Separation anxiety: mussels self-organize into similar power-law clusters regardless of predation threat cues

Mussels have myriad effects on population, community, and ecosystem processes. Their aggregation behavior is an inducible defense that links non-consumptive effects of predators to benthic spatial pattern formation. Aggregation increases intraspecific competition but can be beneficial due to lower perimeter-related predation and other risks. Mytilus edulis aggregation responses to predation threats have not been investigated outside of Europe. We studied the effects of chemical cues from heterospecifics (predators Carcinus maenas, Nucella lapillus; herbivore Littorina littorea) and conspecifics (injured and intact M. edulis) on M. edulis aggregation behavior in Maine, USA. Mussels self-organized into fractal power-law spatial patterns like those in the field. Aggregations had lower perimeter:area (P:A) ratios than singletons, despite having more complex, irregular shapes with higher fractal dimensions (D). However, with one exception, no significant differences in aggregation rate, P:A ratio, and D were observed for any chemical cue treatment when compared to no-cue controls. Our experiment revealed higher aggregation rates than reported from similar experiments, leaving little scope for additional aggregation when exposed to chemical cues. We suggest that increased aggregation in response to predation threat is context-dependent: costs outweigh benefits beyond some optimal aggregation size, and mussels in our experiment were at the upper aggregation limit beyond which more aggregation could have negative consequences. Bet-hedging with a power-law distribution of aggregation shapes and sizes may be the optimal spatial strategy, especially if predation and other risks are variable in space and time.