The geometry of resource constraint: An empirical study of the golden snub-nosed monkey

Apposite conceptualization and measurement of resource variation is critical for understanding many issues in ecology, including ecological niches, persistence and distribution of populations, the structure of communities and population resilience to perturbations. We apply the nutritional geometry framework to conceptualize and quantify the responses of a temperate-living primate, the golden snub-nosed monkey Rhinopithecus roxellana to variation in resource quality and quantity and in nutrient requirements associated with seasonal environments. We present a geometric model distinguishing qualitative constraint, quantitative constraint and 'pseudo-constraint' whereby nutrient intakes resemble response to qualitative resource constraint but are in fact driven by variation in nutrient requirements. The model is applied to analyse nutrient intakes recorded in 164 full-day observations of monkeys from two populations, one wild and the other captive, across seasons. Additionally, we recorded the diet of a single animal over 32 consecutive days in the wild. Despite considerable differences in available resources, the captive and wild populations showed marked similarities in nutrient intakes, including indistinguishable amounts and ratios of ingested macronutrients during summer and autumn and strong year-round maintenance of protein compared to seasonally variable fat and carbohydrate intakes. These similarities suggest homeostatically regulated nutritional targets and provide reference points to identify factors driving population differences in macronutrient intake in winter and spring. Our framework enabled us to distinguish examples of quantitative, qualitative and 'pseudo-constraint'. We suggest that this approach can increase the resolution at which resource constraint is conceptualized and measured in ecological studies.