form differs under routine vs. fallback hard-object feeding regimes

Background Components of diet known as fallback foods are argued to be critical in shaping primate dental anatomy. Such foods of low(er) nutritional quality are often non-preferred, mechanically challenging resources that species resort to during ecological crunch periods. An oft-cited example of the importance of dietary fallbacks in shaping primate anatomy is the grey-cheeked mangabey Lophocebus albigena. This species relies upon hard seeds only when softer, preferred resources are not available, a fact which has been linked to its thick dental enamel. Another mangabey species with thick enamel, the sooty mangabey Cercocebus atys, processes a mechanically challenging food year-round. That the two mangabey species are both thickly-enameled suggests that both fallback and routine consumption of hard foods are associated with the same anatomical feature, complicating interpretations of thick enamel in the fossil record. We anticipated that aspects of enamel other than its thickness might differ between Cercocebus atys and Lophocebus albigena. We hypothesized that to function adequately under a dietary regime of routine hard-object feeding, the molars of Cercocebus atys would be more fracture and wear resistant than those of Lophocebus albigena. Methods Here we investigated critical fracture loads, nanomechanical properties of enamel, and enamel decussation in Cercocebus atys and Lophocebus albigena. Molars of Cercopithecus, a genus not associated with hard-object feeding, were included for comparison. Critical loads were estimated using measurements from 2D µCT slices of upper and lower molars. Nanomechanical properties (by nanoindentation) and decussation of enamel prisms (by SEM-imaging) in trigon basins of one upper second molar per taxon were compared. Results Protocone and protoconid critical fracture loads were significantly greater in Cercocebus atys than Lophocebus albigena and greater in both than in Cercopithecus. Elastic modulus, hardness, and elasticity index in most regions of the crown were greater in Cercocebus atys than in the other two taxa, with the greatest difference in the outer enamel. All taxa had decussated enamel, but that of Cercocebus atys uniquely exhibited a bundle of transversely oriented prisms cervical to the radial enamel. Quantitative comparison of in-plane and out-of-plane prism angles suggests that decussation in trigon basin enamel is more complex in Cercocebus atys than it is in either Lophocebus albigena or Cercopithecus cephus. These findings suggest that Cercocebus atys molars are more fracture and wear resistant than those of Lophocebus albigena and Cercopithecus. Recognition of these differences between Cercocebus atys and Lophocebus albigena molars sharpens our understanding of associations between hard-object feeding and dental anatomy under conditions of routine vs. fallback hard-object feeding and provides a basis for dietary inference in fossil primates, including hominins.

Aspects of molar form and dietary proclivities of African colobines

This study investigates aspects of molar form in three African colobine species: Colobus polykomos, Colobus angolensis, and Piliocolobus badius. Our samples of C. polykomos and P. badius are from the Taï Forest, Ivory Coast; our sample of C. angolensis is from Diani, Kenya. To the extent that protective layers surrounding seeds are hard, we predicted that molar features related to hard-object feeding would be more pronounced in Colobus than they are Piliocolobus, as seed-eating generally occurs at higher frequencies in species of the former. We further predicted that among the colobines we studied, these features would be most pronounced in Taï Forest C. polykomos, which feeds on Pentaclethra macrophylla seeds encased within hard and tough seed pods. We compared overall enamel thickness, enamel thickness distribution, absolute crown strength, cusp tip geometry, and flare among molar samples. Sample sizes per species and molar type varied per comparison. We predicted differences in all variables except overall enamel thickness, which we expected would be invariant among colobines as a result of selection for thin enamel in these folivorous species. Of the variables we examined, only molar flare differed significantly between Colobus and Piliocolobus. Our findings suggest that molar flare, an ancient feature of cercopithecoid molars, was retained in Colobus but not in Piliocolobus, perhaps as a result of differences in the seed-eating proclivities of the two genera. Contrary to predictions, none of the aspects of molar form we investigated tracked current dietary differences in seed-eating between the two Colobus species. Finally, we explored the possibility that molar flare and absolute crown strength, when analyzed together, might afford greater differentiation among these colobine species. A multivariate t test of molar flare and absolute crown strength differentiated C. polykomos and P. badius, possibly reflecting known niche divergence between these two sympatric Taï Forest species.

