Emergence of weak-intransitive competition through adaptive diversification and eco-evolutionary feedbacks

Indirect biotic interactionssuch as intransitive competitionare increasingly recognized as being important in shaping ecological patterns in natural systems. Over long time-scales, such indirect interactions may affect the evolution of species phenotypes, which in turn can modify these interactions, thereby begetting eco-evolutionary feedbacks. If indirect intransitive interactions can emerge in situ during lineage diversification, they could profoundly affect species' phenotypic diversity, temporal stability, and subsequent diversification rates. We address these questions by investigating the conditions under which indirect intransitive competition can emerge from a lineage diversifying in sympatry. We use an adaptive dynamics model to study the ecological and evolutionary properties of this lineage under different scenarios where competition for resources between phenotypes varies in strength and (a)symmetry. Results show that weak-intransitive competition can emerge during the sympatric diversification of a single lineage. Weak-intransitivity here refers to situations where species interactions are not perfectly transitive, that is, there is no strict hierarchy in species competitive abilities. The strength of such weak-intransitivity increases when the competition between phenotypes increases in strength and asymmetry. The strength of intransitivity also correlates with other system properties. We notably found that the strength of intransitivity increases with the number of phenotypes, and that greater intransitivity correlates with the evolution of greater functional trait divergences between phenotypes, greater resistance to invasion by new phenotypes but lower resistance to disturbances as well as slower evolutionary rates. Synthesis. This theoretical exploration of the evolution of intransitive competition provides the first formal bridge between the ecological and evolutionary aspects of intransitive competition. We show that, when competitive interactions are strong enough, weak-intransitive competition is more likely to emerge through adaptive diversification than from a random community assembly. Intransitive competition is, therefore, not only restricted to between-species interactions but can also function as a regulator of diversification within species, thereby affecting lineage functional diversity, and ecological and evolutionary stability.