The importance of ingestion rates for estimating food quality and energy intake.

Testing ecological or socioecological models in primatology often requires estimates of individual energy intake. It is a well established fact that the nutrient content (and hence the energy content) of primate food items is highly variable. The second variable in determining primate energy intake, i.e., the ingestion rate, has often been ignored, and few studies have attempted to estimate the relative importance of the two predictors. In the present study individual ingestion rates were measured in two ecologically very different populations of Hanuman langurs (Semnopithecus entellus) at Jodhpur, India, and Ramnagar, Nepal. Protein and soluble sugar concentrations in 50 and 100 food items. respectively, were measured using standardized methods. Variation in ingestion rates (gram of dry matter per minute) was markedly greater among food items than among langur individuals in both populations, but did not differ systematically among food item categories defined according to plant part and age. General linear models (GLMs) with ingestion rate, protein, and soluble sugar content explained 40-80% of the variation in energy intake rates (kJ/min). The relative importance of ingestion rates was either similar (Ramnagar) or much greater (Jodhpur) than the role of sugar and/or protein content in determining the energy intake rates of different items. These results may impact socioecological studies of variation in individual energy budgets, investigations of food choice in relation to chemical composition or sensory characteristics, and research into habitat preferences that measures habitat quality in terms of abundance of important food sources. We suggest a definition of food quality that includes not only the amount of valuable food contents (energy, vitamins, and minerals) and the digestibility of different foods, but also the rate at which the food can be harvested and processed. Such an extended definition seems necessary because time may constrain primates when feeding competition is intense and foraging is risk-prone.