Dental topography and dietary specialization in Papionini primates

Our understanding of primate adaptive evolution depends on appreciating the way in which dental functional morphology affects food processing. The Papionini tribe ofCercopithecoideaprimates shows great dietary versatility and ecological adaptations to resource seasonality across the African and Asian ecosystems, however, there are few studies focusing on the occlusal topography of the bilophodont teeth and the effect of tooth wear in the crown shape. Here, we explore the relationship between wear-related dental functional morphology and dietary ecological constraints within the Papionini. Three-dimensional (3D) polygonal meshes of the upper permanent molar row (M1-3) were obtained in a large papionine sample (838 specimens) of known dietary preferences including species from six genera (Cercocebus,Lophocebus,Macaca,Mandrillus,Papio, andTheropithecus). All the sample was classified in four diet categories and four topographic metrics (orientation patch count rotated, OPCR; Dirichlet normal energy, DNE; occlusal relief, OR; and ambient occlusion, portion de ciel visible, PCV) were measured for each tooth-type according to wear stage (lightly and moderately worn) to determine diet-related interspecific morphological changes with long-term functionality. The results indicate that hard-object feeders (CercocebusandLophocebus) and grass eaters (Theropithecus gelada) exhibit a pattern of occlusal complexity (OPCR), surface curvature (DNE), relief (OR), and morphological wear resistance (PCV) that is significantly different from the omnivores and folivore-frugivore species (MandrillusandMacaca) despite the overall homogeneity of the bilophodont dentition. A multifactorial ANOVA showed that the topographic metrics were sensitive to tooth wear as expected. The results also indicate that the interspecific variability of dental topography of the upper molars reflects dietary specializations rather than phylogenetic proximity. These findings support the hypothesis that evolutionary convergence processes could have affected the Papionini, clustering the hard-object feeders (LophocebusandCercocebus)together in the morphospace, and clearly discriminating this group from the graminivorous and frugivores-folivores.

Controlled feeding experiments with juvenile alligators reveal microscopic dental wear texture patterns associated with hard-object feeding

Dental wear analyses are classically applied to mammals because they have evolved heterodont dentitions for sophisticated mastication. Recently, several studies have shown a correlation between pre-assigned and analytically inferred diet preferences in extant reptiles through dental microwear texture analysis (DMTA), a method using quantitative assessment of microscopic wear marks to reconstruct the diet material properties. The first tentative applications of DMTA to extinct reptiles have followed. However, for large and small mammals, microwear analyses have undergone a long time of ground-truthing through direct feeding observations, stomach content analyses, and feeding experiments. Such data are currently lacking for reptiles, but are necessary to further extend DMTA, especially to Archosauria, as the application to dinosaurs could be of great interest to the scientific community. We herein present data from a pilot feeding experiment with five juvenile American alligators (Alligator mississippiensis). Each individual received a diet of assumed different hardness for ~4 months: crocodylian pellets (control), sardines, quails, rats, or crawfish. All individuals initially received the same pellet diet, and we found them to show similar dental microwear texture patterns before they were switched to their designated experimental diet. From the first feeding bout on, dental microwear textures differed across the diets. The crawfish-feeder showed consistently higher surface complexity, followed by the rat-feeder. Quail- and fish-feeding resulted in similar wear signatures, with low complexity. Fast tooth replacement and selective tooth use likely affected microwear formation, but we were able to detect a general hard (crawfish and rat) versus soft (quail and fish) DMTA signature. Such patterns can support the identification of hard-object feeding in the fossil record.

