Mode of access: World Wide Web. / Thesis (Ph. D.)--University of Hawaii at Manoa, 2005. / Includes bibliographical references (leaves 183-210). / Electronic reproduction. / Also available by subscription via World Wide Web / xv, 210 leaves, bound ill., maps 29 cm

Mussels are important ecological engineers on intertidal rocks where they create habitat that contributes substantially to overall biodiversity. They provide secondary substratum for other free-living, infaunal or epifaunal organisms, and increase the surface area for settlement by densely packing together into complex multilayered beds. The introduction of the alien invasive mussel Mytilus galloprovincialis has extended the upper limit of mussels on the south coast of South Africa, potentially increasing habitat for associated fauna. The aim of this study was to describe the structure of mussel beds, the general biodiversity associated with multi- and monolayered mussel beds of indigenous Perna perna and alien M. galloprovincialis, and to determine the relationship between mussels and epibiotic barnacles. This was done to determine the community structure of associated macrofauna and the role of mussels as biological facilitators. Samples were collected in Plettenberg Bay, South Africa, where M. galloprovincialis dominates the high mussel zone and P. perna the low zone. Three 15 X 15 cm quadrats were scraped off the rock in the high and low zones, and in the mid zone where the two mussel species co-exist. The samples were collected on 3 occasions. In the laboratory mussel-size was measured and sediment trapped within the samples was separated through 75 μm, 1 mm and 5 mm mesh. The macrofauna was sorted from the 1 mm and 5 mm sieves and identified to species level where possible. The epibiotic relationship between mussels and barnacles was assessed by measuring the prevalence and intensity of barnacle infestation and the condition index of infested mussels. Multivariate analysis was used on the mean abundance data of the species for each treatment (Hierarchical clustering, multidimensional scaling, analysis of similarity and similarity of percentages) and ANOVA was used for most of the statistical analyses. Overall, the results showed that tidal height influences the species composition and abundance of associated fauna. While mussel bed layering influenced the accumulation of sediments; it had no significant effect on the associated fauna. Time of collection also had a strong effect. While there was an overlap of species among samples from January, May and March, the principal species contributing to similarity among the March samples were not found in the other two months. The outcomes of this study showed that low shore mussel beds not only supported a higher abundance and diversity of species, but were also the most structurally complex. Although the condition index of mussels did not correlate to the percentage cover of barnacle epibionts, it was also evident that low shore mussels had the highest prevalence. The levels of barnacle infestation (intensity) for each mussel species were highest where it was common and lowest where it was least abundant. This is viewed as a natural artefact of the distribution patterns of P. perna and M. galloprovincialis across the shore. Mussels are more efficient as facilitators on the low mussel zone than the high mussel zone possibly because they provide habitats that are more effective in protecting the associated macrofauna from the effects of competition and predation, than they are at eliminating the effects of physical stress on the high shore. Although mussels create less stressful habitats and protect organisms from the physical stress of the high shore, there are clear limitations in their ability to provide ideal habitats. The biological associations in an ecosystem can be made weak or strong depending on the external abiotic factors and the adaptability of the affected organisms.

Epibiosis is an ecological relationship that has been described as one of the closest possible associations in marine ecosystems. In the space limited rocky intertidal, mussel beds provide important secondary space for barnacles. The epibiotic relationship between mussels and barnacles on the south-east coast of South Africa was considered at different scales, from large-scale, natural patterns of epibiosis on the rocky shore, to fine-scale settlement choices of barnacles and the effects on the condition and growth rates of individual mussels. Mussel and barnacle assemblages were generally stable over a 12-month period. The tracking of individual mussels with and without barnacle epibionts resulted in a significant increase in mortality rate of mussels with epibionts over 12 months (two-way ANOVA, p = 0.028). Barnacles on rocks, as well as on mussels, were also tracked with no significant effect of substratum on mortality of barnacles (two-way ANOVA, p = 0.119). Prevalence and intensity of barnacle infestations was also examined in relation to coastline topography on two co-occurring mussel species, the indigenous Perna perna and invasive Mytilus galloprovincialis. The results were complex, but bay status had significant effects on prevalence and intensity for both mussel species, depending on the time and zone. The effect of bay in relation to time was particularly relevant for M. galloprovincialis (four-way nested ANOVA, Season X Site(Bay): p = 0.0002), where summer prevalence was higher than that of winter in bays, regardless of zone, while in open coast sites, the effect of season was only significant in the mid zone. Patterns of intensity generally showed higher values in summer. Substratum preference by barnacles was investigated by recording settlement, survival and mortality of Chthamalus dentatus barnacles on various treatments. There was a strong preference for the rock-like plastic substratum by primary settlers (pair-wise tests of PERMANOVA: Dead < Rock mimic (p = 0.0001); Replica < Rock mimic (p = 0.019) and Live < Rock mimic (p = 0.0001)). This indicates that barnacles settle on mussel shells only as a secondary choice and that micro-topography is an important variable in barnacle settlement. The effect of barnacle epibiosis on condition index and growth of P. perna and M. galloprovincialis was also examined as a direct indication of the health of mussels subjected to the biological stress of epibiosis. Although not significant (PERMANOVA: P. perna: p(perm) = 0.890; M. galloprovincialis: p(perm) = 0.395), growth for both mussel species was slower for barnacle-infested individuals in summer, which is the main growing season for mussels in the region. Results from condition index calculations, however, showed no negative impacts of epibiotic barnacles (three-way ANCOVA: P. perna: p = 0.372; M. galloprovincialis: p = 0.762). Barnacle epibionts create a new interface between the mussel and its environment and this interaction can affect other members of the community. The possibility of the barnacle epibiont causing increased drag also needs further investigation. Biological processes operating within a wide range of physical stressors drive the interactions on the rocky shore, such as epibiosis. Overall, the results of this study suggest that the epibiotic relationship between mussels and barnacles on the south-east coast of South Africa does not significantly affect the mussel species present and that barnacles only use mussel shells as a secondary choice of substratum.