Summary: Dynamical systems containing heteroclinic cycles and networks can be invoked as models of intransitive competition between three or more species. When populations are assumed to be well-mixed, a system of ordinary differential equations (ODEs) describes the interaction model. Spatially extending these equations with diffusion terms creates a system of partial differential equations which captures both the spatial distribution and mobility of species. In one spatial dimension, travelling wave solutions can be observed, which correspond to periodic orbits in ODEs that describe the system in a steady-state travelling frame of reference. These new ODEs also contain a heteroclinic structure. For three species in cyclic competition, the topology of the heteroclinic cycle in the well-mixed model is preserved in the steady-state travelling frame of reference. We demonstrate that with four species, the heteroclinic cycle which exists in the well-mixed system becomes a heteroclinic network in the travelling frame of reference, with additional heteroclinic orbits connecting equilibria not connected in the original cycle. We find new types of travelling waves which are created in symmetry-breaking bifurcations and destroyed in an orbit flip bifurcation with a cycle between only two species. These new cycles explain the existence of ‘defensive alliances’ observed in previous numerical experiments. We further describe the structure of the heteroclinic network for any number of species, and we conjecture how these results may generalise to systems of any arbitrary number of species in cyclic competition.
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