Fine-scaled climate variation in equatorial Africa revealed by modern and fossil primate teeth

Variability in resource availability is hypothesized to be a significant driver of primate adaptation and evolution, but most paleoclimate proxies cannot recover environmental seasonality on the scale of an individual lifespan. Oxygen isotopecompositions (δ18O values) sampled at high spatial resolution in the dentitions of modern African primates (n = 2,352 near weekly measurements from 26 teeth) track concurrent seasonal precipitation, regional climatic patterns, discrete meteorological events, and niche partitioning. We leverage these data to contextualize the first δ18O values of two 17 Ma Afropithecus turkanensis individuals from Kalodirr, Kenya, from which we infer variably bimodal wet seasons, supported by rainfall reconstructions in a global Earth system model. Afropithecus' δ18O fluctuations are intermediate in magnitude between those measured at high resolution in baboons (Papio spp.) living across a gradient of aridity and modern forest-dwelling chimpanzees (Pan troglodytes verus). This large-bodied Miocene ape consumed seasonally variable food and water sources enriched in 18O compared to contemporaneous terrestrial fauna (n = 66 fossil specimens). Reliance on fallback foods during documented dry seasons potentially contributed to novel dental features long considered adaptations to hard-object feeding. Developmentally informed microsampling recovers greater ecological complexity than conventional isotope sampling; the two Miocene apes (n = 248 near weekly measurements) evince as great a range of seasonal δ18O variation as more time-averaged bulk measurements from 101 eastern African Plio-Pleistocene hominins and 42 papionins spanning 4 million y. These results reveal unprecedented environmental histories in primate teeth and suggest a framework for evaluating climate change and primate paleoecology throughout the Cenozoic.

3D surface scan dataset for the publication "Controlled feeding experiments with juvenile alligators reveal microscopic dental wear patterns associated with hard-object feeding"

3D surface scan dataset for the publication "Controlled feeding experiments with juvenile alligators reveal microscopic dental wear patterns associated with hard-object feeding"

Sagittal crest morphology decoupled from relative bite performance in Pleistocene tapirs (Perissodactyla: Tapiridae).

Bite force is often associated with specific morphological features, such as sagittal crests. The presence of a pronounced sagittal crest in some tapirs (Perissodactyla: Tapiridae) was recently shown to be negatively correlated with hard-object feeding, in contrast with similar cranial structures in carnivorans. The aim of this study was to investigate bite forces and sagittal crest heights across a wide range of modern and extinct tapirs and apply a comparative investigation to establish whether these features are correlated across a broad phylogenetic scope. We examined a sample of 71 specimens representing 15 tapir species (5 extant, 10 extinct) using the dry-skull method, linear measurements of cranial features, phylogenetic reconstruction, and comparative analyses. Tapirs were found to exhibit variation in bite force and sagittal crest height across their phylogeny and between different biogeographical realms, with high-crested morphologies occurring mostly in Neotropical species. The highest bite forces within tapirs appear to be driven by estimates for the masseter-pterygoid muscle complex, rather than predicted forces for the temporalis muscle. Our results demonstrate that relative sagittal crest height is poorly correlated with relative cranial bite force, suggesting high force application is not a driver for pronounced sagittal crests in this sample. The divergent biomechanical capabilities of different contemporaneous tapirids may have allowed multiple species to occupy overlapping territories and partition resources to avoid excess competition. Bite forces in tapirs peak in Pleistocene species, independent of body size, suggesting possible dietary shifts as a potential result of climatic changes during this epoch.

Biomechanical correlates of zygomaxillary-surface shape in papionin primates and the effects of hard-object feeding on mangabey facial form

