Interactions between wave action and grazing control the distribution of intertidal macroalgae
                
                        Journal article, 2006
                
            
                    
                        Canopy-forming macroalgae are key species on temperate rocky shores. However, there is a lack of understanding of how the relative balance of physical and biological factors controls the establishment and persistence of intertidal macroalgae. Here we present an integrated study of the relative importance of wave-induced forces and grazing for the recruitment and Survival of the canopy-forming intertidal macroalgae Fucus vesiculosus and F. spiralis. A set of overtopped breakwaters provided a nearly unconfounded gradient in wave exposure between seaward and landward sides. A biomechanical analysis was performed based on empirical measurements of maximum drag forces in breaking waves, a model of long-term maximum wave height, and the breaking stress of Fucus spp. The estimated maximum flow speed (7-8 m/s) on the seaward side of the breakwaters was predicted to completely dislodge or prune Fucus spp. larger than similar to 10 cm, while dislodgment was highly unlikely on the landward side for all sizes. Experimental transplantation of Fucus spp. supported the biomechanical analysis but also suggested that mechanical abrasion may further limit survival in wave-exposed locations. Experimental removal of the limpet Patella vulgata, which was the principal grazer at this site, resulted in recruitment of Fucus spp. on the seaward side. We present a model of limpet grazing that indicates that limpet densities > 5-20 individuals/m(2) provide a proximate mechanism preventing establishment of Fucus spp., whereas wave action > 2 m/s reduces persistence through dislodgment and battering. In a conceptual model we further propose that recruitment and survival of juvenile Fucus spp. are controlled indirectly by wave exposure through higher limpet densities at exposed locations. This model predicts that climate change, and in particular an increased frequency of storm events in the northeast Atlantic, will restrict fucoids to more sheltered locations.
                    
                    
                            
                                wave exposure
                            
                            
                                dislodgment
                            
                            
                                SURVIVAL
                            
                            
                                Fucus spp.
                            
                            
                                PHYSICAL DISTURBANCE
                            
                            
                                transplantation
                            
                            
                                limpets
                            
                            
                                climate change
                            
                            
                                breaking waves
                            
                            
                                DISLODGMENT
                            
                            
                                hydrodynamic drag
                            
                            
                                CANOPY
                            
                            
                                SURF-ZONE
                            
                            
                                grazing
                            
                            
                                WATER MOTION
                            
                            
                                breakwaters
                            
                            
                                FUCUS-GARDNERI
                            
                            
                                Ulva sp.
                            
                            
                                LIMPET PATELLA-VULGATA
                            
                            
                                COMMUNITIES
                            
                            
                                Patella vulgata
                            
                            
                                ROCKY SHORES