The East African hominin Paranthropus boisei was characterized by a suite of craniodental features that have been widely interpreted as adaptations to a diet that consisted of hard objects that required powerful peak masticatory loads. These morphological adaptations represent the culmination of an evolutionary trend that began in earlier taxa such as Australopithecus afarensis, and presumably facilitated utilization of open habitats in the Plio-Pleistocene. Here, we use stable isotopes to show that P. boisei had a diet that was dominated by C 4 biomass such as grasses or sedges. Its diet included more C 4 biomass than any other hominin studied to date, including its congener Paranthropus robustus from South Africa. These results, coupled with recent evidence from dental microwear, may indicate that the remarkable craniodental morphology of this taxon represents an adaptation for processing large quantities of low-quality vegetation rather than hard objects.C4 photosynthesis | C3 photosynthesis T he East African hominin Paranthropus boisei possessed large and low-cusped postcanine dentition, large and thick mandibular corpora, and powerful muscles of mastication, which are generally believed to be adaptations for a diet of nuts, seeds, and hard fruit (1-3). This notion emerged from interpretations of P. boisei's morphology, but gained indirect support from dental microwear studies of its congener, Paranthropus robustus; these concluded that wear on the molars of South African Paranthropus was consistent with its having ingested and chewed small, hard food items, if not as primary resources, then at least as fallback foods (4-6). Although some have suggested that the craniodental morphology of P. boisei is consistent with the consumption of tough rather than hard foods (7,8), this idea has been largely eschewed by most workers. Thus, when a recent study using dental microwear texture analysis revealed no evidence for the consumption of hard foods by P. boisei (9), it challenged decades of received wisdom, and underscored the need for independent lines of paleodietary evidence.Stable carbon isotope analysis has proven a powerful tool for testing hypotheses about the diets of extinct herbivorous mammals (10, 11). It is based on the idea that carbon isotope compositions vary predictably between plant foods [e.g., plants using the C 3 photosynthetic pathway (most dicotyledonous plants including trees, shrubs, forbs, herbs) and those using the C 4 pathway (predominantly tropical grasses and sedges, which are monocotyledonous plants)], and further that dietary carbon remains locked in tooth enamel even after millions of years (10). Carbon isotope studies of P. robustus from South Africa indicated that it consumed some plants using C 4 photosynthesis such as tropical grasses or sedges, but were also consistent with most of its dietary carbon (approximately 70%) having been derived from the C 3 food items favored by extant chimpanzees (Pan troglodytes) such as tree fruits (12, 13). In contrast, stable isotopes measuremen...