Extant African papioninans are distinguished from macaques by the presence of excavated facial fossae; however, facial excavation differs among taxa. Mangabeys (Cercocebus, Rungwecebus, and Lophocebus) exhibit fossae that invade the zygomatic forming pronounced suborbital fossae (SOFs). Larger-bodied Papio, Mandrillus, and Theropithecus have lateral rostral fossae with minimal/absent suborbital fossae. Because prior studies have shown that mangabeys exhibit adaptations to anterior dental loading (e.g., palatal retraction), it is plausible that mangabey SOFs represent structural accommodation to masticatory-system shape rather than facial allometry, as commonly hypothesized. We analyzed covariation between zygomaxillary-surface shape, masticatory-system shape, and facial size in 141 adult crania of Macaca fascicularis, Papio kindae, Cercocebus, and Lophocebus. These taxa represent the range of papionin SOF expression while minimizing size variation (narrow allometry). Masticatory-system landmarks (39) registered palate shape, bite points, masticatory muscle attachments, and the temporomandibular joint. Semilandmarks (450) captured zygomaxillary-surface shape. Following Procrustes superimposition with semilandmark sliding and principal components analyses, multivariate regression was used to explore allometry, and two-block partial least-squares analyses (within-configuration and separate-blocks) were used to examine covariation patterns. Scores on principal components 1–2 and the first partial least-square (PLS1) separate mangabeys from Macaca and Papio. Both zygomaxillary-surface shape and masticatory-system shape are correlated with size within taxa and facial morphotypes; however, regression distributions indicate morphotype shape differences are non-allometric. PLS1 accounts for ∼95% of shape covariance (p < 0.0001) and shows strong linear correlations (r-PLS = ∼0.95, p < 0.0001) between blocks. Negative PLS1 scores in mangabeys reflect deep excavation of the suborbital malar surface, palatal retraction, and anterior displacement of jaw adductor muscles and the temporomandibular joint. Neither PC1 nor PLS1 scores ordinate specimens by facial size. Taken together, these results fail to support the allometric hypothesis but suggest that mangabey zygomaxillary morphology is closely linked with adaptations to hard-object feeding.

Dental chipping supports lack of hard-object feeding in Paranthropus boisei

Dental chipping supports lack of hard-object feeding in Paranthropus boisei

Quantification of enamel decussation in capuchin monkeys

Introduction & Objective – An intricate suite of highly complex enamel has led robust capuchin monkeys to be uniquely capable in breaking down hard-object foods as compared to other cebid monkeys, and in routinely breaking down extremely mechanically challenging foods as compared to other capuchins. The goal of this study is to compare dental enamel complexity in images of dental thin sections from a sample of robust vs. gracile capuchins using Image Compression Ratio (ICR) analysis. We hypothesize that: 1) robust capuchins will exhibit an overall pattern of higher ICR, and therefore more complex enamel, compared to gracile capuchins, and 2) the difference in ICR between the two groups will be greater for anterior as compared to posterior teeth. Materials and Methods – 113 images of thin sections from three gracile capuchin species and from unidentified species of robust capuchins, Sapajus spp were captured using a Leica-Leitz DMRX/E Universal Microscope configured with a Marzhauser motorized stage, phase contrast, and circularly polarizing filters. The images were then digitally processed to delete non-enamel components from the images and any other factor that can affect the compression ratio. After various compressions, the ICR values were compared across our images using standard non-parametric Kruskal-Wallis and Mann Whitney tests. Results – As for the first hypothesis, we found no significant difference in ICR between robust and gracile capuchins across all teeth (p = 0.35) using Mann-Whitney U tests. Likewise, we found no significant difference in ICR for all tooth categories when all gracile capuchin species were examined separately rather than assessed as one category (Kruskal Wallis, p = 0.48), nor was there a significant difference among tooth types (canines, premolars, and molars) in the total dataset (p = 0.36). As for the second hypothesis, we separately compared anterior (canines) and posterior teeth (premolars and molars) across the same taxonomic groupings. We once again found no difference in ICR between robust and gracile capuchins for the posterior dentition (p = 0.28). However, the anterior tooth sample did display a significant difference between the two groups (p = 0.04). Conclusion – Our ICR results indicate that premolars and molars among gracile and robust capuchins exhibit similar degrees of enamel complexity, but that canines in robust capuchins display increased complexity reflecting high specialization in this group for hard object feeding in the anterior masticatory apparatus. Significance/Implication – These findings provide further evidence to the hypothesis that the need to break apart obdurate foodstuffs with the anterior – as opposed to posterior – teeth has driven much of the specialized craniodental anatomy seen in robust capuchins. Future work including further sampling may help assess the degree in which different species of robust capuchins differ from one another. The findings of this research may also lead us to assess differences among the robust species, considering previous studies suggesting that tool use in some species may lessen the need for generation of high anterior bite forces compare capuchins to other primate hard object feeders such as the pitheciines and Paranthropus, the latter of which would be particularly useful for assessing patterns of hard object feeding in hominin evolution.

