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Nutritional factors are among the most important influences on primate food choice. We examined the influence of macronutrients, minerals, and secondary compounds on leaf choices by members of a foli-frugivorous population of eastern black-and-white colobus-or guerezas (Colobus guereza)-inhabiting the Kakamega Forest, Kenya. Macronutrients exerted a complex influence on guereza leaf choice at Kakamega. At a broad level, protein content was the primary factor determining whether or not guerezas consumed specific leaf items, with eaten leaves at or above a protein threshold of ca. 14% dry matter. However, a finer grade analysis considering the selection ratios of only items eaten revealed that fiber played a much greater role than protein in influencing the rates at which different items were eaten relative to their abundance in the forest. Most minerals did not appear to influence leaf choice, though guerezas did exhibit strong selectivity for leaves rich in zinc. Guerezas avoided most leaves high in secondary compounds, though their top food item (Prunus africana mature leaves) contained some of the highest condensed tannin concentrations of any leaves in their diet. Kakamega 674 Fashing, Dierenfeld, and Mowry guerezas periodically traveled great distances to exploit rare foods (bark from exotic Myrtaceae trees and soil) outside their normal home ranges. Our results suggest that these journeys were driven by the fact that these rare foods contained exceptionally high sodium concentrations, a mineral believed to be deficient in the guereza's usual diet. Lastly, our results are consistent with the pattern established across other Paleotropical rain forests in which colobine biomass can be predicted by the protein-to-fiber ratio in mature leaves. Of the 8 rain forests for which the relevant data are available, Kakamega features the second highest mature leaf protein-to-fiber ratio as well as the second highest colobine biomass.

Protein limitation has been considered a key factor in hypotheses on the evolution of life history and animal communities, suggesting that animals should prioritize protein in their food choice. This contrasts with the limited support that food selection studies have provided for such a priority in nonhuman primates, particularly for folivores. Here, we suggest that this discrepancy can be resolved if folivores only need to select for high protein leaves when average protein concentration in the habitat is low. To test the prediction, we applied meta-analyses to analyze published and unpublished results of food selection for protein and fiber concentrations from 24 studies (some with multiple species) of folivorous primates. To counter potential methodological flaws, we differentiated between methods analyzing total nitrogen and soluble protein concentrations. We used a meta-analysis to test for the effect of protein on food selection by primates and found a significant effect of soluble protein concentrations, but a non-significant effect for total nitrogen. Furthermore, selection for soluble protein was reinforced in forests where protein was less available. Selection for low fiber content was significant but unrelated to the fiber concentrations in representative leaf samples of a given forest. There was no relationship (either negative or positive) between the concentration of protein and fiber in the food or in representative samples of leaves. Overall our study suggests that protein selection is influenced by the protein availability in the environment, explaining the sometimes contradictory results in previous studies on protein selection. Am. J. Primatol. 79:e22550, 2017. © 2016 Wiley Periodicals, Inc.

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