Changing webs—Variation of complex networks over a tidal cycle in an intertidal rocky reef

Abstract

Incorporating temporal variation in models is one of the most important challenges in food web research. One of the environments where time causes profound changes is the intertidal zone, where the immersion-emersion cycle drastically changes the abiotic and biotic conditions. Intertidal rocky shores have been intensively studied, however the variation in the complex food web network that occurs during a tidal cycle remains undescribed. Highly resolved food web networks were assembled for an intertidal reef depicting the food web during low and high tide, and with and without tide pools. It was concluded that high tide adds new species to the web, but it does not add complexity since network connectance was not changed. This occurs because incoming species are mostly highly generalist fish, which add many new links to the web. Tide pools, however, add not only diversity but also complexity. Webs were dominated by intermediate species, with the proportion of top consumers fluctuating throughout the tidal cycle, being lowest during low tide and highest at high tide, due to the incoming larger vertebrate predators. Consumer taxa outnumbered resource taxa, except at low tide when pools are present. Mean trophic level was lowest at low tide (2.3) and highest at high tide with pools (2.6). Omnivory was high and showed little change. “Chain”, the number of links connecting top to basal species, was stable but low. This implies that disturbance can rapidly travel bottom-up or top-down through predator-prey links. The increased connectance given by the addition of tide pools likely increases robustness to disturbances, an important feature in coastal areas so often impacted by human action.