Dietary ecology of Alaskan polar bears (Ursus maritimus) through time and in response to Arctic climate change.

Arctic climate change poses serious threats to polar bears (Ursus maritimus) as reduced sea ice makes seal prey inaccessible and marine ecosystems undergo bottom-up reorganization. Polar bears' elongated skulls and reduced molar dentition, as compared to their sister species the grizzly bear (Ursus arctos), are adaptations associated with hunting seals on sea ice and a soft, lipid-rich diet of blubber and meat. With significant declines in sea ice, it is unclear if and how polar bears may be altering their diets. Clarifying polar bear dietary responses to changing climates, both today and in the past, is critical to proper conservation and management of this apex predator. This is particularly important when a dietary strategy may be maladaptive. Here, we test the hypothesis that hard-food consumption (i.e., less preferred foods including bone), inferred from dental microwear texture analysis, increased with Arctic warming. We find that polar bears demonstrate a conserved absence of hard-object feeding in Alaska through time (including approximately 1000years ago), until the 21st century, consistent with a highly conserved and specialized diet of soft blubber and flesh. Notably, our results also suggest that some 21st-century polar bears may be consuming harder foods (e.g., increased carcass utilization, terrestrial foods including garbage), despite having skulls and metabolisms poorly suited for such a diet. Prior to the 21st century, only polar bears with larger mandibles demonstrated increased hard-object feeding, though to a much lower degree than closely related grizzly bears which regularly consume mechanically challenging foods. Polar bears, being morphologically specialized, have biomechanical constraints which may limit their ability to consume mechanically challenging diets, with dietary shifts occurring only under the most extreme scenarios. Collectively, the highly specialized diets and cranial morphology of polar bears may severely limit their ability to adapt to a warming Arctic.

Quantification of enamel decussation in gracile and robust capuchins (Cebus, Sapajus, Cebidae, Platyrrhini).

Multiple behavioral and biomechanical analyses have demonstrated that capuchin monkeys (Cebus and Sapajus) are specialized for breaking down hard-object foods as compared to other cebid monkeys. In addition to a complex suite of craniodental adaptations, it has specifically been demonstrated that capuchins possess highly complex dental enamel, with extensive Hunter-Schreger banding and other decussation, that likely serve as an adaptation to resist crack propagation during hard-object feeding. Furthermore, it has been demonstrated that robust capuchins (Sapajus spp., formerly Cebus apella) demonstrate further adaptation for hard-object feeding than other capuchins, routinely breaking down extremely mechanically challenging foods. However, there has been no comparison of dental enamel complexity in robust versus gracile capuchins, to assess whether the dental enamel in Sapajus follows this same pattern of further specialization. Therefore, this study compares dental enamel complexity in images of dental thin sections from a sample of robust versus gracile capuchins using image compression ratio (ICR) analysis. ICR is a variable that correlates with enamel complexity, such that higher ICR values are indicative of increased complexity in the form of enamel decussation. We found no significant difference between robust and gracile capuchins when assessing all teeth in our sample together, however, we did find that robust capuchins have significantly higher ICR values than gracile capuchins for canine teeth, specifically. Our results support prior studies suggesting that robust capuchins are specialized to generate increased masticatory loads with their anterior dentition, specifically, as compared to gracile species.

Sumatran orangutan diets in the Late Pleistocene as inferred from dental microwear texture analysis

