The fundamental role of competition in the ecology and evolution of mutualisms.

Mutualisms are interspecific interactions that yield reciprocal benefits. Here, by adopting a consumer-resource perspective, we show how considering competition is necessary in order to understand the evolutionary and ecological dynamics of mutualism. We first review the ways in which competition shapes the ecology of mutualisms, using a graphical framework based on resource flows rather than net effects to highlight the opportunities for competition. We then describe the known mechanisms of competition and show how it is a critical driver of the evolutionary dynamics, persistence, and diversification of mutualism. We argue that empirical and theoretical research on the ecology and evolution of mutualisms will jointly progress by addressing four key points: (i) the existence and shape of physiological trade-offs among cooperation, competition, and other life-history and functional traits; (ii) the capacity for individuals to express conditional responses to variation in their mutualistic and competitive environment; (iii) the existence of heritable variation for mutualistic and competitive traits and their potentially conditional expression; and (iv) the structure of the network of consumer-resource interactions in which individuals are embedded.