Recent studies have brought renewed attention to the importance of complex species interactionsânotably intransitive interactionsâto patterns of plant community diversity. One underappreciated avenue through which intransitivity can occur is through cyclic population dynamics. Although such cyclic intransitive relationships have been extensively studied in predatorâprey systems, evidence of their importance in competitive communities, notably plant communities, is more limited. Most studies of coexistence in plant communities assume fixedâpoint coexistence even while utilising models that allow for cyclic population dynamics.
In this paper, we explore the potential for densityâdependent, cyclic population dynamics and intransitivity in a model for annual plants. We then examine how these densityâdependent cycles impact mutual invasibility and ultimately stable coexistence between plant species pairs. We do this using data collected from four coâoccurring annual plant species living in natural wildflower communities in SW Western Australia. To maximise the number of biologically plausible pathways by which coexistence mediated by densityâdependent cyclic intransitivity can occur, we use an annual plant model that allows for competitive direct interactions, facilitative direct interactions and higherâorder interactions between species.
Results from our empirically parameterised model suggest that monocultures of all four focal species can have cyclic solutions with periodicity <1 under sunny (âopenâ) or shaded field conditions. Cyclic patterns drive variation in annual abundance patterns, with stable solutions for persistent monocultures and invasibility potential (the capacity of one population to invade another) common. Mutual invasibility in the face of cyclic population dynamics was found for just one of six species pairs, only under open environmental conditions. Our results illustrate the potential for cyclic intransitivity to both drive and prevent stable coexistence in environmentally heterogeneous biological communities.
Synthesis. We provide analytical and empirical evidence that coexistence in competitive communities (annual plants) can be achieved under nonâequilibrium circumstances, through densityâdependent cyclic intransitivity. Our results suggest that cyclic population dynamics may be common and important for coexistence dynamics in some types of communities. In such communities, the exploration of stable coexistence should, therefore, include consideration of cyclic as well as fixedâpoint equilibria for maximal accuracy.