Experimental confirmation of multiple community states in a marine ecosystem.

Small changes in environmental conditions can unexpectedly tip an ecosystem from one community type to another, and these often irreversible shifts have been observed in semi-arid grasslands, freshwater lakes and ponds, coral reefs, and kelp forests. A commonly accepted explanation is that these ecosystems contain multiple stable points, but experimental tests confirming multiple stable states have proven elusive. Here we present a novel approach and show that mussel beds and rockweed stands are multiple stable states on intertidal shores in the Gulf of Maine, USA. Using broad-scale observational data and long-term data from experimental clearings, we show that the removal of rockweed by winter ice scour can tip persistent rockweed stands to mussel beds. The observational data were analyzed with Anderson's discriminant analysis of principal coordinates, which provided an objective function to separate mussel beds from rockweed stands. The function was then applied to 55 experimental plots, which had been established in rockweed stands in 1996. Based on 2005 data, all uncleared controls and all but one of the small clearings were classified as rockweed stands; 37% of the large clearings were classified as mussel beds. Our results address the establishment of mussels versus rockweeds and complement rather than refute the current paradigm that mussel beds and rockweed stands, once established, are maintained by site-specific differences in strong consumer control.

PROPAGATION OF SCALE-DEPENDENT EFFECTS FROM RECRUITS TO ADULTS IN BARNACLES AND SEAWEEDS.

The interplay between local and large spatial scale processes in open systems is often dependent upon ecological context and species specific factors such as longevity, dispersal capability, or vulnerability to predation. When disturbance clears patches in open systems, the successful reestablishment of adult colonizers and the trajectory of succession may depend upon both the scale of the disturbance event and scale of life history characteristics. Here we examine the links between the size of a disturbance event and long term patterns of variation in recruitment, density, and percent cover in a relatively short-lived but long-range disperser, the acorn barnacle Semibalanus balanoides, and a relatively long lived and short range disperser, the fucoid alga Fucus vesiculosus. Further, a model of linked scale-dependent processes is developed and we explore how long term patterns of scale-dependent recruitment are related to successful establishment of adults. Recruitment, densities, and cover were monitored for five years for both species in a single experiment using clearings of different sizes spread over two north-facing and two south-facing bays on Swan's Island, Maine, USA. Barnacle recruitment was particularly variable in small clearings and was lower overall at more interior sites within bays. While local-scale factors also strongly influenced patterns of variation in fucoid recruitment, fucoids dominate surface cover in large clearings and in south-facing bays. Recruitment was a poor predictor of density, whereas density itself was a good predictor of percent cover for both species. These results indicate that scale dependencies, location-specific factors, and life history traits contribute to patterns of community development on rocky shores and may ultimately determine whether an open patch converges to or diverges from its initial community state.