Phase shift facilitation following cyclone disturbance on coral reefs.

While positive interactions have been observed to influence patterns of recruitment and succession in marine and terrestrial plant communities, the role of facilitation in macroalgal phase shifts is relatively unknown. In December 2012, typhoon Bopha caused catastrophic losses of corals on the eastern reefs of Palau. Within weeks of the typhoon, an ephemeral bloom of monospecific macroalgae (Liagora sp.) was observed, reaching a peak of 38.6% cover in February 2013. At this peak, we observed a proliferation of a second macroalgal species, Lobophora variegata. Lobophora was distributed non-randomly, with higher abundances occurring within the shelter of Liagora canopies than on exposed substrates. Bite rates of two common herbivorous fish (Chlorurus sordidus and Ctenochaetus striatus) were significantly higher outside canopies (2.5- and sixfold, respectively), and cage exclusion resulted in a significant increase in Lobophora cover. Experimental removal of Liagora canopies resulted in a 53.1% decline in the surface area of Lobophora after 12 days, compared to a 51.7% increase within canopies. Collectively, these results indicate that Liagora canopies act as ecological facilitators, providing a 'nursery' exclusion zone from the impact of herbivorous fish, allowing for the establishment of understory Lobophora. While the ephemeral Liagora bloom had disappeared entirely 9 months post-typhoon, the facilitated shift to Lobophora has persisted for over 18 months, dominating ~40% of the reef substrate. While acute disturbance events such as typhoons have been suggested as a mechanism to reverse algal phase shifts, our results suggest that typhoons may also trigger, rather than just reverse, phase shifts.