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Previous studies have hypothesized that wave-induced drag forces may constrain the size of intertidal organisms by dislodging or breaking organisms that exceed some critical dimension. In this study, we explored constraints on the size of the articulated coralline alga Calliarthron, which thrives in wave-exposed intertidal habitats. Its ability to survive depends critically upon its segmented morphology (calcified segments separated by flexible joints or ;genicula'), which allows otherwise rigid fronds to bend and thereby reduce drag. However, bending also amplifies stress within genicula near the base of fronds. We quantified breakage of genicula in bending by applying known forces to fronds until they broke. Using a mathematical model, we demonstrate the mitigating effect of neighboring fronds on breakage and show that fronds growing within dense populations are no more likely to break in bending than in tension, suggesting that genicular morphology approaches an engineering optimum, possibly reflecting adaptation to hydrodynamic stress. We measured drag in a re-circulating water flume (0.23-3.6 m s(-1)) and a gravity-accelerated water flume, which generates jets of water that mimic the impact of breaking waves (6-10 m s(-1)). We used frond Reynolds number to extrapolate drag coefficients in the field and to predict water velocities necessary to break fronds of given sizes. Laboratory data successfully predicted frond sizes found in the field, suggesting that, although Calliarthron is well adapted to resist breakage, wave forces may ultimately limit the size of intertidal fronds.

To break a coralline: mechanical constraints on the size and survival of a wave-swept seaweed