The combined effect of canopy shading and sea urchin grazing on recruitment in kelp forest (Laminaria hyperborea)

The aim of this study was to determine the impact of sea urchin grazing ( Echinus esculentus) and canopy shading on the recruitment of the kelp Laminaria hyperborea in mid-Norway. A spatially variable distribution of sea urchins was observed, and recruitment processes were studied both after disturbance, caused by kelp harvesting removal of the canopy kelps, and in pristine kelp forests. The combination of sea urchin density and the density of canopy-forming kelps had the strongest influence on the density of small kelps in pristine kelp forest, suggesting that both grazing from sea urchins and shading from the canopy contributed to the mortality of small kelps. High densities of small kelps ( > 20 m(-2)) were only found in pristine kelp forest together with <= 6 canopy-forming kelps m(-2) or < 3 sea urchins m(-2) on average. However, within the observed range of sea urchin densities these had no effect on the density of large, canopy-forming kelps. Large L. hyperborea were apparently not subject to grazing. In addition, only a small number of surviving kelp recruits was needed to maintain the density of canopy-forming kelps, as L. hyperborea specimens may survive many years. These conditions result in high stability of the kelp forest. A different picture was seen after kelp harvesting, when high recruitment and survival of recruits are the conditions for rapid restoration of the kelp vegetation. After removal of the canopy-forming plants, the kelp recruits were temporarily released from high density-dependent mortality due to shading. Some influence of sea urchin grazing on the density of recruits was observed, but this was small compared with the strong canopy effect. However, the accumulated impact of grazing during a period of time had a strong overall effect on the regrowth of kelp. After 2.5 years the accumulated biomasses at the harvested stations were strongly related to average sea urchin density, and a density of between 4 and 5 sea urchins m(-2) resulted in very little biomass accumulation. This suggests that the L. hyperborea kelp forest vegetation has a high degree of stability, but shows less resilience after disturbance, when exposed to moderate sea urchin grazing.