Footprint evidence of early hominin locomotor diversity at Laetoli, Tanzania.

Bipedal trackways discovered in 1978 at Laetoli site G, Tanzania and dated to 3.66 million years ago are widely accepted as the oldest unequivocal evidence of obligate bipedalism in the human lineage1-3. Another trackway discovered two years earlier at nearby site A was partially excavated and attributed to a hominin, but curious affinities with bears (ursids) marginalized its importance to the paleoanthropological community, and the location of these footprints fell into obscurity3-5. In 2019, we located, excavated and cleaned the site A trackway, producing a digital archive using 3D photogrammetry and laser scanning. Here we compare the footprints at this site with those of American black bears, chimpanzees and humans, and we show that they resemble those of hominins more than ursids. In fact, the narrow step width corroborates the original interpretation of a small, cross-stepping bipedal hominin. However, the inferred foot proportions, gait parameters and 3D morphologies of footprints at site A are readily distinguished from those at site G, indicating that a minimum of two hominin taxa with different feet and gaits coexisted at Laetoli.

Enamel chipping and its ecological correlates in African papionins: Implications for hominin feeding behavior.

African papionins are classic paleoecological referents for fossil hominins. Enamel chips on the teeth of baboons and hominins are argued to represent responses to similar dietary habits; however, a comprehensive analysis of modern papionin chipping is lacking, leaving open the question of analog suitability. Here, we investigate patterns of antemortem enamel chipping across a diverse set of African papionin species occupying a range of ecological niches. We compare papionin chipping frequencies to estimates for Plio-Pleistocene hominins to address hypotheses of habitat and/or dietary similarities. Antemortem chips in seven African papionin species were scored on intact postcanine teeth (P3-M3) using established protocols. Chip size was scored on a tripartite scale. Papio hamadryas and Papio ursinus-two common paleoecological referents-display higher levels of chipping than Plio-Pleistocene hominin taxa (Australopithecus and Paranthropus) posited to have similar dietary habits. Papio populations occupying dry or highly seasonal habitats accumulate more large chips than Papio taxa occupying more mesic habitats, and terrestrial papionins chip their teeth more often than closely related taxa occupying arboreal niches. Chipping is present on the teeth of all Plio-Pleistocene hominins; however, chipping in baboons (P. ursinus and P. hamadryas) consistently exceeds most hominin taxa. Chipping frequencies on their own do not reliably sort taxa into major dietary groupings. We conclude that the large differences in chipping frequency may instead reflect habitat use and food processing idiosyncrasies. Less chipping in Plio-Pleistocene hominin teeth compared to modern Papio is more likely attributable to differences in dental morphology rather than diet.

Grit and consequence.

Grit is implicated in several biological phenomena-it wears teeth, it fractures teeth, it drives tooth evolution, it elicits complex manual manipulations-any one of which could be described as a central topic in evolutionary anthropology. But what is grit? We hardly know because we tend to privilege the consequences of grit (it is abrasive) over its formal features, all but ignoring crucial variables such as mineral composition, material properties, and particle geometry (size, angularity), not to mention natural variation in the habitats of primates and their food surfaces. Few topics have animated so much debate and invited such cool indifference at the same time. Our goal here is to shine a light on grit, to put a philosophical lens on the nature of our discourse, and to call attention to large empirical voids that should be filled and folded into our understanding of primate natural history and evolution.

Oral processing, sexual selection, and size variation in the circumorbital region of Colobus and Piliocolobus.

OBJECTIVES: The function of the browridge in primates is a subject of enduring debate. Early studies argued for a role in resisting masticatory stresses, but recent studies have suggested sexual signaling as a biological role. We tested associations between circumorbital form, diet, oral processing, and social behavior in two species of colobus monkey-the king colobus (Colobus polykomos) and western red or bay colobus (Piliocolobus badius). MATERIALS AND METHODS: We quantified circumorbital size and dimorphism in a sample of 98 crania. Controlling for age and facial size, we tested whether variation in circumorbital morphology can be explained by variation in diet, oral processing behavior, masticatory muscle size, and mating system. To contextualize our results, we included a broader sample of facial dimorphism for 67 anthropoid species. RESULTS: Greater circumorbital thickness is unrelated to the stresses of food processing. King colobus engages in longer bouts of anterior tooth use, chews more per ingestive event, and processes a tougher diet, yet circumorbital thickness of C. polykomos is reduced compared to P. badius. Differences in circumorbital development do not vary with wear or facial size. Greater sexual dimorphism is present in P. badius; comparisons across anthropoids indicated patterns of circumorbital dimorphism were decoupled from overall size dimorphism. CONCLUSIONS: The expanded circumorbits of male red colobus monkeys evolved in response to intense male-male competition. This hypothesis is consistent with the pattern across anthropoid primates and highlights the underappreciated role of sexual selection in shaping the primate face.

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.

Does Oxygen Stable Isotope Composition in Primates Vary as a Function of Vertical Stratification or Folivorous Behaviour?

Stable isotopes of oxygen often vary within a community of primates. For example, folivorous monkeys that forage in the upper reaches of the forest canopy tend to evince high δ18O values, whereas those that prefer the understory tend to have lower δ18O values. Given that leaves also have high δ18O values, particularly higher in the canopy, there is uncertainty as to which behavioural variable - vertical stratification or folivory - is the primary determinant of variation in δ18O values. Here, we explore further δ18O values from the Taï Forest monkeys (n = 7 species; n = 33 individuals) by examining the interaction between diet and vertical stratification, thereby allowing us to differentiate the effects of each covariate. We found that δ18O values varied as a function of mean canopy height, but not folivory, resolving uncertainty about the primary cause of δ18O variation. This outcome revolves largely, but not entirely, on the behaviours of Procolobus verus, a highly folivorous but understory forager. Relatively elevated values in Cercopithecus diana, a frugivorous but middle-to-high canopy forager, raises the possibility that plant reproductive tissues (e.g., fruits, flowers) may be increasingly sensitive to evaporative fractionation at higher forest canopy levels. Overall, our results further affirm the value of using δ18O values to estimate the vertical behaviour of primate species in a fossil assemblage.

Downclimbing and the evolution of ape forelimb morphologies.

The forelimbs of hominoid primates (apes) are decidedly more flexible than those of monkeys, especially at the shoulder, elbow and wrist joints. It is tempting to link the greater mobility of these joints to the functional demands of vertical climbing and below-branch suspension, but field-based kinematic studies have found few differences between chimpanzees and monkeys when comparing forelimb excursion angles during vertical ascent (upclimbing). There is, however, a strong theoretical argument for focusing instead on vertical descent (downclimbing), which motivated us to quantify the effects of climbing directionality on the forelimb kinematics of wild chimpanzees (Pan troglodytes) and sooty mangabeys (Cercocebus atys). We found that the shoulders and elbows of chimpanzees and sooty mangabeys subtended larger joint angles during bouts of downclimbing, and that the magnitude of this difference was greatest among chimpanzees. Our results cast new light on the functional importance of downclimbing, while also burnishing functional hypotheses that emphasize the role of vertical climbing during the evolution of apes, including the human lineage.