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Dietary shifts and corresponding morphological changes can sometimes evolve in succession, not concurrently—an evolutionary process called behavioral drive. Detecting behavioral drive in the fossil record is challenging because it is difficult to measure behaviors independently from corresponding morphologies. To solve this problem, we focused on a puzzling behavior in the fossil record of some primates: eating graminoid plants. We report carbon and oxygen isotope ratios from fossil cercopithecid monkeys and integrate the data into a view of hominin dietary evolution, finding that changes in graminivorous behavior preceded corresponding changes in dental morphology by ~700,000 years. Decoupling diets and morphologies in time was conducive to determining when and to exploring why dietary changes helped to propel human evolution. 

Few animals have the cognitive faculties or prehensile abilities needed to eliminate tooth-damaging grit from food surfaces. Some populations of monkeys wash sand from foods when standing water is readily accessible, but this propensity varies within groups for reasons unknown. Spontaneous food-washing emerged recently in a group of long-tailed macaques (Macaca fascicularis) inhabiting Koram Island, Thailand, and it motivated us to explore the factors that drive individual variability. We measured the mineral and physical properties of contaminant sands and conducted a field experiment, eliciting 1282 food-handling bouts by 42 monkeys. Our results verify two long-standing presumptions: monkeys have a strong aversion to sand and removing it is intentional. Reinforcing this result, we found that monkeys clean foods beyond the point of diminishing returns, a suboptimal behavior that varied with social rank. Dominant monkeys abstained from washing, a choice consistent with the impulses of dominant monkeys elsewhere: to prioritize rapid food intake and greater reproductive fitness over the long-term benefits of prolonging tooth function. 

1. Silica is crucial to terrestrial plant life and geochemical cycling on Earth. It is also implicated in the evolution of mammalian teeth, but there is debate over which type of siliceous particle has exerted the strongest selective pressure on tooth morphology. 2. Debate revolves around the amorphous silica bodies (phytoliths) present in plants and the various forms of siliceous grit—that is, crystalline quartz (sand, soil, dust)—on plant surfaces. The problem is that conventional measures of silica often quantify both particle types simultaneously. 3. Here we describe a protocol that relies on heavy-liquid flotation to separate and quantify siliceous particulate matter in the diets of herbivores. The method is reproducible and well suited to detecting species- or population-level differences in silica ingestion. In addition, we detected meaningful variation within the digestive tracts of cows, an outcome that supports the premise of ruminal fluid ‘washing’ of siliceous grit. 4. We used bootstrap resampling to estimate the sample sizes needed to compare species, populations or individuals in space and time. We found that a minimum sample of 12 individuals is necessary if the species is a browser or as many as 55 if the species is a grazer, which are more variable. But a sample size of 20 is adequate for detecting statistical differences. We conclude by suggesting that our protocol for differentiating and quantifying silica holds promise for testing competing hypotheses on the evolution of dental traits.