Disturbance and organisms on boulders : II. Causes of patterns in diversity and abundance.

The tops of intertidal boulders on the east coast of Australia may carry a half-dozen algae, while the undersides may support a dozen or more species of sessile animals; mobile forms such as molluscs, crustaceans, and echinoderms are also common. These organisms are disturbed when boulders are moved by waves or buried in sand. Experiments were done to test the hypothesis that these disturbances are responsible for the reduced diversity and abundance of the assemblages on the undersides of small boulders. Boulders on one shore were bolted to the substratum to prevent movement by waves. In two separate experiments on another shore, boulders with and without attached organisms were buried in a few centimetres of sand. All experiments included relevant controls and were done high and low on the shore using rocks of several sizes.The assemblages of algae developing on the tops of all stabilised boulders, regardless of size, were similar to those on boulders free to roll indicating that, in contrast to results elsewhere, disturbance by waves was too infrequent or slight to affect these species. The tops of rocks were also rarely buried in sand and the main influences on the community in this situation were apparently exposure at low-tide and grazing gastropods.More species, however, did settle or survive on the undersides of rocks which were free of sand or could not be moved by waves; thus disturbances were important in this situation. In the absence of disturbance low on the shore, much or all space was occupied and sessile species such as sponges and ascidians overgrew other forms and reduced diversity. This did not happen higher on the shore and here diversity was simply an increasing function of rock-size. Overall disturbance played a similar role in all places - it killed organisms and created free space - but the final effects on the community varied depending upon the species present and the actions of other factors.