Hidden layers of density dependence in consumer feeding rates.

Functional responses relate a consumer's feeding rates to variation in its abiotic and biotic environment, providing insight into consumer behaviour and fitness, and underpinning population and food-web dynamics. Despite their broad relevance and long-standing history, we show here that the types of density dependence found in classic resource- and consumer-dependent functional-response models equate to strong and often untenable assumptions about the independence of processes underlying feeding rates. We first demonstrate mathematically how to quantify non-independence between feeding and consumer interference and between feeding on multiple resources. We then analyse two large collections of functional-response data sets to show that non-independence is pervasive and borne out in previously hidden forms of density dependence. Our results provide a new lens through which to view variation in consumer feeding rates and disentangle the biological underpinnings of species interactions in multi-species contexts.