An aerial view of an intertidal mussel bed often reveals large scale striped patterns aligned perpendicular to the direction of the tide; dense bands of mussels alternate periodically with near bare sediment. Experimental work led to the formulation of a set of coupled partial differential equations modelling a mussel-algae interaction, which proved pivotal in explaining the phenomenon. The key class of model solutions to consider are one-dimensional periodic travelling waves (wavetrains) that encapsulate the abundance of peak and trough mussel densities observed in practice. These solutions may, or may not, be stable to small perturbations, and previous work has focused on determining the ecologically relevant (stable) wavetrain solutions in terms of model parameters. The aim of this paper is to extend this analysis to two space dimensions by considering the full stripe pattern solution in order to study the effect of transverse two-dimensional perturbations-a more true to life problem. Using numerical continuation techniques, we find that some striped patterns that were previously deemed stable via the consideration of the associated wavetrain solution, are in fact unstable to transverse two-dimensional perturbations; and numerical simulation of the model shows that they break up to form regular spotted patterns. In particular, we show that break up of stripes into spots is a consequence of low tidal flow rates. Our consideration of random algal movement via a dispersal term allows us to show that a higher algal dispersal rate facilitates the formation of stripes at lower flow rates, but also encourages their break up into spots. We identify a novel hysteresis effect in mussel beds that is a consequence of transverse perturbations.
Keywords: Activator-inhibitor; Mussels; Mussel–algae interaction; Pattern formation; Self-organisation; Tranverse perturbations; Turing–Hopf bifurcation; Two-dimensional stability.