Why overgrowth of intertidal encrusting algae does not always cause competitive exclusion

Encrusting algae are well-known to be able, for long periods, to withstand shading and overgrowth by other organisms. How this is achieved remains a mystery. It had been proposed that connections with unshaded (non-overgrown) parts of the thallus may allow transfer of nutrients to the shaded part. From this model, I proposed and tested the hypothesis that shaded patches of the intertidal red alga, Hildenbrandia rubra, would survive overgrowth longer, or better, when connected to unshaded thallus than when experimentally separated from surrounding alga. Experimental treatments were shading (black or transparent 80 mm perspex discs or no cover) and scraping (scraped around the disc to remove contacts, a control for effects of scraping, no treatment). The 9 orthogonal combinations of cover and scraping were applied to 3 independent, random replicates (i.e. 27 plots) in each of four randomly chosen sites.In all 4 sites, over 13 months, shaded H. rubra survived in greater abundance (as % cover) where in contact with surrounding thallus. In one site, there was no effect of shading unless the thallus was isolated. In two sites, shading reduced cover, but was more deleterious where the thallus was isolated. In the fourth site, there were artefacts due to a perspex cover, but still less cover of alga where it was isolated.This encrusting alga can withstand a long period of complete shading, provided there is connection to unshaded thallus. Interpreting or predicting overgrowth interactions in terms of competitive outcomes is therefore dependent on consideration of whether the overgrown species is actually being affected. It also depends on the duration of overgrowth and, as shown here, the extent to which connectivity with unshaded thallus is effective at preventing or reducing any consequences. Observations and experiments that do not ascertain these are difficult to interpret.