Orangutan (Pongo spp.) fossils have been found throughout much of Pleistocene Southeast Asia. Today, Pongo is restricted to three living species on the islands of Borneo and Sumatra. They are predominately frugivores, and during fruit masting events their diets can consist entirely of fruit. However, food procurement strategies between masting events differ between islands and thus species. Sumatran orangutans (Pongo abelii) are largely able to fall back on non-masting fruit species, such as figs, such that they can maintain higher quality diets compared to their Bornean counterparts (Pongo pygmaeus), who ingest more lower quality and tougher foods between masting events. However, diets of fossil orangutans remain largely unknown. We investigated the diets of Sumatran orangutans from the Pleistocene deposits of the Padang Highlands, western Sumatra, using dental microwear texture analysis. We tested whether the diets of the fossil species were similar to those of Bornean orangutans, all great apes generally, and macaques that occupy similar habitats to modern orangutans, in order to gain insights into palaeoenvironments present in western Sumatra. We found that fossil orangutans consumed foods less tough than modern Bornean Pongo pygmaeus, and thus presumably more fleshy fruits (absent of significant seed mastication). We found no differences in the amount of hard object feeding between modern and fossil orangutans. Taken together, we suggest that Pleistocene orangutans from western Sumatra had similar diets to Pongo abelii, indicating that masting events were not a significant contributing factor in their intake of fruit. Finally, we suggest anisotropy values in orangutan dental microwear may be an important marker for fruit masting events in the Pleistocene.

Clarifying relationships between cranial form and function in tapirs, with implications for the dietary ecology of early hominins

Paleontologists and paleoanthropologists have long debated relationships between cranial morphology and diet in a broad diversity of organisms. While the presence of larger temporalis muscle attachment area (via the presence of sagittal crests) in carnivorans is correlated with durophagy (i.e. hard-object feeding), many primates with similar morphologies consume an array of tough and hard foods—complicating dietary inferences of early hominins. We posit that tapirs, large herbivorous mammals showing variable sagittal crest development across species, are ideal models for examining correlations between textural properties of food and sagittal crest morphology. Here, we integrate dietary data, dental microwear texture analysis, and finite element analysis to clarify the functional significance of the sagittal crest in tapirs. Most notably, pronounced sagittal crests are negatively correlated with hard-object feeding in extant, and several extinct, tapirs and can actually increase stress and strain energy. Collectively, these data suggest that musculature associated with pronounced sagittal crests—and accompanied increases in muscle volume—assists with the processing of tough food items in tapirs and may yield similar benefits in other mammals including early hominins.

Enamel chipping in Taï Forest cercopithecids: Implications for diet reconstruction in paleoanthropological contexts

Antemortem enamel chipping in living and fossil primates is often interpreted as evidence of hard-object feeding (i.e., ‘durophagy’). Laboratory analyses of tooth fracture have modeled the theoretical diets and loading conditions that may produce such chips. Previous chipping studies of nonhuman primates tend to combine populations into species samples, despite the fact that species can vary significantly in diet across their ranges. Chipping is yet to be analyzed across population-specific species samples for which long-term dietary data are available. Here, we test the association between enamel chipping and diet in a community of cercopithecid primates inhabiting the Taï Forest, Ivory Coast. We examined fourth premolars and first molars (n = 867) from naturally deceased specimens of Cercocebus atys, Colobus polykomos, Piliocolobus badius,Procolobus verus, and three species of Cercopithecus. We found little support for a predictive relationship between enamel chipping and diet across the entire Taï monkey community. Cercocebus atys, a dedicated hard-object feeder, exhibited the highest frequencies of (1) chipped teeth and (2) chips of large size; however, the other monkey with a significant degree of granivory, Co. polykomos, exhibited the lowest chip frequency. In addition, primates with little evidence of mechanically challenging or hard-food diets—such as Cercopithecus spp., Pi. badius, and Pr. verus—evinced higher chipping frequencies than expected. The equivocal and stochastic nature of enamel chipping in the Taï monkeys suggests nondietary factors contribute significantly to chipping. A negative association between canopy preference and chipping suggests a role of exogenous particles in chip formation, whereby taxa foraging closer to the forest floor encounter more errant particulates during feeding than species foraging in higher strata. We conclude that current enamel chipping models may provide insight into the diets of fossil primates, but only in cases of extreme durophagy. Given the role of nondietary factors in chip formation, our ability to reliably reconstruct a range of diets from a gradient of chipping in fossil taxa is likely weak.

Unexpected hard-object feeding in Western lowland gorillas.

