Ecological interactions on a rocky shore : the control of macroalgal distribution by intertidal grazers

摘要

The aim of the present study was to determine the potential impact of intertidal grazers on the distribution of macro algae on the south coast of South Africa. Particular attention was paid to the large patellid limpet, Patella oculus, which is found thoughout the intertidal zone. Studies of gut contents revealed that Patella oculus was capable of ingesting not only the thallus of foliose (eg. Ulva spp.) and encrusting coralline macroalgae, but also spores and diatoms. The inclusion of these relatively small particles in the diet was surprising, as electron micrographs of the radula of P.oculus revealed that it is typically docoglossan in structure. Such radulae are thought to be poorly suited for collecting small food particles. Sand made up a significantly higher proportion of the gut contents than other particles at all shore heights, which suggests that P.oculus might be capable of excavating the rocky substratum, or of sweeping up sand, while searching for food. Analysis of the gut contents of other local herbivorous molluscs, was also carried out. These species included the winkles, Oxystele variegata and O.sinensis, and the small pulmonate limpets, Siphonaria concinna, S.capensis, and S.serrata. The guts of all species contained mainly spores and diatoms, although small fragments of Ulva sp. were found. The population structure of Patella oculus was investigated at two sites, Cannon Rocks and Old Woman's River. At Cannon Rocks, mean shell length of low-shore animals was significantly lower than that of both mid- and high-shore animals, while at Old Woman's River, no significant difference was found among shore heights. A regression equation for In (shell length) vs In (dry weight) was calculated, and based on length data, the biomass density (g dry mass.m⁻²) of P.oculus at Old Woman's River was estimated. Values ranged from 2.8 on the low- and midshore to 0.37 on the high-shore. A manipulative field experiment was used to determine the impact of mesograzers and macrograzers (such as Patella oculus) on the distribution of intertidal macro algae on the mid- and low-shore at Old Woman's River. Grazers were excluded using mesh cages (mesh size = 3mm), in two separate experiments, one in winter and the other in spring. Percentage cover of macroalgal species and sessile invertebrates was estimated at approximately 6 week intervals for up to 3 months. MANOV A showed that treatments did not significantly affect cover of macroalgae or barnacles during winter. However, towards the end of the spring experiment (midshore only) cover of barnacles and green foliose turfs did increase in those plots from which mesograzers and/or macro grazers were excluded. The failure of the statistical tests to detect significant differences at some time intervals may have been caused by high levels of variation among replicates. This suggests that factors other than grazing are of overriding importance in determining the distribution of local macroalgae. The existence of a possible symbiotic relationship between Patella oculus and the red foliose alga, Gelidium pristoides, was investigated. The availability of various substratum types, including rock, limpet shells, barnacles etc., and the proportion of the total cover of G.pristoides on each, was calculated. It was shown that a significantly higher proportion of the alga grew on limpet shells, although the availability of this substratum type was low. It is thought that the aggressive behaviour of P.oculus prevents all but juvenile Patella longicosta from grazing on its shell, thus providing a refuge from grazing for G.pristoides.