Recent evidence suggests that several extinct primates, including contemporaneous Paranthropus boisei and Theropithecus oswaldi in East Africa, fed largely on grasses and sedges (i.e., graminoids). As the only living primate graminivores, gelada monkeys (Theropithecus gelada) can yield insights into the dietary strategies pursued by extinct grass- and sedge-eating primates. Past studies of gelada diet were of short duration and occurred in heavily disturbed ecosystems. We conducted a long-term study of gelada feeding ecology in an intact Afroalpine ecosystem at Guassa, Ethiopia. Geladas at Guassa consumed ≥56 plant species, ≥20 invertebrate species, one reptile species, and the eggs of one bird species over a 7-year period. The annual diet consisted of 56.8% graminoid parts, 37.8% forb parts, 2.8% invertebrates, and 2.6% other items, although geladas exhibited wide variability in diet across months at Guassa. Edible forbs were relatively scarce at Guassa but were strongly selected for by geladas. Tall graminoid leaf and tall graminoid seed head consumption correlated positively, and underground food item consumption correlated negatively, with rainfall over time. Geladas at Guassa consumed a species-rich diet dominated by graminoids, but unlike geladas in more disturbed habitats also ate a diversity of forbs and invertebrates along with occasional vertebrate prey. Although graminoids are staple foods for geladas, underground food items are important "fallback foods." We discuss the implications of our study, the first intensive study of the feeding ecology of the only extant primate graminivore, for understanding the dietary evolution of the theropith and hominin putative graminivores, Theropithecus oswaldi and Paranthropus boisei.

Despite intensive study in humans, responses to dying and death have been a neglected area of research in other social mammals, including nonhuman primates. Two recent reports [Anderson JR, Gillies A, Lock LC. 2010. Pan thanatology. Current Biology 20:R349-R351; Biro D, Humle T, Koops K, Souse C, Hayashi M, Matsuzawa T. 2010. Chimpanzee mothers at Bossou, Guinea carry the mummified remains of their dead infants. Current Biology 20:R351-R352] offered exciting new insights into behavior toward dying and dead conspecifics in our closest living relatives-chimpanzees. Here, we provide a comparative perspective on primate thanatology using observations from a more distant human relative-gelada monkeys (Theropithecus gelada)-and discuss how gelada reactions to dead and dying groupmates differ from those recently reported for chimpanzees. Over a 3.75-year study period, we observed 14 female geladas at Guassa, Ethiopia carrying dead infants from 1 hr to ≥48 days after death. Dead infants were carried by their mothers, other females in their group, and even by females belonging to other groups. Like other primate populations in which extended (>10 days) infant carrying after death has been reported, geladas at Guassa experience an extreme climate for primates, creating conditions which may favor slower rates of decomposition of dead individuals. We also witnessed the events leading up to the deaths of two individuals and the responses by groupmates to these dying individuals. Our results suggest that while chimpanzee mothers are not unique among primates in carrying their dead infants for long periods, seemingly "compassionate" caretaking behavior toward dying groupmates may be unique to chimpanzees among nonhuman primates (though it remains unknown whether such "compassionate" behavior occurs outside captivity).

Chewing efficiency has been associated with fitness in mammals, yet little is known about the behavioral, ecological, and morphological factors that influence chewing efficiency in wild animals. Although research has established that dental wear and food material properties independently affect chewing efficiency, few studies have addressed the interaction among these factors. We examined chewing efficiency, measured as mean fecal particle size, as a function of seasonal shifts in diet (and corresponding changes in food fracture toughness) in a single breeding population of a grazing primate, the gelada monkey, at Guassa, Ethiopia. We also measured dental topographic traits (slope, angularity, and relief index) and relative two- and three-dimensional shearing crest lengths in a cross-sectional wear series of gelada molars. Chewing efficiency decreased during the dry season, a pattern corresponding to the consumption of foods with higher fracture toughness. Older individuals experienced the most pronounced decreases in chewing efficiency between seasons, implicating dental wear as a causal factor. This pattern is consistent with our finding that dental topographic metrics and three-dimensional relative shearing crest lengths were lowest at the last stage of wear. Integrating these lines of behavioral, ecological, and morphological evidence provides some of the first empirical support for the hypothesis that food fracture toughness and dental wear together contribute to chewing efficiency. Geladas have the highest chewing efficiencies measured thus far in primates, and may be analogous to equids in their emphasis on dental design as a means of particle size reduction in the absence of highly specialized digestive physiology.

Paleoanthropologists have long argued-often contentiouslyabout the climbing abilities of early hominins and whether a foot adapted to terrestrial bipedalism constrained regular access to trees. However, some modern humans climb tall trees routinely in pursuit of honey, fruit, and game, often without the aid of tools or support systems. Mortality and morbidity associated with facultative arboreality is expected to favor behaviors and anatomies that facilitate safe and efficient climbing. Here we show that Twa hunter-gatherers use extraordinary ankle dorsiflexion (>45°) during climbing, similar to the degree observed in wild chimpanzees. Although we did not detect a skeletal signature of dorsiflexion in museum specimens of climbing hunter-gatherers from the Ituri forest, we did find that climbing by the Twa is associated with longer fibers in the gastrocnemius muscle relative to those of neighboring, nonclimbing agriculturalists. This result suggests that a more excursive calf muscle facilitates climbing with a bipedally adapted ankle and foot by positioning the climber closer to the tree, and it might be among the mechanisms that allow hunter-gatherers to access the canopy safely. Given that we did not find a skeletal correlate for this observed behavior, our results imply that derived aspects of the hominin ankle associated with bipedalism remain compatible with vertical climbing and arboreal resource acquisition. Our findings challenge the persistent arboreal-terrestrial dichotomy that has informed behavioral reconstructions of fossil hominins and highlight the value of using modern humans as models for inferring the limits of hominin arboreality.Australopithecus | human pygmy phenotype | phenotypic plasticity