Gorilla diets are characterized by large amounts of fruit and tough fibrous plant material. Hard-object feeding is not generally associated with this genus as the high crests on their molar teeth would be at risk of damage from the mechanically challenging woody endocarp. This study aims to demonstrate that at least one population of western lowland gorillas are seasonal hard-object feeders, orally processing the seeds of Coula edulis. Feeding behavior of habituated western lowland gorillas and phenology of fruiting trees was observed over a 4-year period to determine the extent they exploited the seeds of C. edulis. Additionally, the endocarps of C. edulis were subjected to testing to determine their mechanical properties. Our results demonstrate that during the fruiting season (January, February, and December) gorillas consistently opened the seeds of C. edulis using their postcanine dentition. The protective endocarp is composed of a very stiff material, presenting a substantial mechanical challenge to a gorilla. However, the high ratio between elastic modulus and toughness will facilitate brittle, cataclysmic fracture of the seed shell given a high enough load. Although a rich energy source, C. edulis likely tax gorilla dentitions to their upper limit. The rarity of such behavior at sites where it could be observed may indicate a degree of social learning or culture driving its occurrence. This shows a greater breadth of gorilla diets than previously described and suggests gorillas may be a useful model for interpreting the dietary mechanics that necessitated robust craniodental morphology in australopiths.

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.

Primitive Old World monkey from the earliest Miocene of Kenya and the evolution of cercopithecoid bilophodonty

Old World monkeys (Cercopithecoidea) are a highly successful primate radiation, with more than 130 living species and the broadest geographic range of any extant group except humans. Although cercopithecoids are highly variable in habitat use, social behavior, and diet, a signature dental feature unites all of its extant members: bilophodonty (bi: two, loph: crest, dont: tooth), or the presence of two cross-lophs on the molars. This feature offers an adaptable Bauplan that, with small changes to its individual components, permits its members to process vastly different kinds of food. Old World monkeys diverged from apes perhaps 30 million years ago (Ma) according to molecular estimates, and the molar lophs are sometimes incompletely developed in fossil species, suggesting a mosaic origin for this key adaptation. However, critical aspects of the group's earliest evolution remain unknown because the cercopithecoid fossil record before ∼18 Ma consists of only two isolated teeth, one from Uganda and one from Tanzania. Here we describe a primitive Old World monkey from Nakwai, Kenya, dated at ∼22 Ma, that offers direct evidence for the initial key steps in the evolution of the cercopithecoid dentition. The simple dentition and absence of bilophodonty in the Nakwai monkey indicate that the initial radiation of Old World monkeys was first characterized by a reorganization of basic molar morphology, and a reliance on cusps rather than lophs suggests frugivorous diets and perhaps hard object feeding. Bilophodonty evolved later, likely in response to the inclusion of leaves in the diet.

New Specimens of the Late Cretaceous Metatherian Eodelphis and the Evolution of Hard-Object Feeding in the Stagodontidae

The Stagodontidae include the largest metatherians known from the Cretaceous of North America. Of the recognized species of the stagodontid genus Eodelphis, E. cutleri is larger and has a more robust dentary, more inflated premolars, and third premolars specialized for crushing, as opposed to the more gracile E. browni. These differences have led to the hypothesis that an E. cutleri-like ancestor gave rise to Didelphodon—another, mostly younger, stagodontid, which has been interpreted as a durophagous predator-scavenger. If correct, E. cutleri would be expected to show more morphological adaptation toward durophagy than E. browni does. Here, we describe two new dentary fossils referable to E. browni and test the evolutionary hypothesis by applying beam theory to estimate bending force capabilities of 22 dentaries of Cretaceous stagodontids and other metatherians. The resulting diversity of bending force profiles of the sampled dentaries implies that Cretaceous metatherians had a wide range of feeding behaviors. Among the stagodontids, E. cutleri has a mediolateral bending force profile of the dentary that is more similar to that of Didelphodon than it is to that of E. browni; whereas its dorsoventral bending force profile is more similar to that of E. browni. These results indicate that anteriorly the dentary of E. cutleri was capable of resisting high torsional stresses from hard-object feeding but lacked other dorsoventral buttressing associated with exceptionally high bite forces of Didelphodon. Our results imply that some morphological changes associated with durophagy evolved twice within this clade, independently in E. cutleri and Didelphodon.